Improved Native Grasses and

Establishment Methods for

Use on Military Training Lands

Antonio J. Palazzo, Susan E. Hardy, and Kevin B. Jensen October 2003

Approved for public release; distribution is unlimited. ERDC/CRREL TR-03-20 Cold Regions Research and Engineering Laboratory

US Army Corps

of Engineers®

Engineer Research and

Development Center
ERDC/CRREL TR-03-20

October 2003

Approved for public release; distribution is unlimited.

Prepared for STRATEGIC ENVIRONMENTAL RESEARCH AND DEVELOPMENT PROGRAM

Improved Native Grasses and

Establishment Methods for

Use on Military Training Lands

Antonio J. Palazzo, Susan E. Hardy, and Kevin B. Jensen

Cold Regions Research and Engineering Laboratory

72 Lyme Road

Hanover, New Hampshire 03755
ABSTRACT

The objective of this project was to develop more wear-resistant plants and evaluate the relationships

between military training and plant injury, regrowth, and wear resistance. Through plant breeding, we

were able to improve traits related to resiliency and establishment in introduced and native species of

range-land grasses. We selected for early spring growth, increased seedling vigor, improved tiller and

rhizome development after disturbance, and resistance to abiotic and biotic stresses. Our improved plant

materials will be ecologically compatible at the military sites because they were developed from collections of species native to or previously seeded at these sites. We made advances in relating molecular

markers to plant characteristics and in using DNA fingerprinting techniques to characterize genetic diversity. We used markers to identify species and plants that can grow better at low temperatures. We now

have the tools to assess the genetic differences and similarities in commercial and natural seed sources,

enabling land managers to select seed sources that will ensure genetic compatibility with existing populations. Our tank traffic studies showed that naturalized, introduced species are more tolerant and recover

more rapidly under repeated tracking than native plants. However, two improved native species, western

wheatgrass and Snake River wheatgrass, showed promise as stabilization species because of their ability

to colonize damaged areas. Our studies on what we call “ecological bridges” confirm that we can select

seed mixtures that will establish more rapidly than all-native mixes and will ultimately lead to healthy

and persistent stands of native plants. The species in the seed mixtures and the equipment needed are

readily available, and the seeding can be done in one application, thus saving money. Our improved

germplasm will make these seeding mixes even more desirable.

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Citation of trade names does not constitute an official endorsement or approval of the use of such commercial products.

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be construed as an official Department of the Army position unless so designated by other authorized documents.

DESTROY THIS REPORT WHEN IT IS NO LONGER NEEDED. DO NOT RETURN TO THE ORIGINATOR.
Improved Native Grasses and Establishment Methods iii 

CONTENTS 

Terminology.........................................................................................................vii

Scientific Names ................................................................................................... ix

Preface .................................................................................................................. x

1 PROJECT BACKGROUND........................................................................... 1

DoD Relevance ............................................................................................... 1

Benefits............................................................................................................ 2

Scientific Background ..................................................................................... 3

2 OBJECTIVES ................................................................................................. 6

Objective (a): Breeding Resilient Plants ......................................................... 6

Objective (b): Soil Compaction and Root Studies .......................................... 7

Objective (c): Establishment Techniques........................................................ 7

3 TECHNICAL APPROACH............................................................................ 8

Approach (a): Breeding Resilient Plants ......................................................... 8

Approach (b): Soil Compaction and Root Studies ........................................ 11

Approach (c): Establishment Techniques...................................................... 12

4 PLANT BREEDING ACCOMPLISHMENTS............................................. 13

Plant Surveys and Collections....................................................................... 13

Preliminary Evaluation Trials to Identify Promising Species ....................... 14

Non-Invasiveness of Introduced Species....................................................... 22

Breeding Program ......................................................................................... 25

Improved Species .......................................................................................... 27

Molecular Marker Development ................................................................... 46

5 SOIL COMPACTION AND ROOT STUDIES ACCOMPLISHMENTS.... 54

Measuring Soil Compaction: Preliminary Soil Penetrometer Studies........... 54

Greenhouse Root Studies .............................................................................. 58

Tracking Study .............................................................................................. 61

6 METHODS FOR ESTABLISHING NATIVES AND FIGHTING 

INVASIVE WEEDS............................................................................... 77

Establishment of Natives on Sandy Soils at Fort Drum ................................ 77

Native Wheatgrass Establishment in Cheatgrass-Infested Areas at Yakima 80

Mixed Native and Introduced Seedings at Fort Carson................................. 80
iv ERDC/CRREL TR-03-20 

Forage Kochia as an Ecological Bridge ........................................................ 84

7 CONCLUSIONS........................................................................................... 86

Improved Plant Materials .............................................................................. 86

Tracking Studies............................................................................................ 87

Establishment of Natives............................................................................... 87

Benefits.......................................................................................................... 87

REFERENCES .................................................................................................... 89

APPENDIX A. PUBLICATIONS AND PRESENTATIONS ORIGINATING 

FROM SERDP PROJECT...................................................................... 96

APPENDIX B. DATA ON NON-INVASIVENESS OF INTRODUCED 

SPECIES............................................................................................... 100

FIGURES 

Figure 1. Adapted area for SERDP-select germplasms ........................................3

Figure 2. Plant breeding cycle ..............................................................................9

Figure 3. Comparison of populations of slender wheatgrass ..............................44

Figure 4. Genetic variation in bluebunch wheatgrass cultivars and a multipleorigin polycross ...........................................................................................50

Figure 5. UPGMA analysis of AFLP variation within and among Sandberg 

bluegrass populations collected near Mountain Home, Idaho, and 

Yakima Training Center, and the cultivars Canbar canby bluegrass and 

Sherman big bluegrass ................................................................................51

Figure 6. Venn diagram representation of genetic variation in Sandberg 

bluegrass populations collected near Mountain Home, Idaho, and 

Yakima Training Center, and the cultivars Canbar canby bluegrass and 

Sherman big bluegrass ................................................................................52

Figure 7. Using the Rimik CP20 cone penetrometer in the field ........................54

Figure 8. Soil compaction measurements at Fort Drum .....................................55

Figure 9. Effect of the compaction–depth relationship on soil moisture 

and plant growth .........................................................................................56

Figure 10. Soil compaction measurements at Fort Carson .................................57

Figure 11. Soil compaction measurements on dry and moist soils 

at Yakima Training Center ..........................................................................57

Figure 12. Elongation of roots after germination in growth pouches for five 

species compared to the invasive weed cheatgrass......................................62 

Figure 13. Layout of the tracking study ..............................................................63
Improved Native Grasses and Establishment Methods v 

Figure 14. Soil compaction measured before and immediately after zero, 

one, two, and four passes of an M1A1 tank on the Yakima Training 

Center Snake A and B plots on 19 May 1999 .............................................64

Figure 15. Ecological bridge concept .................................................................79

Figure 16. Percent annual and biennial weeds in Turkey Creek plots 

planted with the standard Fort Carson mix or with a core native mix 

plus an introduced grass ..............................................................................82

Figure 17. Percent native grasses in Turkey Creek plots planted with 

the standard Fort Carson mix or with a core native mix plus an 

introduced grass ..........................................................................................83

TABLES 

Table 1. Precipitation and soil type at breeding nursery sites .............................13

Table 2. Precipitation and soil types at species evaluation sites .........................14

Table 3. Plant species used in evaluation trials ...................................................15

Table 4. Stand establishment of perennial grasses at the Fort Carson Turkey 

Creek site, seeded in the fall of 1994 ..........................................................16

Table 5. Stand establishment of perennial native and introduced grasses 

at the Fort Carson South Boundary site, seeded in the fall of 1996 ............18

Table 6. Seeding vigor ratings and percent stand of native and introduced 

perennial Triticeae grasses during stand establishment (1995) and 

subsequent years on Snake sites at Yakima Training Center ......................19 

Table 7. Mean values of desired traits for four SERDP-select native 

germplasms compared to the “original” populations of known cultivars ...26

Table 8. Improved traits and current status of SERDP-select germplasms ........28 

Table 9. Growth of CD-II and Hycrest seedlings at low temperature 

in a growth chamber ...................................................................................32

Table 10. Growth of CD-II and Hycrest seedlings at a field site near 

Logan, Utah. ................................................................................................32

Table 11. Growth of RoadCrest and Ephraim crested wheatgrass in a 

greenhouse study .........................................................................................34

Table 12. Characteristics of the Siberian wheatgrass selections compared 

to the entire original breeding population ...................................................35

Table 13. Characteristics of bluebunch wheatgrass selections compared 

to the original breeding population .............................................................38

Table 14. Seedling emergence rate of western wheatgrass from a 

6.3-cm-deep planting ..................................................................................40

Table 15. SERDP-select Snake River wheatgrass cycle-1 compared to 

the mean of the base population ..................................................................41
vi ERDC/CRREL TR-03-20 

Table 16. Results of trials comparing the percent stand of Yakima 

populations with the commercial variety of western yarrow ......................46

Table 17. Soil compaction rates selected for this study ......................................58

Table 18. Stem and root length measurements of hard fescue grown in 

rhizotrons containing either a sandy or silty soil originating from 

Fort Drum, New York, and Yakima Training Center, Washington ............59

Table 19. Length, surface area, and diameter of roots and length and 

surface area of leaves of six native plants grown in rhizotron tubes 

containing a Plainfield sandy soil ...............................................................60

Table 20. Cone resistance measurements on Yakima Training Center plots 

subjected to zero, one, two, or four passes of an M1A1 tank .....................65

Table 21. Mean gap between any sown plants in seeded rows of vegetation 

on adjacent sites at Yakima Training Center ..............................................66

Table 22. Percent target species in seeded rows of vegetation on adjacent 

sites at Yakima Training Center ..................................................................68

Table 23. Forage yield as shown by mean dry weight of sown species on 

adjacent sites at Yakima Training Center ...................................................70

Table 24. Mean percent Sandberg bluegrass in sown species on adjacent 

sites at Yakima Training Center ..................................................................72

Table 25. Percent bare ground in plots of sown species on adjacent sites 

at Yakima Training Center ..........................................................................73

Table 26. Percent cheatgrass encroachment on adjacent sites at 

Yakima Training Center .............................................................................75

Table 27. Percent cheatgrass in establishment study plots with two row 

spacings at Yakima Training Center in 2000 ..............................................81

Table 28. Core native mix plus introduced species for mixed seedings 

at Fort Carson ..............................................................................................81

Table 29. Fort Carson standard mix at seeding rate increased to match 

the core-mix-plus-introduced species treatment .........................................82
Improved Native Grasses and Establishment Methods vii 

TERMINOLOGY 

Apomictic. Apomixis is a complex form of asexual reproduction that occurs 

extensively in Poa (bluegrass) species. In an apomictic species, the seed produced is genetically identical to the seed-bearing parent. 

Breeder’s, foundation, and certified seed. Breeder’s seed is produced from 

the last cycle of selection. This seed is used to produce foundation seed, which, 

in turn, is used to establish certified seed fields from which seed is produced for 

commercial sale. 

Cultivar vs. germplasm vs. selected class germplasm. Plant materials 

developed in this project will be released as cultivars, germplasms, or selectedclass germplasms. Selected-class germplasm refers to a native species that has 

not been tested prior to formal release; it may be selected for a certain trait (such 

as germination), or it may be a multiple-origin population that has not undergone 

any selection. A germplasm (pre-cultivar) can be a single genotype or a collection of multiple genotypes from multiple origins that are unique for a given character. A cultivar (sometimes called a variety) is a population that is unique for 

selected traits and that has undergone multiple cycles of selection and extensive 

testing in multiple locations. A cycle refers to a complete generation from seed to 

plant (selection) to seed. Cultivars are genetically stable. 

Full sib and half sib. In breeding, when you can control both parents, the 

offspring are full sibs. When only one of the parents is known, the offspring are 

half sibs. 

Introduced vs. naturalized vs. native species. In this report, we use the 

term introduced to represent species not indigenous to North America. Many of 

the introduced plant materials on western rangelands, including those in this 

project, originated in Central Asia, where they occur in very diverse ecosystems. 

The superior stand establishment characteristics, hardiness, adaptability, persistence under grazing, seed availability, seed cost, and productivity of introduced 

perennial species compared with indigenous native species have been documented in many regions (Barker et al. 1977, Vallentine 1978, Kilcher and 

Looman 1983, Lawrence and Ratzlaff 1989). Like their native counterparts, 

introduced grasses have the capacity to sort by natural selection and improve 

their adaptation to the environmental conditions on sites where they are seeded. 

As a result, many of the introduced species included in the project are naturalized, having existed in stands for over 50 years. These naturalized species have 

co-existed with native flora on North American rangeland (both private and public) for years. Within this report, we use only the terms introduced and native, 

based on the species’ origin. 
viii ERDC/CRREL TR-03-20 

Invasive. For the purposes of this study, we define an invasive species as an 

introduced species that will spread beyond the areas it currently inhabits and prevent the establishment of desired perennial plants. We do not agree with definitions of invasive that equate it to any introduced or exotic species. Tiller and rhizome development and seedling encroachment through seed dispersal are 

indicators of potential invasiveness. Except for RoadCrest crested wheatgrass, 

which is moderately rhizomatous and is best suited for cantonments and roadsides, we did not use any introduced species displaying these characteristics. On 

the other hand, rhizome development is valuable in desired native species for 

land stabilization and reclamation of disturbed lands. We worked with some rhizomatous natives to improve the establishment and persistence of desired 

species. 
Improved Native Grasses and Establishment Methods ix 

SCIENTIFIC NAMES 

Common name Scientific name Range relative to U.S. 

Alfalfa Medicago sativa Introduced

Basin wildrye Leymus cinereus (Scribn. & Merr.) Á. Löve Native (western U.S.) 

Beardless wildrye Leymus triticoides (Buckley) Pilg. Native (western U.S.) 

Big sagebrush Artemisia tridentata Native 

Bluebunch wheatgrass Pseudoroegneria spicata (Pursh) A. Löve Native (western U.S.) 

Cheatgrass Bromus tectorum L. Introduced invasive weed 

Crested wheatgrass 

(Fairway type) 

Agropyron cristatum (L.) Gaertn. Introduced 

Crested wheatgrass 

(Standard type) 

Agropyron desertorum (Fisch. ex Link) 

Schult. 

Introduced 

Forage kochia Kochia prostrata sp. virescens Introduced shrub 

Hard fescue Festuca brevipila R. Tracey Introduced 

Kentucky bluegrass Poa pratensis L. Native (northern U.S.) 

Little bluestem Schizachyrium scoparium (Michx.) Nash 

var. scoparium] 

Native 

Medusahead rye Taeniatherum asperum (Simonk.) Nevski Introduced invasive weed 

Purple needlegrass Nassella pulchra (Hitchc.) Barkworth Native (California) 

Russian wildrye Psathyrostachys juncea (Fisch.) Nevski Introduced 

Sandberg bluegrass Poa sandbergii Vasey or 

Poa secunda J. Presl 

Native (western U.S.) 

Sheep fescue Festuca ovina L. Introduced 

Siberian crested wheatgrass Agropyron fragile (Roth) P. Candargy Introduced

Slender wheatgrass Elymus trachycaulus (Link) Gould 

ex Shinners 

Native 

Snake River wheatgrass Elymus wawawaiensis ined. Native (northwestern U.S.) 

Switchgrass Panicum vergatum L. Native 

Tall fescue Festuca arundinacea Schreb. Introduced 

Thickspike wheatgrass Elymus lanceolatus (Scribn. & J. G. Sm.) 

Gould

Native 

Weeping lovegrass Eragrostis curvula (Schad.) Nees Introduced 

Western wheatgrass Pascopyrum smithii (Rydb.) Á. Löve Native (western U.S.) 

Western yarrow Achillea millefolium L. Native forb 
x ERDC/CRREL TR-03-20 

PREFACE 

This report was prepared by Antonio J. Palazzo, Research Agronomist, and 

Susan E. Hardy, Research Program Specialist, Environmental Sciences Branch, 

U.S. Army Engineer Research and Development Center (ERDC), Cold Regions 

Research and Engineering Laboratory (CRREL), Hanover, New Hampshire, and 

by Dr. Kevin B. Jensen, Research Geneticist, U.S. Department of Agriculture – 

Agricultural Research Service (USDA–ARS) in Logan, Utah. 

This project was a cooperative effort primarily with the USDA–ARS in 

Logan, Utah, including research geneticists Dr. Kay Asay (retired) and Dr. Steve 

Larson, plant breeding specialist Dr. Blair L. Waldron, and rangeland specialists 

W. Howard Horton and R. Deane Harrison. The principal investigator at ERDCCRREL was Antonio J. Palazzo, with supporting work by Timothy J. Cary and 

Susan E. Hardy. Nancy Perron and Troy Arnold provided technical support. 

James Wuebben, CRREL Acting Director, and J.-C. Tatinclaux, Chief, Environmental Sciences Branch, also helped with field work. 

Supporting work was also performed by Dr. David Huff, a specialist in plant 

genetics and root growth at Pennsylvania State University, and by Alan Anderson 

and Dr. Richard Gebhart, specialists in land capability models at the Construction 

Engineering Research Laboratory (ERDC-CERL) in Champaign, Illinois. Field 

work was assisted by George Savoy, Jeff Linn, and James Kulbeth from Fort 

Carson; Pete Nissan, Clark Reames, and Brian Cochran from Yakima Training 

Center; and Tom Lent and Ian Warden at Fort Drum. Dr. Kay Asay, USDAARS, Logon, Utah (retired), and Dr. C. Richard Lee, ERDC Environmental 

Laboratory (retired), reviewed the manuscript. 

The project was funded primarily by the Strategic Environmental Research 

and Development Program (SERDP), with additional funding from Stuart Canon 

and Ted Reid of Army Forces Command (FORSCOM) and from the Army BT25 

program. The authors thank Dr. Robert Holst and Dr. Femi Ayorinde, SERDP 

Conservation Program Managers present and past, respectively, and Brad Smith, 

SERDP Office Director, for their support during this program. 

The Commander and Executive Director of ERDC is Colonel James R. 

Rowan, EN. The Director is Dr. James R. Houston. 
Improved Native Grasses and Establishment Methods 

for Use on Military Training Lands 

ANTONIO J. PALAZZO, SUSAN E. HARDY, AND KEVIN B. JENSEN 

1 PROJECT BACKGROUND 

DoD Relevance 

The Department of Defense (DoD) must constantly balance its military mission and its commitment to stewardship on millions of hectares of federal land. 

These military training lands are some of the most intensely used public lands in 

the United States. The military mission requires that vegetation, primarily 

grasses, be as resilient as possible to military training exercises to maintain realism and control soil erosion. 

Throughout the DoD, land stewardship and management of natural resources 

fall under the Sikes Act of 1960, which promotes “effective planning, development, maintenance, and coordination of wildlife, fish, and game conservation and 

rehabilitation in military reservations on military lands.” The Army implements 

the Sikes Act through the Integrated Training Area Management (ITAM) program in place at over 60 installations. Through the ITAM, consideration of natural resources management objectives are integrated with land warfare training 

requirements. Today, all of the military departments are completing biological 

inventories on their lands, building on efforts that were initially driven by the 

National Environmental Policy Act and the Endangered Species Act. Integrated 

Natural Resources Management Plans (INRMPs)—the tool for implementing 

ecosystem management—provide balanced and coordinated consideration of 

various perspectives, including forestry, agronomy, soil science, pest management, livestock grazing, and fish and wildlife management (Gibb 2002). 

In the future the military faces increasingly difficult land management challenges. Proposed weapons systems will be longer-range, requiring more land for 

training exercises. As technology improves, training and testing requirements 

change. Complicating this challenge is the impact of continuing development, 

especially urbanization, outside the boundaries of military installations. As 

population growth and urban expansion continue, landscapes around facilities 
2 ERDC/CRREL TR-03-20 

will be further degraded, and additional pressures are likely to be brought to bear 

on native species, biological communities, and the ecological processes that sustain them. In some cases this growing pressure is likely to intensify demands that 

military installations and other federal lands take on even greater responsibilities 

for biodiversity conservation (Keystone Center 1996). 

The DoD has approximately 10 million hectares (25 million acres) of land to 

manage in the continental United States (CONUS). Within this area the Army 

has 4.5 million hectares (11 million acres) of training lands under the ITAM program, and the greatest expense in the ITAM budget is Land Repair and Maintenance (LRAM). The Air Force has over 3.8 million hectares (9.5 million acres) 

of land in CONUS. While each DoD organization produces its own unique 

impacts to the land related to its mission, all federal land managers (military and 

others) need to consider threatened and endangered habitats and follow Executive 

Orders on native plants and invasive weed control. 

The goal of this project is to increase the efficiency of lands for military use 

by understanding the effects of training on plant establishment and growth and by 

enhancing plant resiliency through improved selective breeding programs to 

develop new plant materials.

Benefits 

We anticipate that this work will improve the return on the military’s investment in the ITAM Program, which is budgeted at about $50 million annually.*

Along with guidance for mitigation methods and improved seedling establishment, this work will provide more-resilient plants that will help increase training 

opportunities on existing training areas while protecting environmental resources 

and wildlife habitat. These plant materials will be ecologically compatible with 

the various areas because they were developed on and from collections made 

near the sites. Lower costs in land maintenance are anticipated because of 

increased persistence and less frequent seedings. Thus, we will increase the value 

and use of current training areas, offer reduced unit training costs, and enhance 

DoD mission-related environmental activities. 

Within the range of distribution for the new germplasms (Fig. 1), we identified 42 DoD facilities, including over 525,000 hectares (1.3 million acres) of 

Army and Air Force land. The new germplasms are also appropriate for other 

federal, state, or local agencies; highway rights-of-way; mine spoils; rangelands; 

and other disturbed areas. 

 

*

 Personal communication with T. Macia, Team Leader for Training Lands at Chief of 

Staff Army, G3 Training Support Division, August 2002. 
Improved Native Grasses and Establishment Methods 3 

National Guard Sites

Air Force Sites

Army Sites

Navy Sites

Figure 1. Adapted area for SERDP-select germplasms. All plant 

materials were collected within this area. 

Scientific Background 

There is limited knowledge on the relationships between military training 

and plant injury, regrowth, and wear resistance. Before this study, no plant 

breeding or selection had been conducted to increase the wear resistance of plants 

for military training-land rehabilitation. 

Resilience is defined as the ability of a vegetative system to recover after 

disturbance and return to its original state (Doe et al. 1999). Military training 

exercises often destroy the vegetation, which leads to soil erosion (Halvorson et 

al. 2001). Based on ecological studies conducted in the shortgrass prairie, Shaw 

and Diersing (1990) reported that a two-year rotation might not be long enough 

for adequate plant recovery. Hinchman et al. (1990) concluded that it took up to 

three years after seeding to return shortgrass-prairie land to a condition suitable 

for resuming training. 

Training by tracked vehicles disaggregates and compacts soil, which 

increases the effects of erosive forces (Diersing et al. 1988). Shaw and Diersing 

(1990) found that tracking decreased plant basal and litter cover. Ayers et al. 

(1990) noted that the greatest soil compaction occurred in the upper 15 cm of the 

soil profile. 

Shaw and Diersing (1990) concluded that increased training intensity was 

correlated to the frequency of noxious weeds. A common weed problem on western military lands is cheatgrass, which was introduced to North America at the 
4 ERDC/CRREL TR-03-20 

end of the 19th century and has become a major threat to the ecological balance 

across much of the western rangelands (Young and McLain 1997). This noxious 

weed, as well as others, fuels recurrent fires that increase seed production for 

cheatgrass and eventually remove native plant materials from plant communities. 

Limited progress has been made in rehabilitating land infested with cheatgrass 

and other noxious weeds (Asay et al. 2001). 

Seeding perennial, cool-season grasses on rangelands is a valuable tool for 

restoring burned and disturbed areas and managing noxious weeds (Vallentine 

1971). The use of introduced perennial grasses, however, has been the focus of 

considerable debate among environmental, political, research, and user groups. 

Some contend that introduced grasses reduce biodiversity by displacing native 

species and disrupt the ecology and aesthetics of the plant community. This 

rationale has gained considerable support and has led to policies restricting the 

use of introduced plant materials on public lands. In particular, introduced 

Triticeae grasses have fallen into disfavor because many early seedings were 

made as large-scale monocultures that were unappealing and had limited biological diversity (Marlette and Anderson 1986). Despite this disfavor, the superior 

stand establishment characteristics, hardiness, adaptability, seed availability, seed 

cost, and productivity of introduced perennial species compared to native species 

have been documented for many regions (Barker et al. 1977, Kilcher and 

Looman 1983, Lawrence and Ratzlaff 1989, Vallentine 1978). Like their native 

counterparts, introduced grasses have the capacity to sort by natural selection and 

improve their adaptation to environmental conditions. Indeed many introduced 

grasses are more tolerant of frequent defoliation and disturbances because they 

have been subjected to similar conditions longer than our natives. These advantages have led to continued use of Triticeae grasses such as crested wheatgrass, 

intermediate wheatgrass, and Russian wildrye on federal lands (Richards et al. 

1997). Introduced cool-season perennial grasses have proven effective in competing with annual weedy grasses (Whitson and Koch 1998, Hull and Stewart 

1948) and forbs (Cronin and Williams 1966). More recently they have been 

found to compete favorably with some of the perennial invasive noxious weeds 

(Bottoms and Whitson 1998, Ferrell et al. 1998, Borman et al. 1991, Hubbard 

1975, Larson and McInnis 1989). 

The success of native plant establishment is partially related to root growth in 

a competitive environment. In our earlier studies (Palazzo and Brar 1997, 

Palazzo and Lee 1997, Brar and Palazzo 1995a, 1995b), we reported that plant 

root and leaf growth and plant resiliency were influenced by differences in soil 

texture, soil moisture, cultivars and species, and zinc-contaminated soil. Root 

growth is closely associated with plant resiliency. We postulated that if we could 

identify populations possessing improved root growth and other traits promoting 
Improved Native Grasses and Establishment Methods 5 

resiliency to military activities, we could selectively breed improved populations 

that would withstand the stresses on DoD training lands more successfully. Such 

improved plant materials should have wide adaptability to other areas with lowmaintenance vegetation needs, such as roadsides, dams, embankments, and other 

conservation projects. 
6 ERDC/CRREL TR-03-20 

2 OBJECTIVES 

The overall objectives of this project were to: 

(a) Breed improved native and introduced plant germplasms that have 

increased persistence and establishment characteristics under conditions 

created by military training activities; 

(b) Understand the effects of training on soil compaction, plant injury, and 

regrowth; and 

(c) Evaluate seeding methodology to better establish native and introduced 

grasses in mixed stands and still control invasive weeds. 

By combining plant and soil data, we can enhance existing land management 

models to include the effects of training on soil compaction, plant injury, and 

regrowth, which will provide methods for mitigation and rehabilitation. Land 

users will be able to make knowledgeable choices concerning plant selection and 

site rehabilitation procedures to reduce soil erosion. 

The project began with evaluation trials to identify the most promising 

species and with basic research on genetic markers for traits associated with 

resiliency on low-maintenance, high-use military training areas. The applied 

research phase took the identified species through several cycles of selection to 

develop germplasms that possess the desired traits. In the demonstration phase of 

the project, the resiliency of new plant materials is being evaluated through controlled tests on military training lands at Yakima Training Center, Washington, 

and Camp Guernsey, Wyoming. 

Objective (a): Breeding Resilient Plants 

Our main goal was to develop improved, training-resilient plants, especially 

grasses, that would be adapted to a wide range of environmental conditions in the 

semiarid, temperate regions of the U.S. Our intention was to improve existing 

plant materials through standard selection processes enhanced by identification 

of genetic markers. We would not use species that were not already on the site 

nor would we perform any genetic engineering. 

We worked with both native species and introduced species that had become 

naturalized on the training areas. Our goal was to help land managers satisfy the 

requirements for native plants, while also cost-effectively restoring training land 

as rapidly as possible to prevent erosion and reduce downtime. We included the 

introduced species in the program because they establish more rapidly and we 

believed that they could serve as a nurse crop for the establishment of the slower-
Improved Native Grasses and Establishment Methods 7 

developing natives. As the project progressed, we placed increased emphasis on 

improving native germplasms to address changing regulatory decisions. 

The desired characteristics of the new germplasms were early spring growth, 

rapid seedling establishment, improved tiller and rhizome development after 

disturbance, and resistance to abiotic and biotic stresses. At the same time, we 

wanted to ensure that newly developed germplasms would not themselves 

become invasive and move beyond the target habitats. 

Our new native plant germplasms should provide improved plant persistence 

on military lands at a reduced environmental risk with respect to habitat loss and 

soil erosion. These plant materials will be ecologically compatible with the areas 

where they will be used, as well as with similar climatic areas, because they were 

developed from collections made from these sites. 

Objective (b): Soil Compaction and Root Studies 

The second objective was to conduct field and greenhouse studies to quantify 

the degree of soil compaction that occurs during training and relate this soil condition to root injury in plants with known resilience. The most critical variable is 

the effect of soil compaction on root growth. Increased soil compaction leads to 

reduced soil aeration and shallow root growth, which reduces the capability of 

the plants to regrow after injury. Knowledge of the effects of soil compaction on 

plant growth will help us provide better habitats with less soil erosion. 

Objective (c): Establishment Techniques 

As we placed increasing emphasis on native species, we added a third objective of developing cost-effective seeding techniques to establish viable stands of 

natives. Native species are generally much slower to establish than are introduced 

species. Before a native grass can become established, erosion or noxious weeds 

often become costly problems to land managers. Our goal was to find costeffective methods that would rapidly protect the soil and permit the eventual 

establishment of resilient native species. We want to reduce the costs of land 

repair and maintenance while simultaneously reducing downtime on military 

training areas. 
8 ERDC/CRREL TR-03-20 

3 TECHNICAL APPROACH 

For the three main objectives, the general approach was as follows: 

(a) For plant breeding and germplasm development, we surveyed existing 

plants on military facilities and selected the most promising species. The most 

promising populations of these species were either used directly for seed increase 

without selection or included in the breeding program, where they underwent two 

cycles of selection for desired characteristics such as improved seed germination, 

seedling vigor, rhizome development, persistence, drought tolerance, and weed 

suppression. Simultaneously, genetic markers associated with desired traits were 

identified for future breeding programs. The final steps in this part of the 

approach were the testing and release of the new germplasms. 

(b) For plant root growth and tracking, we studied root growth in the greenhouse and established tracking studies on existing grasses in the field. 

(c) To improve establishment and inhibit invasive weeds, we evaluated the 

use of seed mixtures at three field locations. 

In our final technology transfer phase of the project, we have provided data 

for several land management models, demonstrated the new germplasms and our 

seeding mixtures on three military facilities, and begun to implement a business 

plan that will ultimately make the improved seeds available to land users. 

Approach (a): Breeding Resilient Plants 

Assessment 

The first step was to assemble a broad genetic base and select the best accessions and clonal lines for the desired characteristics. We surveyed representative 

DoD installations to identify the most promising introduced and native species as 

well as the characteristics associated with resiliency in those species. Through 

research funded by Army Forces Command (FORSCOM), we had already identified resilient plant species at several training sites using field surveys and the 

data generated by the ERDC-CERL Integrated Training Area Management 

(ITAM)–Land Condition Trend Analysis (LCTA) program. The LCTA program, 

which includes monitoring of vegetation dominance on lands subjected to military training, provided information on the resiliency, persistence, and adaptation 

of plant species. While making field collections, we identified plant characteristics that are critical to soil conservation, plant survival, and training resiliency. 

These traits included a vigorous ground cover; a deep, fibrous root system; tiller 

and rhizome development; initiation of growth early in the spring; persistence 
Improved Native Grasses and Establishment Methods 9 

under environmental extremes (temperature and drought); and rapid seedling and 

plant establishment. 

To assemble the best lines for the introduced breeding populations, we 

looked for the desired characteristics among the plant materials already assembled in nurseries at USDA-ARS and Pennsylvania State University. To assemble 

native breeding populations, we collected plants with the desired traits from 

training sites and other conservation lands. 

We planted the assembled promising species in seeded evaluation trials at 

Yakima Training Center and Fort Carson during the fall of 1994 and 1996. More 

than 60 cultivars, breeding lines, and plant accessions directly from the training 

sites were included in the Yakima trials. Over 40 cultivars, breeding lines, and 

material indigenous to the training sites were included at Fort Carson. Visual 

percent stand and vigor ratings were taken throughout 1995, 1996, 1997, and 

1998. From the nurseries and the seeded evaluation trials, we selected the most 

promising species for the breeding process. 

Development 

The next step was to make crosses among the best lines to produce progeny, 

conduct parent–progeny tests to initiate the second breeding cycle, and continue 

the breeding cycle as necessary. We used traditional plant-breeding techniques 

to combine desired traits into breeding populations. Figure 2 shows a typical 

Identify plant species

• Plant surveys 

• Evaluation Trials 

Assemble genetic base from 

• Surveyed plants 

• Breeding program 

Release germplasm 

Cross to 

develop strains 

Select best 

parental lines 

Establish source

nurseries 

Repeat cycle

Evaluate and select 

superior plants from 

source nurseries 

Evaluate

progenies

Figure 2. Plant breeding cycle. 
10 ERDC/CRREL TR-03-20 

breeding program from plant surveys to germplasm release. Under normal conditions, it takes two to three years to complete one cycle of selection. The first 

year is a field establishment year with no selection. The second and third years 

are for selecting traits and harvesting seed. If hybridization between selected 

clones is desired, it can take an additional year to isolate the hybrid and increase 

seed. With the exception of conducting plant surveys and assembling a broad 

genetic base (the boxes at the top of Figure 2), the breeding cycle is repeated 

until the population has become genetically stable for the improved traits; each 

cycle increases the frequency of genes conditioning increased adaptation or 

specific traits. At some point after selection, the population is genetically stable 

for the desired traits from one generation to the next and may be released as a 

germplasm or, with more intense testing, as a cultivar. 

We typically took plants through two cycles of selection. We selected 

parental lines by analyzing measurable traits such as stand establishment vigor, 

rate of tillering and rhizome development, vegetative vigor, and seed yield 

potential. Native grasses with seed dormancy problems were screened for more 

rapid germination. For future use we used Random Amplified Polymorphic DNA 

(RAPD) or Amplified Fragment Length Polymorphic (AFLP) DNA analyses to 

identify genetic markers for desirable traits in some of the species. 

Some new germplasms were not subjected to selection pressure. Instead, they 

were assembled from broad ecological ranges, resulting in a broader genetic base 

than in any of the individual populations and the potential to be better adapted to 

a wide range of different ecotypes. 

Test and evaluation 

We then tested new strains in replicated seeded trials and in soil compaction 

trials in the field. We also evaluated the new strains to ensure that they would not 

become invasive, and we analyzed the genetic diversity of the new strains. 

Prior to formal release, cycle-2 breeding populations are established in 

seeded trials under a range of environmental conditions. Data obtained from 

these trials are used to write up the formal release notices, which include areas of 

adaptation. Seed from cycle-2 populations—breeder’s seed—is also used to 

establish foundation seed-increase fields. Seed from those fields will be released 

commercially as certified seed. 

We established an Independent Review Panel to assess whether the introduced species of interest are a threat to invade habitats other than those currently 

occupied by the species. For this review we evaluated older sites at Yakima 

Training Center (YTC) that contained the same species as those in the breeding 
Improved Native Grasses and Establishment Methods 11 

program. We evaluated the species for spread (through either vegetative tillering 

or reseeding) after 4–19 years of growth by counting plants growing outside the 

plots where they were planted. The Independent Review Panel met at YTC to 

view the sites, discuss their findings, and make a definitive statement on the 

potential invasiveness of the plant materials. 

We used both RAPD and AFLP DNA analyses to compare the genetic diversity of several of the new strains to their currently available counterparts. 

Approach (b): Soil Compaction and Root Studies 

Assessment 

We used cone penetrometers to assess the degree of soil compaction that 

occurs on training lands at three facilities: Yakima Training Center, Fort Carson, 

and Fort Drum. A variety of lands were assessed, including training areas, lightly 

used dirt roads, and bivouac areas. We also used the cone penetrometer to study 

the influence of frost on alleviating soil compaction. 

Development 

We conducted controlled studies to support the field tracking studies. In the 

greenhouse we compressed soil inside rhizotron tubes using compaction rates 

taken from the field measurements in the assessment step above, and we evaluated the root growth of plants grown in the compressed soil. In environmental 

chambers we used growth pouches to study the effect of various temperatures on 

early growth and root elongation of military-important species. 

Test and evaluation 

We conducted simulated tracking exercises at Yakima Training Center on 

our original evaluation plots, four years after they were established. The plots 

were tracked with a tank making no, one, two, or four passes. We used the cone 

penetrometer to record soil compaction values before and immediately after 

tracking and then again a year later. We used point-frame techniques to analyze 

the different species for resiliency evaluated after the tank passes, measuring 

gaps between target species, percent target species, dry matter yield, bluegrass 

encroachment, percent bare ground, and cheatgrass invasion on and off the 

tracked area. 
12 ERDC/CRREL TR-03-20 

Approach (c): Establishment Techniques 

Native grasses are usually more difficult to establish than their introduced 

counterparts because of seed dormancy, poor seedling vigor, and reduced tolerance to defoliation, grazing, and traffic. While improving these traits in our 

SERDP-select natives, we also investigated seeding methods to further enhance 

their establishment and prevent the encroachment of invasive weeds. We are 

using non-invasive introduced grasses as what we call an “ecological bridge” to 

the establishment of native grasses. 

At Fort Drum we tested and demonstrated a seed mixture of introduced and 

native species to obtain a native meadow on sandy (92% sand), infertile soils. 

At Yakima Training Center we evaluated the establishment of native grasses 

using a mixture of native and introduced wheatgrasses. We seeded various combinations of the native ‘Secar’ Snake River and ‘Goldar’ Bluebunch wheatgrasses 

with the introduced ‘Vavilov’ Siberian wheatgrass, which has been shown to 

inhibit growth of the noxious weed cheatgrass (Asay et al. 2001). Each grass was 

planted alone, in a mixture with one or two other grasses, or in alternating rows 

with one of the other grasses. Plots were evaluated for the percent cheatgrass 

growing after one and two years. 

At Fort Carson we evaluated a native grass mix planted with one of five 

introduced grass add-ons. We compared those mixes with the standard Fort 

Carson seed mix. The stands were evaluated after two, three, and four years for 

species composition and percent ground cover. 
Improved Native Grasses and Establishment Methods 13 

4 PLANT BREEDING ACCOMPLISHMENTS 

Plant Surveys and Collections 

We conducted vegetation surveys at Fort Carson, Yakima Training Center 

(YTC), and Fort Drum to identify the most promising native and introduced 

species and to determine plant characteristics that are important for adaptability 

in a training land environment. To identify specific collection sites, we reviewed 

soil and ecological site data from the USDA Natural Resources Conservation 

Service (USDA-NRCS). 

For the native grasses of interest, we collected seeds and clonal materials 

from a broad ecological range at Yakima Training Center, Fort Carson, and Fort 

Drum. At Yakima Training Center we made 134 collections of native species in 

1994 and 1995, including the grasses bluebunch wheatgrass, Sandberg bluegrass, 

and basin wildrye, and the forb western yarrow. At Fort Carson we made 166 

collections of natives including western wheatgrass, blue grama, sideoats grama, 

and Indian ricegrass. At Fort Drum we collected fine fescues, hairgrass, little 

bluestem, and switchgrass. 

For the introduced species we used known germplasm lines in collections at 

space-planted nurseries at the Utah State University Blue Creek Experimental 

Farm in Box Elder County, Utah, approximately 80 km northwest of Logan, 

Utah. We also collected related species from Asia. We did some preliminary 

testing and selection at a space-planted nursery at Exit 11 at Yakima Training 

Center. Table 1 gives the annual precipitation and soil types at the two nursery 

sites. 

Table 1. Precipitation and soil type at breeding nursery sites.

Location Site 

Annual 

precipitation (mm) Soil type 

Yakima, Washington Exit 11 200–250 Benwy series 

Blue Creek, Utah 200–375 Parley’s silt loam 

We identified the important characteristics to be seedling vigor, tiller and 

rhizome development, resistance to drought and plant pests, and growth at low 

temperatures. Seedling vigor was identified as the most important of the these 

desired characteristics for returning land to training as soon as possible and for 

fighting invasive weeds. 
14 ERDC/CRREL TR-03-20 

Preliminary Evaluation Trials to Identify Promising Species 

At the beginning of the program, many more species were examined and 

eventually dropped based on adaptive characteristics as the most promising 

species became apparent. We began with evaluation trials at two U.S. Army sites 

to help identify promising germplasm. The evaluation trials to assess stand 

establishment were conducted at Fort Carson in south-central Colorado and at the 

Yakima Training Center (YTC) in south-central Washington. The trials were 

established during the fall of 1994 and 1996. The seeded evaluation sites at the 

facilities differed in soil type and precipitation, which allowed for evaluation of 

germplasm over different environmental regimes (Table 2). 

Table 2. Precipitation and soil types at species evaluation sites. 

Location Site 

Annual 

precipitation (mm) Soil type 

Yakima, Washington Snake A and B 200–250 Benwy series 

Fort Carson, Colorado Turkey Creek 300–350 Rizozo-Neville Complex 

South Boundary 175–225 Minnequa-Manvel 

loams 

Two evaluation trials were conducted at Fort Carson: Turkey Creek and 

South Boundary. Both sites received less than normal precipitation during our 

trials, especially in the latter years. Turkey Creek receives 325 mm of average 

annual precipitation; however, there were only 265, 159, 274, and 120 mm in 

1999, 2000, 2001, and 2002, respectively. South Boundary receives 200 mm of 

average annual precipitation, but only 134, 141, 197, and 115 mm were received 

in 1999, 2000, 2001, and 2002, respectively. The soil type at Turkey Creek is a 

Rizozo-Neville Complex [fine-loamy, mixed (calcareous), mesic Ustic 

Torriorthent], and the area is dominated by piñon pine, juniper, blue grama, and 

western wheatgrass. At South Boundary the soils are Minnequa-Manvel loams 

[fine silty, mixed (calcareous), mesic Ustic Torriorthents], and the dominant 

vegetation is blue grama and galletta. 

At YTC, evaluation trials were conducted on adjacent Snake River sites A 

and B. The YTC location received 353, 375, 175, and 193 mm in 1995, 1996, 

1997, and 1998, respectively. The soil type is a Benwy series (fine-loamy, mixed 

superactive, mesic Calciargidic Argixerolls), and the area is dominated by bluebunch wheatgrass and Sandberg bluegrass, with some basin wildrye in low areas. 

The Fort Carson plots were seeded as dormant seedings during late October 

1994 at Turkey Creek and in early August 1996 at South Boundary. Native and 
Improved Native Grasses and Establishment Methods 15 

introduced grasses were evaluated on Turkey Creek and South Boundary. At 

YTC, plots were seeded during late October 1994 as a dormant seeding; native 

and introduced grasses were evaluated on Site A and natives on Site B. 

Seeding was accomplished on mechanically prepared, weed-free seedbeds 

with a drill equipped with double-disk furrow openers and depth band regulators. 

Seeds were placed from 1.25 to 2.0 cm below the soil surface at a rate of 

approximately one seed per centimeter. Individual plots consisted of drilled rows 

spaced 30 cm apart. Plot sizes were 1.5 by 15 m. Plots were arranged in a 

randomized complete block with four replications at each of the two sites. The 

seeding procedures are fully described by Asay et al. (2001). 

The percent stand establishment was determined annually on a visual basis at 

all sites; however, stand data in 1997 were determined on the basis of plant 

counts in two 1-m2

 sampling areas per plot. Seedling vigor was also determined 

at the YTC sites using ratings of 1 to 9, with 1 for the worst plot and 9 for the 

best plot. All data were subject to analysis of variance using ANOVA and General Linear Model (GLM) procedures, and mean separations were made on the 

basis of a least significant difference (LSD) test at the 0.05 probability level 

(SAS Institute Inc. 1994). 

Table 3 shows the species used in the evaluation trials. Over 60 cultivars, 

breeding lines, and material collected from the training sites were included in the 

Yakima trials. Over 40 cultivars, breeding lines, and material indigenous to the 

training sites were included at Fort Carson. 

Table 3. Plant species used in evaluation trials. 

Native grasses Introduced grasses 

Bluebunch wheatgrass Thurbers needle grass Lovegrass Crested wheatgrass 

Snake River wheatgrass Green needle grass Blue grama Siberian wheatgrass 

Wheatgrass hybrid Idaho fescue Side oats grama Intermediate wheatgrass 

Western wheatgrass Sheep fescue Buffalograss Russian wildrye 

Slender wheatgrass Sandberg bluegrass Prairie sandreed grass Giant wildrye 

Thickspike wheatgrass Canby bluegrass Little bluestem 

Bottlebrush squirreltail Indian ricegrass Sand dropseed 

Basin wildrye Needle and thread grass Galleta grass 

Native forbs and legumes 

Introduced forbs, 

legumes, and shrubs 

Western yarrow Spiny hopsage Bitterbrush Forage kochia 

Small burnett Fourwing saltbrush Globemallow Cicer milk vetch 

 Alfalfa 
16 ERDC/CRREL TR-03-20 

Results at Turkey Creek, Fort Carson 

The Turkey Creek site has fertile soils, and most of the introduced species 

established well (Table 4). In the first year the three Russian wildrye cultivars did 

not establish as well as Siberian or crested wheatgrass cultivars; stands of all 

introduced entries were similar after the first year. ‘Rosana’ western wheatgrass 

and the two thickspike wheatgrass cultivars established better than the other 

native plant entries but initially not as well as all but two of the introduced 

Table 4. Stand establishment of perennial grasses at the Fort Carson Turkey Creek site, seeded in the fall of 1994. 

Entry 1997 1998 1999 2000 2001 

 ——————— Mean % stand —————— 

Introduced entries 

Crested wheatgrass 

CD-II crested wheatgrass 100 97 94 89 99 

RoadCrest crested wheatgrass 91 100 100 92 93 

Siberian wheatgrass 

P27 Siberian wheatgrass 84 100 94 91 90 

Vavilov Siberian wheatgrass 83 97 88 80 95 

Kazakhstan Siberian wheatgrass 73 94 78 69 83 

Russian wildrye 

Tetraploid Russian wildrye 68 94 91 78 93 

Mankota Russian wildrye 46 91 88 72 89 

Bozoisky Russian wildrye 30 84 81 64 86 

Native entries 

Rosana western wheatgrass 54 81 100 97 100 

Thickspike wheatgrass 

Sodar thickspike wheatgrass 55 91 84 80 93 

E27 thickspike wheatgrass 52 81 75 66 77 

Indian ricegrass 

Paloma Indian ricegrass 31 56 47 30 28 

T593 Indian ricegrass 13 38 31 19 28 

Nezpar Indian ricegrass 26 25 19 23 12 

CSU10 Indian ricegrass 9 22 16 16 25 

Needle and thread grass 5 31 47 45 51 

Alma blue grama 0 0 31 22 15 

Vaughan sideoats grama 0 0 19 19 15 

LSD (0.05) entries 8 16 18 16 14 
Improved Native Grasses and Establishment Methods 17 

entries. The stand increase of those three natives was due to rhizome development. In the remaining years those three leading native plant entries had stand 

values similar to the introduced species. None of the other native entries attained 

a stand greater than 50%. 

Results at South Boundary, Fort Carson 

The soils at South Boundary are shallower, much drier, and less fertile than 

those at the Turkey Creek site. Except for the two intermediate wheatgrass 

entries, the introduced species had high percent stand values after one year 

(Table 5). The percent stand values for the intermediate wheatgrasses never 

improved over the next four years. The best-performing introduced species were 

the four entries of Russian wildrye, ‘RoadCrest’ crested wheatgrass, and Vavilov 

Siberian wheatgrass. For native plant entries, ‘Barton’ and ‘Rodan’ western 

wheatgrass and ‘Pryor’ slender wheatgrass had the best initial stand values. In the 

final two years the three western wheatgrass entries were the only native entries 

with stand values greater than 14%. With its strong early establishment, Pryor 

slender wheatgrass may prove useful in mixes with other slower-establishing 

natives such as the western wheatgrasses. Pryor may be a possible ecological 

bridge providing cover and protecting the soil, giving the other natives time to 

establish (for discussion of the ecological bridge concept, see the later section on 

“Methods for Establishing Natives and Fighting Invasive Weeds”). 

Results at Yakima Training Center 

Annual precipitation during stand establishment (353 mm in 1995 and 375 

mm in 1996) was substantially above normal, and good stands were obtained for 

all of the introduced and native wheatgrasses. Differences between introduced 

and native Triticeae grasses were less definite at this site, which is dominated by 

bluebunch wheatgrass (Table 6). Vavilov and P-27 Siberian wheatgrass demonstrated the best stand establishment of the introduced grasses. Among the native 

grasses the Snake River and bluebunch wheatgrasses, bluegrasses, Bannock 

thickspike wheatgrass, and most of the thickspike wheatgrass hybrids established 

well and maintained their stands for the duration of the trial. Secar Snake River 

wheatgrass was particularly well adapted to this site, and in 1998 this cultivar had 

stands equivalent to or significantly better than any other entry at the location. 

At the Snake A site, good stands were generally obtained for all introduced 

grasses during the year of establishment in 1995, and stands were maintained 

during subsequent years through 1998 (Table 6a). Vavilov Siberian wheatgrass 

established faster than other grasses evaluated. In 1998, stands of Vavilov Siberian wheatgrass were similar to P-27 (P>0.05); however, stands of all introduced 
18 ERDC/CRREL TR-03-20 

Table 5. Stand establishment of perennial native and introduced grasses 

at the Fort Carson South Boundary site, seeded in the fall of 1996. 

Entry 1997 1998 1999 2000 2001 

 ——————— Mean % stand —————— 

Introduced entries 

Russian wildrye 

Bozoisky 63 93 94 81 91 

Tetraploid 59 88 78 70 88 

Syn A 66 88 84 78 86 

Mankota 72 78 88 80 85 

Crested wheatgrass 

CD-II 69 41 47 37 30 

RoadCrest 88 94 91 83 91 

Siberian wheatgrass 

Vavilov 81 81 78 64 78 

Kazak 56 69 63 52 55 

Intermediate wheatgrass 

Luna 28 9 16 7 3 

Rush 34 6 13 1 1 

Native entries 

Western wheatgrass 

Barton 63 50 72 69 88 

Rodan 63 44 53 42 58 

Rosana 41 34 53 50 78 

Bottlebrush squirreltail 34 6 3 4 2 

Sandhollow 25 9 9 4 3 

Thickspike wheatgrass 

E-27 25 3 6 3 4 

Sodar 28 13 3 8 14 

Pryor slender wheatgrass 72 38 31 7 0 

Sand lovegrass 31 3 16 0 0 

Mean 53 45 47 39 44 

LSD (0.05) 18 23 20 15 18 
Improved Native Grasses and Establishment Methods 19 

Table 6a. Seeding vigor ratings and percent stand of native and introduced perennial Triticeae 

grasses during stand establishment (1995) and subsequent years on Snake A site at Yakima 

Training Center. (This site was used for the tracking study in 1999; subsequent data are included 

in the tracking section of this report.) 

Entry 

Seedling vigor z

May 1995 

% Standy

1995 

% Stand 

1996 

% Stand 

1997 

% Stand 

1998 

Snake A introduced entries 

Siberian wheatgrass 

Vavilov Siberian wheatgrass T 9 99 100 100 100 

P-27 Siberian wheatgrass T 9 96 82 83 88 

Kazak Siberian wheatgrass T 7 75 66 80 72 

Mean 8 90 83 88 87 

Crested wheatgrass 

Hycrest T 7 94 82 69 79 

Ephraim T 8 77 69 71 75 

Mean 7 85 75 70 77 

Russian wildrye 

Bozoisky T 5 67 57 55 63 

Tetraploid T 4 50 25 38 38 

Mean 4 59 41 46 50 

Intermediate wheatgrass (cv. Luna)T 4 61 72 77 78 

Snake A native entries 

Wildryes 

Leymus-1 hybrid 1 8 0 3 7 

Basin wildrye (Yakima) T 3 35 13 34 38 

Mean 1 21 7 19 22 

Bluegrasses 

Canby bluegrass T 8 69 70 60 82 

Sandberg bluegrass 7 49 50 51 54 

Mean 7 59 53 32 68 

Indian ricegrass 

Nezpar Indian ricegrass 6 60 10 0 10 

T553 Indian ricegrass 1 11 3 0 16 

Mean 3 35 7 0 13 

Western wheatgrass (cv. Rosana)T 3 33 35 32 41 

Bottlebrush 

Squirrel tail 87 3 41 41 0 0 

Squirrel tail 89 6 55 47 0 0 

Mean 4 48 44 0 0 

Miscellaneous species 

Needle and thread grass 2 14 0 8 6 

SL hybrid (thickspike X bluebunch) 4 46 57 34 50 

Mean 2 30 28 21 28 

LSD (0.05)–Entries 1 19 14 12 14 

LSD (0.05)–Species 2 20 18 17 17 

z

 Seedling vigor ratings 9 May 1995, following a dormant fall seeding in 1994; 1=poor, 9=best seedling vigor. y

 Percent stand based on visual ratings. 

T

 Entries included in the tracking study. 
20 ERDC/CRREL TR-03-20 

Table 6b. Seeding vigor ratings and percent stand of native and introduced perennial Triticeae 

grasses during stand establishment (1995) and subsequent years on Snake B site at Yakima 

Training Center. (This site was used for the tracking study in 1999; subsequent data are included 

in the tracking section of this report.) 

Entry 

Seedling vigor z

May 1995 

% Standy

1995 

% Stand 

1996 

% Stand 

1997 

% Stand 

1998 

Snake B native entries 

Bluebunch wheatgrass 

Goldar T 7.0 67 56 49 75 

Whitmar T 6.3 66 72 62 75 

ACC-238 T 6.0 70 53 53 75 

Yakima collection T 6.5 59 38 45 69 

Mean 6.4 66 55 52 73 

Snake River wheatgrass 

Secar T 7.3 69 94 92 97 

ACC-707 T 7.0 75 88 75 91 

EVT-572 T 7.5 73 91 87 100 

Mean 7.3 72 91 85 96 

Thickspike wheatgrass and hybrids 

Bannock T 7.3 73 72 53 75 

Sodar 6.5 64 41 35 41 

E-20 (thickspike X Snake River) T 7.0 72 78 80 88 

E-27 (thickspike X Snake River) T 6.5 70 81 69 81 

SL hybrid (thickspike X bluebunch) 4.3 52 22 23 28 

Mean 6.3 66 59 52 63 

LSD (0.05)–Entries 1 14 16 11 10 

LSD (0.05)–Species 0.6 8 15 12 13 

z

 Seedling vigor ratings 9 May 1995, following a dormant fall seeding in 1994; 1=poor, 9=best seedling vigor. 

y

 Percent stand based on visual ratings. 

T

 Entries included in the tracking study. 

grasses were greater than 70% with the exception of Bozoisky-Select and Tetraploid (Table 6a). Vavilov Siberian wheatgrass, ‘Ephraim’ crested wheatgrass, 

and ‘Luna’ intermediate wheatgrass maintained their stands throughout the study. 

Of the native grasses at the Snake A site, the bluegrasses, especially the 

‘Canby’ cultivar, maintained or increased in percent stand from 1995 to 1998. 

The SL hybrid (thickspike wheatgrass × bluegrass) also did about as well as 

Canby bluegrass, which was the poorest performer of the bluegrasses. Seedlings 
Improved Native Grasses and Establishment Methods 21 

of ‘Nezpar’ Indian ricegrass established relatively well during 1995 following the 

late-fall seeding; however, stands of this caespitose species declined from 60% in 

1995 to 10% in 1998. All other native grasses at the Snake A site were difficult 

to establish and subsequently had relatively poor stands throughout the study 

(Table 6a). 

At the Snake B site, seedlings of Snake River wheatgrass had significantly 

(P<0.05) higher vigor ratings in 1995 than did bluebunch wheatgrass and thickspike wheatgrass (Table 6b). With the exception of wildland-collected bluebunch 

wheatgrass from the Yakima Training Center (69%), Sodar thickspike wheatgrass (41%), and the SL hybrid (28%), all native grasses had stands in excess of 

70% by 1998. Stands of the rhizomatous native grasses (Bannock thickspike and 

hybrids between thickspike and Snake River wheatgrass) were maintained at an 

acceptable level during the study. 

Evaluation Trial Conclusions 

These evaluation trials helped us identify the most promising species for 

germplasm improvement, and they supported our hypothesis that mixed seedings 

of introduced and native species might provide the best method for establishing 

natives and preventing encroachment by invasive weeds. Most of the introduced 

species established better than did the native species at both facilities. Any differences between entries of the introduced species lasted only one year, while the 

native grass entries usually took two to three years to attain a stand greater than 

50%. 

On sites with severe water limitations (<300 mm per year) or infertile soils, 

the native grasses were more difficult to establish, less vigorous, and less persistent than their introduced counterparts (Siberian and crested wheatgrass and Russian wildrye). At Yakima, where moisture conditions were more favorable, the 

native grasses established and persisted relatively well compared to the introduced entries. Although difficult to establish, stands of the rhizomatous native 

western wheatgrass increased each season after establishment. Most wheatgrasses are bunchgrasses, but western wheatgrass has rhizomes and the ability to 

develop an excellent ground cover. With its hard seed coat and persistency, western wheatgrass looked like a good candidate for selective breeding for improved 

rhizome or tiller characteristics. 

These findings suggest that adapted introduced grasses be considered along 

with native grasses as a component of seed mixtures on environmentally harsh 

sites that have been burned, infested with competitive weedy species, or otherwise degraded. An improper choice of plant materials would perpetuate degradation of soil resources, especially on sites that are dominated by weedy annual 
22 ERDC/CRREL TR-03-20 

species such as cheatgrass and medusahead rye. Because introduced grasses generally establish better than natives in such conditions, the best choice may be to 

begin by planting introduced grasses or a mixture of introduced and native 

grasses (Asay et al. 2001). 

All introduced grasses in these trials exist in biologically diverse ecosystems 

in their native habitats, and rarely, if ever, are they invasive. On the Palouse 

rangelands near Yakima, bluebunch wheatgrass has successfully co-existed with 

Siberian wheatgrass on sites that were seeded more than 15 years ago (Asay et al. 

2001). At the Snake A and B sites in these trials, the fully established native 

grasses will be competitive with and may largely replace the introduced grasses. 

Non-grasses that looked promising included the introduced shrub forage 

kochia and the native forb western yarrow. Forage kochia is a good candidate for 

use as a firebreak and on ranges. Western yarrow will add diversity to seed mixtures of grasses. 

Non-Invasiveness of Introduced Species 

While our current emphasis is on improving the resilience of native species 

on training lands, the research program began by looking primarily at introduced 

or naturalized species. Our first three releases are introduced species, and inprogress program reviewers were concerned about the effects of introducing 

these germplasms in native ecosystems. The reviewers wanted assurance that the 

introduced germplasms we develop will not dominate lands currently inhabited 

by native species or prevent the return of native plants in the future. Similarly the 

training-land managers want to avoid any potential regulatory problems (invasive 

plants, soil erosion, etc.) that might interfere with their training needs. 

To address this concern we invited a panel of independent reviewers from the 

U.S. Environmental Protection Agency, The Nature Conservancy, the USDANRCS, and another USDA-ARS office to review the materials we were evaluating at Yakima Training Center (Appendix B; Palazzo et al. 1999). We asked the 

review panel to assess whether the germplasms used to develop the new cultivars 

have the potential to invade habitats other than those currently occupied by the 

species. This group met in May 1999 and evaluated the mature Snake River A 

and B sites (described in the previous section), examined several sites previously 

seeded to introduced grasses, and reviewed our data. 

The Invasiveness Question 

We believe that extensive invasion (through seed dispersal or vegetative 

growth) by the new plant materials we are developing is unlikely because we 
Improved Native Grasses and Establishment Methods 23 

limited our breeding studies to bunch-type species, all of which are already 

present on the training lands. Most of our work has been with natives; the three 

introduced species we have worked with—crested wheatgrass, Siberian wheatgrass, and Russian wildrye—were most likely originally introduced on the property before it became a military installation. 

For the purposes of this study we defined an invasive species as an introduced species that will spread substantially beyond the areas it currently inhabits. 

The characteristics we were concerned about as indicators of potential invasiveness in introduced species are tiller and rhizome development and seedling 

encroachment through seed dispersal. (This study was concerned with the potential invasiveness of our introduced germplasms. Rhizome development is 

valuable in desired native species for land stabilization and reclamation of 

disturbed lands.) 

Summary of Data on the Spread of Introduced Species 

We evaluated the spread of the introduced species we used to develop new 

cultivars. We did not see any obvious vegetative spread by the introduced 

species, which are predominantly bunch-type grasses. For spread via escaped 

seeds, we measured seven sites: two four-year-old research sites and five areas 

seeded 13–19 years earlier. We found very few plants of crested or Siberian 

wheatgrass outside of the areas where they were planted, and after 19 years the 

greatest spread was only two plants found 12 m (40 ft) from the seeded plot. In 

one research site (Snake A) the native bluebunch wheatgrass was successfully 

invading the stands of introduced and native plants sown there four years ago. In 

the old seeding sites, Sandberg bluegrass (another native) was found to be 

invading the seedings of the wheatgrasses. Sandberg bluegrass is an understory 

plant that adds to the biological diversity of the site, and it is part of our breeding 

program (Palazzo et al. 1999; see Appendix B for the data collected for the 

review meeting). 

Discussion and Conclusions on the Spread of Introduced Species 

There is a great difference between the introduced, naturalized species that 

have been present on Yakima Training Center since ranchers used the land and 

the aggressive, exotic species that invade and exclude existing species from the 

community. Although some of the introduced grasses at Yakima appeared to be 

outside the area where they were originally sown, there was a greater movement 

of surrounding native vegetation into the sown plots. The introduced species 

were not forming monocultures nor did they appear to be “invading” new 
24 ERDC/CRREL TR-03-20 

habitats. It was evident that native grasses such as bluebunch wheatgrass and 

Sandberg bluegrass were moving into the seeded introduced areas. 

Participants on the review panel discussed the relative merits and disadvantages of native species compared to non-natives. Many also looked for assurance 

that the introduced species, which did not cause problems in the intended habitat, 

would not escape to other habitats and become noxious invaders. It was agreed 

that crested and Siberian wheatgrass had not established monocultures, nor were 

they invading other habitats in the area. With that in mind, participants weighed 

the other options available (such as the exclusive use of natives) and found those 

options would not be as successful in achieving all the goals of the project, 

primarily due to the slow establishment rate of the native plants. Introduced 

species are important to the ecosystem because they prevent the loss of fertile 

topsoil and help keep weeds out of disturbed areas until the natives can reestablish in those areas. There are no rapidly establishing native species that can 

provide the same degree of soil protection and weed prevention in the more arid 

areas around Yakima. 

Crested and Siberian wheatgrass appear to be excellent choices for rehabilitating areas subject to recurrent disturbance by providing soil protection and 

invasive weed suppression. Where the species volunteered outside seeded areas, 

they appeared to be filling in gaps rather than displacing native grasses. Crested 

and Siberian wheatgrass tend to appear early in the succession in low-disturbance 

areas. Late-seral native species were actively colonizing such sites where a seed 

source was available and disturbance was infrequent and of limited severity. 

The Independent Review Panel concluded that the plants we are using to 

develop wear-resistant cultivars for military training lands were not encroaching 

on other plant communities and were not establishing monocultures. The panel 

felt that continued long-term monitoring of some of the sites would be valuable, 

both to monitor the spread of the introduced plants and to investigate the competitiveness of crested and Siberian wheatgrasses planted with native species 

such as Snake River, western, and bluebunch wheatgrass and Sandberg 

bluegrass. 

We were encouraged that our work with seeding mixtures of native and 

introduced plants may lead to grass stands that are dominated by native species. 

The later section on “Methods for Establishing Natives and Fighting Invasive 

Weeds” describes how we are continuing to evaluate seeding mixtures so that we 

can find the best combination to maximize the establishment of natives and 

diminish the need for persistent non-natives. 
Improved Native Grasses and Establishment Methods 25 

Breeding Program 

Based on data from our field surveys, the LCTA program, and the seeded 

evaluation trials, we selected the following species to carry forward in our 

breeding program to develop the new germplasms; the right-hand column shows 

the current status of each species in our breeding program. 

Introduced 

Crested wheatgrass (SERDP-select RoadCrest and CD-II commercial 

releases) 

Siberian wheatgrass (SERDP-select) 

Russian wildrye (SERDP-select Tetra-1 public release) 

Native 

Bluebunch wheatgrass (P-7 multi-line public release and SERDP-select) 

Snake River wheatgrass (SERDP-select) 

Basin wildrye (SERDP-select) 

Western wheatgrass (two SERDP-select populations) 

Slender wheatgrass (two SERDP-select populations) 

Sandberg bluegrass (SERDP multi-line) 

Western yarrow (SERDP multi-line) 

The basic breeding process was described earlier in this report and illustrated 

in Figure 2. We used the following selection process for most of the species we 

developed in this program: 

Cycle 0 1. Transplant a broad range of materials to the field. 

 2. Evaluate plants for desired traits, such as stand establishment 

vigor, rate of tillering and rhizome development, vegetative vigor, 

and seed yield potential, and mark the best plants. 

 3. Allow selected plants to open pollinate with all plants or control 

pollination so that only the selected plants cross with each other. 

 4. Harvest seed from the selected plants. 

Cycle 1 1. Plant seeds harvested from selected clones 5 cm (2 in.) deep in 

cones in the greenhouse. 
26 ERDC/CRREL TR-03-20 

 2. Monitor germination and select the seedlings that emerge first and 

are the most vigorous. 

 3. Plant selected seedlings in the field to produce seed for the next 

cycle of selection. 

 4. Evaluate plants for vigor, etc. and mark the best plants (this may 

take two to three years). 

 5. Harvest seed from selected plants. 

This example includes one cycle of selection for vegetative vigor and other 

morphological traits in the field and one cycle for seedling vigor, as monitored in 

the greenhouse. With each cycle of selection, we assume we are accumulating 

favorable alleles (genes) that promote improved seedling vigor. Because emergence from a deep planting is a greenhouse test of seedling vigor, field testing is 

also necessary to validate that progress is being made from the selection process. 

Subsequent cycles repeat Cycle 1 until we have produced populations that show 

improvements over the original populations and that are genetically stable for the 

desired traits from one generation to the next. In our program, breeding populations were typically subjected to two cycles of selection. Table 7 illustrates how 

we were able to demonstrate improvements in certain traits when the original 

populations of known cultivars were compared with those of the selected 

populations. 

Table 7. Mean values of desired traits for four SERDP-select native germplasms compared to the “original” populations of known cultivars. 

Seedling emergence 

rate/day (mean) 

Seed weight 

(mean g/50 seeds) 

Total seed yield 

(mean g/plant) 

 Original Selected Original Selected Original Selected 

Bluebunch wheatgrass 1.2 2.6 0.2 0.3 5.9 8.3 

Western wheatgrass 2.3 3.9 

Snake River wheatgrass 2.3 3.9 11.8 16.3 

Basin wildrye 0.9 1.3 

Some new native germplasms were not subjected to selective pressure. 

Instead they were assembled from a range of ecotypes, resulting in a broader 

genetic base than in any of the individual populations and creating populations 

that should more readily adapt to changing stresses. Broad-based germplasms of 

Sandberg bluegrass, bluebunch wheatgrass (P-7), and western yarrow were 

developed by assembling seed collected from plant populations found on representative ecosystems throughout the training bases. To maintain the broad 
Improved Native Grasses and Establishment Methods 27 

genetic base within each germplasm, we did not subject these populations to 

intentional selection. Therefore, these germplasms are not specifically adapted to 

any given site on the base, but they are widely adapted to the training base as a 

whole. This type of germplasm may not be the most vigorous or the most productive for a given environment, but it is more likely to result in a successful 

seeding after military disturbance throughout the training base. Because these 

native species are important for recolonization or understory growth of disturbed 

sites, it is imperative that reseeding is successful. 

A second population of bluebunch wheatgrass (Yakima collections) also 

originated from a broad-based population that was then selected for seed germination, seedling and plant vigor, and emergence from a deep seeding depth. 

Similarly one of our two populations of slender wheatgrass seed originated as a 

broad-based population from several Fort Carson sites and was then selected for 

improved seed germination, seedling vigor, and plant vigor. 

Once the cycles are completed so that a genetically and morphologically stable germplasm is developed, our research phase is complete. In the next phase, 

we begin the seed increase process, and we register the new germplasm in the 

journal Crop Science. The registration communicates to others that we have a 

new source-identified or selected-class germplasm or a new cultivar. The seed 

increase, accomplished in production fields in Utah, is done both to produce seed 

for large-scale demonstrations (the seed harvest shown in Table 8 below) and to 

produce breeder’s seed. The seed harvested for large-scale demonstration usually 

amounts to about 150–200 kg/year (330–440 lb/year). The “Transition Plan” 

section of this report describes our large-scale demonstrations. We also increase 

seed to produce breeder’s seed to begin the initial production of seed for the 

newly released germplasms or cultivars. For breeder’s seed, seed is produced in 

large enough quantity (25 kg; 50–60 lb) to provide seed for sale. The typical 

process is for a middleman, such as a crop improvement association, to acquire 

the breeder’s seed from the researcher and use that seed to produce larger quantities of foundation seed. That foundation seed is, in turn, sold to commercial seed 

companies, who use it to produce certified seed for sale directly to users. 

Improved Species 

Table 8 summarizes the new traits developed in each new germplasm. 

Details for each species are given in the sections below. 
28 ERDC/CRREL TR-03-20 

Table 8. Improved traits and current status of SERDP-select germplasms. 

Introduced 

selections Original traits Traits of improved populations 

Seed 

harvest 

Release 

date

Russian wildrye 

RWR-Tetra-1 

Poor seedling vigor

Selected for improved seed germination and 

seedling vigor, increased plant height, longer and 

wider leaves, increased seedling emergence, 

heavier seeds, improved water-use efficiency 

 1997 

Crested wheatgrass 

CD-II Selected for increased growth under low 

temperatures, drought resistance, easy 

establishment 

 1996 

RoadCrest 

Moderate growth in 

low temperatures; 

few rhizomes 

Selected for low-maintenance turf with moderate 

rhizome development; suitable for gunnery ranges 

and roadside plantings; early spring growth 

 1998 

Siberian 

wheatgrass 

Moderate seedling 

vigor 

Selected for seedling vigor, plant color, vegetative 

vigor, seed yield, drought tolerance, early spring 

green-up 

2003–

2004 

2004 

Native selections Original traits Traits of improved populations 

Seed 

harvest 

Release 

date

Bluebunch 

wheatgrass 

 

P-7 Bluebunch 

wheatgrass 

Hard to establish; 

sensitive to grazing 

A broad-based multi-line population with no 

selection pressure applied 

 2001 

Bluebunch 

wheatgrass 

 A broad-based population selected for seed 

germination; seedling and plant vigor; emergence 

from a deep seeding 

2004 2004 

Western 

wheatgrass 

 

Turkey Creek 

population 

Strongly 

rhizomatous 

Selected for plant and seedling vigor, increased 

germination, seed yield 

2002–

2004 

2003 

South Boundary 

population 

 Selected for plant and seedling vigor, increased 

germination, seed yield 

2002–

2004 

2003 

Snake River 

wheatgrass 

Seedling vigor Selected for increased seedling vigor and seed yield 2003–

2004 

2004 

Basin wildrye Poor seedling vigor Selected for improved seed germination and 

seedling vigor 

2003–

2004 

2003 

Slender 

wheatgrass 

Poor persistence 

Bunch-type 

population 

 Broad-based population selected for emergence 

from a deep planting depth; improved plant vigor 

2002–

2004 

2004 

Rhizomatous 

population 

 Selected for same as above plus rhizome 

development 

2003–

2004 

2004 

Sandberg 

bluegrass 

Early establishment 

after a disturbance 

A broad-based multi-line population with no 

selection pressure applied 

2003–

2004 

2004 

Western yarrow 

(a forb) 

 A broad-based multi-line population with no 

selection pressure applied 

2002–

2004 

2003 
Improved Native Grasses and Establishment Methods 29 

Russian Wildrye (Psathyrostachys juncea; Introduced) 

Introduction. Russian wildrye is a long-lived bunchgrass that is exceptionally tolerant of cold and drought. Most of the forage produced is contained in the 

basal leaves, which grow rapidly in the spring as long as soil moisture is available. In North America it has been seeded most often on arid and semiarid 

rangelands of the Northern Great Plains and Intermountain regions in areas 

receiving between 150 and 300 mm of precipitation annually. It is better adapted 

to grazing than most grasses and is equal to crested wheatgrass in its ability to 

withstand drought. Because of its dense and vigorous growth, Russian wildrye 

competes effectively against undesirable plants within stands once it is established. Within the Great Basin, Russian wildrye is adapted to sagebrush, mountain brush, and juniper piñon sites, and it is useful on soils too alkaline for crested 

wheatgrass. The diploid cultivar Bozoisky is an excellent dryland bunchgrass that 

is drought resistant. 

Limitations. The use of Russian wildrye is often limited by its relatively 

slow seedling growth and development. Stands are often difficult to obtain, particularly under severe soil-water stress and where seeds have been planted too 

deeply. Russian wildrye does not tolerate spring flooding and is generally not 

well adapted to moist cool areas in the Great Basin. 

SERPD Russian wildrye. Asay et al. (1996) showed that by doubling the 

chromosome number in Russian wildrye from 2n = 14 to 2n = 28, the plants will 

have larger seeds and better seedling vigor than the currently used diploids. In the 

SERDP program, we developed a 28-chromosome tetraploid from Bozoisky Russian wildrye that has improved seed germination and seedling vigor.

Registration of RWR-Tetra-1 Tetraploid (2n = 28). RWR-Tetra-1 Russian 

wildrye [Psathyrostachys juncea (Fisch.) Nevski] germplasm (Reg. no. GP-75, 

PI 599302) was released in 1997 (Jensen et al. 1998). This 26-line composite was 

released as source material for genetic studies and for the development of 

improved cultivars of tetraploid (2n = 4x = 28) Russian wildrye. Most Russian 

wildryes found in natural populations are diploids (2n = 2x = 14) (Asay et al. 

1996). 

RWR-Tetra-1 traces to ten parental accessions that were initially obtained by 

K.H. Asay, D.A. Johnson, and M.D. Casler during a collecting expedition to 

Kazakhstan in 1988. Four of these accessions (AJC538, AJC539, AJC540, and 

AJC601) were identified as natural tetraploids, and the remaining lines (AJC595, 

AJC596, AJC597, AJC598, AJC599, and AJC600) were described as induced 

tetraploids. The ten parental tetraploid accessions of Russian wildrye were 

donated by the N.I. Vavilov Institute of Plant Industry (VIR), St. Petersburg, 
30 ERDC/CRREL TR-03-20 

Russia. The accessions have been entered in the National Plant Germplasm 

System (NPGS) as PI numbers 565063 to 565072. 

The ten parental accessions (Cycle-0) were evaluated from 1989 to 1992 at 

the Utah State University (USU) Evans Experimental Farm in Logan, Utah, 

(330–558 mm annual precipitation) and at the USU Blue Creek Experimental 

Farm (mean annual precipitation of 369 mm) in northwestern Utah. The parental 

accessions have significantly heavier seeds, greater seedling vigor, taller stature, 

and longer and wider leaves than standard diploid Russian wildrye cultivars and 

the tetraploid cultivar Tetracan. The parental accessions also have better water 

use efficiency than the diploid cultivars as determined by C-isotope discrimination measurement. 

At the Blue Creek Experimental Farm, open-pollinated seed from selected 

clones of the parental accessions was screened for seedling emergence from deep 

(7.6-cm) plantings in the greenhouse. The Cycle-1 population consisted of 20 

half-sib families (a total of 2160 plants) that traced to open-pollinated seed of 

selected accessions AJC538, AJC539, AJC540, AJC596, AJC597, AJC598, 

AJC599, AJC600, and AJC601. The Cycle-1 population was established in 1992 

as a space-planted nursery at the Evans Experimental Farm. Selection for vegetative vigor, seed yield, 100-seed weight, and seedling emergence from deep (7.6-

cm) seedings in the greenhouse was practiced among and within half-sib families. Seed was harvested only from selected plants to create the Cycle-2 population. Undesirable plant types were culled prior to pollination. The Cycle-2 population consisting of 40 half-sib families was established in nurseries on semiarid 

sites (annual precipitation 340 mm) near Nephi, Utah, and on the Curlew Grasslands in northwestern Utah (annual precipitation 150–250 mm) to initiate the 

third cycle of selection. 

The 40 half-sib families from the Cycle-2 population were evaluated for 100-

seed weight and rate of emergence from a 7.6-cm-deep planting. The half-sib 

families were significantly greater (P < 0.05) for both characteristics than were 

the diploid cultivars tested. Using a selection index that included forage yield, 

total seed weight, and individual seed weight (double weighted), equal amounts 

of open-pollinated seed from 26 selected half-sib families were combined to generate the broad-based RWR-Tetra-1 germplasm. 

RWR-Tetra-1 is highly cross-pollinated and behaves meiotically as an autotetraploid (2n = 4x = 28) (genome constitution NsNsNsNs) with a high frequency 

of multivalents typical of those observed in artificially induced tetraploids of 

Russian wildrye (Wang and Berdahl 1990). It is the first release of tetraploid 

Russian wildrye germplasm that includes naturally occurring tetraploid plants in 

its parentage. Previous work, including that conducted with the cultivar Tetracan 
Improved Native Grasses and Establishment Methods 31 

(Dhindsa and Slinkard 1963, Lawrence et al. 1990), has been limited to 

tetraploids artificially induced from diploids. Morphologically, RWR-Tetra-1 is 

taller than the Vinall, Cabree, and Tetracan cultivars and has longer leaves than 

Vinall. RWR-Tetra-1 has heavier seeds than existing diploid cultivars. Spike 

characteristics are similar to diploid Russian wildrye. 

Crested Wheatgrass (Agropyron cristatum and A. desertorum; Introduced) 

Introduction. Crested wheatgrass is an extremely long-lived, droughttolerant, bunch to moderately rhizomatous range grass that is adapted to a wide 

range of ecological sites and zones receiving as little as 200–250 mm of annual 

precipitation. In North America it is particularly well adapted to the northern and 

central Great Plains and the more arid intermountain region. It is very winter 

hardy and has an extensive root system that gives it excellent drought resistance. 

Vegetative growth is greater from mid-April to mid-June and declines rapidly by 

early July. It has very vigorous seedlings, making it relatively easy to establish 

under harsh environmental conditions. With seedling vigor and times of rapid 

growth coinciding with those of invasive annual weeds, crested wheatgrass is a 

natural competitor. Crested wheatgrass does well on most fertile soils, including 

heavy clay soils, and it is fairly tolerant of highly alkaline soils. 

There are three recognized morphological types of crested wheatgrass: fairway (A. cristatum), standard (A. desertorum), and Siberian (A. fragile). The fairway type has short, broad spikes that taper at the top, and, compared to the standard type, it has smaller seeds, grows shorter, and has finer leaves and stems. The 

spikes on the standard type are longer than on the fairway type, but they vary in 

shape from comb-like to oblong. The spikes of the Siberian type are longer and 

much narrower than on the standard and fairway types. (Throughout this document we have grouped the Siberian crested wheatgrasses separately from the 

fairway- and standard-type crested wheatgrasses.) 

Limitations. In general, crested wheatgrass is not adapted to moist soils and 

cool, short growing days with high humidity. It will tolerate short periods of 

spring flooding that do not exceed 7–10 days, but it is intolerant of soils with 

high water tables. Crested wheatgrass is less tolerant of soil salinity than tall 

wheatgrass, western wheatgrass, and quackgrass. Reduced plant vigor and poor 

stands are likely at elevations above 1980 m (6500 ft) in the Great Basin. 

SERPD crested wheatgrass. We have released two new crested wheatgrass 

cultivars: CD-II and RoadCrest. 

Registration of CD-II Crested Wheatgrass. CD-II (Reg. no. CV-24, PI 

594024) crested wheatgrass is a 10-clone synthetic derived from the cultivar 
32 ERDC/CRREL TR-03-20 

Hycrest, which is a hybrid between the induced tetraploid Agropyron cristatum

(L.) Gaertner and the natural tetraploid Agropyron desertorum (Fisch. ex Link) 

Schultes. CD-II was released on 25 January 1996 (Asay et al. 1997). 

A breeding program was initiated in 1985 to improve the Hycrest breeding 

population. The base population was derived from 100 clonal lines, which were 

selected from a Hycrest foundation seed-increase block consisting of 40,000 

spaced plants. Selection was based primarily on vegetative vigor and the absence 

of purple leaves during the early spring, tolerance to diseases and insects, and 

leafiness. 

The 100 clonal lines were evaluated a second time in a 10-replicate crossing 

block for the same vegetative characters as well as for individual seed weight and 

emergence of polycross seedlots from deep seedings. Polycross progenies from 

30 selected clonal lines were bulked in equal quantities to form a breeding population, which was advanced through two additional breeding cycles of selection 

for leafiness, vegetative vigor, and seedling vigor. Polycross seed from 10 clonal 

lines selected from the final breeding cycle was bulked to form breeder’s seed. 

CD-II produces significantly more forage at low temperatures in the growth 

chamber than Hycrest (Table 9). The seedling vigor of CD-II on a field site near 

Logan, Utah, was significantly greater than Hycrest (Table 10). CD-II has been 

distinguished from Hycrest on the basis of Random Amplified Polymorphic 

DNA (RAPD) fingerprinting profiles (Hu et al. 2001) (see the section on 

“Molecular Marker Development: Markers for Cold Tolerance and Early Spring 

Growth” later in this report). 

Table 9. Growth of CD-II and Hycrest seedlings 

at low temperature in a growth chamber. 

Cultivar 

Grams / plot 

10 ̊C (50 ̊F) 

Grams / plot

5 ̊C (41 ̊F)

Hycrest 3.34 4.94 

CD-II 5.39 6.17 

Table 10. Growth of CD-II and Hycrest seedlings at a field site near 

Logan, Utah. 

Seedling height at 

6 weeks (cm) 

Vigor rating 

(1 to 9) 

Stand rating

 (1 to 9) Dry weight (g) 

Hycrest 5.7 6.5 7.5 104 

CD-II 10.4 8.5 8.5 117 

LSD (0.05) 1.0 0.6 1.2 23 
Improved Native Grasses and Establishment Methods 33 

The CD-II cultivar produces from 670 to 900 kg seed/ha on sites receiving 

400–450 mm of annual precipitation with no supplemental irrigation. CD-II produces abundant forage during the spring and early summer, and it is recommended for semiarid range sites in the Intermountain Region and Great Plains 

receiving 200–450 mm of annual precipitation at altitudes up to 2200 m. When 

drilled under dryland range conditions, a seeding rate of 7–9 kg/ha is 

recommended. 

Registration of RoadCrest Crested Wheatgrass. RoadCrest crested 

wheatgrass (Reg. no. CV-25, PI 606546), a rhizomatous cultivar of crested 

wheatgrass [Agropyron cristatum (L.) Gaertn.], was released on 2 June 1998 

(Asay et al. 1999). It was derived from two accessions originally collected at 

Dikmen Ankara and Güvercinlik Ankara, Turkey, and provided to the USDAARS Forage and Range Research Unit by Dr. Esvet Acikgoz of Turkey. Rhizomatous plants, observed in these accessions during their initial evaluation on a 

semiarid range site in central Utah, were selected and established in a crossing 

block to develop the parental germplasm. These plant materials were then subjected to two cycles of selection based largely on progeny evaluation for 

increased rhizome development, fine leaf texture, short growth stature, and 

improved seedling vigor. Breeder’s seed was compounded from selected 

polycross seedlots in the final breeding cycle. 

RoadCrest is long-lived and is significantly more rhizomatous than any other 

crested wheatgrass included in evaluation trials, including the cultivar Ephraim, 

which is the only other rhizomatous cultivar of crested wheatgrass to be released. 

RoadCrest produces less biomass and is 15–25% shorter in stature than foragetype crested wheatgrass cultivars (Table 11). The cultivar has been evaluated on 

roadsides and in turf trials in Utah, Colorado, and Washington. Seedling vigor 

and drought resistance of RoadCrest compares favorably with other crested 

wheatgrasses cultivars, including Hycrest, CD-II, Fairway, and Nordan. RoadCrest is significantly easier to establish and initiates growth earlier in the spring 

than other turf and low-maintenance turf grasses, including Kentucky bluegrass, 

‘Sodar’ thickspike wheatgrass, tall fescue, and hard fescue. Leaf color intensity 

and turf quality of RoadCrest are not as good as Kentucky bluegrass, tall fescue, 

and perennial ryegrass when the latter grasses are grown under optimum conditions. RoadCrest greens up early in the spring and remains green until midsummer on temperate sites similar to Logan, Utah. Plants then go dormant until 

temperatures decline in the fall. This characteristic varies according to summer 

temperatures and annual precipitation. Summer dormancy is much less prominent 

at elevations above 1500 m. 
34 ERDC/CRREL TR-03-20 

Table 11. Growth of RoadCrest and Ephraim 

crested wheatgrass in a greenhouse study. The 

differences between the two grasses were significant 

at the 0.01 probability level for both plant height and 

crown width.

 Plant height (cm) Crown width (cm)

RoadCrest 72 66 

Ephraim 85 59 

RoadCrest has good potential as a low-maintenance turf grass on semiarid 

sites (250–500 mm of annual precipitation) such as roadsides, low-use playing 

fields, walking trails, lawns, and other non-irrigated cantonment areas, as well as 

on ammunition storage bunkers in training areas. As with other crested wheatgrass cultivars, supplemental irrigation can be detrimental if the total water application (precipitation plus irrigation) exceeds 550 mm annually. RoadCrest was a 

top performer in the dry conditions at the South Boundary site at Fort Carson. 

RoadCrest is a natural tetraploid (2n = 4x = 28) and is fully interfertile with 

tetraploid cultivars of crested wheatgrass (Agropyron cristatum and A. 

desertorum), Siberian crested wheatgrass (A. fragile), and the crested wheatgrass 

hybrid cultivars Hycrest and CD-II. Established isolation distances from these 

cultivars must be maintained in certified seed fields. The cultivar produced 560 

kg of seed/ha when grown in rows 0.9 m apart with moderate supplemental 

irrigation and 50 kg N/ha applied in the fall. At 100% purity, there are approximately 530,000 seeds/kg. 

Siberian Wheatgrass (Agropyron fragile; Introduced) 

Introduction. The Siberian type of crested wheatgrass occupies sites where 

standard crested wheatgrass will grow, but it is particularly noted for its ability to 

establish quickly on drier, sandy soils (200–260 mm annual precipitation). The 

Siberian type of crested wheatgrass is awnless, has finer leaves, and retains its 

greenness and palatability later into the summer than standard- and fairway-type 

crested wheatgrasses discussed in the previous section. 

Limitations. Siberian wheatgrass has poor seedling vigor along with the 

same limitations as the other crested wheatgrasses (see the previous section). 

SERDP-select Siberian wheatgrass. This germplasm originated from the 

cultivar Vavilov (Asay et al. 1995) and from collections from western 

Kazakhstan. The parental germplasm for Vavilov was derived from accessions 

originally received from the N.I. Vavilov Institute of Plant Industry (VIR); the 
Improved Native Grasses and Establishment Methods 35 

Stavropol Botanical Garden, Stavropol, Russia; the Eskisehir Plant Breeding 

Station, Eskisehir, Turkey; and selections from the cultivar P-27. In addition, it 

included selected lines from Kazakhstan that were originally evaluated under the 

SERDP program for seed yield, seed weight, seedling vigor, forage yield, and 

military suitability (Jensen et al. 2000). 

Cycle-0 was initiated by screening seedlings of Vavilov and Kazakhstan 

collections for emergence from a 6.7-cm-deep seeding (an estimate of seedling 

vigor). These plants were established at Yakima Training Center (832 plants), 

Lakeside, Utah (600 plants), and Snowville, Utah (600 plants), in 1999. Based on 

plant vigor, total seed weight (g/plant), 100-seed weight (g), and rate of seedling 

emergence from a 7.6-cm-deep planting, 50 plants (15 Yakima, 29 Lakeside, and 

6 Snowville) were selected for generation advance. Table 12 compares the performance of the selected plants (50; Cycle-1) with the average of the entire 

breeding population for the above characteristics. 

Table 12. Characteristics of the Siberian wheatgrass selections compared 

to the entire original breeding population. 

Original population 

mean 

Selected population 

mean 

Seedling emergence rate (seedlings/day) 2.2 3.1 

Seed weight (g/100 seeds) 0.29 0.33 

Total seed yield (g/plant) 26 25 

Clones from the Cycle-1 polycross block (50) were established in a seed 

increase block at the Blue Creek Research Station in northern Utah in 2000. Bulk 

seed, which is available to the Army for seed increase, was harvested for testing 

in 2001 and 2002. In addition, 10 clones from Cycle-1 were selected based on 

performance and polycrossed in the greenhouse winter of 2001-02 to initiate 

Cycle-2. Cycle-2 progeny were screened for their ability to emerge from a 7.6-

cm-deep planting. Superior genotypes (1800 plants) were established at the 

Nephi Research Station (Nephi, Utah) in 2002 for further selection for seedling 

vigor and plant vigor. This is an excellent-looking population with good seedling 

and plant vigor. A formal release of Cycle-2 is expected in 2004. SERDP-select 

Siberian wheatgrass Cycle-2 was established in three demonstration nurseries in 

Colorado, Washington, and Utah during the fall of 2002. 
36 ERDC/CRREL TR-03-20 

Bluebunch Wheatgrass (Pseudoroegneria spicata; Native) 

Introduction. Bluebunch wheatgrassis a cross-pollinated, predominantly 

diploid species widely distributed in the IntermountainWest. It is a long-lived, 

drought-tolerant native bunchgrass that begins growth early in the spring and 

again with the onset of fall rains. It is an important native grass on Palouse 

Prairie and Intermountain sagebrush sites, and it is widely adapted geographically from high elevations in the Sonoran Desert north to Washington. It is 

adapted to dry mountain slopes, sagebrush, ponderosa pine, mountain-brush, and 

juniper-piñon ranges receiving 250–350 mm of precipitation annually. Bluebunch wheatgrass is adapted to coarse-textured soils, but it will establish and persist on deep, well-drained loamy soils. In the Palouse Prairie of eastern Washington, bluebunch wheatgrass is often associated with Idaho fescue on deeper 

soils and Sandberg bluegrass on shallower soils. In the Great Basin, bluebunch 

wheatgrass occurs as a co-dominant species with big sagebrush on arid sites that 

receive 120–400 mm of annual precipitation. At Yakima Training Center we 

have observed it to be a natural invader of lands sown to introduced species (see 

Table B1 in Appendix B, “Background Data for Non-Invasiveness of Introduced 

Species”). 

Limitations. Even though bluebunch wheatgrass recovers rapidly after limited grazing, it is not tolerant of heavy or repeated grazing or military training 

during the growing season. Because of its poor seedling vigor, bluebunch wheatgrass requires several years for stands to reach full productivity. 

SERDP bluebunch wheatgrass. To improve the genetic diversity and 

adaptability of bluebunch wheatgrass, we have developed two new germplasms 

(P-7 and SERDP-select) from polycrosses of multiple collections. The P-7 

release is a multiple-origin polycross with no selection pressure applied. The 

SERDP-select germplasm originated from a multiple-origin polycross, and we 

are selectively breeding in that population for improved seed germination, seedling and plant vigor, and emergence from a deep seeding. 

Registration of P-7 Bluebunch Wheatgrass Germplasm. P-7 bluebunch 

wheatgrass [Pseudoroegneria spicata (Pursh) A. Löve] germplasm (Reg. no. GP7, PI 619629) was released on 28 February 2001 as a Selected Class Germplasm 

(Jones et al. 2002). This class of prevariety germplasm is eligible for seed certification under guidelines developed by the Association of Official Seed Certifying 

Agencies (2001). 

P-7 is a multiple-origin polycross generated by intermating two cultivars and 

23 open-pollinated, native-site collections from Washington, Oregon, Nevada, 

Utah, Idaho, Montana, and British Columbia. The two cultivar populations were 

Whitmar and Goldar, which both originated in southeasternWashington. Whit-
Improved Native Grasses and Establishment Methods 37 

mar is an awnless cultivar developed from a population collected near Colton, 

Whitman County, Washington, and Goldar is an awned cultivar developed from a 

population collected near Anatone, Asotin County, Washington; they were 

released by the USDA-NRCS in 1946 and 1989, respectively (Hein 1958, Gibbs 

et al. 1991). Nine of the remaining populations were collected by T.A. Jones (PI 

537368, Pollock,Idaho; PI 537370, Riggins, Idaho; PI 598821, Wawawai Park, 

Washington; PI 537374, Steptoe Butte, Washington; PI 537375, Durkee, Oregon; 

PI 537378, Lone Mountain Junction, Nevada; PI 516185, Seneca, Oregon; PI 

537388, Dayton, Washington; PI 563870, Green Canyon, Utah), seven were collected by K.H. Asay (PI 563872, New Meadows, Idaho; PI 563867, Colton, 

Washington; PI 563868, Wawawai Road, Washington; PI 563874, Wawawai 

Park, Washington; PI 562050, Wawawai Park, Washington; PI 598816, Connell, 

Washington; PI 562056, Lind, Washington), and seven were obtained from miscellaneous sources (PI 595192, WawawaiRoad, Washington; PI 595193, Almota 

Road, Washington; PI 595196, Darby, Montana; PI 236670, Slocan, British 

Columbia; P-3, Grande Ronde River, Oregon; P-5, unknown; KJ-10, Salina Canyon, Utah). Twenty-four of the constituent populations are diploid (2n = 2x = 14) 

and one (PI 537374) is tetraploid (2n = 4x = 28). The inclusion of the tetraploid 

PI 537374 in the polycross was inadvertent. The representation of this tetraploid

is expected to decline dramatically through generations of seed increase. Therefore, P-7 can be considered to be predominately diploid—the dominant ploidy 

level of bluebunch wheatgrass.

Sixteen of the P-7’s 25 component populations are predominately awned and 

nine are predominately awnless. Because the awnless state is dominant and the 

awned state is recessive in bluebunch wheatgrass and its relatives, P-7 individuals 

are predominately awnless.

P-7 is intended to provide genetic diversity within a single germplasm for 

semiarid to mesic sites where bluebunch wheatgrass was an original component 

of the vegetation. Our genetic studies confirm that P-7 displays greater genetic 

variation than the Goldar or Whitmar cultivars (Larson et al. 2000; see the section on “Molecular Marker Development: Genetic Variation” later in this report). 

Breeder’s seed of P-7 was bulked across the 25 populationsin direct proportion to their seed yield in 1995 in a replicated test at the Blue Creek Experimental 

Farm in northwestern Utah. Generation-0 (the separate 25 populations), G-1 (first 

intermating), and G-2 (second intermating) will be maintained by the USDAARS Forage and Range Research Laboratory, Logan, Utah. G-2 seed will be 

made available to growers for production of G-3 and G-4 generations of seed 

(third and fourth intermating) by the Utah Crop Improvement Association. To 

limit genetic shift, the sale of P-7 seed beyond generation G-4 is expressly prohibited.
38 ERDC/CRREL TR-03-20 

SERDP-Select Bluebunch Wheatgrass. A second SERDP-select bluebunch 

population was derived from approximately 22 collections near Yakima, Washington. The original collections were evaluated at Yakima Training Center (Exit 

11) in a space-planted nursery (3750 plants). This base population was screened 

for vegetative vigor, and seed from 78 superior clones was harvested in 1996 

(Cycle-1). Forty of these progeny lines were subsequently selected on the basis 

of total seed yield (g/plant), 50-seed weight (g), and emergence from a 7.6-cmdeep seeding. These lines were then established in a 1600-plant nursery (four 

replications) at the Blue Creek Research Station in 1997 to initiate the next cycle 

of field screening. Open-pollinated seed was harvested from 53 clones selected 

from this nursery on the basis of superior plant vigor. Subsequent screening for 

total seed weight (g/plant), 50-seed weight (g), and emergence from a 7.6-cmdeep seeding reduced the 53 plants to five for generation advance (Table 13). 

Open-pollinated seed from the five selected lines was screened for seedling 

vigor, with superior genotypes being established in a seed increase field in northern Utah in 2001. Seed for demonstrations was harvested in 2002. A Selected 

Class Germplasm release is expected in 2004. 

Table 13. Characteristics of bluebunch wheatgrass selections compared 

to the original breeding population. 

Original 

population mean 

Selected 

population mean 

Seedling emergence rate (seedlings/day) 1.24 2.58 

Seed weight (g/50 seeds) 0.23 0.30 

Total seed yield (g/plant) 5.91 8.32 

Western Wheatgrass (Pascopyrum smithii; Native) 

Introduction. Western wheatgrass is native to North America and is distributed throughout the western two-thirds of the United States and Canada. It is an 

important rangeland and pasture grass in the central and northern Great Plains 

and Intermountain Region. It is a long-lived, late-maturing, rhizomatous, coolseason grass with coarse, blue-green leaves. Because of its sod-forming characteristics, western wheatgrass is well adapted for stabilizing soils on sites subject 

to excessive erosion as well as on rangelands disturbed by military training 

activities, surface-mining operations, construction, overgrazing, brush control, 

and fires. Western wheatgrass is winter hardy and can survive drought. It is 

adapted to a wide range of soils, but it prefers the heavy and somewhat alkaline 

soils characteristic of shallow lakebeds or intermittent watercourses that receive 
Improved Native Grasses and Establishment Methods 39 

excess surface drainage. It requires moderate to high soil moisture content and is 

most adapted to regions receiving 250–350 mm of annual precipitation. When the 

annual precipitation exceeds 500 mm, western wheatgrass usually becomes very 

aggressive. Western wheatgrass is commonly associated with blue grama and the 

needlegrasses of the Great Plains and with bluebunch and thickspike wheatgrasses and various shrubs of the Great Basin, where it is an important constituent of spring, summer, and early fall ranges. At arid and semiarid sites, western 

wheatgrass is often confused morphologically with thickspike wheatgrass. Its 

elevation range is 300–2750 m (1000–9000 ft). 

Limitations. Western wheatgrass germinates poorly, has a high rate of seed 

dormancy, has low seedling vigor, and often requires two or three years to reach 

well-established stands. As a result, plants are initially scattered in stands, but 

because of its aggressive rhizome development, it may eventually dominate the 

site within three to four years. Western wheatgrass is a poor seed producer. 

SERDP-Select Western Wheatgrass. This population originated from collections near and on Fort Carson, Colorado, from germplasm accessions, and 

from already existing cultivars commonly used in reseeding projects. We are 

developing two populations: one each from the Turkey Creek and South Boundary areas of Fort Carson. 

The Turkey Creek germplasm includes parental material collected from Fort 

Carson (WW117FC), germplasm related to the cultivar Rodan (D2945), and 

Rosana, a well-proven cultivar. The current population has undergone one cycle 

of selection for seed yield, plant vigor, and general adaptation to the Turkey 

Creek area; and two cycles of selection for increased germination and seedling 

vigor. Initially, individual plants of a large number of germplasm collections and 

cultivars were evaluated at the Turkey Creek Recreation Area for two years. In 

1998, open-pollinated seed was harvested from the best plants that were selected 

for adaptation to the area, moderate spread, vigorous growth, and abundant seed 

production. The harvested seed was planted in a Logan, Utah, greenhouse 6.3 cm 

deep and evaluated for rate of emergence (Table 14). The 50 fastest-emerging 

seedlings of the top four plants were transplanted to containers and, in 1999, 

transplanted into a field-crossing block near Logan, Utah. In 2001, seed was harvested from the crossing block and screened a second time for the ability to 

emerge from a deep planting. Two thousand seedlings with the fastest emergence 

were selected (500 from each parental source) and used to establish a seed 

increase field in 2001. Seed was harvested in 2002 and used for test plantings and 

for evaluating the gain from selection for improved seedling vigor. 
40 ERDC/CRREL TR-03-20 

Table 14. Seedling emergence rate of western wheatgrass from a 6.3-cmdeep planting. 

Original 

population mean 

Selected 

population mean 

Seedling emergence rate (seedlings/day) 2.3 3.9 

Total seedling emergence 31.6 52.9 

The South Boundary germplasm was selected for improved seedling vigor 

and adaptation to the South Boundary area. It initially comprised a selection of 

seven original plants that survived the harsh, dry conditions found at the South 

Boundary evaluation plot area. These plants originated from the cultivars Arriba 

and Rosana and from the D2945 germplasm described above. The seven plants 

were selected from a large evaluation nursery as being the most vigorous plants 

of the few surviving entries. Clones of each selected plant were transplanted to a 

field crossing block near Logan, Utah, in 1999. In 2001, seed was harvested from 

the crossing block and screened as described above for the ability to emerge from 

a deep planting, demonstrating improved germination and seedling vigor. Two 

thousand seedlings with the fastest emergence were selected (approximately 300 

from each parental source) and used to establish a seed increase field in 2001. 

Seed was harvested in 2002 and used for test plantings and for evaluating the 

gain from selection for improved persistence and seedling vigor. 

Snake River Wheatgrass (Elymus wawawaiensis; Native) 

Introduction. Snake River wheatgrass is very well adapted in the northwest, 

especially at Yakima Training Center. It is a long-lived, perennial bunchgrass 

that is native to the valleys of the Snake River and its tributaries in Washington 

and northern Idaho. Morphologically this taxon is almost identical to bluebunch 

wheatgrass, but genetically it is similar to thickspike wheatgrass. It is adaptable 

to most areas suitable for bluebunch wheatgrass but is more vigorous and productive. Snake River wheatgrass has been successfully established on sites that 

receive as little as 200 mm of annual precipitation. At many sites, Snake River 

wheatgrass grows in grass mixtures with unawned forms of bluebunch wheatgrass. At such sites, Snake River wheatgrass is easily distinguished by its awns. 

Limitations. Limitations for Snake River wheatgrass are similar to those for 

bluebunch wheatgrass. Because of its poor seedling vigor, Snake River wheatgrass requires several years for stands to reach full productivity. During seedling 

establishment, plants should be allowed to reach maturity before being subjected 

to grazing or military training activities. 
Improved Native Grasses and Establishment Methods 41 

SERDP-Select Snake River Wheatgrass. The SERDP-select Snake River 

wheatgrass population originated from ‘Secar’ and two native collections near 

Yakima, Washington. Based on results from the evaluation trials in 1998, the 

cultivar Secar accessions Acc:707 and T572 were selected and established in a 

3096-plant space-planted source nursery. Space plants of Secar, Acc:707, and 

T572 had previously been screened for rate of emergence from a 7.6-cm-deep 

planting. Based on increased plant vigor, open-pollinated seed from 302 plants 

(10% selection), which included plants from all three accessions, were harvested 

in 1999 (Cycle-1). Based on total seed weight (g/plant) and emergence rate from 

a 7.6-cm-deep seeding, 36 lines were selected and clones removed from the 

Yakima Training Nursery in November 1999 (Table 15). 

Table 15. SERDP-select Snake River wheatgrass Cycle-1 compared to the 

mean of the base population. 

Beginning 

population mean 

SERDP-selected 

population mean 

Seedling emergence rate (seedlings/day) 2.3 3.9 

Total seed yield (g/plant) 11.75 16.25 

Cycle-1 clones were established (1080 space plants) in a seed increase block 

at the Blue Creek Research Station in northern Utah in 2000. Bulk seed, which is 

available to the Army for seed increase, was harvested for testing in 2001 and 

2002. In addition, eight clones selected from Cycle-1 were polycrossed in the 

greenhouse during the winter of 2001-02 to initiate Cycle-2. Cycle-2 progeny 

were screened for their ability to emerge from a 7.6-cm-deep planting. Superior 

genotypes (2200 plants) were established at the Nephi Research Station in 2002 

to facilitate further selection for seedling and plant vigor. This is an excellentlooking population with good seedling and plant vigor. A formal release is 

expected in 2004. SERDP-select Snake River Cycle-1 was established in three 

demonstration nurseries in Colorado, Washington, and Utah during the fall of 

2002. A release as a Selected Class Germplasm is expected in 2004. 

Basin Wildrye (Leymus cinereus; Native) 

Introduction. Basin wildrye is a hardy, robust, long-lived perennial bunchgrass that is native throughout the western United States. It is tall and coarse, and 

it is highly palatable early in the spring but becomes very unpalatable with 

maturity. It is used as forage by livestock and wildlife. Poor seedling vigor usually results in sparse stands, but it is one of the highest-producing grasses once 

established. It can be used as a component in mixtures with other adapted plants 
42 ERDC/CRREL TR-03-20 

to reseed rangelands, mine spoils, highway rights-of-way, or other disturbed 

areas. 

Basin wildrye is widely distributed throughout British Columbia to Saskatchewan, south to California, and east to Arizona and Colorado. It is adapted to 

the fine-textured soils in valley bottoms, along roadsides and streams, and in gullies. In sagebrush ecosystems it is often found on riverbanks or waterways, in 

ravines, and on other sites with a high water table. It is adapted to areas with an 

average annual precipitation of more than 350 mm. It is moderately tolerant of 

alkaline and saline soils. Basin wildrye is often associated with other wheatgrasses, rabbitbrush, and sagebrush. 

Limitations. As with other wildrye species, Basin wildrye is usually difficult 

to establish. Poor seed development along with low germination and seedling 

survival has limited its usefulness for range revegetation. New seedlings should 

not be grazed or harvested until the late summer or fall of the second growing 

season. 

Genetic marker studies. Inheritance studies with Basin and beardless wildrye hybrids found markers associated with leaf growth and carbohydrate content, 

suggesting that these two traits associated with early spring growth could be 

simultaneously improved (Larson et al. 1999; see the section on “Molecular 

Marker Development: Markers for Cold Tolerance and Early Spring Growth” 

later in this report). 

SERDP-Select Basin Wildrye. This germplasm originated from 14 collections made on the Yakima Training Center, Washington. The original collections 

were evaluated at the Exit 11 site at Yakima Training Center in a space-planted 

nursery (700 plants). Based on increased plant vigor, open-pollinated seed from 

118 plants, which included all 14 collections, was harvested in 1996 (Cycle-1). 

Based on total seed weight (g/plant), 50-seed weight (g), and emergence from a 

7.6-cm-deep seeding, seed from 40 plants was selected and established at the 

Blue Creek Research Station in northern Utah (1600 space plants/four replications) in 1997. Open-pollinated seed of 53 Cycle-2 plants was selected from the 

Blue Creek Nursery based on superior plant vigor. Subsequent screening for total 

seed weight (g/plant) and emergence rate from a 7.6-cm-deep seeding reduced 

the 118 plants to 18 for generation advance. Open-pollinated seed from the 18 

selected plants was screened for seedling vigor (Cycle-2), with superior genotypes being established in a seed increase field in northern Utah in 2002. Seed 

will be available for the Army and demonstration nurseries by 2003. A Selected 

Class Germplasm release is expected in 2004. 
Improved Native Grasses and Establishment Methods 43 

Slender Wheatgrass (Elymus trachycaulus; Native) 

Introduction. Slender wheatgrass is a short-lived, native bunchgrass with 

good seedling vigor and moderate palatability. It is widely adapted throughout 

the western United States and Canada, where it is found growing at elevations of 

1370–3660 m (4,500–12,000 ft) along dry to moderately wet roadsides, streambanks, meadows, and woodlands from the valley bottoms to subalpine and alpine 

elevations in aspen and open coniferous forests. It is most common in Montana 

and Wyoming, where it has been reported between 900 and 2135 m (3000 and 

7000 ft). The salinity tolerance of slender wheatgrass is similar to that of Russian 

wildrye and less than that of tall wheatgrass. The seed is larger than most wheatgrasses and is easily seeded with conventional seed drills. 

This self-fertilizing bunchgrass was the first native grass to be widely used in 

revegetation programs in the western United States and Canada. Slender wheatgrass tolerates a wide range of conditions and adapts well to high-altitude ranges 

and more favorable sites on aspen and tall-mountain brush regions. Because of its 

rapid seedling germination and establishment, moderate salt tolerance, and compatibility with other species, slender wheatgrass is a valuable component in 

erosion-control and mine-land reclamation seed mixes. Slender wheatgrass is 

most commonly used as a component of a mix in burned or disturbed-site rehabilitation, where it serves as a cover crop, eventually giving way to longer-lived 

perennials. 

Limitations. Slender wheatgrass is less drought tolerant than most of the 

wheatgrasses, including crested and bluebunch wheatgrass. It prefers loams and 

sandy loams in areas receiving at least 350 mm of annual precipitation, but it 

does not tolerate waterlogged soils. Slender wheatgrass is relatively short lived. It 

yields well for the first three to four years, but then the stand density decreases 

rapidly. It is not as competitive with weeds as other wheatgrasses, but it is shade 

tolerant. Slender wheatgrass is highly susceptible to the grass billbug (Sphenophorus spp.), which can severely reduce stands. Slender wheatgrass is not resistant to close grazing, and continual heavy grazing reduces stands quickly. 

SERDP-Select Slender Wheatgrass. We are developing two populations, a 

bunch type and a rhizomatous type. The bunch-type population originated from 

seven collections made on Fort Carson, Colorado, in 1994. An initial spaceplanted nursery (560 plants) was established at the South Boundary site at Fort 

Carson in 1995. Based on visible plant vigor, 210 plants were selected in 1997. 

These were established as clones at Snowville, Utah, in 1998, and seed was harvested in 1999 to generate a multi-line germplasm. A seed increase field of this 

multi-line composite was established in 2000, and large-scale seed harvests were 

completed in 2001 and 2002; we established three demonstration nurseries in 
44 ERDC/CRREL TR-03-20 

Colorado, Washington, and Utah during the fall of 2002. We currently have several hundred pounds of this germplasm available for the DoD demonstrations. 

This bunch-type germplasm is an excellent-looking population with good seedling and plant vigor. A Selected Class Germplasm release is expected in 2004. 

The rhizomatous population originated from a multi-plant collection near 

Pike’s Peak in Colorado. This collection is unique in that it exhibited a limited 

degree of rhizome development. Early in the 1990s, we cycled this population for 

persistence and rhizome development. As part of the SERDP project, we established a space-planted nursery (2250 plants), which had been screened for emergence from a 7.6-cm-deep planting in 2000. In 2001, 34 plants were selected 

based on rhizome development (Fig. 3), persistence, and seed production. In 

2002, self-pollinated seed of these 34 plants were once again screened for rate of 

emergence from a 7.6-cm-deep planting and established at two locations in 

southern and northern Utah. A Cycle-2 seed increase with increased seedling 

vigor and germination was established in the spring of 2002. The development of 

a rhizomatous native that has the ability to establish quickly, reduce erosion, and 

allow for the establishment of more-persistent natives is important in the restoration process. A Selected Class Germplasm release is expected in 2004. 

Figure 3. Comparison of populations of slender wheatgrass. The selected 

population on the left has rhizomes and the unselected population on the 

right lacks rhizomes. 
Improved Native Grasses and Establishment Methods 45 

Sandberg Bluegrass (Poa sandbergii; Native) 

Introduction. Sandberg bluegrass is an apomictic native grass and is an 

important understory plant of the sagebrush–bluebunch wheatgrass communities 

found on the Yakima Training Center. It is a perennial bunchgrass that grows in 

small tufts no more than 0.3 m (1 ft) in diameter. It is widely distributed throughout the western range where annual precipitation is 350–500 mm, and it is somewhat tolerant of heavy trampling. Once established, Sandberg bluegrass is perhaps the most drought tolerant of the bluegrasses. It is adapted to mountains, 

uplands, and semi-deserts on lighter-textured and stony loam soils. It inhabits a 

wide range in elevations, being found as low as 300 m (1,000 ft) in Washington 

and as high as 3,660 m (12,000 ft) in the mountains of northern New Mexico. 

Limitations. The seed viability of Sandberg bluegrass is poor, and forage 

quality declines rapidly as it matures. It reproduces apomictically (the progeny 

can be exactly like the mother plant), suggesting that, over time, populations may 

become genetically distinct and be adapted only to specific environments. 

SERDP Sandberg Bluegrass. This germplasm originated from 28 ecotypes

from 26 locations at the Yakima Training Center. The plants were transplanted to 

a field near Logan, Utah. Seed was harvested from those plants and used to 

establish a seed-increase block near Logan. Because we combined the germplasm 

from the different ecotypes, this broad-based mix of apomictic genotypes should 

be better adapted to a wider range of environments. Molecular DNA tests (Larson et al. 2001; see the section on “Molecular Marker Development: Genetic 

Variation” later in this report) confirmed that a similar broad-based collection of 

Sandberg bluegrass has a large amount of genetic diversity but that it is still 30% 

different from Sandberg bluegrass germplasm originating near Boise, Idaho. 

Seed was being increased in 2002 and 2003, and a Selected Class Germplasm 

release is expected in 2004. 

Western Yarrow (Achillea millefolium; Native) 

Introduction. In addition to the grasses, we also have western yarrow in the 

improved breeding program. It is a drought-tolerant, native forb that provides a 

good complement to grass stands. In areas of natural plant disturbance, western 

yarrow establishes readily and increases over time. Western yarrow is one of the 

most widely distributed herbaceous species in the western United States, and it 

grows in a wide variety of habitats ranging from aspens and open forests to sagebrush zones. It avoids dense shade, is comparatively drought resistant, and flourishes in sandy and gravelly loam soils. 
46 ERDC/CRREL TR-03-20 

Limitations. Sources of commercial seed are limited, although this may 

become less of a problem, as it is gaining in popularity as a revegetation species. 

SERDP Western Yarrow. This germplasm originated from 28 collections 

made within Yakima Training Center. It is a broad-based population with no 

selection pressure having been applied. Our early trials at the Yakima Training 

Center suggested that western yarrow collected there outperformed commercially 

available western yarrow (Table 16). After these tests, we collected western 

yarrow seed from 26 Yakima Training Center locations representing six 

ecological sites. Plants were started from the collected seed and established in a 

seed-increase field. A formal release as a Selected Class Germplasm is expected 

by 2004. 

Table 16. Results of trials comparing the percent stand of Yakima 

populations with the commercial variety of western yarrow. 

% stand of western yarrow 1995 1996 1997 1998 

Commercial variety 38 25 4 0 

Yakima ecotype 21 44 44 56 

Molecular Marker Development 

We leveraged SERDP funding with a research project funded under the BT25 Program “Genetic Characterization of Native Plants in Cold Regions” to 

identify genetic markers that relate to plant characteristics necessary for resiliency and to analyze the genetic diversity of the new germplasms. Using genetic 

markers can save time, since potential plants produced during the breeding process can be selected for desired genes rather than having to be grown over a twoyear period to identify and select the desired traits. 

In our earlier work we used the Random Amplified Polymorphic DNA 

(RAPD) technique to identify the markers (Huff et al. 1998). We found four 

markers for early spring growth in Hycrest crested wheatgrass (Hu et al. 2001) 

and a marker for fast seed germination in western wheatgrass (Liu et al. 1997). 

This latter species, which is a widely distributed, long-lived, sod-forming grass, 

has strong rhizomatous growth in sandy soil, an important characteristic for plant 

resiliency on military training lands (Asay and Knowles 1985). 

We used flow cytometry to differentiate and identify fine fescue (Festuca) 

species (Huff and Palazzo 1998). Fine fescues comprise a group of plants that are 

morphologically similar and have fine to very fine leaf texture. Identification and 
Improved Native Grasses and Establishment Methods 47 

classification is very difficult in such species, which creates problems for programs such as the LCTA monitoring system. We found that we can separate 

morphologically similar fine fescue species using flow cytometry, which essentially counts the number of chromosomes in plant tissue. The DNA content 

among the ten species we examined was highly positively correlated with 

observed or reported chromosome numbers. 

As part of this SERDP project we continued to identify markers for important characters, including growth at low temperatures and early spring growth. As 

the project progressed, we updated our analysis techniques from RAPD and flow 

cytometry to the Amplified Fragment Length Polymorphic (AFLP) procedure. 

AFLP is much more reproducible than is the RAPD technique. For the genetic 

variance studies, we also used analysis of molecular variance (AMOVA) to 

determine the significance in genetic differences between populations (Excoffier 

et al. 1992). 

Markers for Cold Tolerance and Early Spring Growth 

We used AFLP techniques to find relationships between genetic markers and 

early spring growth in wildrye (Leymus sp.) and crested wheatgrass—two coolseason grasses used widely on military lands in the west. Early spring growth is 

important because most of the soil erosion on military bases occurs in the early 

spring prior to a plant’s active growth stage. A plant usually initiates early spring 

growth by using the stored soluble carbohydrates (CHO) that were produced the 

previous fall and were carried over during the winter or dormant months. CHO 

compounds can also be used as cryoprotectants to protect plants against winterkill. Therefore, CHO compounds including anthocyanins are useful phenotypic 

markers for genetic and molecular studies of plant gene expression during cold 

stress and acclimation (Christie et al. 1994) and may facilitate the selection of 

genotypes with higher concentrations of soluble CHO (Chatterton et al. 1989, 

1990). We found genetic relationships between low-temperature soluble CHO 

accumulations, anthocyanin coloration, and plant growth and development traits 

in wildrye (Leymus sp.) (Larson et al. 1999) and crested wheatgrass (Hu et al. 

2000, 2001). 

Wildrye. This grass has a circumpolar distribution. A total of 870 mapped 

AFLP markers were used to detect quantitative trait loci (QTL) controlling soluble carbohydrate accumulations, anthocyanin coloration, and growth characteristics in a segregating population derived from open-pollinated Basin wildrye 

(Leymus cinereus) × beardless wildrye (L. triticoides) hybrids. Positive trait correlations and pleiotropic gene effects were detected for soluble carbohydrate 

accumulations and anthocyanin coloration. Findings suggest that genetic 
48 ERDC/CRREL TR-03-20 

manipulations in five QTL regions on linkage groups 1A, 1B, 2A, 4Xm, and 5Ns 

could improve fructan and anthocyanin synthesis (Larson et al. 1999). 

In further inheritance studies on an F3 population (132 replicated clones) of 

the same basin wildrye × beardless wildrye hybrids, we examined three traits: 

leaf growth, leaf carbohydrate abundance, and anthocyanin expression. Using 

AFLP markers, we found one DNA marker negatively associated with all three 

traits, one marker positively associated with growth and carbohydrate, one 

marker positively associated with carbohydrate and anthocyanin, and at least four 

unique DNA marker groups for each trait. The results suggest that leaf growth 

and carbohydrate content can be simultaneously improved, extending the growing season and improving the resilience of these plants. 

Crested Wheatgrass. For this plant we identified AFLP markers associated 

with anthocyanin coloration and upright vs. prostrate growth. Crested wheatgrass 

is well adapted to many semiarid steppe and prairie environments and was widely 

planted on military lands. The SERDP-select CD-II crested wheatgrass cultivar 

was selected from the cultivar Hycrest on the basis of vigorous vegetative growth 

and green leaf coloration during the early spring. Using AFLP markers we found 

that the two cultivars are distinguishable, although they remain closely related. 

CD-II plants have lower anthocyanin coloration (greater green leaf coloration), 

more upright growth, wider leaves, and earlier spring vegetative growth than 

Hycrest plants. It is possible to make further gains in the CD-II line for early 

spring growth with the aid of marker-assisted selection (Hu et al. 2000, 2001). 

Genetic Variation 

We studied the genetic variation of several grasses to help classify and confirm the improved characteristics or increased diversity of the SERDP-select 

germplasms. At the same time we looked at the diversity of commercially available plants to answer questions on the genetic diversity of purchased seeds and to 

help land managers make seeding decisions. A greater diversity enables a species 

to be more resilient or more adaptable to various environmental stresses, and our 

intent, especially in the selected-class germplasms, was to increase the genetic 

diversity in the improved germplasms. Some ecologists, however, may be concerned that the agronomic approach of cultivar development to select or breed 

plants with specific characteristics creates plant materials that may lack the 

genetic diversity to maintain their adaptation in dynamic environments. They 

believe that non-local seed sources may not have adapted genomes, or that nonlocal sources may result in genetic pollution. To put these concerns to rest, we 

examined the genetic diversity of natural populations and known cultivars of 

bluebunch wheatgrass, bluegrass, little bluestem, and western wheatgrass. 
Improved Native Grasses and Establishment Methods 49 

Bluebunch Wheatgrass. This cool-season, perennial grass is native to semiarid regions of western North America. During our review of the invasiveness of 

the introduced species under development, we found that bluebunch wheatgrass 

was successfully moving into seeded stands of introduced species (Palazzo et al. 

1999; see Table B1 in Appendix B). One of the two new bluebunch wheatgrass 

germplasms we developed, P-7, is the result of multiple collections and is 

intended to provide genetic diversity within a single germplasm for semiarid to 

mesic sites where bluebunch wheatgrass was an original component of the vegetation. P-7 is a multiple-origin polycross generated by intermating 23 openpollinated, native-site collections from Washington, Oregon, Nevada, Utah, 

Idaho, Montana, and BritishColumbia, along with the only cultivars currently 

available for purchase, Whitmar and Goldar. Although they are morphologically 

distinct—Goldar has awns and Whitmar does not—the commercially available 

cultivars originate from collections of populations less than 83 km apart in southeastern Washington. 

We used the AFLP method to quantify and compare genetic variation within 

and between the Goldar and Whitmar cultivars and the SERDP P-7 multipleorigin polycross. More AFLP alleles (99) were found to be unique to P-7—

present in P-7 but absent in Goldar and Whitmar—than were found to be unique 

to Whitmar(59) or Goldar (49). P-7 also had fewerfixed loci (233) than Whitmar 

(385) or Goldar (318). Overall within-population variation as shown by 

nucleotide-sequence diversity [π ± SE(1000)] was greater for P-7 (38.7 ± 1.6) 

than for Whitmar (34.2 ± 1.5) or Goldar (33.9 ± 1.5). Between-population variation as shown by the average net nucleotide-sequence divergence (dA) was 0.3 ± 

0.2 between P-7 and Goldar, 1.3 ± 0.2 between P-7 and Whitmar, and 2.6 ± 0.3 

between Goldar and Whitmar. Therefore, P-7 is genetically intermediate between 

the two cultivars but more similar to Goldar than to Whitmar. Figure 4 illustrates 

these relationships among the P-7, Goldar, and Whitmar populations. We also 

found that the genetic diversity of the two named cultivars, collected more than 

50 years ago, is stable, indicating that it is likely that our multi-origin cross will 

maintain its greater genetic diversity (Larson et al. 2000). 

Bluegrasses. Bluegrasses are dominant perennial grasses in the understory 

steppe vegetation of western North America and are important for creating biological and genetic diversity and plant resiliency on military training lands. We 

used the AFLP method to measure nucleotide sequence variation within and 

between four commonly recognized forms of Poa secunda grasses: two cultivars, 

‘Sherman’ big bluegrass and ‘Canbar’ canby bluegrass, and two broad-based 

Sandberg bluegrass sources, one collected from 19 sites at Yakima Training 

Center (YTC) and one collected from Mountain Home (MH) near Boise, Idaho 

(Larson et al. 2001). The Canbar and Sherman plants were grown from
50 ERDC/CRREL TR-03-20 

Multiple-origin polycross 

Goldar

Whitmar

Figure 4. Genetic variation in bluebunch 

wheatgrass cultivars and a multiple-origin 

polycross (Larson et al. 2000). 

seed obtained from the USDA-NRCS Plant Materials Center in Pullman, 

Washington. Relative DNA contents were compared between individual plants 

within and between populations, DNA fingerprinting was conducted using the 

AFLP technique, and the degree of polymorphism within populations was 

quantified using the Shannon–Weaver diversity index (Larson et al. 2001). 

Relationships between individual genotypes were analyzed by cluster analysis 

using the unweighted pair-group (UPGMA) procedure in NTSYS-PC (Rohlf 

1992). 

Of the 22 plants analyzed from each cultivar, one fixed genotype was 

detected in the Sherman big bluegrass and only three genotypes were detected in 

Canbar canby bluegrass. Although several identical genotypes were also detected 

with the YTC and MH Sandberg bluegrass populations, genetic diversity within 

these natural Sandberg populations is much greater than within Sherman or 

Canbar, as represented in Figures 5 and 6. These results show a high degree of 

variation within, but little divergence between, the two natural Sandberg bluegrass populations collected from sites nearly 600 km apart. If these populations 

have similar adaptation potential, then efforts to develop genetically diverse, 

natural germplasm sources for Sandberg bluegrass might be streamlined (Larson 

et al. 2001). 

Western Wheatgrass. This grass is an ecologically dominant, native, perennial range grass in the northern Great Plains (Hart et al. 1996) that extends 

throughout much of temperate North America (Hitchcock 1951). We used AFLP 
Improved Native Grasses and Establishment Methods 51 

Figure 5. UPGMA analysis of AFLP variation within and among Sandberg 

bluegrass populations collected near Mountain Home (MH), Idaho, and 

Yakima Training Center (YTC), and the cultivars Canbar canby bluegrass 

and Sherman big bluegrass. The numbers in parentheses indicate identical 

genotypes. (From Larson et al. 2001.) 
52 ERDC/CRREL TR-03-20 

YTC MH 

Canbar 

Sherman 

Figure 6. Venn diagram representation of genetic 

variation in Sandberg bluegrass populations collected near Mountain Home (MH), Idaho, and Yakima 

Training Center (YTC), and the cultivars Canbar 

canby bluegrass and Sherman big bluegrass. 

techniques to look at the genotypic identification of this species. The populations 

analyzed were either source-identified single-origin accessions obtained from the 

National Plant Germplasm System or multiple-origin germplasms. We tested the 

similarity among individual accessions, which had come from different geographic locations, and we compared rates of DNA variation between singleorigin accessions and the multiple-origin germplasms. Individual plants from 36 

of 39 source-identified single-origin germplasm were different from each other; 

in other words, there was little genetic similarity among the single-origin populations. Three of the four multiple-origin accessions did not cluster by accession 

and displayed higher rates of DNA variation than the single-origin accessions; 

these populations show genetic variation greater than that found in any one geographic area (Larson et al. 2003). These results suggest that it might not be wise 

to use a single-origin source from outside the area being revegetated. The 

multiple-origin accessions, however, with their wider genetic variations, would 

be better able to adapt and survive when used in a broader range of areas. 

Little Bluestem. We also studied the genetic variability of plants important 

in the eastern United States, especially those relevant to Fort Drum in their 

LRAM Program. Little bluestem is a native, warm-season, perennial bunchgrass 
Improved Native Grasses and Establishment Methods 53 

that ranges throughout much of central and eastern North America, from Canada 

to Mexico. As a native species, little bluestem is increasingly being used for restoration projects to enhance the biodiversity of locally disturbed sites. We used 

RAPD markers to estimate the comparative genetic variation within and among 

four native populations of little bluestem. Genotypes were collected from highand low-fertility sites in both New Jersey (forest biome) and Oklahoma (grassland biome) and propagated in the greenhouse. We used AMOVA to estimate 

variance components for the RAPD phenotypes, partitioning the total variation 

among individuals within populations, between populations within a biome, and 

between biomes. Even though most genetic variation resided within populations, 

significant differences were detected between sites within each biome. Furthermore, genetic distances between high- and low-fertility sites within biomes were 

equal to or greater than biome distances. Therefore, in this wide-ranging and 

highly variable species, analysis suggests that extreme local site differences in 

fertility and ecological history promote genetic differentiation equal to or greater 

than geographic differentiation (Huff et al. 1998). 
54 ERDC/CRREL TR-03-20 

5 SOIL COMPACTION AND ROOT STUDIES 

ACCOMPLISHMENTS 

During maneuvers on training lands, military vehicles damage vegetation and 

compact soil. Vegetation damage includes shearing of leaves and damage to plant 

crowns and roots. While all three are related to plant regrowth and resiliency, 

crown and root damage are directly related to plant persistence and adaptation. 

Soil compaction changes the soil surface and reduces infiltration, which increases 

the volume and period of surface water runoff and increases soil erodibility. To 

understand the effects of training on soil compaction, plant injury, and regrowth, 

we measured soil compaction on several training lands subjected to various 

training events, and we examined plants in a controlled greenhouse environment 

and after simulated training exercises in the field. We also examined the resiliency after tracking of several of the species we are working with. 

Measuring Soil Compaction: Preliminary Soil Penetrometer Studies 

To select realistic soil compaction rates to use in our greenhouse study, we 

surveyed various areas on three military facilities. We used two types of cone 

penetrometers to measure the degree of soil compaction that occurs after land use 

at Yakima Training Center and Forts Carson and Drum. For most of the measurements, we used a Rimik CP20 cone penetrometer (Fig. 7), which measures 

compaction as a unit of resistance, MPa; the higher the MPa value, the higher the 

compaction of the soil. For some measurements at YTC, we used a different 

Figure 7. Using the Rimik CP20 cone 

penetrometer in the field. 
Improved Native Grasses and Establishment Methods 55 

Figure 8. Soil compaction measurements at 

Fort Drum. Higher resistance values indicate 

greater compaction. 

penetrometer that measures compaction as g cm–3; again, the higher the value, the 

greater the compaction. Both of these instruments can measure compaction continuously below the surface of the soil. The areas at each location were selected 

for various degrees of training intensity and included high- and low-use areas and 

bivouac sites. At each facility, we sampled two predominant soil types: sandy or 

silty texture. 

At Fort Drum we measured a grassy training area and lightly and heavily 

used travel corridors (Fig. 8). We found that land use was directly related to soil 

compaction at the soil surface to a depth of 5 cm (Cary et al. 2001, Palazzo et al. 

2001). The more the land was traveled on, the greater the compaction of soil near 

the surface. 

Our data on untracked sandy soils in the grassy areas at Fort Drum showed 

that as soil depth increases, the soil becomes more compacted and therefore has a 

greater water-holding capacity (Fig. 8). This is the opposite of what is normally 

described in the literature (Halvorson et al. 2001), which says that soil compaction decreases with depth. It is very hard to establish plants on sandy soils, such 

as those at Fort Drum with an average of 92% sand content, because of the low 

moisture-holding capacity of sandy soils. Plants sown on sandy soils begin to 

grow and then dry out and die, and the area then has to be reseeded. Our data 

suggest that we could establish plants on the Fort Drum sandy soils if we could 

find plants that root quickly and can extend their roots down to the moister areas 

in the soil at 10–15 cm deep (Fig. 9). We examined potential grasses in the 

greenhouse (see next section) and then developed an appropriate seed mixture for 
56 ERDC/CRREL TR-03-20 

0 1.0 2.0 3.0 4.0

Resistance (MPa)

0

-5

-10

-15

-20

-25

-30

-35

Depth (cm)

Soil moisture front

More compacted

Less compacted

0 1.0 2.0 3.0 4.0

Resistance (MPa)

0

-5

-10

-15

-20

-25

-30

-35

Depth (cm)

Soil moisture front

More compacted

Less compacted

Figure 9. Effect of the compaction–depth relationship on soil moisture and 

plant growth. As the soil depth increases at Fort Drum, the soil becomes 

more compacted and therefore has a greater water-holding capacity. To 

establish plants on the sandy soils at Fort Drum, we need to use plants that 

root quickly and can extend their roots down to the moister areas in the 

soil. 

use at Fort Drum (see “Methods for Establishing Natives and Fighting Invasive 

Weeds” section of this report). 

At Fort Carson we measured soil compaction under various military land 

uses. The Fort Carson results agreed with those found at Fort Drum, where land 

use increased soil compaction on fine-textured soils (Fig. 10). 

At YTC we looked at the effect of land use and soil moisture content on soil 

compaction and the effect of freeze–thaw cycling on soil compaction alleviation. 

Soil compaction increased in the more heavily used areas, especially in moist 

soils. The greater the soil moisture, the greater the increase in soil compaction 

when military vehicles are driven across the soils (Fig. 11). At YTC we also 

looked at soil compaction and the resiliency of plant species after one to four 

tank passes (see the “Tracking Study” section below). 
Improved Native Grasses and Establishment Methods 57 

Figure 10. Soil compaction measurements at 

Fort Carson. Higher resistance values indicate greater compaction. 

Figure 11. Soil compaction measurements on dry and moist soils at Yakima 

Training Center. Higher bulk density values indicate greater compaction. 
58 ERDC/CRREL TR-03-20 

We prepared a literature review on soil freeze–thaw cycling and its effect on 

soil compaction published in a book chapter by the Army Research Office (Gatto 

et al. 2001). The manuscript described the importance of incorporating the effects 

of freeze–thaw on soil compaction and strength, runoff, and erosion into soil erosion models. It also reports that soil freeze–thaw cycling reduces soil compaction 

and soil strength, increases infiltration, and changes the geometry of vehicle ruts 

and natural rills.

Greenhouse Root Studies 

In 1998 we began a series of greenhouse studies. The first greenhouse studies 

evaluated how compaction affects the root growth of species in soils originating 

from three military bases. In two other greenhouse studies, we evaluated the differences between SERDP-select germplasms and existing cultivars. The compaction studies and the first germplasm evaluation, which examined root and 

stem growth, were carried out in rhizotron tubes. The second germplasm evaluation examined early root growth in growth pouches. The SERDP-select germplasms we used in these studies were in the early development stage. 

Soil Compaction Study 

For this study we compacted soils in rhizotrons where the amount of root 

growth of the grasses can be measured and observed. The rhizotrons are clear 

acrylic tubes (4.8-cm i.d., 60-cm long, 6-mm wall thickness) closed at the base 

with an aluminum cap containing holes for drainage and tightened with a steel 

clamp. The tubes were placed at a 15 ̊ angle in an enclosed wooden box (Brar et 

al. 1990). The sides, bottom, and top of the wooden box were covered with 2.5-

cm-thick polystyrene sheets to decrease temperature fluctuations and to prevent 

exposure of roots to direct sunlight in the greenhouse. 

We used four soils from two facilities (YTC and Fort Drum) with three compaction rates for each soil. We used a sandy soil and a silty soil from each of the 

facilities; the compaction rates (high, medium, and low) were based on the measurements made in the field (Table 17). The grasses used were hard fescue (introduced) and switchgrass (native). 

Table 17. Soil compaction rates selected for this 

study. These rates were based on field measurements.

 Compaction rates (MPa) 

Installation High Medium Low 

Fort Drum 4.0 2.0 1.0 

Yakima Training Center 4.0 3.0 1.5 
Improved Native Grasses and Establishment Methods 59 

The soils were steam-sterilized for 24 hours to kill any viable seed before 

they were placed in the tubes. The soil compaction levels were attained by filling 

each tube incrementally with equal volumes of soil and compressing the soil with 

a wooden dowel having a diameter that matched the inside of the tube. The level 

of compaction of the soil in the tube was measured with the Rimik CP20 cone 

penetrometer, and the compression was continued until the desired level of compaction was obtained. 

The grasses were sown, two plants per tube, on 10 December 1999 and harvested on 15 February 2000, 67 days after planting. We cut the plants at the root 

and shoot interface and washed them with distilled water, using a paper towel to 

remove excess surface water from the shoots. The roots were washed free of soil 

over a screen by both spraying and immersion in water as described by Brar and 

Palazzo (1995a). The total root length was measured with an AgVision root and 

leaf imaging system (Decagon Devices, Inc., Pullman, Washington). Analyses of 

variance of the data were conducted using SAS version 8.0 (SAS Institute, Cary, 

North Carolina), and the means were separated using least significant differences 

at the 0.05 level of probability. 

We anticipated that an increase in soil compaction would restrict root growth 

in each of the soil types. However, we did not have any consistent root growth 

patterns for any of the soil types. The only consistent difference was that the Fort 

Drum soils produced the lowest amounts of leaf and root growth in terms of 

length at all compaction levels (Table 18). No consistent differences were found 

between the soil types from the various locations; we think the lack of correlation 

between root growth and soil compaction is due to the stronger influence of 

optimal plant growth conditions in the greenhouse. 

Table 18. Stem and root length measurements of hard fescue grown 

in rhizotrons containing either a sandy or silty soil originating from 

Fort Drum, New York, and Yakima Training Center, Washington. 

 Fort Drum soils YTC soils 

Soil type 

Compaction 

level 

Stem length 

(cm) 

Root length 

(cm) 

Stem length 

(cm) 

Root length 

(cm) 

Sand 

 Low 86 700 953 2013 

 Medium 119 846 1279 2700 

 High 94 456 1501 3002 

Silt 

 Low 456 2030 1959 4479 

 Medium 478 1941 3269 6563 

 High 911 3293 3332 6162 
60 ERDC/CRREL TR-03-20 

Shoot and Root Growth Study 

To evaluate shoot and root growth of grasses adapted to arid, western areas 

of the United States, we again used slant-tube rhizotrons. Each tube was filled 

incrementally with a Plainfield sandy soil (mixed, mesic Typic Udipsamments). 

The soil was settled by applying 200 mL of distilled water and tapping the tube. 

We used three native SERDP-select germplasms available as of 1998 along 

with a standard cultivar for each. The grasses studied were: ‘Goldar’ and Yakima 

bluebunch wheatgrass, ‘Magnar’ and Yakima basin wildrye, and ‘Secar’ and 

Yakima Snake River wheatgrass. The seeds were primed in distilled water for four 

days. Five seeds of each grass type were planted about 10 mm deep in each tube. 

The SERDP-select germplasms responded similarly to or slightly better than 

the commercially available cultivars (Table 19). An added benefit of the SERDPselect germplasms is that they have a wider genetic base without any loss of 

growth rate (see section on “Molecular Marker Development”). 

Table 19. Length, surface area, and diameter of roots and length and surface area of leaves of six native plants grown in rhizotron tubes containing a Plainfield sandy soil. 

 Root data Leaf data 

Species and variety 

Length 

(cm) 

Surface 

area 

(cm2

) 

Average 

diameter 

(mm) 

Length 

(cm) 

Surface 

area 

(cm2) 

Snake River wheatgrass 

SERDP-select (Yakima) 278.4 20.7 0.2352 79.3 17.5 

Secar 271.3 19.9 0.2335 52.6 10.8 

Bluebunch wheatgrass 

SERDP-select (Yakima) 119.3 18.1 0.4779 119.3 18.1 

Goldar 120.8 23.9 0.6800 120.8 23.9 

Basin wildrye 

SERDP-select (Yakima) 471.6 44.3 0.2989 139.3 37.0 

Magnar 268.9 24.6 0.2905 7.28 20.8 

Early Root Growth Study 

We compared early root growth of five SERDP-select germplasms available 

as of 1998 to currently available cultivars using a growth-pouch technique (Brar 

et al. 1990). The grasses studied were: ‘Goldar’ and SERDP-select Yakima bluebunch wheatgrass, ‘Vavilov’ and SERDP-select P-27 and Yakima Siberian 

wheatgrass, ‘Magnar’ and SERDP-select Yakima basin wildrye, ‘Secar’ and 
Improved Native Grasses and Establishment Methods 61 

SERDP-select Yakima Snake River wheatgrass, and ‘Bottlebrush’ and SERDPselect Yakima squirreltail. We also studied cheatgrass to compare this invasive 

species with the SERDP-select species under development. The SERDP-select 

grasses studied were still in the early stages of the selection process. 

The seeds were primed in distilled water for four days. Ten seeds of each 

entry were selected randomly and placed in the growth pouches. The growth 

pouches were placed in the dark in environmental chambers at 20oC for 16 days. 

Root lengths were measured on days 5, 8, 11, and 16 after seeding with an 

AgVision root and leaf imaging system (Decagon Devices, Inc., Pullman, 

Washington). Analyses of variance (ANOVA) of the data were conducted using 

SAS version 8.0 (SAS Institute, Cary, North Carolina), and the means were separated using least significant differences at the 0.05 level of probability. 

For the introduced species the SERDP Siberian wheatgrass entries had 

slower root elongation than did the known cultivar (Fig. 12a). For the native species under development, SERDP basin wildrye had faster root elongation than its 

known cultivar (Fig. 12b), but the SERDP Snake River wheatgrass, bluebunch 

wheatgrass, and bottlebrush squirreltail entries did not (Fig. 12c–e). The two 

native wheatgrasses were similar to the commercially available cultivars. Bottlebrush squirreltail was dropped from the breeding program shortly after this study 

as it did not perform well here or in field evaluations. 

Tracking Study 

In May 1999 we conducted tank traffic studies to determine plant resiliency 

in our evaluation plots at Yakima Training Center. We worked with Colorado 

State University and ERDC-CERL to determine the resiliency of the species we 

are breeding using the Tracked Vehicle Impact Miles (TVIM) approach. 

For the tracking we used the evaluation plots at the Yakima Snake River 

Sites A and B (Table 6) four years after the plots were seeded. Because of poor 

stands, only 21 of the 30 entries were included in the tracking study. On 19 May 

1999 the plots were subjected to traffic using an M1A1 tank to make zero, one, 

two, and four passes that were perpendicular to the species plots (Fig. 13). The 

control was designated as the 1-m-wide area adjacent to the track. The tracking 

techniques we used at YTC were developed in TVIM studies conducted by 

ERDC-CERL and the Center for the Ecological Management of Military Lands 

(CEMML). All of the tank passes in our study occurred on the same day, which 

was intended to represent different training intensities for a single training event. 

We focused on a single event since one of the goals of the Army Strategic Planning ATTACC model (Anderson et al. 2001) is to place a monetary value on the 

degradation occurring after a training activity. 
62 ERDC/CRREL TR-03-20 

e. Bottlebrush Squirreltail (native)

Cheatgrass

Bottlebrush Squirreltail

SERDP

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

Days after seeding

5 11 16 8

b. Basin wildrye (native)

0

50

100

150

200

250

300

350

400

450

500

Days after seeding

Root Length (cm)

‘Magnar’

SERDP

Cheatgrass

5 11 16 8

a. Siberian wheatgrass (introduced)

Days after seeding

0

100

200

300

400

500

600

700

‘Vavilov’

Cheatgrass

SERDP

P-27

Root Length (cm)

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

c. Snake River wheatgrass (native)

5 11 16 8

‘Secar’

Cheatgrass

SERDP

Days after seeding

0

100

200

300

400

500

600

‘Goldar’

Cheatgrass

d. Bluebunch wheatgrass (native)

SERDP

5 11 16 8

Days after seeding

Root Length (cm)

e. Bottlebrush Squirreltail (native)

Cheatgrass

Bottlebrush Squirreltail

SERDP

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

Days after seeding

5 11 16 8

b. Basin wildrye (native)

0

50

100

150

200

250

300

350

400

450

500

Days after seeding

Root Length (cm)

‘Magnar’

SERDP

Cheatgrass

5 11 16 8

a. Siberian wheatgrass (introduced)

Days after seeding

0

100

200

300

400

500

600

700

‘Vavilov’

Cheatgrass

SERDP

P-27

Root Length (cm)

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

c. Snake River wheatgrass (native)

5 11 16 8

‘Secar’

Cheatgrass

SERDP

Days after seeding

0

100

200

300

400

500

600

‘Goldar’

Cheatgrass

d. Bluebunch wheatgrass (native)

SERDP

5 11 16 8

Days after seeding

Root Length (cm)

Cheatgrass

Bottlebrush Squirreltail

SERDP

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

Days after seeding

5 11 16 8

Cheatgrass

Bottlebrush Squirreltail

SERDP

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

Days after seeding

5 11 16 8 0

50

100

150

200

250

300

350

400

450

500

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

Days after seeding

5 11 16 8

b. Basin wildrye (native)

0

50

100

150

200

250

300

350

400

450

500

Days after seeding

Root Length (cm)

‘Magnar’

SERDP

Cheatgrass

5 11 16 8

b. Basin wildrye (native)

0

50

100

150

200

250

300

350

400

450

500

0

50

100

150

200

250

300

350

400

450

500

Days after seeding

Root Length (cm)

‘Magnar’

SERDP

Cheatgrass

‘Magnar’

SERDP

Cheatgrass

5 11 16 8

a. Siberian wheatgrass (introduced)

Days after seeding

0

100

200

300

400

500

600

700

‘Vavilov’

Cheatgrass

SERDP

P-27

Root Length (cm)

a. Siberian wheatgrass (introduced)

Days after seeding

0

100

200

300

400

500

600

700

‘Vavilov’

Cheatgrass

SERDP

P-27

Root Length (cm)

0

100

200

300

400

500

600

700

0

100

200

300

400

500

600

700

‘Vavilov’

Cheatgrass

SERDP

P-27

Root Length (cm)

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

c. Snake River wheatgrass (native)

5 11 16 8

‘Secar’

Cheatgrass

SERDP

Days after seeding

0

50

100

150

200

250

300

350

400

450

500

0

50

100

150

200

250

300

350

400

450

500

Root Length (cm)

c. Snake River wheatgrass (native)

5 11 16 8

‘Secar’

Cheatgrass

SERDP

Days after seeding

0

100

200

300

400

500

600

‘Goldar’

Cheatgrass

d. Bluebunch wheatgrass (native)

SERDP

5 11 16 8

Days after seeding

Root Length (cm)

0

100

200

300

400

500

600

0

100

200

300

400

500

600

‘Goldar’

Cheatgrass

d. Bluebunch wheatgrass (native)

SERDP

5 11 16 8

Days after seeding

Root Length (cm)

Figure 12. Elongation of roots after

germination in growth pouches for five

species compared to the invasive weed

cheatgrass. For each species, we tested

seeds from the early development of the

SERDP-select germplasm (as of 1998)

along with seeds of a known cultivar. 
Improved Native Grasses and Establishment Methods 63 

1 pass

2 passes

4 passes

Figure 13. Layout of the tracking study. The tracking passes by the M1A1 

tank were perpendicular to the direction of the seeded plots. 

Compaction data were recorded with a cone resistance penetrometer before 

tracking, immediately after tracking, and five months later in October 1999. The 

cone resistance penetrometer measures compaction at the surface in centimeters 

of deflection. Lower readings indicate a greater degree of resistance to penetration and, therefore, more compaction. 

Plant resiliency data were collected on September 2001, two years after the 

tracking. The degree of resiliency was measured by comparing the differences 

between treatments. Variables measured along a 3-m transect within the plot 

include the length of gaps that lacked the target species (species seeded), the 

percent target species, the forage yield, the percent encroachment of Sandberg 

bluegrass, the percent bare ground, and cheatgrass invasion. Forage yield was 

determined in September 2001 by harvesting 1 m2

 from the control and the 

tracked area. The percent dry matter was determined from plot samples, and 

forage yields expressed as grams per plot. All other variables were visually 

estimated. 

Variable means across tracks were analyzed as a randomized complete block 

design. F-tests of main effects and interactions were made, considering all main 

effects as fixed. All data were subjected to analyses of variance using GLM procedures (SAS Institute Inc. 1994), and mean separations were made on the basis 

of least significant differences at the 0.05 probability level (SAS Institute Inc. 

1994). 

Compaction Results 

The data recorded immediately after tracking show that the two- and fourpass treatments resulted in the greatest soil compaction near the soil surface 

(Table 20, Fig. 14). Five months after tracking, the compaction measurements 

were similar to those taken immediately after tracking in two of the three tracking treatments in each plot. Compaction was reduced over the summer months in 
64 ERDC/CRREL TR-03-20 

0

1

2

3

4

5

6

7

0124

Number of Tank Passes

Cone 

Deflection 

(cm)

Before tracking

Immediately after tracking

5 months after tracking

0

1

2

3

4

5

6

7

0124

Number of Tank Passes

Cone 

Deflection 

(cm)

Before tracking

Immediately after tracking

5 months after tracking

0

1

2

3

4

5

6

7

0124

Number of Tank Passes

Cone 

Deflection 

(cm)

Before tracking

Immediately after tracking

5 months after tracking

a. Snake A plots. 

0

1

2

3

4

5

6

0124

Number of Tank Passes

Cone 

Deflection 

(cm)

Before tracking

Immediately after tracking

5 months after tracking

0

1

2

3

4

5

6

0124

Number of Tank Passes

Cone 

Deflection 

(cm)

Before tracking

Immediately after tracking

5 months after tracking

0

1

2

3

4

5

6

0124

Number of Tank Passes

Cone 

Deflection 

(cm)

Before tracking

Immediately after tracking

5 months after tracking

b. Snake B plots. 

Figure 14. Soil compaction measured before and immediately after 

zero, one, two, and four passes of an M1A1 tank on the Yakima 

Training Center Snake A and B plots on 19 May 1999. An additional 

measurement was made in October 1999. Higher cone deflections 

indicate less compaction. 
Improved Native Grasses and Establishment Methods 65 

Table 20. Cone resistance measurements on Yakima Training Center 

plots subjected to zero, one, two, or four passes of an M1A1 tank. 

 Deflection (cm) 

0 passes 

(control) 1 pass 2 passes 4 passes 

Snake A 

Before tracking 5.64 5.61 5.97 5.87 

Immediately after tracking 5.64 4.36 4.52 3.83 

Five months later 5.82 5.37 4.52 4.07 

Snake B 

Before tracking 4.86 5.02 4.84 5.08 

Immediately after tracking 4.86 4.85 3.84 4.01 

Five months later 5.04 4.57 4.73 4.08 

the one-pass treatment in Snake A and the two-pass treatment in Snake B. Alan 

Anderson, ERDC-CERL, used this compaction data in his SERDP Project 

“Improved Units of Measure For Training and Testing Area Carrying Capacity 

Estimation (CS1102)” to add parameters to equations for developing the 

ATTACC model on management of military lands (Anderson et al. 2001). 

Plant Resiliency Results 

Gaps Between Sown Species. The gaps in the rows of plants were measured 

to obtain information on plant density. The greater the number of gaps, the lower 

the density of plants present. 

At the Snake A site, with the exception of Vavilov and Kazakhstan Siberian 

wheatgrass and Rosana western wheatgrass, the number of track passes had little 

effect on the length of gaps between the target species within the plot (Table 21). 

However, even though not significant in most entries, there was a general trend 

towards reduced stands as the number of track passes increased. The rhizomatous, native Rosanna western wheatgrass expressed moderate tolerance to 

repeated tracking, showing a significant increase in gap length only at the onepass treatment. The wildryes had the greatest gaps in all tracking treatments 

compared to the other grasses. This was in part due to their poorer stand establishment and their inability to tolerate any level of tracking. 

Of the native species studied at Snake B, only Yakima bluebunch wheatgrass 

and E-27 hybrid were affected by the different number of track passes (Table 21). 

In addition to establishing better, Snake River wheatgrasses, especially Secar and 

EVT-572, persisted better under repeated tracking than bluebunch and thickspike 

wheatgrasses. The bluebunch wheatgrass cultivar Whitmar was the most persis- 
66 ERDC/CRREL TR-03-20 

Table 21. Mean gap between any sown plants in seeded rows of vegetation on adjacent sites (Snake A and B) at Yakima Training Center. 

 Length (cm) 

Entry Control 1 Pass 2 Passes 4 Passes 

LSD 

(0.05) 

Snake A: Introduced 

Crested wheatgrass 

Hycrest 4.6 6.5 5.8 6.5 ns 

Ephraim 3.8 6.0 4.3 5.3 ns 

Mean 4.2 6.3 5.1 5.9 

Siberian wheatgrass 

Vavilov 2.6 2.5 4.8 4.8 2.1 

P-27 2.8 3.8 3.8 6.3 ns 

Kazakhstan 3.3 3.0 6.0 8.3 4.2 

Mean 2.9 3.1 4.9 6.5 

Russian wildrye 

Bozoisky 9.2 11.3 8.8 10.3 ns 

Tetra-1 10.1 11.5 9.8 11.8 ns 

Mean 9.7 11.4 9.3 11.1 

Intermediate wheatgrass (cv. Luna) 5.6 7.8 7.0 7.0 ns 

LSD (0.05) Introduced 1.5 4.7 4.1 2.8 

Snake A: Native 

Basin wildrye (Yakima) 10.3 12.5 12.0 12.8 ns 

Canby bluegrass 3.8 4.8 3.3 3.0 ns 

Western wheatgrass (cv. Rosana) 4.1 8.7 5.5 4.5 3.6 

LSD (0.05) Native 1.7 5.5 4.8 1.1 

Snake B: Native 

Bluebunch wheatgrass 

Goldar 4.9 7.8 7.3 7.0 ns 

Whitmar 4.8 8.3 7.5 4.8 ns 

ACC-238 5.4 8.0 7.3 10.8 ns 

Yakima 6.0 10.8 6.8 8.5 4.0 

Mean 5.3 8.7 7.2 7.8 

Snake River wheatgrass 

Secar 2.8 7.5 4.5 4.3 ns 

ACC-707 3.2 6.8 5.3 6.5 ns 

EVT-572 2.5 4.5 3.8 4.5 ns 

Mean 2.8 6.3 4.5 5.1 

Thickspike wheatgrass and hybrids 

Bannock 4.3 7.8 6.0 7.0 ns 

E-20 (Snake River × thickspike wheatgrass) 3.8 6.0 6.3 6.0 ns 

E-27 (Snake River × thickspike wheatgrass) 3.5 5.0 3.5 6.8 1.6 

Mean 3.9 6.3 5.3 6.6 

LSD (0.05) Native entries 1.7 3.2 ns ns 

LSD (0.05) Native species 1.6 ns 1.4 1.3 
Improved Native Grasses and Establishment Methods 67 

tent under tracking of the bluebunch wheatgrasses. There were significant differences between entries, but differences were greater between the species than 

within. 

Percent target species. When we measured the percent target (sown) species 

in each plot, there was a general trend towards a decline in target species from 

the control to the four-pass treatment for all entries and species (Table 22). 

At the Snake A site, however, that trend was significant only for the Siberian 

wheatgrass entries, Ephraim crested wheatgrass, and intermediate wheatgrass 

(Table 22). There were no significant declines among the three native entries, and 

both Canby bluegrass and Rosana western wheatgrass were among the most 

persistent at the four-pass treatment. In fact, Rosana western wheatgrass was 

equal to or significantly better than all the entries for all treatment passes. This 

persistence of western wheatgrass is likely the result of aggressive rhizome 

development, particularly after disturbance, in western wheatgrass. Among the 

introduced entries there were no significant changes in composition across all 

treatments for Hycrest crested wheatgrass and the two Russian wildryes (Table 

22). At each tracking treatment there were significant differences between the 

entries and species. As previously reported, the wildryes were the poorest to 

establish and subsequently had the lowest percent target entries in the study. 

Although western wheatgrass and canby bluegrass fared well at the Snake A 

site, the native grasses at Snake B were generally less persistent under repeated 

tracking. The entries least affected by multiple tracking were Goldar and ACC238 bluebunch wheatgrass and E-20 hybrid wheatgrass (Table 22). Within each 

treatment, entries differed significantly (P<0.05) in the percent target entries 

within each plot, but differences among species were significant only at the fourpass treatment. At the four-pass treatment only, the percent target species for the 

Snake River wheatgrasses was significantly better than bluebunch wheatgrasses 

(Table 22). 

There appears to be an association between the original stand establishment 

and the degree to which plants recover after tracking. In nearly all cases, those 

with increased stands at the time of tracking had recovery or persistence rates 

higher than those with reduced stands. The exception was western wheatgrass, 

which has the ability through rhizome development to colonize disturbed areas 

without the use of seeds (Jensen et al. 2001). The data suggest that a plant’s ability to germinate quickly with strong seedlings and become established is perhaps 

the most critical characteristic in selecting plants for resiliency after tracking. 
68 ERDC/CRREL TR-03-20 

Table 22. Percent target species in seeded rows of vegetation on adjacent 

sites (Snake A and B) at Yakima Training Center.

 % sown species 

Entry Control 1 Pass 2 Passes 4 Passes 

LSD 

(0.05) 

Snake A: Introduced 

Crested wheatgrass 

Hycrest 17.5 17.5 18.0 13.5 ns 

Ephraim 19.5 18.0 16.3 11.5 6.7 

Mean 18.5 17.8 17.1 12.5 

Siberian wheatgrass 

Vavilov 22.8 23.8 17.8 15.0 4.9 

P-27 21.6 20.8 18.5 12.5 5.6 

Kazakhstan 19.5 20.0 13.0 9.3 5.7 

Mean 21.3 21.5 16.4 12.3 

Russian wildrye 

Bozoisky 11.8 9.8 12.3 7.3 ns 

Tetra-1 9.7 9.5 9.3 4.3 ns 

Mean 10.7 9.6 10.8 5.8 

Intermediate wheatgrass (cv. Luna) 17.2 12.5 14.0 12.3 2.3 

Snake A: Native 

Basin wildrye (Yakima) 12.5 8.5 7.5 4.8 ns 

Canby bluegrass 16.0 20.5 15.0 11.5 ns 

Western wheatgrass (cv. Rosana) 16.0 17.3 16.3 16.8 ns 

LSD (0.05) Entries 2.6 9.8 8.2 3.6 

LSD (0.05) Species 2.8 9.7 9.2 3.7 

Snake B: Native 

Bluebunch wheatgrass 

Goldar 15.2 9.3 10.8 7.5 ns 

Whitmar 18.0 8.5 10.3 7.3 6.9 

ACC-238 17.5 11.5 10.8 5.8 ns 

Yakima 15.9 6.5 11.3 9.0 5.5 

Mean 16.6 8.9 10.8 7.4 

Snake River wheatgrass 

Secar 22.4 9.8 15.8 14.3 5.2 

ACC-707 17.7 10.0 13.3 7.0 10.3 

EVT-572 21.4 12.0 18.3 9.5 8.1 

Mean 20.5 10.6 15.8 10.3 

Thickspike wheatgrass and hybrids 

Bannock 16.9 11.0 13.0 5.0 2.0 

E-20 (Snake River × thickspike wheatgrass) 17.2 10.0 12.0 10.3 ns 

E-27 (Snake River × thickspike wheatgrass) 20.8 14.8 17.3 8.8 8.1 

Mean 19.0 12.4 14.6 9.5 

LSD (0.05) Native entries 4.8 4.7 6.9 7.1 

LSD (0.05) Native species ns ns ns 2.2 
Improved Native Grasses and Establishment Methods 69 

Forage Yield. As an estimate of plant vigor after tracking, forage yield is 

important, but it should not be solely related to the ability of plants to reduce soil 

erosion. There was a general decline in forage yield of all entries and species 

with repeated tracking (Table 23). 

Although forage yield decreased with tracking for all species at Snake A, the 

effects were significant only for Siberian wheatgrasses, Ephraim crested wheatgrass, and Rosana western wheatgrass (Table 23). All three Siberian wheatgrasses had statistically significant declines in forage yield across tracking treatments, with the Kazakhstan cultivar having significantly poorer yield (77% 

reduction) than the other two Siberian wheatgrass cultivars (Table 23). The fourpass treatments of Vavilov Siberian wheatgrass and Rosana western wheatgrass 

(the native) had less than 40% difference in forage yield when compared to the 

control. Within each pass treatment and the control, there were significant differences between entries and species for forage yield (Table 23). The forage yield of 

Hycrest crested wheatgrass had no significant decline across treatments, and it 

was significantly better than or equal to the yield of all other entries. 

The effects of tracking were more severe on native entries at the Snake B site 

than on introduced entries at Snake A. The most significant declines in forage 

yield occurred at the one-pass treatment at this site (Table 23). Entries showing 

significant (greater than 60%) declines in forage yield from the control to the 

four-pass treatment were Goldar and ACC-238 bluebunch wheatgrass, EVT-572 

Snake River wheatgrass, and E-27 Snake River × thickspike wheatgrass crosses 

(Table 23). Regardless of the treatment, there were no differences between species means for bluebunch wheatgrasses, Snake River wheatgrasses, and thickspike wheatgrasses in forage yield (Table 23). Despite having higher forage 

yields on the control plots at the Snake B, the native grasses were more adversely 

affected by tracking pressure than the introduced grasses at Snake A. 

Sandberg Bluegrass Invasion. We measured Sandberg bluegrass invasion 

in each of the sown rows to see if a native bluegrass would be able to move into 

the plots. Sandberg bluegrass is a relatively low-producing perennial bunchgrass 

that grows as small tufts no more than 0.3 m (1 ft) in diameter. It is widely distributed throughout the western range, where it is considered an important range 

grass for soil stabilization and forage for wildlife and livestock. Once established, 

Sandberg bluegrass is perhaps the most drought tolerant of the bluegrasses. If 

seeded species will allow the invasion of Sandberg bluegrass into disturbed sites, 

it will increase desirable ground cover and reduce erosion. Encroachment of 

Sandberg bluegrass into existing plots was estimated as a percentage of the total 

ground cover within the plot. 
70 ERDC/CRREL TR-03-20 

Table 23. Forage yield as shown by mean dry weight (g) of sown species 

on adjacent sites (Snake A and B) at Yakima Training Center. 

 Dry weight (g) 

Entry Control 1 Pass 2 Passes 4 Passes 

LSD 

(0.05) 

Snake A: Introduced 

Crested wheatgrass 

Hycrest 44.3 37.3 42.3 36.3 ns 

Ephraim 30.2 19.8 17.8 12.3 4.0 

Mean 37.2 28.5 30.0 25.3 

Siberian wheatgrass 

Vavilov 46.5 33.8 31.8 30.3 11.6 

P-27 45.2 44.0 41.5 25.5 18.3 

Kazakhstan 37.0 22.3 23.8 8.5 19.3 

Mean 43.9 33.3 32.3 21.4 

Russian wildrye 

Bozoisky 17.8 12.3 16.8 7.3 ns 

Tetra-1 17.5 17.5 14.0 5.8 ns 

Mean 17.7 14.9 15.4 6.5 

Intermediate wheatgrass (cv. Luna) 25.4 13.5 21.0 18.3 ns 

Snake A: Native 

Basin wildrye (Yakima) 41.6 18.5 22.5 11.3 ns 

Canby bluegrass 2.9 3.3 2.0 1.8 ns 

Western wheatgrass (cv. Rosana) 14.5 7.0 9.0 8.8 6.4 

LSD (0.05) Entries 15.5 21.2 12.1 7.6 

LSD (0.05) Species 16.7 20.2 14.0 8.1 

Snake B: Native 

Bluebunch wheatgrass 

Goldar 59.1 30.0 28.8 18.8 25.9 

Whitmar 69.9 38.5 29.3 24.8 ns 

ACC-238 74.5 33.8 32.8 14.8 33.5 

Yakima 68.6 27.3 38.0 29.0 ns 

Mean 68.1 32.4 32.2 21.8 

Snake River wheatgrass 

Secar 65.2 28.3 40.3 33.3 ns 

ACC-707 65.8 36.0 38.5 29.0 ns 

EVT-572 75.6 30.8 51.5 20.5 14.8 

Mean 68.9 31.7 43.4 27.6 

Thickspike wheatgrass and hybrids 

Bannock 24.8 18.3 12.8 8.5 ns 

E-20 (Snake River × thickspike wheatgrass) 55.6 29.0 37.5 31.5 22.3 

E-27 (Snake River × thickspike wheatgrass) 93.8 38.3 56.0 35.5 31.6 

Mean 56.6 26.9 31.6 21.1 

LSD (0.05) Native entries 23.7 18.3 25.5 23.5 

LSD (0.05) Native species ns ns ns ns 
Improved Native Grasses and Establishment Methods 71 

The one-pass treatment at the Snake A site resulted in a significant increase 

in the frequency of Sandberg bluegrass observed in the seeded plots of P-27 

Siberian wheatgrass, Bozoisky Russian wildrye, and Rosana western wheatgrass 

(Table 24). However, by the four-pass treatment, invasion of Sandberg bluegrass 

declined below or were the same as the control observations. Western wheatgrass 

spreads by rhizome development, thereby controlling erosion; however, as it 

grows, it creates an open or loose sod that allows for the invasion of bluegrass 

regardless of the tracking activity. 

At the control treatment on the Snake B site, differences in percent Sandberg 

bluegrass encroachment among entries and species were not significant, and the 

one- and two-pass treatments had little or no effect on the level of Sandberg with 

the plots (Table 24). By the four-pass treatment the invasion of Sandberg had 

increased in most plots, which corresponds to a decline in target entry species at 

the four-pass treatment. The largest significant percentages of Sandberg bluegrass occurred at the four-pass treatment with the entries Goldar (4.8%) and 

ACC-238 (4.3%) bluebunch wheatgrass and Bannock (4.3%) thickspike wheatgrass. The level of tracking had little effect on Sandberg bluegrass invasion in 

plots of Whitmar and Yakima bluebunch wheatgrass, Secar Snake River wheatgrass, and E-27 thickspike × Snake River wheatgrass hybrid. 

There was very little difference between species within tracking treatments 

for Sandberg bluegrass invasion. The average invasion of Sandberg bluegrass 

was 3.9% of the total stand for the introduced species and 2.4% for the natives. 

Thus, the native Sandberg bluegrass is able to invade introduced grass stands as 

readily as it does stands of native grasses. 

Percent Bare Ground. At Snake A, species differed significantly in percent 

bare ground only at the four-pass treatment. Rosana western wheatgrass, which 

spreads by rhizomes, had the lowest levels of bare ground compared to the control with the four-pass treatment (Table 25). Five of the eight introduced entries, 

including all the Siberian wheatgrass entries, had significant increases in bare 

ground after tracking by the two- and four-pass treatments. Of the remaining 

introduced entries, all the Russian wildrye entries and Hycrest crested wheatgrass 

showed no significant differences in percentage of bare ground at any level of 

tracking (Table 25). Among the natives at Snake A, Rosana western wheatgrass 

and Yakima basin wildrye showed significant increases in bare ground at the 

two- and four-pass treatments, respectively, while Canby bluegrass had no significant increases in bare ground at any treatment. 

At Snake B, half of the native entries had significant increases in bare ground 

by the two- or four-pass treatments and half did not. There were significant 

increases in bare ground for all of the bluebunch wheatgrasses except ACC-238, 
72 ERDC/CRREL TR-03-20 

Table 24. Mean percent Sandberg bluegrass in sown species on adjacent 

sites (Snake A and B) at Yakima Training Center.

 % Sandberg bluegrass 

Entry Control 1 Pass 2 Passes 4 Passes 

LSD 

(0.05) 

Snake A: Introduced 

Crested wheatgrass 

Hycrest 2.3 3.8 3.3 1.8 ns 

Ephraim 3.8 5.0 3.8 2.5 ns 

Mean 3.0 4.4 2.5 2.1 

Siberian wheatgrass 

Vavilov 3.3 3.3 2.8 2.3 ns 

P-27 3.5 5.0 3.8 1.8 2.1 

Kazakhstan 4.3 4.8 3.0 2.8 ns 

Mean 3.7 4.3 3.2 2.3 

Russian wildrye 

Bozoisky 5.1 9.0 4.3 2.0 1.6 

Tetra-1 4.1 6.3 4.0 3.5 ns 

Mean 4.6 7.6 4.1 2.8 

Intermediate wheatgrass (cv. Luna) 4.0 6.5 4.8 3.3 ns 

Snake A: Native 

Basin wildrye (Yakima) 4.5 5.5 3.3 2.8 ns 

Canby bluegrass 1.9 1.3 1.3 1.0 ns 

Western wheatgrass (cv. Rosana) 2.9 6.7 3.8 2.9 2.9 

LSD (0.05) Entries 2.4 4.9 2.7 2.0 

LSD (0.05) Species 2.1 ns 2.8 ns 

Snake B: Native 

Bluebunch wheatgrass 

Goldar 2.3 1.3 1.8 4.8 3.1 

Whitmar 1.8 1.5 2.0 2.0 ns 

ACC-238 2.3 0.6 2.5 4.3 0.7 

Yakima 2.3 2.3 2.0 2.3 ns 

Mean 2.2 1.5 2.1 3.3 

Snake River wheatgrass 

Secar 1.5 1.1 2.0 1.8 ns 

ACC-707 2.7 1.8 1.8 3.5 1.0 

EVT-572 1.8 1.3 1.3 3.0 1.6 

Mean 2.0 1.5 1.7 2.8 

Thickspike wheatgrass and hybrids 

Bannock 1.8 1.3 1.8 4.3 1.0 

E-20 (Snake River × thickspike wheatgrass) 1.9 1.0 2.0 3.3 1.9 

E-27 (Snake River × thickspike wheatgrass) 2.0 2.0 1.8 2.8 ns 

Mean 2.0 1.5 1.8 3.4 

LSD (0.05) Native entries ns 1.3 ns 1.4 

LSD (0.05) Native species ns ns ns 0.6 
Improved Native Grasses and Establishment Methods 73 

Table 25. Percent bare ground in plots of sown species on adjacent sites 

(Snake A and B) at Yakima Training Center.

 % bare ground 

Entry Control 1 Pass 2 Passes 4 Passes 

LSD 

(0.05) 

Snake A: Introduced 

Crested wheatgrass 

Hycrest 55.8 67.5 58.0 69.0 ns 

Ephraim 56.2 63.0 62.5 75.5 14.7 

Mean 56.0 65.2 60.2 72.2 

Siberian wheatgrass 

Vavilov 57.2 58.2 67.0 74.3 7.5 

P-27 53.8 57.5 60.0 71.0 15.6 

Kazakhstan 52.5 55.5 66.2 72.5 3.0 

Mean 54.5 57.1 64.4 72.6 

Russian wildrye 

Bozoisky 60.7 61.0 66.0 76.3 ns 

Tetra-1 61.8 62.0 69.5 77.2 ns 

Mean 61.2 61.5 67.7 76.7 

Intermediate wheatgrass (cv. Luna) 52.0 62.0 63.3 67.5 13.4 

Snake A: Native 

Basin wildrye (Yakima) 58.3 66.0 67.5 78.2 13.8 

Canby bluegrass 55.8 55.3 57.0 72.3 ns 

Western wheatgrass (cv. Rosana) 53.7 54.0 63.8 63.0 6.2 

LSD (0.05) Entries 8.4 ns 9.7 6.9 

LSD (0.05) Species ns ns ns 7.1 

Snake B: Native 

Bluebunch wheatgrass 

Goldar 63.9 77.7 73.8 77.5 11.4 

Whitmar 64.7 76.9 76.5 86.2 15.3 

ACC-238 62.9 78.9 75.9 79.4 ns 

Yakima 63.3 75.2 74.0 82.8 19.4 

Mean 63.8 77.8 75.4 81.0 

Snake River wheatgrass 

Secar 60.3 73.9 70.2 73.9 7.4 

ACC-707 59.3 71.5 72.0 79.2 ns 

EVT-572 57.3 69.9 62.2 78.3 ns 

Mean 59.8 72.7 71.1 76.6 

Thickspike wheatgrass and hybrids 

Bannock 56.5 64.8 64.3 68.5 ns 

E-20 (Snake River × thickspike wheatgrass) 60.7 74.0 70.7 75.7 ns 

E-27 (Snake River × thickspike wheatgrass) 57.9 66.7 66.0 84.0 13.7 

Mean 58.6 69.4 67.5 72.1 

LSD (0.05) Native entries ns 12.3 12.5 11.0 

LSD (0.05) Native species 
74 ERDC/CRREL TR-03-20 

Secar Snake River wheatgrass, and the E-27 Snake River × thickspike wheatgrass 

hybrid (Table 25). The remaining Snake River wheatgrasses, Bannock thickspike 

wheatgrass, and the E-20 Snake River × thickspike wheatgrass hybrid had no 

significant increases in bare ground. 

Cheatgrass Invasion. Cheatgrass has become a major threat to the ecological balance, resource conservation, and productivity of western rangelands 

(Young and McLain 1997). This noxious weed fuels recurrent fires that eventually remove all of the native woody plant components of plant communities and, 

hence, perpetuate cheatgrass dominance and continued burning (Asay et al. 

2001). Therefore, we examined cheatgrass encroachment to see how it was 

affected by tracking across the different sown species. 

At both Snake A and B sites among all entries, cheatgrass invasion was 

low—never more than 2.3% and usually lower than 1%—and there were few 

significant differences among entries or species or level of tracking (Table 26). 

Tracking Study Conclusions 

Tracking generally increased the size of gaps between plants, increased the 

amount of bare ground, and reduced the number of sown plants. The reduction of 

forage yield due to tracking was greater for the native species than for the introduced species; however, the actual weights of plants were similar between the 

two types of species in the tracking treatments. The introduced species had lower 

reductions in gaps and weights than did the natives, suggesting that, in most 

cases, the introduced species were more tolerant or recovered more rapidly under 

repeated tracking. The entries that showed the greatest resiliency to tracking 

appear to be those with the best establishment rate prior to tracking or those that 

possess rhizomes. In most cases the more rapidly establishing introduced species 

(Table 6) were more tolerant to or recovered more rapidly after repeated tracking. 

For the wildryes, which are introduced species that did not establish well, tracking reduced the number of plants, caused lower yields, and increased the gaps 

between species. Among the natives, Snake River wheatgrasses and western 

wheatgrass exhibited the most resiliency across the different measurements. The 

presence of rhizomes in the native western wheatgrass appeared to aid resiliency 

by helping plants spread into damaged areas. 

The native species Sandberg bluegrass successfully invaded both the native 

and introduced sites after tracking. Invasion of this early colonizing species can 

help promote native plant stands, satisfy federal requirements, reduce soil erosion, and provide ground cover while the slower-growing perennials are establishing. 
Improved Native Grasses and Establishment Methods 75 

Table 26. Percent cheatgrass encroachment on adjacent sites (Snake A 

and B) at Yakima Training Center.

 % cheatgrass 

Entry Control 1 Pass 2 Passes 4 Passes 

LSD 

(0.05) 

Snake A: Introduced 

Crested wheatgrass 

Hycrest 0.8 0.0 0.8 1.0 0.5 

Ephraim 0.9 0.5 1.3 0.5 ns 

Mean 0.9 0.3 1.0 0.8 

Siberian wheatgrass 

Vavilov 0.0 0.0 0.3 0.3 ns 

P-27 0.6 0.5 0.3 1.5 0.9 

Kazakhstan 0.7 2.3 0.3 1.0 ns 

Mean 0.4 0.9 0.3 0.9 

Russian wildrye 

Bozoisky 1.4 2.3 1.0 1.0 ns 

Tetra-1 1.8 2.0 0.8 1.0 ns 

Mean 1.6 2.1 0.9 1.0 

Intermediate wheatgrass (cv. Luna) 0.6 1.0 0.5 0.8 ns 

Snake A: Native 

Basin wildrye (Yakima) 1.3 1.3 1.8 1.5 ns 

Canby bluegrass 0.8 1.3 0.0 0.3 ns 

Western wheatgrass (cv. Rosana) 1.1 0.0 1.8 1.3 ns 

LSD (0.05) Entries 1.5 ns 1.2 ns 

LSD (0.05) Species ns ns 0.7 ns 

Snake B: Native 

Bluebunch wheatgrass 

Goldar 0.6 0.8 1.0 0.5 ns 

Whitmar 0.5 0.5 0.6 0.5 ns 

ACC-238 0.2 0.4 0.5 1.0 0.6 

Yakima 0.5 0.8 1.0 0.3 ns 

Mean 0.4 0.6 0.8 0.6 

Snake River wheatgrass 

Secar 0.3 0.6 0.4 0.5 0.1 

ACC-707 0.3 0.8 1.1 0.6 0.7 

EVT-572 0.2 0.4 0.6 0.4 ns 

Mean 0.3 0.6 0.7 0.5 

Thickspike wheatgrass and hybrids 

Bannock 1.2 1.8 1.8 1.3 ns 

E-20 (Snake River × thickspike wheatgrass) 0.5 0.8 1.1 0.6 ns 

E-27 (Snake River × thickspike wheatgrass) 0.4 0.6 0.8 0.4 ns 

Mean 0.7 1.1 1.2 0.8 ns 

LSD (0.05) Native entries 0.5 0.7 1.1 ns 

LSD (0.05) Native species 0.3 0.4 ns ns 
76 ERDC/CRREL TR-03-20 

The greater yields by the natives in the controls, the apparent resiliency of 

some of the natives such as Snake River and western wheatgrass, and the ability 

of Sandberg bluegrass to invade introduced grass stands suggest that natives can 

be used successfully on military training lands. The success of natives on tracked 

lands can likely be improved by mixed seedings of appropriate native entries and 

rapidly establishing introduced species. 
Improved Native Grasses and Establishment Methods 77 

6 METHODS FOR ESTABLISHING NATIVES AND 

FIGHTING INVASIVE WEEDS 

To maintain good land stewardship, we need to establish stands of native 

plants. With their rapid growth rate soon after germination, annual invasive or 

noxious weeds often prevent native plants from establishing. We believe that 

native plants can be established if they can initiate root growth more quickly after 

germination or if they can be protected from the competitive or allelopathic 

mechanisms of the noxious weeds. Along with the genetic improvements in the 

SERDP-select natives for establishment and seedling vigor, we investigated 

seeding methods to further enhance the ability of our improved germplasms to 

establish viable native plant stands as rapidly as possible. 

Introduced grasses establish more quickly, but where we have established 

strong native grass stands at both Yakima Training Center and Fort Drum, we 

have seen the natives moving into adjacent non-native grass swards. For this reason, we believe it is cost-effective to seed a mixture of non-natives with natives. 

The non-natives can provide quick, interim protection until the natives become 

sufficiently established. This use of non-natives to serve as a short-term ecological bridge to help establish healthy native stands should, ultimately, lead to a 

decrease in the need for non-native stands to provide protection for soil and wind 

erosion in moderately used training areas. 

This section reports on three studies in three climatic areas to test our 

hypothesis about using introduced plants as ecological bridges to establish native 

plants. We also discuss our work with forage kochia, which, with its potential for 

fire suppression, may also serve as an ecological bridge to allow non-fire-tolerant 

natives to become established. A potential native ecological bridge in some geographic areas is Pryor slender wheatgrass, which did well in the first year of our 

early evaluation studies at South Boundary, Fort Carson, and then died out (Table 

5). 

Establishment of Natives on Sandy Soils at Fort Drum 

On sandy soils at Fort Drum we planted mixed seedings of weeping lovegrass and fine fescues with the desired native switchgrass. Sandy soils are a 

problem on many northeastern DoD lands, including Fort Drum, New York, and 

numerous U.S. Army Corps of Engineers civil works sites with dam embankments, mine tailing sites, borrow pit areas, and roadsides. These soils are difficult 

to rehabilitate because of their low moisture- and nutrient-holding capacities, the 

short growing season, and the drying effects of winds. 
78 ERDC/CRREL TR-03-20 

We wanted to develop a cost-effective native vegetative cover for sandy soils 

that is quick to establish, long lasting, and low maintenance. Suitable species are 

available for reseeding these soils, but there are restrictions in terms of seeding 

season, time required for establishment, and length of persistence. Studies in 

plant establishment have shown that both warm- and cool-season grasses have a 

place in rehabilitating sandy soils (Schwendiman and Hawk 1975, Gaffney and 

Dickerson 1987). Cool-season grasses are easier to establish than warm-season 

grasses, they withstand some traffic, and they do not produce as much biomass as 

warm-season grasses, thus reducing the fire hazard. Their disadvantages are that 

they do better when seeded in the fall and their persistence is suspect after five 

years without additional care such as fertilization. With their long-term survival, 

native warm-season grasses are the species of choice for revegetating sandy, 

infertile soils. However, they can be sown only in the spring, and they can be 

very slow to establish, sometimes taking several years to reach a suitable cover to 

prevent soil erosion. At Fort Drum we explored methods of establishing an ecological bridge that will allow training within one year after seeding and also will 

allow the establishment of warm-season native plants over time. 

We first looked at the factors that will optimize the success of native plants 

on sandy soil, including soil compaction and early root growth of native and 

introduced plants (Brar and Palazzo 1995a,b, Palazzo 1994), which will ease 

plant establishment by allowing plants to exploit moisture at greater soil depths. 

We then established a large research site at Fort Drum where we applied varying 

rates of liquid cow manure and tested seeding mixtures. In the seeding mixtures 

we used varying rates of weeping lovegrass along with fescues and the desired 

warm-season native switchgrass. Weeping lovegrass is a tropical perennial that 

establishes very quickly but is not winter hardy this far north. We hypothesized 

that weeping lovegrass and the fescues (introduced species) would provide sufficient soil cover to allow the switchgrass to establish successfully. 

Fort Drum soil is a Plainfield sandy loam (92% sand, 6% silt, and 2% clay). 

Random sampling of soils throughout the site yielded a mean pH of 5.5, medium 

to low phosphorus, and low potassium. Before beginning our study, we amended 

the soil with fertilizer nutrients N applied at 56 kg/ha (50 lb/acre) and P2O5 and 

K2O applied at 112 kg/ha (100 lb/acre) each. Dolomitic limestone was applied at 

2240 kg/ha (2000 lb/acre). 

For treatments we applied liquid cow manure at rates of 0, 22,400, 44,800, 

and 89,600 kg/ha (0, 10, 20, and 40 tons/acre), and we seeded the weeping lovegrass at 1.12, 3.36, and 5.6 kg/ha (1, 3, and 5 lb/acre). The species used in the 

mixtures with weeping lovegrass were the native ‘Blackwell’ switchgrass and the 

introduced ‘Reliant’ hard fescue and ‘Azure’ sheep fescue. The sites were seeded 

the day after the manure application at a depth of about 1.3 cm (0.5 in.) with a 
Improved Native Grasses and Establishment Methods 79 

no-till seeder. The site was divided into four large plots, each measuring 13.7 by 

106.7 m, or about 441 m2

 (45 by 350 ft, or 15,750 ft2), and there were four replications within each treatment. Using permanent transects and a point-frame sampler, we recorded the percent total vegetative cover, dead vegetative cover, live 

introduced cover, and native vegetative cover. 

The manure slurry provided a mulching effect that prevented drying of the 

newly established plants, allowing time for them to develop roots long enough to 

reach into the deeper soil layers. All grass species appeared to grow better where 

the cow manure was applied, and good soil cover was obtained over the entire 

area in the initial season through the strong growth of weeping lovegrass. The 

weeping lovegrass established quickly at all three rates sown, providing rapid 

control of wind and water erosion and allowing the land to be opened for training 

in about one year. The seeding provided a vegetative cover of greater than 85% 

in the first year with the manure applications, quickly protecting the soil and 

moisture. Establishment was not as quick where manure was not applied, but it 

eventually established a good cover in the first year. As a warm-climate annual, 

the lovegrass died back after the first year, allowing the fescues to come in. After 

four years, switchgrass dominated the stand (Fig. 15). 

Weeping lovegrass shows promise as a nurse crop for improving the establishment of slow-growing native species and for short-term rehabilitation of sites 

that receive intensive use, such as airborne drop zones and bivouac areas. This 

technique of planting natives and non-natives together for a transition into a 

native stand is now an accepted practice at Fort Drum. 

Weeping 

lovegrass Fescues Switchgrass

Years

Cover

Weeping 

lovegrass Fescues Switchgrass

Years

Cover

Figure 15. Ecological bridge concept. Weeping lovegrass 

acts as a nurse crop on sandy soils at Fort Drum, allowing 

fescues and eventually the desired native grass, switchgrass, to become established. 
80 ERDC/CRREL TR-03-20 

Native Wheatgrass Establishment in Cheatgrass-Infested Areas at Yakima 

At Yakima Training Center we tested the hypothesis that introduced Siberian 

wheatgrass could act as an ecological bridge to the establishment of the native 

bluebunch wheatgrass in a cheatgrass-infested area. As previously mentioned, 

cheatgrass is a major threat to ecological balance, resource conservation, and 

productivity. It forms a closed system and fuels recurrent fires, which enhances 

the production of seed by cheatgrass and further perpetuates its dominance. A 

logical means of controlling cheatgrass is to replace it with a perennial grass, but 

there has been limited success replacing the cheatgrass with native grasses. The 

introduced crested and Siberian wheatgrasses have shown potential in inhibiting 

cheatgrass growth and may be used as an ecological bridge to reduce the amount 

of cheatgrass, allowing the establishment of natives. 

Disturbed sites at Yakima Training Center were seeded in November 1998. 

We used a drill equipped with double-disk furrow openers and depth-band regulators to seed on mechanically prepared, weed-free seedbeds. Seeds were placed 

from 1.25 to 2.0 cm below the soil surface at a rate of 100 pure live seeds per 

linear meter. Individual plots consisted of four drilled rows with either 25- or 35-

cm row spacings. We used Snake River wheatgrass (native), bluebunch wheatgrass (native), and Vavilov Siberian wheatgrass (introduced) in the following 

treatments, with four replications of each treatment: 

• A monoculture of each of the three grasses; 

• A binary seed mixture of Vavilov with each native grass; and 

• Alternating rows of Vavilov with each native grass. 

Table 27 shows percent of cheatgrass in each plot in 2000. Plots with 

Vavilov had lower amounts of cheatgrass. For instance, significant reductions in 

cheatgrass occurred when Vavilov was planted in alternating rows with bluebunch or Snake River wheatgrass, compared to each of those natives planted 

without Vavilov. In the Vavilov and bluebunch combinations, Vavilov allowed 

bluebunch to get established. The next step is to see if Vavilov will allow bluebunch to persist, especially through drought (Asay et al. 2001). 

Mixed Native and Introduced Seedings at Fort Carson 

In the second study we evaluated mixtures of native and introduced grasses 

in plantings at Turkey Creek, Fort Carson, Colorado. This study was dormantseeded in the fall of 1997. The treatments involved a core mix of native grasses 

plus one of five introduced grasses. The core native mix is shown in Table 28; 

the introduced species was either Bozoisky Russian wildrye, SERDP-select 

Tetra-1 Russian wildrye, SERDP-select RoadCrest crested wheatgrass, Vavilov 
Improved Native Grasses and Establishment Methods 81 

Table 27. Percent cheatgrass in establishment study plots with two row 

spacings at Yakima Training Center in 2000 (two years after establishment). 

 % Cheatgrass 

Grasses 25-cm spacing 35-cm spacing Mean 

Bluebunch wheatgrass 53 65 59 

Snake River wheatgrass 35 78 57 

Vavilov Siberian wheatgrass 7 30 19 

Bluebunch/Snake River Mix 57 70 64 

Bluebunch/Snake River alternating rows 30 52 41 

Bluebunch/Vavilov mix 17 47 32 

Bluebunch/Vavilov alternating rows 30 52 41 

Snake River/Vavilov mix 25 48 36 

Snake River/Vavilov alternating rows 25 48 36 

Bluebunch/Snake River/Vavilov mix 16 50 33 

Mean 32 56 44 

LSD (0.05) 23 19 15 

Table 28. Core native mix plus introduced species for mixed seedings at 

Fort Carson. 

Species kg/ha lb/acre % of mix 

Core native mix 

Barton western wheatgrass 4.5 4 26 

Pryor slender wheatgrass 2.2 2 14 

Nezpar Indian ricegrass 1.1 1 6 

Vaughan sideoats grama 1.1 1 6 

Critana thickspike wheatgrass 2.2 2 13 

Lovegrass 0.6 0.5 3 

Blue grama 1.1 1 6 

Introduced grass 4.5 4 26 

Total 17.3 15.5 100 

Siberian wheatgrass, or SERDP-select CD-II crested wheatgrass. For comparison 

the Fort Carson standard mix (Table 29) was also seeded at increased rates to 

match the above treatments. The plots were evaluated in 1999, 2000, and 2001 

for species composition, percent ground cover, percent annual and biennial 

weeds, percent introduced grasses, and percent natives. 
82 ERDC/CRREL TR-03-20 

Table 29. Fort Carson standard mix at seeding rate increased to match 

the core-mix-plus-introduced-species treatment. 

Species kg/ha lb/acre % of mix 

Barton western wheatgrass (native) 8.8 7.9 51 

Vaughan sideoats grama (native) 4.9 4.4 28 

Alkali sacaton (native) 0.5 0.4 3 

Sand dropseed (native) 0.3 0.3 2 

Nordan crested wheatgrass (introduced) 2.0 1.8 11 

Ladak alfalfa (introduced) 0.8 0.7 5 

Total 17.3 15.5 100 

After three years all mixes resulted in stands with less than 5% weeds. In all 

three years the mixes with the crested or Siberian wheatgrasses had the fewest 

weeds, with less than 15% of the stand each year of the study (Fig. 16). The 

weeds in the plots containing the Russian wildryes decreased with time: after the 

first year, the Bozoisky plot had about 19% weeds and the Tetra-1 plot had 32% 

weeds, with both decreasing to about 3% after three years. The Fort Carson mix 

plot had over 60% weeds after the first year, but it also decreased to about 5% 

after three years. 

0

10

20

30

40

50

60

70

1999 2000 2001 1999 2000 2001 1999 2000 2001 1999 2000 2001 1999 2000 2001 1999 2000 2001

Bozoisky RWR Tetra-1 RWR Ft. Carson Mix RoadCrest CWG Vavilov SWG CD-2 CWG

T reatment/Year

% of Plot Biomass

Figure 16. Percent annual and biennial weeds in Turkey Creek plots planted with the standard Fort Carson mix or with a core native mix plus an introduced grass (as named on the 

treatment axis). RWR = Russian wildrye, CWG = crested wheatgrass, SWG = Siberian 

wheatgrass. 
Improved Native Grasses and Establishment Methods 83 

Percent Native Grasses

0

10

20

30

40

50

60

70

80

90

100

1999 2000 2001 1999 2000 2001 1999 2000 2001 1999 2000 2001 1999 2000 2001 1999 2000 2001

Bozoisky RWR Tet ra-1 RWR Ft. Carson Mix RoadCrest CWG Vavilov SWG CD-2 CWG

Treatment/Year

% o f Plo t B iomass

Sand dropseed

Lovegrass

Indian Ricegrass

Blue grama

Side Oats Grama

Thickspike Wheatgrass

Slender Wheatgrass

Western Wheatgrass

Figure 17. Percent native grasses in Turkey Creek plots planted with the standard Fort 

Carson mix or with a core native mix plus an introduced grass (as named on the treatment 

axis). RWR = Russian wildrye, CWG = crested wheatgrass, SWG = Siberian wheatgrass. 

After three years, native grasses predominated in the plots with the Fort Carson mix and the core mix with Russian wildrye, while the introduced wheatgrasses predominated with the core mixes containing the crested or Siberian 

wheatgrass (Fig. 17). Thus, the mixes that inhibited the weedy species in the first 

year or two—the core mixes with crested or Siberian wheatgrass—resulted in the 

lowest establishment of native plants. Core mixtures containing either Bozoisky 

or Tetra-1 Russian wildrye had at least 60% natives in the stand from the first 

year on, and those stands had three native species present in nearly equal numbers. The Fort Carson mix resulted in the greatest number of natives after three 

years, but the natives established more slowly, from about 30% to 90% over the 

three years, and the mix produced much less diversity among the natives species, 

with western wheatgrass as the only prominent native. The Fort Carson mix 

contained some crested wheatgrass but at a lower rate (11%) than for the core 

mix with crested wheatgrass (26%). It might be interesting to try different rates 

of crested wheatgrass to find the optimum inhibition of weeds while ultimately 

producing a diverse native stand. 

From these results, there appears to be several options depending on the 

objective: 

• If the objective is rapid control of soil erosion and weeds in areas with 

frequent disturbance, then the core mix plus crested wheatgrass is the 
84 ERDC/CRREL TR-03-20 

best choice. Although fewer natives become established with this mix, it 

establishes rapidly and provides the most complete weed control and soil 

stabilization. 

• If there are four or more years between disturbances (or if disturbances 

are light) and erosion and weeds are not problems, then the Fort Carson 

mix with western wheatgrass predominating is the optimum choice. 

There is the opportunity for noxious weed invasion during the first two 

years when the Fort Carson mix is used, but after four years it results in a 

stand predominantly of the native western wheatgrass. 

• For control of soil erosion and weeds while allowing general buildup of 

diverse native grasses, then the best choice is the core mix plus Russian 

wildrye. 

Forage Kochia as an Ecological Bridge 

Forage kochia is an introduced, perennial shrub that has the potential for fire 

suppression. We evaluated forage kochia accessions for vigor and greenness at 

several sites in Utah. We initially had this species in the improved breeding program but eventually dropped it because we were not successful in selecting for 

improved seed viability. Forage kochia remains of interest in our program for its 

possible role in fire suppression and, hence, its potential as an ecological bridge. 

Many scientists and rangeland managers consider forage kochia to be a prime 

candidate for use in western range rehabilitation and fire prevention (Harrison et 

al. 2002). By stopping the fire cycle, forage kochia may act as an ecological 

bridge to fight invasive weeds and support native plant establishment. Fire is 

associated with increased invasion of cheatgrass by reducing and eliminating 

native perennial plant communities and increasing seed set in cheatgrass. Forage 

kochia can be used in “green strips” to contain wildfires, and it is competitive 

enough to help stop the spread of invasive weeds such as cheatgrass. Forage 

kochia seeded at 1.1 kg/ha (1 lb/acre) with a mix of grasses provides good forage 

and habitat for wildlife and livestock; however, forage kochia seeded as a monoculture at 3.4–6.7 kg/ha (3–6 lb/acre) can rapidly provide a good firebreak. Firebreaks can be established using lower rates, such as 1.1 kg/ha, but it may take six 

or more years to become fully established to the point where it can stop fires. 

While controlling fires is a major effort at many western military facilities, 

there is concern that this non-native species will invade native stands of perennial 

grasses. We surveyed the dominance of this species in its native land of Kazakhstan and included discussions with Russian and Kazakhstan scientists (Harrison 

et al. 2000). Forage kochia was never observed to be dominant in the major plant 

communities of sagebrush, saltbush, winterfat, crested and Siberian wheatgrass, 
Improved Native Grasses and Establishment Methods 85 

and needlegrass. Nowhere did this species invade from disturbed sites into adjacent perennial plant communities on the Kazakhstan steppes. We found that, 

many years after planting, forage kochia had not spread into perennial plant 

communities. However, it will recruit into disturbed sites and alkali slick spots 

and stabilize these areas, which is a better choice over cheatgrass and other invasive weeds (Harrison et al. 2000). Forage kochia is good for insects and wildlife 

such as sage grouse and other birds, deer, and elk, whereas cheatgrass creates a 

relatively barren and closed ecosystem. Soil is the most important commodity, 

and its loss through erosion is irreplaceable; therefore, it may be beneficial to use 

an introduced species such as forage kochia to preserve the soil for future restoration to more desired natives. 
86 ERDC/CRREL TR-03-20 

7 CONCLUSIONS 

We developed more wear-resistant plants and a knowledge base of the relationships between military training and plant injury, regrowth, and wear resistance to help land managers make knowledgeable choices concerning plant 

selection and site rehabilitation procedures to reduce soil erosion, promote 

training, and protect environmentally sensitive areas. Although our original work 

was with introduced plants, we adapted our approach to include more native species in response to Presidential Order 13148 on using native species and to 

increased interest in using native species to prevent encroachment of invasive 

weeds. 

Improved Plant Materials 

Using plant breeding techniques we were able to improve traits related to 

resiliency and establishment in introduced and native species of rangeland 

grasses. In our improved plant materials, we were able to select for early spring 

growth, increased seedling vigor, improved tiller and rhizome development after 

disturbance, and resistance to abiotic and biotic stresses. We have made four 

releases to date, and our latest release, the native P-7 bluebunch wheatgrass, has 

been widely accepted by other federal land managers because of its increased 

genetic diversity. 

An Independent Review Panel, convened to address concerns about the invasiveness of the introduced plants we used to develop wear-resistant cultivars for 

military training lands, found that those plants were not a threat to diverse, native 

ecosystems. The introduced germplasms we worked with did not encroach into 

other plant communities and did not establish monocultures. Our improved plant 

materials are ecologically compatible to the military sites because they were 

developed on and from collections of species existing at these sites. Military 

installations and other federal landowners can use both our introduced and native 

germplasms with no fears of adverse effects on ecosystem diversity. 

We made significant advances in relating molecular markers to both plant 

characteristics and genetic diversity. We used markers to identify species and 

individual plants that can grow better at low temperatures. For genetic diversity 

we now have the tools to assess the genetic differences and similarities in commercial and natural seed sources. Land managers will be able to confidently 

select the most cost-effective seed sources that will ensure genetic compatibility 

with existing populations. 
Improved Native Grasses and Establishment Methods 87 

Tracking Studies 

Our improved species show promise for better resilience on training lands. 

Our tank traffic studies showed that introduced species are more tolerant and 

recover more rapidly under repeated tracking than native plants. However, two of 

the natives we are working with, western wheatgrass and Snake River wheatgrass, showed promise as stabilization species since they are able to spread into 

damaged areas. We also observed that the native species Sandberg bluegrass, 

another of the species we are improving, successfully invaded sites containing 

introduced species after tracking. The greater yields by the natives in the controls 

indicates that, as native plants are developed with improved training resilience, 

they can be used successfully in combination with introduced species to obtain 

native plant swards. The tracking study data suggest that a plant’s ability to germinate quickly with strong seedlings and become established is perhaps the most 

critical character in selecting plants for resiliency after tracking, and that is, 

indeed, one of the traits that we emphasized in the breeding program. 

Establishment of Natives 

Our studies on ecological bridges show that we can make intelligent choices 

on selecting seed mixtures that will allow for earlier land use for training and still 

end up in the out years with a native plant stand. The species in the seed mixtures 

and the equipment used are readily available, and the seeding is accomplished in 

one application, thus saving money. Our improved germplasms will make these 

seeding mixes even more desirable. This ecological bridge concept has been very 

well received at several invited technical presentations. Our methods can be further enhanced to provide greater cost-effective benefits by combining our findings with those of other SERDP projects on invasive weed control: “Application 

of Hyperspectral Techniques To Monitoring and Management of Invasive Weed 

Infestation” (Ustin, CS-1143), “Exotic Annual Grasses in Western Rangelands: 

Predicting Resistance and Resilience of Native Ecosystems to Invasion” (Belnap, 

CS-1144), and “Integrated Control and Assessment of Knapweed and Cheatgrass 

on Department of Defense Installations” (Paschke, CS-1145). 

Benefits 

Our work is already providing a better return on the military investment in 

the ITAM program, through seed mixtures in use at three installations. Our new 

plant materials and seeding techniques can already provide improved plant persistence on all military lands at a reduced environmental risk with respect to 

habitat loss and soil erosion. As native grass stands are established more quickly 

and at lower seeding costs, training lands will have decreasing downtime, further 
88 ERDC/CRREL TR-03-20 

adding to cost savings for facilities. The various facets of this project are already 

increasing the value and use of current training areas, offering reduced unittraining costs, and enhancing DoD mission-related environmental activities. 
Improved Native Grasses and Establishment Methods 89 

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Doe III, W.W., R.B. Shaw, R.G. Bailey, D.S. Jones, and T.E. Macia (1999) 

Locations and environments of U.S. Army training and testing lands: An ecoregional framework for assessment. Federal Facilities Environmental Journal,

Autumn. 

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inferred from metric distances among DNA haplotypes: Applications to human 

mitochondrial DNA restriction data. Genetics, 131: 479–491. 

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(Euphorbia esula) control with several grass species. Weed Technology, 12: 374–

380. 

Gaffney, F.B. and J.A. Dickerson (1987) Species selection for revegetating 

sand and gravel mines in the Northeast. Journal of Soil and Water Conservation,

42: 358–361. 

Gatto, L.W., J.J. Halvorson, D.K. McCool, and A.J. Palazzo (2001) Effects of 

freeze–thaw cycling on soil erosion. In Landscape Erosion and Evolution Modeling (R.S. Harmon and W.W. Doe III, ed.). New York: Kluwer Academic/ 

Plenum Publishers, Chapter 3, p. 29–55. 

Gibb, D.M. (2002) DoD and ecosystem management—Hitting a moving target. 

Proceedings, Partners in Environmental Technology Technical Symposium and 

Workshop, Washington, D.C., p. 92. 

Gibbs, J.L., G. Young, and J.R. Colson (1991) Registration of ‘Goldar’ bluebunch wheatgrass. Crop Science, 31: 1708. 

Halvorson, J.J., D.K. McCool, L.G. King, and L.W. Gatto (2001) Soil compaction and over-winter changes to tracked-vehicle ruts, Yakima Training Center, Washington. Journal of Terramechanics, 38: 133–151. 

Hardy, S.E., and A.J. Palazzo (2002) Report on the workshop on new grass 

germplasms and invasive weed control. ERDC/CRREL Special Report SR-02-2, 

Cold Regions Research and Engineering Laboratory, Hanover, New Hampshire. 

(http://www.crrel.usace.army.mil/techpub/CRREL_Reports/reports/SR02-2.pdf). 

Harrison, R.D., N.J. Chatterton, B.L. Waldron, B.W. Davenport, A.J. 

Palazzo, W.H. Horton, and K.H. Asay (2000) Forage kochia—Its compatibility 

and potential aggressiveness on Intermountain rangelands. Utah Agricultural 

Experiment Station Research Report 162, Utah State University, Logan, Utah. 

Harrison, R.D., B.L. Waldron, K.B. Jensen, R. Page, T.A. Monaco, W.H. 

Horton, and A.J. Palazzo (2002) Forage kochia helps fight range fires. Rangelands, 24 (5): 3–7. 
92 ERDC/CRREL TR-03-20 

Hart, R.H., W.D. Willms, and M.R. George (1996) Cool-season grasses in 

rangelands. In Forages, Volume I: An Introduction to Grassland Agriculture

(M.E. Heath et al., ed.). Fifth edition. Ames, Iowa: Iowa State University Press, 

p. 357–381. 

Hein, M.A. (1958) Registration of varieties and strains of grasses. Wheatgrasses 

(Agropyron spp.). III. Whitmar beardless wheatgrass (Reg. No. 4). Agronomy 

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(1990) Rehabilitation of military training areas damaged by tracked vehicles at 

Fort Carson, CO. Facilities Engineering Applications Program, FEAP-TR-N91/01, Fort Belvoir, Virginia. 

Hu, Z.-M., R.R.-C. Wang, S. Larson, A.J. Palazzo, K.H. Asay, and N.J. 

Chatterton (2000) Studies on quantitative traits of Hycrest growing at low temperatures using AFLP and RAPD markers. Plant and Animal Genome VIII Conference, San Diego, California.

Hu, Z.-M., R.R.-C. Wang, S. Larson, A.J. Palazzo, K.H. Asay, and N.J. 

Chatterton (2001) Selection response for molecular markers associated with 

anthocyanin coloration and low-temperature growth traits in crested wheatgrasses. Canadian Journal of Plant Science, 81: 665–671. 

Hubbard, W.A. (1975) Increased range forage production by reseeding and the 

chemical control of knapweed. Journal of Range Management, 28: 406–407. 

Huff, D.R., and A.J. Palazzo (1998) Fine fescue species determination by laser 

flow cytometry. Crop Science, 38: 445–450. 

Huff, D.R., J. Quinn, B. Higgins, and A.J. Palazzo (1998) RAPD variation 

among little bluestem [Schizachyrium scoparium (Michx.) Nash] populations 

from sites of high and low fertility in forest and grassland biomes. Molecular 

Ecology, 7: 1591–1597. 

Hull, Jr., A.C., and G. Stewart (1948) Replacing cheatgrass by reseeding with 

perennial grass on southern Idaho ranges. Agronomy Journal, 40: 694–703. 

Jensen, K.B., K.H. Asay, D.A. Johnson, W.H. Horton, A.J. Palazzo, and N.J. 

Chatterton (1998) Registration of RWR-Tetra-1 Tetraploid wildrye germplasm. 

Crop Science, 38: 1405. 
Improved Native Grasses and Establishment Methods 93 

Jensen, K.B., K.H. Asay, D.A. Johnson, and B.J. Li (2000) Characterization of 

Siberian wheatgrass germplasm (Agropyron fragile) from Kazakhstan, U.S.S.R. 

(Poaceae: Triticeae). Journal of Range Management, 53: 347–352. 

Jensen, K.B., W.H. Horton, R. Reed, and R.E. Whitesides (2001) Intermountain planting guide. Extension Publication AG510, Utah State University, Logan, 

Utah. 

Jones, T.A., S. R. Larson, D.C. Nielson, S.A. Young, N.J. Chatterton, and 

A.J. Palazzo (2002) Registration of P-7 bluebunch wheatgrass germplasm. Crop 

Science, 42: 1754–1755. 

Keystone Center (1996) Keystone Center policy dialogue on Department of 

Defense (DoD) Biodiversity Management Strategy: Final report. Keystone, Colorado: The Keystone Center. (www.denix.osd.mil/denix/Public/ES-Programs/ 

Conservation/Strategy/ strategy.html).

Kilcher, M.R. and J. Looman (1983) Comparative performance of some native 

and introduced grasses in southern Saskatchewan. Journal of Range Management, 36: 654–657. 

Larson, L.L., and M.L. McInnis (1989) Impact of grass seedings on established 

density of diffuse knapweed and yellow star thistle. Northwest Science, 63: 162–

166. 

Larson, S.R., T.A. Jones, Z.-M. Hu, R.R.-C. Wang, P.A. Harrison, D.C. 

Nielson, A.J. Palazzo, and N.J. Chatterton (1999) QTL studies of lowtemperature growth in Leymus wildryes. Agronomy Abstracts, American Society 

of Agronomy, Salt Lake City, Utah. 

Larson, S.R., T.A. Jones, Z.-M. Hu, A.J. Palazzo, and C.L. McCracken

(2000) Genetic diversity of bluebunch wheatgrass cultivars and a multiple-origin 

polycross. Crop Science, 40: 1142–1147. 

Larson, S.R., B.L. Waldron, S. Monsen, L. St. John, A.J. Palazzo, C.L. 

McCracken, and R.D. Harrison (2001) AFLP variation in agamospermous and 

dioecious bluegrasses of western North America. Crop Science, 41: 1300–1305. 

Larson, S.R., A.J. Palazzo, and K.B. Jensen (2003) Maintenance of DNA 

variation within and among western wheatgrass cultivars and accessions. Crop 

Science (Accepted). 

Lawrence, T., and C.D. Ratzlaff (1989) Performance of some native and introduced grasses in a semiarid region of western Canada. Canadian Journal of Plant 

Science, 69: 251–254. 
94 ERDC/CRREL TR-03-20 

Lawrence, T.A., A.E. Slinkard, C.D. Ratzlaff, N.W. Holt, and P.G. Jefferson

(1990) Tetracan, Russian wildryegrass. Canadian Journal of Plant Science, 70:

311–313. 

Liu, X.L., R.R.-C. Wang, K.B. Jensen, A.J. Palazzo, N.J. Chatterton, and 

K.H. Asay (1997) RAPD and STS markers for fast-germinating western wheatgrass and salina wildrye. Abstract, American Society of Agronomy meeting, 

Anaheim, California, October 26–31, p.148. 

Marlette, G.M., and J.E. Anderson (1986) Seed banks and propagule dispersal 

in crested wheat. Journal of Applied Ecology, 23: 161–175. 

Palazzo, A.J. (1994) Establishment and persistence of cool- and warm-season 

grasses on sandy soils. Special Report 94-31, Cold Regions Research and Engineering Laboratory, Hanover, New Hampshire. 

Palazzo, A.J., and G. Brar (1997) Screening of 12 Festuca cultivars for rapid 

root development. Journal of Turfgrass Science, 2(1): 15–25. 

Palazzo, A.J., and C.R. Lee (1997) Root growth and metal uptake of plants 

grown on Zn contaminated soils as influenced by soil treatment and plant species. In Proceedings of Fourth International Conference on the Biogeochemistry 

of Trace Elements, Berkeley, California, June 1997. 

Palazzo, A.J., S.E. Hardy, and K. Taylor (1999) Report of independent review 

panel meeting: Evaluation of naturalized species being used for new cultivar 

development. Contract Report CON 154, Cold Regions Research and Engineering Laboratory, Hanover, New Hampshire. 

Palazzo, A.J., T.J. Cary, and K.H. Asay (2001) Planting techniques to establish 

native plant meadows. Tri-Services DoD Pest Management Workshop, Jacksonville, Florida. 

Richards, R.T., J.C. Chambers, and R. Christopher (1997) Use of native 

plants on federal lands: Policy and practice. Journal of Range Management, 51:

625–632. 

Rohlf, F.J. (1992) NTSYS-pc. Numerical taxonomy and multivariate analysis 

system (version 2.02). Exeter Software, Setauket, New York. 

SAS Institute Inc. (1994) SAS/STAT Users Guide, ver. 6. Fourth edition. Cary, 

North Carolina. 

Schwendiman, J.L., and V.B. Hawk (1975) Other grasses for the North and 

West. In Forages (M.E. Heath et al., ed.). Ames, Iowa: Iowa State University 

Press, p. 231–242. 
Improved Native Grasses and Establishment Methods 95 

Shaw, R.B., and V.E. Diersing (1990) Tracked vehicle impacts on vegetation at 

the Pinyon Canyon Maneuver Site, Colorado. Journal of Environmental Quality,

19: 234–243. 

Vallentine, J.F. (1971) Range Development and Improvements. Provo, Utah: 

Brigham Young University Press. 

Vallentine, J.F. (ed.) (1978) U S.–Canadian Range Management, 1935-1977: A 

Selected Bibliography on Ranges, Pastures, Wildlife, Livestock, and Ranching. 

Phoenix, Arizona: Oryx Press, p. 130–134. 

Wang, R.R.-C., and J.D. Berdahl (1990) Meiotic associations at metaphase I in 

diploid, triploid, and tetraploid Russian wildrye [Psathyrostachys juncea (Fisch.) 

Nevski]. Cytologia, 55: 639–643. 

Whitson, T.D., and D.W. Koch (1998) Control of downy brome (Bromus tectorum) with herbicides and perennial grass competition. Weed Technology, 12:

391–396. 

Young, J.A., and J.L. McLain (1997) Grazing livestock as a management tool 

in the intermountain west. Proceedings, American Society of Animal Science, 

Western Section, 48: 3–7. 
96 ERDC/CRREL TR-03-20 

APPENDIX A. PUBLICATIONS AND PRESENTATIONS 

ORIGINATING FROM SERDP PROJECT 

The following publications relate to research funded by the Strategic Environmental Research and Development Program (SERDP) project, CS-1103, 

“Identify Resilient Plant Characteristics and Develop Wear-Resistant Plant Cultivars for Use on Military Training Lands.” 

Peer-reviewed Journals or Papers 

Asay, K.H., N.J. Chatterton, K.B. Jensen, R.R.-C. Wang, D.A. Johnson, 

W.H. Horton, A.J. Palazzo, and S.A. Young (1997) Registration of ‘CD-II’ 

crested wheatgrass. Crop Science, 37: 1023. 

Asay, K.H., W.H. Horton, K.B. Jensen, and A.J. Palazzo (2001) Merits of 

native and introduced Triticeae grasses on semiarid rangelands. Canadian Journal of Plant Science, 81: 45–52. 

Asay, K.H., K.B. Jensen, W.H. Horton, D.A. Johnson, N.J. Chatterton, and 

S.A. Young (1999) Registration of RoadCrest crested wheatgrass. Crop Science,

39: 1535. 

Harrison, R.D., B.L. Waldron, K.B. Jensen, R. Page, T.A. Monaco, W.H. 

Horton, and A.J. Palazzo (2002) Forage kochia helps fight range fires. Rangelands, 24(5): 3–7. 

Hu, Z.-M., R.R.-C. Wang, S. Larson, A.J. Palazzo, K.H. Asay, and N.J. 

Chatterton (2001) Selection response for molecular markers associated with 

anthocyanin coloration and low-temperature growth traits in crested wheatgrasses. Canadian Journal of Plant Science, 81: 665–671. 

Huff, D.R., J. Quinn, B. Higgins, and A.J. Palazzo (1998) RAPD variation 

among little bluestem [Schizachyrium scoparium (Michx.) Nash] populations 

from sites of high and low fertility in forest and grassland biomes. Molecular 

Ecology, 7: 1591–1597. 

Jensen, K.B., K.H. Asay, D.A. Johnson, W.H. Horton, A.J. Palazzo, and N.J. 

Chatterton (1998) Registration of RWR-Tetra-1 tetraploid. Crop Science, 38:

1403. 

Jensen, K.B., K.H. Asay, D.A. Johnson, and B.J. Li (2000) Characterization of 

Siberian wheatgrass germplasm (Agropyron fragile) from Kazakhstan, U.S.S.R. 

(Poaceae: Triticeae). Journal of Range Management, 53: 347–352. 
Improved Native Grasses and Establishment Methods 97 

Jones, T.A., S.R. Larson , D.C. Nielson, S.A. Young, N.J. Chatterton, and 

A.J. Palazzo (2002) Registration of P-7 bluebunch wheatgrass germplasm. Crop 

Science, 42: 1754–1755. 

Larson, S.R., A.J. Palazzo, and K.B. Jensen (2003) Maintenance of DNA 

variation within and among western wheatgrass cultivars and accessions. Crop 

Science (Accepted). 

Larson, S.R., T.A. Jones, Z.-M. Hu, A.J. Palazzo, and C.L. McCracken

(2000) Genetic diversity of bluebunch wheatgrass cultivars and a multiple-origin 

polycross. Crop Science, 40: 1142–1147. 

Larson, S.R., B.L. Waldron, S. Monsen, L. St. John, A.J. Palazzo, C.L. 

McCracken, and R.D. Harrison (2001) AFLP variation in agamospermous and 

dioecious bluegrasses of western North America. Crop Science, 41: 1300–1305. 

Palazzo, A.J., and S.E. Hardy (1998) Department of Defense evaluates genetic 

diversity on military training lands and breeds new plants for Army training 

grounds. Diversity, 14: 28–30. 

Technical Reports 

Asay, K.H., K.B. Jensen, T.A. Jones, B.L. Waldron, A.J. Palazzo, D.A. Johnson, W.H. Horton, and N.J. Chatterton (2001) Breeding native and introduced 

grasses and legumes for the Northern Plains area. Society of Range Management, 

Kilo, Hawaii (Invited paper). 

 Hardy, S.E., and A.J. Palazzo (2002) Report on the workshop on new grass 

germplasms and invasive weed control. ERDC/CRREL Special Report SR-02-2, 

Cold Regions Research and Engineering Laboratory, Hanover, New Hampshire. 

(http://www.crrel.usace.army.mil/techpub/CRREL_Reports/reports/SR02-2.pdf) 

Harrison, R.D., N.J. Chatterton, B.L. Waldron, B.W. Davenport, A.J. 

Palazzo, W.H. Horton, and K.H. Asay (2000) Forage kochia—Its compatibility 

and potential aggressiveness on Intermountain rangelands. Utah Agricultural 

Experiment Station Research Report 162, Utah State University, Logan, Utah. 

Palazzo, A.J., S.E. Hardy, and K. Taylor (1999) Report of independent review 

panel meeting: Evaluation of naturalized species being used for new cultivar 

development. Contract Report CON 154, Cold Regions Research and Engineering Laboratory, Hanover, New Hampshire. 
98 ERDC/CRREL TR-03-20 

Conference/Symposium Proceedings Papers 

Huff, D.R., A.J. Palazzo, M. van der Grinten, and T. Lent (2000) DNA 

marker variation among populations of native and commercial little bluestem 

[Schizachyrium scoparium (Michx.) Nash]. Proceedings of the Northeast Native 

Grass Conference, November 17–19, 1999, Baltimore, Maryland, p. 162. Published by the USDA-Agricultural Research Service and USDA-Natural 

Resources Conservation Service, Beltsville, Maryland. 

Palazzo, A.J., T. Cary, T. Lent, and D. Huff (2000) Revegetation of sandy 

soils in cold regions. Proceedings of the Northeast Native Grass Conferenc,

November 17–19, 1999, Baltimore, Maryland, p. 235. Published by the USDAAgricultural Research Service and USDA-Natural Resources Conservation Service, Beltsville, Maryland. 

Published Technical Abstracts 

Asay, K.H., K.B. Jensen, T.A. Jones, B.L. Waldron, and A.J. Palazzo (2000) 

Breeding grasses for disturbed military training lands. Abstracts, American Society of Agronomy, Minneapolis, Minnesota, p. 105. 

Cary, T.J., A.J. Palazzo, and K.H. Asay (2001) Innovative techniques to establish native plants. Presented at the Tenth Annual ITAM Workshop on Military 

Land Rehabilitation and Management, 27–31 August, Nashville, Tennessee, p. 

45. 

Hu, Z.-M., R.R.-C. Wang, S. Larson, A.J. Palazzo, K.H. Asay, and N.J. 

Chatterton (2000) Studies on quantitative traits of Hycrest growing at low temperatures using AFLP and RAPD markers. Abstracts, Plant and Animal Genome 

VIII Conference, Jan. 9–12, 2000, San Diego, California. 

Huff, D.R. (2000) Genetic characterization of heterogeneous plant populations. 

Abstracts, Plant and Animal Genome VIII Conference, Jan. 9–12, 2000, San 

Diego, California. 

Larson, S.R., T.A. Jones, Z.-M. Hu, R.R.-C. Wang, P.A. Harrison, D.C. 

Nielson, A.J. Palazzo, and N.J. Chatterton (1999) QTL studies of lowtemperature growth in Leymus wildryes. Abstracts, American Society of Agronomy, Salt Lake City, Utah, p. 155. 

Palazzo, A.J. (1999) Identify resilient plant characteristics and develop wearresistant plant cultivars for use on military training lands. Partners in Environmental Technology: Technical Symposium and Workshop, 30 November–2 

December, 1999, Arlington, Virginia, p. 85. 
Improved Native Grasses and Establishment Methods 99 

Palazzo, A.J., K.H. Asay, and K.B. Jensen (2000) Identify resilient plant characteristics and develop wear-resistant plant cultivars for use on military training 

lands. Partners in Environmental Technology: Technical Symposium and Workshop, 28–30 November, 2000, Arlington, Virginia, p. 86. 

Palazzo, A.J., K.H. Asay, K.B. Jensen, and B.L. Waldron (2001) Identify 

resilient plant characteristics and develop wear-resistant plant cultivars for use on 

military training lands. Partners in Environmental Technology: Technical Symposium and Workshop, 27–29 November, 2001, Arlington, Virginia, p. 97. 

Palazzo, A.J., T.J. Cary, K.H. Asay, K. Jensen, B. Waldron, and C. Reames 

(2001) Establishing native plants on western military lands. Agronomy Abstracts, 

Charlotte, North Carolina. 

Palazzo, A.J., K. B Jensen, and B.L.Waldron (2002) Identify resilient plant 

characteristics and develop wear-resistant plant cultivars for use on military 

training lands. Partners in Environmental Technology: Technical Symposium and 

Workshop, 27–29 November, 2001, Arlington, Virginia, p. 102. 

Palazzo, A.J., K. B Jensen, and B.L.Waldron (2003) Identify resilient plant 

characteristics and develop wear-resistant plant cultivars for use on military 

training lands. Partners in Environmental Technology: Technical Symposium and 

Workshop, 27–29 November, 2001, Arlington, Virginia.

Waldron, B.L., K.B. Jensen, K.H. Asay, and A.J. Palazzo (2001) Genotype by 

location effects on seed traits of western wheatgrass. Agronomy Abstracts, 

Charlotte, North Carolina. 

Published Book Chapters 

Gatto, L.W., J.J. Halvorson, D.K. McCool, and A.J. Palazzo (2001) Effects of 

freeze–thaw cycling on soil erosion. In Landscape Erosion and Evolution Modeling (R.S. Harmon and W.W. Doe III, ed.). New York: Kluwer Academic/ 

Plenum Publishers, Chapter 3, p. 29–55. 

General Interest Publications 

SERDP (2001) Getting to the root of DoD’s revegetation problem: SERDP 

success story. SERDP Information Bulletin, Fall (10): 3,7. 

Wood, M. (2001) Plants that take a lickin’ and keep on tickin’. Agricultural 

Research, 49(10): 10–11. 
100 ERDC/CRREL TR-03-20 

APPENDIX B. DATA ON NON-INVASIVENESS OF 

INTRODUCED SPECIES 

The following data are from our “Report of Independent Review Panel 

Meeting: Evaluation of Naturalized Species Being Used for New Cultivar Development” (Palazzo et al. 1999) held at Yakima Training Center in May 1999. 

To analyze the spread of introduced species, we sampled our research plots 

and areas previously seeded at the Yakima Training Center (YTC) in western 

Washington. These sites included an area to evaluate new germplasm (Snake A 

and B sites), sites that were sown 13–19 years earlier by YTC personnel, and a 

plant breeding site (Exit 11). 

Snake A and B Sites 

The Snake A site contained 1.5- by 15-m (5- by 50-ft) plots of 28 cultivars 

(17 natives and 11 introduced), and the Snake B site contained rows of 12 native 

cultivars. Borders were planted with Hycrest crested wheatgrass (an introduced 

species) (Table B1). All plots were allowed to grow to seed maturity each year. 

The main goal of this review was to evaluate the invasive potential of plants 

in the Snake A and B plots that are being used in the breeding program to 

develop new cultivars. The species being used in the breeding program include 

natives—bluebunch wheatgrass, Snake River wheatgrass, Basin wildrye, and 

Sandberg bluegrass—and three introduced species—crested wheatgrass, Siberian 

wheatgrass, and Russian wildrye. In December 1998 we evaluated the spread of 

the trial plants at the Snake A and B sites by assessing vegetative tillering and 

natural reseeding. 

We examined vegetative spread by observing the plants along the borders of 

the sown areas of the Snake A plots (Snake B plots contained only native species). We did not observe any vegetative tillering by the introduced species outside the sown plots. Vegetative spread did not occur because the plant materials 

we are developing are bunch-type or only moderately rhizomatous species. 

To determine reproductive spread by natural reseeding of an introduced species, we counted the number of crested wheatgrass plants that moved into the plot 

area from the border around the plots. Within each plot we surveyed an area 0–3 

m (0–10 ft) and 3–6 m (10–20 ft) away from the outside border, for a total of 224 

counting locations and a total area of 1650 m2

 (18,000 ft2). 

At the Snake A site we counted 14 plants in the 0- to 3-m zone and 15 plants 

in the 3- to 6-m zone, or a total of 29 plants—which was a very small number of 

plants moving outside of areas where they were planted (Table B1). Interestingly 
Improved Native Grasses and Establishment Methods 101 

Table B1. Number of crested wheatgrass (CWG) and bluebunch wheatgrass (BWG) plants found 

inside the Snake A and B evaluation plots. Numbers are totals of eight replications. 

Common Name Origin 

CWG 

0–3 m in 

CWG 

3–6 m in BWG 

Snake A Entries 

Sandberg bluegrass Native 1 0 3 

Bottlebrush squirreltail Native 2 1 6 

Ephraim crested wheatgrass Introduced — — 0 

Vavilov Siberian crested wheatgrass Introduced — — 0 

Russian wildrye Introduced 0 0 3 

Four-wing saltbush Native 0 0 3 

Alfalfa Introduced 0 1 1 

Russian wildrye Introduced 0 0 1 

Basin wildrye - Yakima collection Native 0 1 9 

Canby bluegrass Native 2 2 0 

P27 Siberian crested wheatgrass Introduced — — 0 

Small burnett Native 0 0 2 

Western yarrow - Yakima collection Native 0 0 0 

Forage kochia Introduced 1 2 2 

Spinihop sage Native 0 3 0 

Hycrest crested wheatgrass Introduced — — 1 

Western wheatgrass Native 1 1 2 

Four-wing saltbush Native 3 0 11 

Indian ricegrass Native 0 0 0 

Altai wildrye Introduced 1 0 0 

Western yarrow Native 0 0 1 

Bluebunch x thickspike wheatgrass Native 0 0 0 

Indian ricegrass Native 1 1 2 

Needle & threadgrass Native 1 3 2 

Bottlebrush squirreltail Native 0 0 6 

Bitterbrush Native 1 0 0 

Intermediate wheatgrass Introduced 0 0 1 

Kazakhstan Siberian crested wheatgrass Introduced — — 0 

Snake B Entries 

E24 thickspike x Snake River wheatgrass hybrid Native 0 0 no data 

Sodar thickspike wheatgrass Native 0 0 no data 

Secar Snake River wheatgrass Native 0 0 no data 

Goldar bluebunch wheatgrass Native 0 0 no data 

E20 thickspike x Snake River wheatgrass hybrid Native 0 0 no data 

Bannock thickspike wheatgrass Native 0 0 no data 

5702 Snake River wheatgrass Native 0 0 no data 

Whitmar bluebunch wheatgrass Native 0 0 no data 

P2 bluebunch wheatgrass Native 0 0 no data 

Yakima bluebunch wheatgrass Native 0 0 no data 

701 Snake River wheatgrass Native 0 0 no data 
102 ERDC/CRREL TR-03-20 

we counted 56 native bluebunch wheatgrass plants that had invaded the sown 

plots. At the Snake B site we did not find any crested wheatgrass plants moving 

into the plots. 

Older Seeded Areas 

Because crested and Siberian wheatgrass are among the most promising species for our resilient cultivar development, we also prepared for the panel meeting by evaluating several training areas at Yakima that had been seeded with 

crested or Siberian wheatgrass 13–19 years ago. At these sites we measured 

along transects both inside the seeded areas and outside in the surrounding native 

range, counting “hits” of the sown species (crested or Siberian wheatgrass), any 

other species present, and bare ground. We took data at 0.3-m (1-ft) intervals 

along 30-m (100-ft) transects that were parallel to the plot boundary. The transects were 6, 12, 18, 24, and 30 m (20, 40, 60, 80, and 100 ft) from the plot 

boundaries. 

Selah-Cold Creek Summit Site 

This site was sown with crested wheatgrass in 1980 and includes deeper soils 

that receive 300 mm of precipitation annually. As shown in Table B2 the sown 

species was dominating this site within the sown area. Outside the sown areas we 

made two sets of transects. Only three plants of crested wheatgrass were found in 

one set of transects, while no crested wheatgrass plants were observed in the 

other (Table B3). Two of the plants were 6 m from the seeded area and the other 

was at 12 m. The native plant, bluebunch wheatgrass, was the dominant vegetative species outside the sown areas. 

Table B2. Number of plants found on transects at the Selah-Cold Creek Summit 

Site inside the area sown with crested wheatgrass in 1980. 

 6 m in 12 m in 18 m in 24 m in 30 m in 

Crested wheatgrass 40 34 34 39 28 

Bluebunch wheatgrass 0 0 2 0 0 

Knapweed 1 3 2 2 3 

Intermediate wheatgrass 0 0 0 0 4 

Lichen 2 1 1 2 2 

Lupine 4 1 0 0 0 

Sandberg bluegrass 6 4 11 3 7 

Yarrow 1 1 0 0 1 

Yellow brush 1 1 1 0 0 

Bare ground 13 22 24 21 17 

Litter 33 33 25 33 38 
Improved Native Grasses and Establishment Methods 103 

Table B3. Number of plants found on transects at the Selah-Cold Creek Summit 

Site outside the area sown with crested wheatgrass in 1980. 

 6 m out 12 m out 18 m out 24 m out 30 m out

Measurement # 1 

Crested wheatgrass 0 0 0 0 0 

Bluebunch wheatgrass 2 45 48 45 34 

Cheatgrass 14 2 0 0 0 

Cusick bluegrass 3 2 0 0 0 

Knapweed 3 0 1 0 0 

Lichen 0 0 0 1 1 

Lupine 4 2 2 3 3 

Miscellaneous species 1 0 0 0 0 

Rabbitbrush 3 4 1 1 4 

Sandberg bluegrass 14 9 11 10 6 

Three-tip sagebrush 0 0 3 9 5 

Yarrow 13 3 0 3 2 

Bare ground 2 4 7 6 6 

Litter 41 29 27 22 39 

Measurement # 2 

Crested wheatgrass 2 1 0 0 0 

Bluebunch wheatgrass 11 12 36 32 20 

Cheatgrass 1 0 0 0 0 

Cusick bluegrass 0 0 0 14 0 

Knapweed 6 4 0 0 0 

Lichen 4 1 2 1 7 

Lupine 2 2 0 3 2 

Miscellaneous spp. 1 2 0 0 0 

Rabbit brush 6 1 0 1 2 

Sandberg bluegrass 15 12 15 3 7 

Three-tip sagebrush 4 1 4 5 13 

Yarrow 2 6 8 9 1 

Bare ground 18 11 8 4 16 

Litter 28 47 27 28 32 
104 ERDC/CRREL TR-03-20 

Moxee Site 

This site was sown with crested wheatgrass and alfalfa in 1980. This was an 

area with less-well-developed soils and 180 mm of annual precipitation, and it 

was a likely site for invasion of noxious species. Within the sown area, both 

crested wheatgrass and the native Sandberg bluegrass were the dominant vegetative types (Table B4). There were small amounts of alfalfa still present after 19 

years. Outside the sown area we made two sets of transects and again found 

Sandberg bluegrass to be a dominant plant, but we found no crested wheatgrass 

in either area (Table B5). The noxious weed cheatgrass predominated in one set 

of measurements outside the sown area, but there were few instances of it within 

the area sown with crested wheatgrass. 

Table B4. Number of plants found on transects at the Moxee Site inside the area 

sown with crested wheatgrass and alfalfa in 1980. 

 6 m in 12 m in 18 m in 24 m in 30 m in 

Crested wheatgrass 22 17 13 20 15 

Alfalfa 4 4 17 2 6 

Bluebunch wheatgrass 0 0 0 0 0 

Cheatgrass 1 1 3 6 11 

Lichen 14 7 14 18 11 

Sandberg bluegrass 25 18 14 6 10 

Bare ground 12 24 7 4 9 

Litter 21 29 32 43 37 

Rock 1 0 0 1 1 
Improved Native Grasses and Establishment Methods 105 

Table B5. Number of plants found on transects at the Moxee Site outside the area 

sown with crested wheatgrass and alfalfa in 1980. 

 6 m out 12 m out 18 m out 24 m out 30 m out 

Measurement # 1 

Crested wheatgrass 0 0 0 0 0 

Bluebunch wheatgrass 0 0 0 0 0 

Cheatgrass 6 25 35 21 20 

Daisy 1 0 0 1 0 

Goats beard 0 0 2 5 3 

Knapweed 4 1 3 2 1 

Lichen 21 9 2 9 9 

Sandberg bluegrass 10 12 17 15 24 

Bare ground 13 13 5 9 3 

Litter 45 40 36 38 39 

Measurement # 2 

Crested wheatgrass 0 0 0 0 0 

Bluebunch wheatgrass 0 0 0 0 0 

Bottlebrush squirreltail 1 0 0 0 0 

Cheatgrass 0 0 19 8 28 

Daisy 0 0 2 1 0 

Goats beard 0 0 1 2 4 

Knapweed 0 0 3 2 1 

Lichen 0 0 21 35 26 

Mustard 0 0 1 0 0 

Sandberg bluegrass 0 0 18 17 11 

Bare ground 0 0 3 7 8 

Litter 0 0 31 28 22 
106 ERDC/CRREL TR-03-20 

Badger Pocket Road near Selah Creek Crossing Site 

This site was sown with Siberian wheatgrass in 1985 and 1986 and has shallow soils and 200 mm of annual precipitation. Siberian wheatgrass dominated 

this site within the sown area (Table B6). We found three Siberian wheatgrass 

plants 6 m from the sown area on only one of the two sets of transects measured 

outside the sown area. Both outside areas were dominated by big sagebrush 

(Table B7). 

Table B6. Number of plants found on transects at the Badger Pocket Road Site 

near Selah Creek Crossing inside the area sown with Siberian wheatgrass in 1985 

and 1986. 

 6 m in 12 m in 18 m in 24 m in 30 m in 

Siberian wheatgrass 18 16 23 21 30 

Bluebunch wheatgrass 0 0 0 0 0 

Lichen 0 6 4 9 8 

Miscellaneous species 0 0 0 0 1 

Russian thistle 0 0 0 0 2 

Sandberg bluegrass 11 4 2 4 3 

Wyoming big sagebrush 1 5 2 5 8 

Bare ground 29 16 26 14 5 

Litter 41 45 43 49 43 
Improved Native Grasses and Establishment Methods 107 

Table B7. Number of plants found on transects at the Badger Pocket Road Site 

near Selah Creek Crossing outside the area sown with Siberian wheatgrass in 

1985 and 1986. 

 6 m out 12 m out 18 m out 24 m out 30 m out 

Measurement # 1 

Siberian wheatgrass 0 0 0 0 0 

Astragalus spp. 0 1 0 0 1 

Bluebunch wheatgrass 0 2 2 4 1 

Bottlebrush squirreltail 2 2 0 0 0 

Cheatgrass 4 2 0 1 6 

Indian ricegrass 1 0 0 0 0 

Knapweed 0 1 0 0 0 

Lichen 1 3 3 11 6 

Miscellaneous species 0 0 0 0 2 

Thurber needlegrass 2 2 1 3 0 

Sandberg bluegrass 3 9 6 10 4 

Wyoming big sagebrush 15 20 23 23 22 

Bare ground 29 26 25 25 17 

Litter 43 32 40 23 39 

Measurement # 2 

Siberian wheatgrass 3 0 0 0 0 

Bluebunch wheatgrass 0 0 0 2 0 

Bottlebrush squirreltail 5 3 1 1 1 

Cheatgrass 1 0 1 4 3 

Indian ricegrass 3 1 0 1 1 

Knapweed 0 0 0 0 1 

Lichen 2 2 3 3 0 

Thurber needlegrass 0 1 1 1 15 

Russian thistle 0 0 0 3 0 

Sandberg bluegrass 10 4 1 5 4 

Wyoming big sagebrush 13 27 13 15 5 

Bare ground 29 30 44 21 15 

Litter 34 32 36 45 55 
108 ERDC/CRREL TR-03-20 

Badger Pocket Road Telephone Cable Planting Site 

This site was sown with crested wheatgrass in 1980 and has moderately deep 

soils and 230 mm of annual precipitation. Crested wheatgrass dominated the 

sown area (Table B8). Of the two sets of transects outside the sown area, we 

observed only a single crested wheatgrass plant 12 m into the native area. Native 

species dominated the area outside the seeding (Table B9). 

Table B8. Number of plants found on transects at the Badger Pocket Road Telephone Cable Planting Site inside the area sown with crested wheatgrass in 1980. 

 6 m in 12 m in 18 m in 24 m in 30 m in 

Crested wheatgrass 22 28 24 36 33 

Astragalus spp. 0 1 1 0 0 

Bluebunch wheatgrass 0 0 0 0 0 

Bottlebrush squirreltail 0 1 2 0 2 

Cheatgrass 1 1 0 0 1 

Cusick bluegrass 0 0 0 1 1 

Fescue 0 0 1 0 0 

Lichen 0 0 0 0 5 

Needle and thread 1 0 1 0 3 

Phlox spp. 1 0 0 1 0 

Rabbitbrush 0 1 0 0 0 

Sandberg bluegrass 0 1 13 4 13 

Sedge 0 0 0 1 0 

Wyoming big sagebrush 13 3 4 2 4 

Bare ground 31 14 17 15 14 

Litter 31 50 37 40 24 
Improved Native Grasses and Establishment Methods 109 

Table B9. Number of plants found on transects at the Badger Pocket Road Telephone Cable Planting Site outside the area sown with crested wheatgrass in 

1980. 

 6 m out 12 m out 18 m out 24 m out 30 m out 

Measurement # 1 

Crested wheatgrass 0 1 0 0 0 

Astragalus spp. 0 0 1 2 2 

Bluebunch wheatgrass 0 5 11 4 1 

Bottlebrush squirreltail 11 5 11 6 7 

Cusick bluegrass 1 1 2 0 1 

Daisy 1 0 0 1 0 

Indian ricegrass 3 0 0 0 1 

Lichen 3 7 3 4 10 

Thurber needlegrass 8 21 21 18 15 

Phlox spp. 0 0 0 1 0 

Rabbitbrush 0 2 0 4 0 

Sandberg bluegrass 21 16 14 20 19 

Wyoming big sagebrush 7 12 1 8 7 

Bare ground 17 14 9 9 19 

Litter 28 16 27 23 18 

Measurement # 2 

Crested wheatgrass 0 0 0 0 0 

Astragalus spp. 0 0 0 2 0 

Bluebunch wheatgrass 1 1 3 0 5 

Bottlebrush squirreltail 4 6 8 7 7 

Cusick bluegrass 2 1 3 2 0 

Indian ricegrass 0 2 0 0 0 

Knapweed 0 0 1 0 0 

Lichen 15 6 7 15 8 

Miscellaneous spp. 0 2 1 0 0 

Thurber needlegrass 23 19 11 6 9 

Rabbit brush 0 0 0 1 2 

Sandberg bluegrass 12 20 12 18 26 

Wyoming big sagebrush 5 6 18 10 9 

Bare ground 8 7 5 13 11 

Litter 30 30 31 26 22 
110 ERDC/CRREL TR-03-20 

Borden Bench Site 

This site was sown with Siberian wheatgrass in 1985 and 1986 and has shallow soils and very low precipitation (130 mm annually). Siberian wheatgrass, 

Sandberg bluegrass, and bare ground dominated within the sown area (Table 

B10). We found no Siberian wheatgrass plants outside the sown area. Sandberg 

bluegrass and cheatgrass dominated the outside native area (Table B11). 

Table B10. Number of plants found on transects at the Borden Bench Site inside 

the area sown with Siberian wheatgrass in 1984. 

 6 m in 12 m in 18 m in 24 m in 30 m in 

Siberian wheatgrass 9 22 15 34 22 

Bluebunch wheatgrass 0 0 0 0 0 

Cheatgrass 3 4 4 4 0 

Knapweed 0 1 0 0 0 

Miscellaneous species 0 1 0 1 0 

Needle and thread grass 0 0 0 0 6 

Sandberg bluegrass 20 9 21 23 16 

Russian thistle 14 2 0 0 0 

Wyoming big sagebrush 0 0 1 0 1 

Bare ground 30 48 49 19 45 

Litter 14 11 9 19 10 

Rock 10 2 1 0 0 
Improved Native Grasses and Establishment Methods 111 

Table B11. Number of plants found on transects at the Borden Bench Site outside 

the area sown with Siberian wheatgrass in 1984. 

 6 m out 12 m out 18 m out 24 m out 30 m out 

Measurement # 1 

Siberian wheatgrass 0 0 0 0 0 

Bluebunch wheatgrass 0 0 0 0 0 

Bottlebrush squirreltail 0 1 0 0 0 

Cheatgrass 10 5 22 24 36 

Daisy 3 0 0 0 0 

Sandberg bluegrass 20 26 17 15 11 

Mustard 0 0 1 0 0 

Russian thistle 1 2 5 3 2 

Stiff sage 0 1 0 0 0 

Bare ground 45 42 40 34 10 

Litter 16 19 15 24 41 

Rock 5 4 0 0 0 

Measurement # 2 

Siberian wheatgrass 0 0 0 0 0 

Cheatgrass 12 14 8 8 10 

Bluebunch wheatgrass 0 0 0 0 0 

Bottlebrush squirreltail 1 0 3 0 0 

Knapweed 0 0 0 1 0 

Needle and thread grass 0 0 1 7 0 

Russian thistle 3 4 0 1 3 

Sandberg bluegrass 20 20 20 22 25 

Yarrow 0 0 0 1 0 

Bare ground 34 38 42 38 40 

Litter 28 22 21 20 19 

Rock 2 2 5 2 3 

Exit 11 Site 

We made no evaluations of spread at the Exit 11 breeding site, but we visited 

this area during the tour with the invasive review panel. During that visit, panel 

members commented that they were seeing more natives coming into the seeded 

areas than introduced seeded plants moving out into the areas of native species. 
112 ERDC/CRREL TR-03-20 

Participants in the Independent Review Panel for Non-Invasiveness of 

Introduced Species 

CRREL Participants: Antonio Palazzo, Tim Cary, Kate Taylor, and James 

Wuebben 

USDA-ARS Participants: Dr. Kay Asay, Dr. Kevin Jensen, Deane Harrison, 

Dr. Blair Waldron, and Dr. Tom Jones. 

Invitees: 

Wendy Connally, The Nature Conservancy, Yakima, Washington 

Dr. Peter Dunwiddie, The Nature Conservancy, Seattle, Washington 

Kerianne Gardner, EPA, Seattle, Washington 

Robert Holst, SERDP, Arlington, Virginia 

Larry Holzworth, USDA-NRCS, Bozeman, Montana 

Dr. Dave Huff, Pennsylvania State University, State College, Pennsylvania 

Robert Jones, U.S. Military Academy, West Point, New York 

Jeff Linn, Fort Carson, Colorado 

Pete Nissen, Yakima Training Center, Washington 

Dr. Gerald Schuman, USDA-ARS, Cheyenne, Wyoming 

Mark Stannard, USDA-NRCS, Pullman, Washington 
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October 2003 Technical Report

Improved Native Grasses and Establishment Methods for

Use on Military Training Lands

Antonio J. Palazzo, Susan E. Hardy, and Kevin B. Jensen

U.S. Army Engineer Research and Development Center

Cold Regions Research and Engineering Laboratory

72 Lyme Road ERDC/CRREL TR-03-20

Hanover, NH 03755-1290

Strategic Environmental Research and Development Program

Standard Form 298 (Rev. 8-98)

Prescribed by ANSI Std. 239.18

Ecological bridge Military lands Plant germplasms Seeding techniques

Invasive weeds Native grasses Rapid plant establishment

Land rehabilitation Native plants Revegetation

The objective of this project was to develop more wear-resistant plants and evaluate the relationships between military training and plant

injury, regrowth, and wear resistance. Through plant breeding, we were able to improve traits related to resiliency and establishment in

introduced and native species of range-land grasses. We selected for early spring growth, increased seedling vigor, improved tiller and

rhizome development after disturbance, and resistance to abiotic and biotic stresses. Our improved plant materials will be ecologically

compatible at the military sites because they were developed from collections of species native to or previously seeded at these sites. We

made advances in relating molecular markers to plant characteristics and in using DNA fingerprinting techniques to characterize genetic

diversity. We used markers to identify species and plants that can grow better at low temperatures. We now have the tools to assess the

genetic differences and similarities in commercial and natural seed sources, enabling land managers to select seed sources that will ensure

genetic compatibility with existing populations. Our tank traffic studies showed that naturalized, introduced species are more tolerant and

recover more rapidly under repeated tracking than native plants. However, two improved native species, western wheatgrass and Snake

River wheatgrass, showed promise as stabilization species because of their ability to colonize damaged areas. Our studies on what we call

“ecological bridges” confirm that we can select seed mixtures that will establish more rapidly than all-native mixes and will ultimately lead

to healthy and persistent stands of native plants. The species in the seed mixtures and the equipment needed are readily available, and the

seeding can be done in one application, thus saving money. Our improved germplasm will make these seeding mixes even more desirable.