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INTRODUCTORY

AUTHORSHIP AND CITATION ABBREVIATION SYNONYMS NRCS PLANT CODE COMMON NAMES TAXONOMY LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
Photos by:	Bill Cook, MI State Univ., www.forestryimages.org. Eastern cottonwood.	W.L. Wagner @ USDA-NRCS PLANTS Database. Rio Grande cottonwood.

AUTHORSHIP AND CITATION:
Taylor, Jennifer L. 2001. Populus deltoides. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/tree/popdel/all.html [].

ABBREVIATION:
POPDEL
POPDELD
POPDELM
POPDELW

SYNONYMS:
Populus deltoides var. occidentalis Rydb.
   = P. d. ssp. monilifera (Ait.) Eckenwalder [126,128]
P. fremontii var. wislizeni S. Wats
   = P. d. ssp. wislizeni (S. Wats.) Dorn [56,126,128,170]
P. monilifera Ait.
   = P. d. ssp. monilifera (Ait.) Eckenwalder [128]
P. sargentii Dode
   = P. d. ssp. monilifera (Ait.) Eckenwalder [88,126,128]
P. wislizeni (S. Wats.) Sarg.
   = P. d. ssp. wislizeni (S. Wats.) Dorn [56,126,128,170]

NRCS PLANT CODE [211]:
PODE3
PODED
PODEM
PODEW

COMMON NAMES:
eastern cottonwood
plains cottonwood
Rio Grande cottonwood
plains poplar

TAXONOMY:
The currently accepted name of eastern cottonwood is Populus deltoides Bartr. ex Marsh. (Salicaceae) [56,128]. It is in the Aigeiros section of Populus [86]. Recognized infrataxa are as follows:

Populus deltoides ssp. deltoides Bartr. ex Marsh.  eastern cottonwood 
Populus deltoides ssp. monilifera (Ait.) Eckenwalder    plains cottonwood [48,126,128]
Populus deltoides ssp. wislizeni (S. Wats.) Eckenwalder   Rio Grande cottonwood [48,56,126,128]

In this species summary eastern cottonwood refers to the species (Populus deltoides) unless otherwise noted as P. d. ssp. deltoides. Plains cottonwood and Rio Grande cottonwood refer to P. d. ssp. monilifera and P. d. ssp. wislizeni, respectively.

Eastern cottonwood produces several hybrids:

Carolina poplar (P. × canadensis Moench); from hybridization with Lombardy cottonwood (P. nigra) [56,57,88]
lanceleaf cottonwood (P. × acuminata Rydb.); afrom hybridization with narrowleaf cottonwood (P. angustifolia) [47,88]
balm-of-Gilead (P. × jackii Sarg.); from hybridization with balsam poplar (P. balsamifera) [47,57,65,88]

LIFE FORM:
Tree

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
In Texas, the cottonwood-tallgrass series is listed as "imperiled globally, very rare, 6 to 20 occurrences (endangered throughout range); imperiled in Texas, very rare, vulnerable to extirpation, 6 to 20 occurrences" [206].

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• ECOSYSTEMS
• STATES
• BLM PHYSIOGRAPHIC REGIONS
• KUCHLER PLANT ASSOCIATIONS
• SAF COVER TYPES
• SRM (RANGELAND) COVER TYPES
• HABITAT TYPES AND PLANT COMMUNITIES

MANAGEMENT CONSIDERATIONS

• WOOD PRODUCTS VALUE
• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• PALATABILITY
• NUTRITIONAL VALUE
• COVER VALUE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES AND VALUES
• OTHER MANAGEMENT CONSIDERATIONS

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

GENERAL BOTANICAL CHARACTERISTICS RAUNKIAER LIFE FORM REGENERATION PROCESSES SITE CHARACTERISTICS SUCCESSIONAL STATUS SEASONAL DEVELOPMENT
Plains cottonwood photo by:	J.S. Peterson@USDA-NRCS PLANTS Database

GENERAL BOTANICAL CHARACTERISTICS:
Eastern cottonwood is a native, deciduous bottomland hardwood [68,121,132,212,225]. Height ranges from 36 to 190 feet (11-57.9 m) [37,47,56,67,120,132,150,151,224]. At maturity (approximately 35 years) [132], diameter at breast height ranges from 10.7 inches to more than 6 feet (27.2-182.9 cm) [7,47,67,132,150,224]. In open areas, eastern cottonwood typically has a large trunk that divides into branches near its base and ascends to form a wide, spreading crown [47,100]. In closed stands, it tends to have a tall, straight, and relatively branch-free bole with a small rounded crown [9]. Life expectancy is approximately 100 to 200 years [9,120,142]. It is dioecious. Female catkins range from 2 to 5.1 inches (5-13 cm) long, and fruit capsules are 0.3 to 0.6 inch (.8-1.5 cm) long [56]. The bark is thick and deeply furrowed with wide, flat ridges [56,199]. The rooting depth averages 100 inches (254 cm) [97], and mature stands can reach 117.6 to 196.8 inches (298.7-499.9 cm) rooting depth [22]. 

Eastern cottonwood is drought tolerant [47,77]. It has been classed as moderately tolerant to water-logged soils [106,110] and is tolerant of short-term inundation [145]. Eastern cottonwood tolerates periodic flooding from January through April, but mortality and growth due to flooding depend on how many events per year, season of year, age class, duration, and depth [89,106,149]. A study using cuttings found that eastern cottonwood survived less than 16 days of complete submergence [109]. Roots die when soaked for more than 1 month and adventitious roots form from dormant buds in the main trunk [110,220].

Plains cottonwood is a quick growing, short-lived deciduous tree [4]. It is smaller than P. d. ssp. deltoides [173] and grows from 10 to 98.4 feet (3-30 m) [4,36,58,124,130,139,173,195,207,223]. It has a diameter at breast height of 5 to 78.7 inches (12.7-200 cm) [4,58,107,124,130,139,173,207]. It is considered the fastest-growing tree in the Great Plains. The life expectancy is about 90 years [173].

Staminate catkins of plains cottonwood range in size from 2 to 3.5 inches (5-9 cm), pistillate catkins are 5.9 to 7.9 inches (15-20 cm) long, and fruits are 0.4 inch (1 cm) in length [173]. Plains cottonwood is susceptible to drought except where water tables are high [4] and drought induced mortality can be high [173]. The average rooting depth is 10 feet (3.1 m), and the longest roots can reach 75 feet (22.9 m) [231].

Rio Grande cottonwood ranges in height from 26.2 to 65.6 feet (8-20 m) [56,59]. It has catkins with pedicels 0.2 to 0.6 inch (.5-1.5 cm) long [56].

RAUNKIAER  LIFE FORM [172]:
Phanerophyte

REGENERATION PROCESSES:
Eastern cottonwood regenerates sexually and vegetatively.

Sexual reproduction: Large seed crops (25 to 28 million seeds/tree/year) [9,22,129] dispersed by wind and water over long distances, are generally produced annually once the trees reach 10 to 15 years of age [9,22,47,132,142,158,173,221,225].

There is no seed dormancy in eastern cottonwood [227,228]. Germination occurs as soon as seeds arrive on a seedbed that is moist, free of vegetation, and in full sunlight [9,23,67,132,173,181]. Seeds are highly viable at dispersal [66,124], but viability decreases rapidly in the absence of a suitable germination environment [66]. Under natural conditions, seeds remain viable for 1 to 2 weeks [22]. Suitable recruitment sites occur naturally as a result of spring flooding [156,173] and scouring by ice [9]. Receding floodwaters leave freshly deposited, exposed alluvium, and seed germination along prairie river floodplains often occurs exclusively on these sites [124,229]. Exposed soil is essential, as young seedlings do not compete well with overtopping vegetation [34,124,171].

Vegetative reproduction: Cottonwood species (Populus spp.) reproduce vegetatively by sprouting from stumps and root crowns, and by the formation of suckers (adventitious shoots on roots ) [47,176,229]. There is asexual reproduction from broken limbs (flood training) and crown breakage [22]. The ability of cottonwoods to sprout declines with age [173].  Cottonwoods in the Aigeiros section of Populus, including eastern cottonwood, do not sprout as readily as cottonwoods in section Tacamahaca. Eastern cottonwood sprouts from the roots and bole after top-kill or damage, but the response is weak [22,86]. Most  suckers arise from suppressed buds embedded in the periderm of undisturbed roots after death or injury of aboveground parts. There is disagreement on the ability or eastern cottonwood to sprout from the bole after being cut [47,154].

Plains cottonwood does not readily form suckers or stem sprouts [87,132], and sprouting is uncommon except in flood-trained shoots [23,86,179]. 

Growth: Eastern cottonwood is the fastest growing native tree in North America [28,47,121,132,212]. It commonly increases 0.7 to 1 inch  (1.69-2.54 cm) in diameter and 5 feet (1.5 m) in height annually up to 10 to 15 years of age, and grows at only a slightly slower rate up to 30 to 35 years of age [28]. In the Mississippi valley, eastern cottonwood can reach 20 inches (50.8 cm) in diameter at breast height and 120 feet (36.6 m) in height at 35 years [28,145].

Growth of young seedlings is rapid [132]. The first 3 weeks of development of seedlings are slow, after which the growth rate rapidly accelerates [145]. Root growth of seedlings is also rapid. Seedlings have been observed to extend taproots 12 to 16 inches (30.5-40.6 cm) and lateral roots 24 inches (61 cm) by the end of their first growing season [222]. These roots serve as anchorage during floods and help to ensure a supply of water during dry periods.

Once established, seedlings are susceptible to damage from flooding. In one instance, 8 days of complete inundation weakened all seedlings, while 16 days resulted in their mortality [112]. Seedlings can survive if flooded less than 50% of the growing season [158]. Inundation does not necessarily have a detrimental effect upon germination [108].

SITE CHARACTERISTICS:
Eastern cottonwood primarily grows on the moist alluvial soil of floodplains and bottomlands [132,145,158,171,229]. It is also found in ravines [16,107,229], along disturbed streams [117], and in low spots of sandy uplands with a high water table [229]. It also found on batture lands, the unprotected areas between the Mississippi River and its levees [145].

Plains cottonwood is found on floodplains and small sandbars in the river beds or large bends where stream flow is dramatically retarded during high water. It is found next to springs that flow long enough to form ponds [4,130,173]. 

Across its range, eastern cottonwood is found from 255 to 6,500 feet (78-1,981 m) [10,12,37,165,169,187], usually 15 to 40 feet (5-12 m) above stream level [90,151,187]. Elevation ranges for plains and Rio Grande cottonwood are given in the following table:

Plains cottonwood		References
CO	3,500 to 6,500 feet (1,067-1,981 m)	[49,163,180]
MT	3,200 to 4,921 feet (975-1,500 m)	[49,208]
NM	5,000 to 6,500 feet (1,524-1,981 m)	[147]
UT	4,500 feet (1,372 m)	[49]
WY	3,500 to 9,000 feet (1,067-2,743 m)	[49,69,162]
Rio Grande cottonwood
AZ	4,000 to 6,500 feet (1,219-1,981 m)	[56]
CO	4,000 to 7,000 feet (1,219-2,134 m)	[49]
NM	1,457 to 6,500 feet (444-1,981 m)	[113,147]
TX	2,800 to 5,000 feet (853-1,524 m)	[49]

The average annual precipitation across the eastern cottonwood range is 13.78 to 55 inches (350-1,397 mm) [125,169]. Average annual precipitation ranges for some states where eastern cottonwood occurs is listed below:

IN	35 to 47 inches (889-1,193.8 mm)	[141]
MI	33.86 inches (860 mm)	[71]
OH	34 inches (863.6 mm)	[18]
TX	14.96 to 33.07 inches (380-840 mm)	[213,214]
ON	37 inches (939.8 mm)	[234]

Eastern cottonwood occurs in a wide range of temperature regimes with extremes ranging from -49 degrees Fahrenheit (-45 ^oC) in winter to 114.8 degrees Fahrenheit (46 ^oC) in summer [125]. The mean annual temperature or annual temperature range for some states is as follows:

MI	48.2 degrees Fahrenheit (9 oC)	[71]
OH	14.5 to 83.8 degrees Fahrenheit (-9.72-28.77 oC)	[18]
TX	59.9 to 67 degrees Fahrenheit (15.5-19.44 oC)	[213,214]
ON	46.4 degrees Fahrenheit (8 oC)	[234]

The annual precipitation varies between 12 and 30 inches (305-762 mm), averaging less than 20 inches (508 mm) across most of the plains cottonwood range. The temperature can range between -50 degrees Fahrenheit (-45.56 ^oC) in January and 115 degrees Fahrenheit in July (46.11 ^oC) [173]. The average annual precipitation and temperature for some states in the plains cottonwood range are given in the table below [4,11,15,20,36,73,82,104,105,124,127,130,146,162,175,182,186,198,208,221,231]:

IN	35 to 47 inches (889-1,193.8 mm)	[141]
MI	33.86 inches (860 mm)	[71]
OH	34 inches (863.6 mm)	[18]
TX	14.96 to 33.07 inches (380-840 mm)	[213,214]
ON	37 inches (939.8 mm)	[234]

Eastern cottonwood tolerates a wide range of soils ranging from coarse sands to clays but grows best on moist, well-drained fine sandy loams or silt [47,90,129,145,151,154,158].

Plains cottonwood is found on sandy soils [124,130] or well-drained soils with a high water table to supply year-round moisture [4]. Availability of moisture is reportedly more significant to plains cottonwood than soil texture or fertility [173].

SUCCESSIONAL STATUS:
Eastern cottonwood is shade intolerant [9,47,151,171,217,225] and a pioneer species that typically establishes on freshly exposed alluvium of sandbars, streambanks, and other floodplain sites [16,23,103,117,124,150,222]. Establishment and dominance may also occur after sandbar willows have stabilized the site [229]. Eastern cottonwood invades unburned prairies in Kansas [1] and old fields and upland sites in the lake states [47,111].

Maintenance of seral, eastern cottonwood-dominated communities depends on either periodic flooding or timber harvest [23,92,155,158]. In the Great Plains, dams and reservoirs which alter the magnitude and frequency of floods, sedimentation rates, and stream meander migration rates have detrimentally affected cottonwood communities by reducing seedling recruitment and survival [23]. In the absence of flooding, succession proceeds, and the shade-intolerant cottonwoods are eventually replaced by more shade-tolerant species [155]. Eastern cottonwood stands are replaced by American elm, sycamore, pecan, sugarberry, boxelder, and sweetgum [103]. The eastern cottonwood/Rocky Mountain juniper community type is an early successional stage in North Dakota. It succeeds to the green ash/western snowberry community if left undisturbed [83].

Plains cottonwood with willow and boxelder represents an early stage on floodplains in Nebraska [4].  The plains cottonwood-willow type progresses to the plains cottonwood-green ash type in North Dakota [124]. It contributes significantly to dune formation on Lake Michigan [38].

SEASONAL DEVELOPMENT:
Time of flowering and leaf emergence for eastern cottonwood varies according to geographic location. Eastern cottonwood flowers during early to late spring, generally from February through May [22,47,68,147,228] and 1 to 2 weeks before leaf initiation [22]. Seed dispersal occurs in early summer [47].  Seed dispersal and germination generally occur in late spring to early- or mid-summer and typically coincide with decreasing flow levels (May through August) [9,22,40,66,181]. Phenological dates for some states and the Great Plains are given below:

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

FIRE REGIMES: Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

Populus deltoides: References

1. Abrams, Marc D.; Gibson, David J. 1991. Effects of fire exclusion on tallgrass prairie and gallery forest communities in eastern Kansas. In: Nodvin, Stephen C.; Waldrop, Thomas A., eds. Fire and the environment: ecological and cultural perspectives: Proceedings of an international symposium; 1990 March 20-24; Knoxville, TN. Gen. Tech. Rep. SE-69. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 3-10. [16627]

2. Adams, Dwight E.; Anderson, Roger C. 1980. Species response to a moisture gradient in central Illinois forests. American Journal of Botany. 67(3): 381-392. [13295]

3. Adams, Dwight E.; Anderson, Roger C.; Collins, Scott L. 1982. Differential response of woody and herbaceous species to summer and winter burning in an Oklahoma grassland. The Southwestern Naturalist. 27: 55-61. [6282]

4. Albertson, F. W.; Weaver, J. E. 1945. Injury and death or recovery of trees in prairie climate. Ecological Monographs. 15: 393-433. [4328]

5. Arno, Stephen F. 1980. Forest fire history in the Northern Rockies. Journal of Forestry. 78(8): 460-465. [11990]

6. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]

7. Bechtold, William A.; Brown, Mark J.; Sheffield, Raymond M. 1990. Florida's Forests, 1987. Res. Bull. SE-110. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station, Forest Inventory and Analysis. 83 p. [14579]

8. Begin, Yves. 1990. The effects of shoreline transgression on woody plants, upper St. Lawrence estuary, Quebec. Journal of Coastal Research. 6(4): 815-827. [17681]

9. Behan, Mark. 1981. The Missouri's stately cottonwoods: How can we save them? Montana Magazine. September: 76-77. [6284]

10. Bell, David T. 1974. Tree stratum composition and distribution in the streamside forest. The American Midland Naturalist. 92(1): 35-46. [10410]

11. Bell, Elbert L.; Hulbert, Lloyd C. 1974. Effect of soil on occurrence of cross timbers and prairie in southern Kansas. Transactions of the Kansas Academy of Science. 77(4): 203-209. [24441]

12. Bellah, R. Glenn; Hulbert, Lloyd C. 1974. Forest succession on the Republican River floodplain in Clay County, Kansas. The Southwestern Naturalist. 19(2): 155-166. [241]

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15. Bjugstad, Ardell J. 1977. Reestablishment of woody plants on mine spoils and management of mine water impoundments: an overview of Forest Service research on the northern High Plains. In: Wright, R. A., ed. The reclamation of disturbed lands. Albuquerque, NM: University of New Mexico Press: 3-12. [4238]

16. Bjugstad, Ardell J.; Girard, Michele. 1984. Wooded draws in rangelands of the northern Great Plains. In: Henderson, F. R., ed. Guidelines for increasing wildlife on farms and ranches: With ideas for supplemental income sources for rural families. Manhattan, KS: Kansas State University, Cooperative Extension Service; Great Plains Agricultural Council, Wildlife Resources Committee: 27B-36B. [4239]

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22. Braatne, Jeffrey H.; Rood, Stewart B.; Heilman, Paul E. 1996. Life history, ecology, and conservation of riparian cottonwoods in North America. In: Steller, R. F., ed. Biology of Populus and its implications for management and conservation. Ottawa, ON: National Research Council of Canada, NRC Research Press: 57-85. [29693]

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28. Bull, Henry; Muntz, H. H. 1943. Planting cottonwood on bottomlands. Bulletin 391. Mississippi State, MS: Mississippi State College, Agricultural Experiment Station. 18 p. [7180]

29. Busch, David E.; Smith, Stanley D. 1993. Effects of fire on water salinity relations of riparian woody taxa. Oecologia. 94: 186-194. [22770]

30. Bush, J. K.; Van Auken, O. W. 1984. Woody species composition of the upper San Antonio River gallery forest. Texas Journal of Science. 36(2&3): 139-148. [12481]

31. Calabrese, Diane M. 1993. A geography of state trees. American Forests. 99(3&4): 34-37. [21053]

32. Carey, Andrew B.; Gill, John D. 1980. Firewood and wildlife. Res. Note 299. Broomall, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 5 p. [9925]

33. Carlson, Jack R. 1992. Selection, production, and use of riparian plant materials for the western United States. In: Landis, Thomas D., technical coordinator. Proceedings, Intermountain Forest Nursery Association; 1991 August 12-16; Park City, UT. Gen. Tech. Rep. RM-211. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 55-67. [20926]

34. Carter, Mason C.; White, E. H. 1971. The necessity for intensive cultural treatment in cottonwood plantations. Circular 189. Auburn, AL: Alabama Agricultural Experiment Station. 11 p. [7177]

35. Cassel, J. Frank; Wiehe, John M. 1980. Uses of shelterbelts by birds. In: DeGraaf, Richard M., technical coordinator. Management of western forests and grasslands for nongame birds: Workshop proceedings; 1980 February 11-14; Salt Lake City, UT. Gen. Tech. Rep. INT-86. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 78-87. [17899]

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37. Cote, Benoit; Dawson, Jeffery O. 1991. Autumnal allocation of phosphorus in black alder, eastern cottonwood, and white basswood. Canadian Journal of Forest Research. 21: 217-221. [14142]

38. Cowles, Henry Chandler. 1899. The ecological relations of the vegetation on the sand dunes of Lake Michigan. Botanical Gazette. 27(2-3): 95-117, 167-202. [11091]

39. Cox, Mike K.; Franklin, William L. 1989. Terrestrial vertebrates of Scotts Bluff National Monument, Nebraska. The Great Basin Naturalist. 49(4): 597-613. [11004]

40. Crouch, Glenn L. 1979. Long-term changes in cottonwoods on a grazed and an ungrazed plains bottomland in northeastern Colorado. Res. Note RM-370. Fort Collins, CO: U.S. Department of of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 4 p. [3496]

41. Cunningham, Richard A. 1975. Provisional tree and shrub seed zones for the Great Plains. Res. Pap. RM-150. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 15 p. [4516]

42. Curtis, John T. 1959. The vegetation of Wisconsin. Madison, WI: The University of Wisconsin Press. 657 p. [7116]

43. Dansereau, Pierre. 1959. Phytogeographia Laurentiana. II. The principal plant associations of the Saint Lawrence Valley. Contributions of the Botanical Institute No. 75. Montreal, PQ: University of Montreal, Botanical Institute. 147 p. [8925]

44. Davidson, A. G.; Prentice, R. M. 1968. Insects and diseases. In: Maini, J. S.; Cayford, J. H., eds. Growth and utilization of poplars in Canada. Departmental Publication No. 1205. Ottawa, ON: Department of Forestry and Rural Development: 116-144. [6505]

45. DeBell, D. S.; Mallonee, E. H.; Alford, L. T. 1975. Effect of nitrogen fertilizer on growth, form, and wood quality of eastern cottonwood. Forest Research Note No. 4. Camas, WA: Crown Zellerbach Corporation. 6 p. [3294]

46. Diamond, David D.; Riskind, David H.; Orzell, Steve L. 1987. A framework for plant community classification and conservation in Texas. Texas Journal of Science. 39(3): 203-221. [24968]

47. Dickmann, Donald I.; Stuart, Katherine W. 1983. The culture of poplars in eastern North America. East Lansing, MI: Michigan State University, Department of Forestry. 168 p. [6317]

48. Diggs, George M., Jr.; Lipscomb, Barney L.; O'Kennon, Robert J. 1999. Illustrated flora of northcentral Texas. Sida Botanical Miscellany No. 16. Fort Worth, TX: Botanical Research Institute of Texas. 1626 p. [35698]

49. Dittberner, Phillip L.; Olson, Michael R. 1983. The plant information network (PIN) data base: Colorado, Montana, North Dakota, Utah, and Wyoming. FWS/OBS-83/86. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 786 p. [806]

50. Dodd, M. B.; Lauenroth, W. K.; Welker, J. 1998. Differential water resource use by herbaceous and woody plant life-forms in a shortgrass steppe community. Oecologia. 117(4): 504-512. [34950]

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