Lesquerella pruinosa - Greene
Frosty Bladderpod
Other English Common Names: Pagosa Springs Bladderpod
Other Common Names: Pagosa Springs bladderpod
Synonym(s): Physaria pruinosa (Greene) O'Kane & Al-Shehbaz
Taxonomic Status: Accepted
Related ITIS Name(s): Lesquerella pruinosa Greene (TSN 23220)
Unique Identifier: ELEMENT_GLOBAL.2.159251
Element Code: PDBRA1N1D0
Informal Taxonomy: Plants, Vascular - Flowering Plants - Mustard Family
Image 12101

Public Domain

 
Kingdom Phylum Class Order Family Genus
Plantae Anthophyta Dicotyledoneae Capparales Brassicaceae Lesquerella
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Lesquerella pruinosa
Conservation Status
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NatureServe Status

Global Status: G2
Global Status Last Reviewed: 25May2006
Global Status Last Changed: 17Jul1986
Rounded Global Status: G2 - Imperiled
Reasons: Endemic to Colorado and New Mexico; known from Archuleta County, and the extreme southern portion of Hinsdale County in Colorado, and one location in Rio Arriba County in New Mexico. Habitat destruction is the biggest threat to Lesquerella pruinosa, especially considering its limited range. Residential growth and development around the city of Pagosa Springs, including a proposed ski resort, threaten occurrences.
Nation: United States
National Status: N2

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Colorado (S2), New Mexico (SNR)

Other Statuses

NatureServe Global Conservation Status Factors

Range Extent Comments: Known from southern Colorado (Archuleta County, and the extreme southern portion of Hinsdale County) and northern New Mexico (Rio Arriba County). Estimated range is at least 588 square kilometers (227 square miles), calculated in 2008 by the Colorado Natural Heritage Program in GIS by drawing a minimum convex polygon around the Colorado occurrences. The New Mexico location needs to be added to this range calculation.

Area of Occupancy: 6-25 4-km2 grid cells
Area of Occupancy Comments: The total occupied habitat is about 1,880 acres. Occurrences without specific information on occupied habitat were considered to occupy 0.5 acre.

Number of Occurrences: 6 - 20
Number of Occurrences Comments: There are 17 principal occurrences documented in the Colorado Natural Heritage Program database. Two of the 17 occurrences have not been observed in over 20 years. The USFS Conservation Assessment (Anderson 2006) reports that the species is known from 22 occurrences, including one newly discovered occurrence in northern Rio Arriba County, New Mexico.

Population Size Comments: Total estimated sum of individuals from the 17 occurrences in the CNHP database is 12,854. Anderson (2006) estimates that the total population of L. pruinosa is between 5,209 and 20,619.

Number of Occurrences with Good Viability/Integrity: Few (4-12)
Viability/Integrity Comments: There are 9 occurrences in Colorado with an A or B rank.

Overall Threat Impact: High
Overall Threat Impact Comments: Residential and commercial developments are considered to be the primary threats to the species at this time (Rondeau et al. 2011). The City of Pagosa Springs lies within the plant's range. Residential growth, development of resort homes, and increased tourism due to the development of a proposed ski resort 20 miles east of Pagosa Springs threaten occurrences. Archuleta County is one of the fastest growing counties in the United States, and future land use plans that have been drafted by Archuleta County do not include adequate provisions for the protection of this species (Anderson 2006).

Populations in the Piedra Valley may be vulnerable to the effects of livestock grazing. Though cattle pose a minimum threat of grazing the plants (they contain chemicals which render the plants unpalatable), cattle grazing tends to promote erosion and up-rooting where plants occur on slopes by severely disturbing the soil (Anderson 1988). These populations are at risk mainly where cattle trails traverse the shale barrens (Neely 1990).

This species is also threatened by off-road vehicle recreation, other recreational activities, energy resource development, exotic species invasions, use of herbicides and pesticides for weed management and range improvement, effects of small population size, prairie dog herbivory, fire, global climate change, and pollution (Anderson 2006).

Short-term Trend: Unknown
Short-term Trend Comments: Unknown

Long-term Trend: Unknown
Long-term Trend Comments: Unknown

Other NatureServe Conservation Status Information

Distribution
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Global Range: Known from southern Colorado (Archuleta County, and the extreme southern portion of Hinsdale County) and northern New Mexico (Rio Arriba County). Estimated range is at least 588 square kilometers (227 square miles), calculated in 2008 by the Colorado Natural Heritage Program in GIS by drawing a minimum convex polygon around the Colorado occurrences. The New Mexico location needs to be added to this range calculation.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map

U.S. & Canada State/Province Distribution
United States CO, NM

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
CO Archuleta (08007), Hinsdale (08053)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
14 Upper San Juan (14080101)+, Piedra (14080102)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A perennial herb with a flowering stem, 1-2.5 dm tall, from basal rosette of leaves. A dense covering of gray hairs gives the leaves and stem a frosted appearance. Flowers are yellow and bloom in May and June.
Technical Description: (The following information is largely from Rollins and Shaw 1973) The stems, 10 to 20 cm long, can be decumbent or erect and arise out of a simple or woody caudex. The basal leaves are 4-8 cm long with suborbicular or obovate blades which can be entire to sinuate or shallowly toothed. The cauline leaves, 0.8-2.3 cm long, are obovate to rhombic and can be entire or shallowly toothed.

The flowers are small and yellow with petals spatulate and expanded at the base. The fruiting inflorescence (infructescences) are dense and elongated. The pedicels are 8-11 mm long and sigmoid. The siliques, 6-9 mm long, are sessile or substipitate and globose to ellipsoid. The seeds are not winged or margined.

Diagnostic Characteristics: Rosettes formed from the abundant basal leaves help to characterize the plants.

LESQUERELLA PRUINOSA is related to and closely resembles L. PINETORUM which is found in similar habitat in New Mexico and Arizona. These species show similarities in the large basal leaves and the elongated infructescences arising from the sigmoid pedicels. The distinctive characteristics of L. PRUINOSA which separate it from L. PINETORUM are the smaller foliar trichomes and the broader basal leaves.

Duration: PERENNIAL, Long-lived
Reproduction Comments: Although little is known about its reproduction, L. PRUINOSA is probably self-incompatible and out-crossing which is consistent with the genus (Anderson 1988, Rollins 1983). Bees and flies are the most likely pollinators (Anderson 1988).

Plants begin to flower by mid-May with fruiting time depending on the elevation. Plants at lower elevations are in late stages of fruiting by the beginning of June, whereas plants at higher elevations are only in early stages of fruiting at that time (Anderson 1988).

Ecology Comments: Seed production does not appear to be a factor limiting populations of this species (Anderson 1988). Based on field observations that noted the presence of old rosettes on the plants, L. PRUINOSA appears to be a long-lived perennial (Anderson 1988). L. PRUINOSA has a long taproot which is adaptive to the erodible Mancos Shale soils (Colorado Native Plant Society 1989). Its limited range suggests that L. PRUINOSA is a relictual species that is disjunct from its closest relatives in New Mexico and Arizona. The plants occur in colonies ranging in size from one to ten acres (Anderson 1988).
Terrestrial Habitat(s): Barrens, Forest/Woodland, Grassland/herbaceous, Savanna, Woodland - Mixed
Habitat Comments: Lesquerella pruinosa is limited to soils derived from Mancos Shale. It is found at elevations ranging from 2,095 to 2,290 m (6810 ft.-7440 ft.). The highest densities tend to be in open clay barrens surrounded by montane grasslands (Anderson 1988). Smaller populations are found in open ponderosa pine (Pinus ponderosa) stands and gambel oak (Quercus gambellii) communities with the numbers apparently decreasing when plants become part of the forest understory (Anderson 1988).

In association with the ponderosa/gambel oak communities, L. pruinosa acts as a climax species, whereas it tends to be a pioneer species on raw shale (Anderson 1988). This is consistent with the report that another rare bladderpod (L. lyrata) from Alabama requires periodic disturbance to sustain the open habitat that it needs (Anonymous 1990). L. pruinosa can also be associated with Douglas fir (Pseudotsuga menziesii) and Englemann Spruce (Picea eugelmanii) communities at the upper limits of its range (Anderson 1988). Jakubos (1985) notes that dominant and subdominant species varied widely between sites, as did species representation at each site. Jakubos notes further study is needed on the effect of forb and grass cover on L. pruinosa densities.

L. pruinosa tended to grow near the edge of the clones of Quercus gambellii, but not in the middle of the clones. Q. gambellii leaf litter did not seem to inhibit the growth of the plants (Jakubos 1985). There was no correlation between aspect and L. pruinosa density. Further study is needed to determine how environmental factors such as aspect, soil pH and chemistry are involved in the distribution of L. pruinosa.

Economic Attributes
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Economically Important Genus: Y
Management Summary
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Stewardship Overview: Several populations should be monitored to determine long-term population trends. Specific biological parameters need to be studied in order to better understand the biology and ecology of the species. The impact of domestic livestock and elk grazing need to be understood. Special designation for populations on U.S. Forest Service ground needs to be pursued.
Restoration Potential: Even though the species' distribution is limited, the existing populations appear to be in good condition.
Preserve Selection & Design Considerations: Element occurrence requires modest acreage protected (low 100's of acres) at each site to encompass existing known populations and provide adequate buffer. A modest amount of buffer is needed to protect against "off site" threats of home construction, road building, and trespass from ORV use. Primary boundary should include all of the plants, with secondary boundary extending to the tops of local ridges which might extend several hundreds of yards beyond the primary boundary. The idea is to protect the local watershed from direct effects of soil erosion. It is probable that the area would need to be fenced to identify area boundaries and be able to manage livestock and to discourage ORV use which is a potential threat.

"Populations" of L. PRUINOSA contain thousands of plants so the adverse effects of inbreeding and demographic stochasticity are not likely to be a problem.

The Piedra Valley Ranch, located 15 miles north of Pagosa Springs, contains significant populations of L. PRUINOSA. The U.S. Forest Service has purchased 2,200 acres of the Ranch with monies appropriated by Congress from the Land and Water Conservation Fund. Another 660 acres may be donated to the Forest Service by the Conservancy and would be managed to protect L. PRUINOSA. A 340-acre portion of the ranch, containing excellent L.PRUINOSA habitat, was sold to a private party. In this case, a conservation easement was donated by the landowners to The Nature Conservancy which gives the Conservancy exclusive management of the L. PRUINOSA habitat area within the property.

Because many of the populations of L. PRUINOSA are on private lands, they receive little or no management or protection (Anderson 1988). In 1988, only ten percent of the L. PRUINOSA plants are on Federal land (Anderson 1988). Forest Service acquisition of portions of the Piedra Valley Ranch and designation of appropriate habitat as a "Special Botanical Area" would facilitate protection of the species.

L. PRUINOSA seems to prefer an intermediate level of disturbance of its habitat. The exact amount and type of disturbance which is best for the plant is unknown. The plants do well on relatively bare Mancos shale slopes that are naturally erosive and which may provide an appropriate level of disturbance for the plants (Carpenter 1992). The plants are not found in dense turf and are less evident near creeks where soils are relatively moist and well vegetated. Cattle have been browsing the Lind site at Piedra Valley for decades and the plants seem to be doing well. The long term effects of cattle grazing are unknown. At least some of the L. PRUINOSA occurrences experience heavy use by elk during the winter. Effects of elk grazing and trampling are unknown (Carpenter 1992).

The Conservancy will work with the U.S. Forest Service to explore various ways of protecting L. PRUINOSA habitat, particularly using Special Interest Area designation.

Management Requirements: The relationships between L. PRUINOSA and livestock and wild ungulate (mainly elk) grazing are not known. L. PRUINOSA does not appear to be eaten readily by cattle or elk (A. Carpenter, personnel observation). It is unknown if trampling by elk and/or cattle directly affects L. PRUINOSA plants. It is unknown if indirect effects of elk or cattle grazing, such as increased soil erosion, soil fertilization or reduction of adjacent vegetation are significant for L. PRUINOSA.

The area inhabited by L. PRUINOSA has been grazed by livestock for decades. The continued existence of seemingly healthy populations of L. PRUINOSA suggests that historic patterns of livestock and wild animal grazing have at least not been highly detrimental to the plants. However, it is possible that grazing generally has modest yet important adverse effects on this species or that certain grazing management practices are detrimental.

It is also unknown to what extent off-road vehicle use might affect L. PRUINOSA plants. Given that these plants usually grow on highly-erodible soils on steep to moderate slopes, is reasonable to expect that ORV use would be detrimental to L. PRUINOSA.

If populations near Pagosa Springs appear to be deteriorating due to development, altered management practices could be implemented, assuming cooperation of the landowners.

The U.S. Forest Service should continue to consider L. PRUINOSA in its environmental analysis. In conjunction with this, the Nature Conservancy will propose that the appropriate areas of the Piedra Valley Ranch acquired by the Forest Service be designated as Special Botanical Areas.

Monitoring Requirements: Plant longevity, seed production, seed bank, annual growth rate, and annual recruitment of seedlings are some of the biological parameters that need to be considered.

To determine the extent of damage by continued development, several populations situated in the outlying residential development area near Pagosa Springs could be monitored periodically. Monitoring is needed to learn more about the life history and population dynamics of the species.

Monitoring would utilize a population approach. Several hundred individual plants in each population would be permanently marked (numbered aluminum tags work well). Marked plants would be visited once annually to determine the plant size, the number of fruits (or some other index of reproductive output), and the mortality rate. In addition, small permanent plots would be set up to monitor seedling establishment. Viability of seeds in the soil would be estimated using small buried bags containing known numbers of live seeds. Data from the monitoring could be used to predict the long-term viability of each monitored population.

Monitoring Programs: No monitoring programs are presently being implemented; one will be implemented by staff from The Nature Conservancy's Colorado Field Office on the Lynd site and possibly on other nearby sites.
Management Research Programs: No research programs are presently under way for L. PRUINOSA.
Management Research Needs: Population-level monitoring could be initiated to investigate the rate of change among the life history stages of seeds, seedlings, juveniles and reproductive individuals. Monitoring could provide field data for the transition probabilities between the above stages. These data could be used with a computer simulation model of a L. PRUINOSA population(s) to determine the life history stage which is most critical to maintaining the population. Suitable computer models are now available, e.g., the RAMAS software by Scott Ferson of Applied Biomathematics. Furthermore, models can be used to assess quantitatively the likelihood that a population will persist over a period of time, e.g., 100 years.

An initial question is how long these plants live. A study of the longevity of L. PRUINOSA would be helpful. If the species has a short life span then studies on the level of recruitment would be important. Also, studies of pollination and seed dispersal would be in order.

This species seems to do well under the effect of intermediate disturbance. The role of disturbance in maintaining good-quality occurrences of L. PRUINOSA could be studied. In particular, is livestock grazing compatible with good-quality occurrences of L. PRUINOSA? A study could be done on lands outside the preserve containing L. PRUINOSA.

Jakubos (1985) found a consistency in slope, topographic position, and geologic substrate. Further studies are needed to determine how these environmental factors are involved in the distribution of L. PRUINOSA. Also, the pollination biology of L. PRUINOSA should be studied.

Population/Occurrence Delineation
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Minimum Criteria for an Occurrence: An element occurrence of this species is defined as any naturally occurring population that is separated by a sufficient distance or barrier from a neighboring population. Good-quality occurrences of Lesquerella pruinosa have been described by Carpenter (1992) as containing numerous plants with many seedlings. The 80-acre Lind site at Piedra Valley, Colorado contains about ten-thousand plants. Good quality occurrences can also contain large plants with numerous leaves that produce flowers and fruit and are widely dispersed across their habitat. The level of seed production in a high-quality occurrence is not known. The longevity of L pruinosa may be ten years or more. Population fluctuation is probably large, 10 percent or more over 3 or 4 years.
Separation Distance for Unsuitable Habitat: 1.61 km
Separation Distance for Suitable Habitat: 3.22 km
Separation Justification: The rationale for this large a separation distance across suitable but apparently unoccupied habitat is that it is likely additional research will find this habitat to be occupied. It can often be assumed that apparently unconnected populations will eventually be found to be more closely connected; these are best regarded as suboccurrences. No information on mobility of pollen and propagules is available on which to base the separation distance for this species.
Date: 27Sep2000
Author: Spackman, S. and D. Anderson
Population/Occurrence Viability
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Excellent Viability: Size: 1000 or more individuals (based on available EOR data). Condition: the occurrence has an excellent likelihood of long-term viability as evidenced by the presence of multiple age classes and evidence of flowering and fruiting, indicating that the reproductive mechanisms are intact. This occurrence should be in a high-quality site with less than 1% cover of exotic plant species and/or no significant anthropogenic disturbance. Landscape Context: the occurrence is surrounded by an area that is unfragmented and includes the ecological processes needed to sustain this species. This plant is well adapted to the easily eroded slopes of mancos shale on which it is found, and erosion may competitively exclude other species that may otherwise invade its habitat.
Good Viability: Size: 100 or more individuals (based on available EOR data). Condition: the occurrence should have a good likelihood of long-term viability as evidenced by the presence of multiple age classes and evidence of flowering and fruiting, indicating that the reproductive mechanisms are intact. Anthropogenic disturbance within the occurrence is minimal. If exotic species are present, they comprise less than 10% of the total ground cover. Landscape Context: the surrounding landscape should contain the ecological processes needed to sustain the occurrence but may be fragmented and/or impacted by humans.
Fair Viability: Size: 10 to 99 individuals (based on available EOR data). Condition: The occurrence may be less productive than the above situations, but is still viable, with multiple age classes and evidence of flowering and fruiting, indicating that the reproductive mechanisms are intact. The occupied habitat is somewhat degraded (exotic plant species make up between 10-50% of the total ground cover and/or there is a moderate level of anthropogenic disturbance). Landscape Context: there may be significant human disturbance, but the ecological processes needed to sustain the species are still intact.
Poor Viability: Size: Less than 10 individuals (based on available EOR data). Condition: little or no evidence of successful reproduction is observed (poor seedling recruitment, no flowering or fruiting observed, or poor age class distribution). Exotic plant species make up greater than 50% of the total ground cover, and/or there is a significant level of human disturbance. Landscape context: the surrounding area is fragmented with many ecological processes no longer intact. The occurrence has a low probability of long-term persistence due to inbreeding depression, natural stochastic events, and its intrinsic vulnerability to human impacts.
Justification: A Rank: Large populations in high quality sites are presumed to contain a high degree of genetic variability, to have a low susceptibility to the effects of inbreeding depression, and to be relatively resilient.

C Rank: EOs not meeting "C"-rank criteria are likely to have a very high probability of inbreeding depression and extirpation due to natural stochastic processes and/or occur in degraded habitat with low long-term potential for survival.

Key for Ranking Species Element Occurrences Using the Generic Approach (2008).
Date: 27Sep2000
Author: Spackman, S. and D. Anderson
Notes: COHP
U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 22May2006
NatureServe Conservation Status Factors Author: Leigh Rouse, COFO, rev. Maybury/Spackman (1996), rev. Spackman, S. and D. Anderson (2000), rev. Neuhaus, K., J. Handwerk, and S. Spackman Panjabi (2006)
Management Information Edition Date: 12May1992
Management Information Edition Author: LEIGH ROUSE, COFO
Element Ecology & Life History Edition Date: 12May1992
Element Ecology & Life History Author(s): LEIGH ROUSE, COFO

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
Help
  • Ackerfield, J. 2012. The Flora of Colorado. Colorado State University Herbarium. 433 pp.


  • Anderson, D.G. (2006, August 29). Lesquerella pruinosa Greene (Pagosa bladderpod): a technical conservation assessment. [Online]. USDA Forest Service, Rocky Mountain Region. Available: http://www.fs.fed.us/r2/projects/scp/assessments/lesquerellapruinosa.pdf [January 2007].

  • Anderson, J.L. 1988. Status report for Lesquerella pruinosa. Unpublished report prepared for the U.S. Fish and Wildlife Service, Golden, Colorado.

  • Anderson, J.L. 1988. USFWS. Status report for Lesquerella pruinosa.

  • Anonymous. 1990. Endangered Species Technical Bulletin. 15(10):11.

  • Colorado Native Plant Society. 1989. Rare Plants of Colorado. Published jointly by Rocky Mountain Nature Association and Colorado Native Plant Society, Estes Park, CO.

  • Colorado Native Plant Society. 1989. Rare plants of Colorado. Rocky Mountain Nature Association, Colorado Native Plant Society, Estes Park, Colorado. 73 pp.

  • Ecology Consultants, Inc. 1978. An illustrated guide to the proposed threatened and endangered plant species in Colorado. U.S. Fish and Wildlife Service, Lakewood, CO. 114 pp.

  • Ferson, S. 1990. RAMAS stage users manual. Applied Biomathematics, 100 N. County Rd., Setauket, NY 11733.

  • Flora of North America Editorial Committee. 2010. Flora of North America North of Mexico. Vol. 7. Magnoliophyta: Salicaceae to Brassicaceae. Oxford University Press, New York. xxii + 797 pp.

  • Heil, K.D., S.L. O'Kane Jr., L.M. Reeves, and A. Clifford, 2013. Flora of the Four Corners Region, Vascular Plants of the San Juan River Drainage; Arizona, Colorado, New Mexico, and Utah. Missouri Botanical Garden Press, St. Louis, Missouri. 1098 pp.

  • Jakubos, B. 1985. A comparison of Lesquerella pruinosa densities under varying habitat site factors. Unpublished report on file at The Nature Conservancy, Colorado Field Office, Boulder, Colorado.

  • Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.

  • Neely, B. 1990. Preserve design for Piedra Valley. Unpublished report on file at The Nature Conservancy, Colorado Field Office, Boulder, Colorado.

  • Neely, B., S. Panjabi, E. Lane, P. Lewis, C. Dawson, A. Kratz, B. Kurzel, T. Hogan, J. Handwerk, S. Krishnan, J. Neale, and N. Ripley. 2009. Colorado Rare Plant Conservation Strategy, Developed by the Colorado Rare Plant conservation Initiative. The Nature Conservancy, Boulder, Colorado, 117 pp.

  • O'Kane, S. L. 1988. Colorado's Rare Flora. Great Basin Naturalist. 48(4):434-484.

  • Payson, E. B. 1921. Monograph of the genus Lesquerella. Annals of the Missouri Botanical Garden, 8:103-236.

  • Rollins, R.C. 1983. Studies in the Cruciferae of western North America. J. Arnold Arboretum 64(4): 491-501.

  • Rollins, R.C., and E.A. Shaw. 1973. The genus Lesquerella (Cruciferae) in North America. Harvard Univ. Press. Cambridge, MA. 288 pp.

  • Rondeau, R., K. Decker, J. Handwerk, J. Siemers, L. Grunau, and C. Pague. 2011. The state of Colorado's biodiversity 2011. Prepared for The Nature Conservancy. Colorado Natural Heritage Program, Colorado State University, Fort Collins, Colorado. 

  • Rouse, L. 1981. Element stewardship abstract for Lesquerella pruinosa. Unpublished report prepared for the Nature Conservancy, Colorado Field Office, Boulder, CO.

  • Spackman, S., B. Jennings, J. Coles, C. Dawson, M. Minton, A. Kratz, C. Spurrier, and T. Skadelandl. 1996. Colorado rare plant field guide. Prepared for the Bureau of Land Management, the U.S. Forest Service, and the U.S. Fish and Wildlife Service by the Colorado Natural Heritage Program, Fort Collins.

  • U. S. Fish and Wildlife Service. 1988. Proposal to determine Astragalus osterhoutii and Penstemon penlandii to be endangered species. Federal Register 53(128): 25181-25185.

  • USDA, NRCS. 2013. The PLANTS Database. National Plant Data Team, Greensboro, NC 27401-4901 USA.

  • Weber, W. A. and R. C. Wittmann. 2012. Colorado Flora, Eastern Slope, A Field Guide to the Vascular Plants, Fourth Edition. Boulder, Colorado. 555 pp.

  • Weber, W. A. and R. C. Wittmann. 2012. Colorado Flora, Western Slope, A Field Guide to the Vascular Plants, Fourth Edition. Boulder, Colorado. 532 pp.

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