Return of the Native: a Look at Select Accessions of North American Lewis Flax Stanley G. Kitchen perennial blue flax for traits associated with stand persistence, commercial seed production, and seedling establishment. Results will be used to develop native flax releases for commercial seed production and for restoration plantings. Abstract—Lewis flax (Linum lewisii) is a widely distributed perennial forb in the Western United States. I compared the performance of native Lewis flax accessions with that of the European cultivar ‘Appar’. In common garden studies, significant differences in survival, vigor, rust resistance, and seed production were observed among ‘Appar’ and 14 Lewis flax accessions. Some native Lewis flax accessions compared favorably to ‘Appar’ for each of these traits. Greenhouse seedling emergence percentages for 12 of 18 Lewis flax accessions were not significantly different than that of ‘Appar’. Early seedling growth for six Lewis flax accessions at three field locations was also comparable to that of ‘Appar’. Hybridization among accessions possessing desired traits may enhance selection for high performance Lewis flax releases. Materials and Methods Study Sites Three field sites were selected to evaluate the growth of Lewis flax in field conditions. Low-elevation sites were selected to test drought tolerance. Brief descriptions of the sites follow. The Orchard site is 32 km southeast of Boise, ID, and 6.5 km south of the Orchard exit on Interstate 84. Elevation is 970 m. The soil is a deep (rooting depth greater than 1.5 m) sandy loam with good to fair drainage and 0 to 2 percent slope. Seventy-eight percent of the 28 cm mean annual precipitation occurs during winter and spring (November to May). Mean annual temperature is 10.1 °C. Principal perennial species of adjacent, somewhat intact communities include Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), Sandberg bluegrass (Poa secunda), and bottlebrush squirreltail (Elymus elymoides). The Nephi site is 13 km southwest of Nephi, UT, on the Utah State University Agricultural Experiment Station’s Nephi farm. Elevation is 1,580 m. The soil is a deep loamy clay on a 0 to 2 percent slope. Mean annual precipitation is 34 cm, with 67 percent of the precipitation occurring from November to May. Mean annual temperature is 9.7 °C. This typical basin big sagebrush (A. tridentata ssp. tridentata) site has been used for testing small grains, alfalfa (Medicago sativa), and numerous forage grasses, forbs, and shrubs. The Skull Valley site is 13 km north of Dugway, UT. Elevation is 1,450 m. The soil is a deep, well-drained sandy loam with a 0 to 5 percent slope. Mean annual precipitation is 18 cm, with 57 percent of the precipitation occurring in winter and spring. Mean annual temperature is 11.5 °C. Few perennials remain on this site due to a history of repeated wildfires and past overgrazing. Before disturbance, vegetation was likely dominated by bottlebrush squirreltail, shadscale (Atriplex confertifolia), Indian ricegrass (Oryzopsis hymenoides), budsage (Artemisia spinescens), and winterfat (Ceratoides lanata). Restoration efforts on disturbed lands in the Intermountain West require appropriate seed mixes to obtain the desired community composition. Adapted forb species are an important component of most of these communities, providing diversity and stability. Familiarity with the traits of adapted forb species and their potential is essential in developing expectations for restoration efforts (Kitchen 1994). Few native forbs have been developed to the point that seed is grown commercially. Cultivars exist for only a few species, including Palmer penstemon (Penstemon palmeri), Rocky Mountain penstemon (P. strictus), northern sweetvetch (Hedysarum boreale), and Louisiana sagewort (Artemisia ludoviciana) (McArthur 1988). ‘Appar’ was released as a broadly adapted cultivar of Lewis flax (Linum lewisii). Favorable traits include good seedling vigor, excellent seed production, and a showy inflorescence (Howard and Jorgensen 1980; Shaw and Monsen 1983). However, recent work (Pendleton and others 1993) has clearly demonstrated that ‘Appar’ belongs to the European species, perennial blue flax (L. perenne), and that it is a distinct species from North American Lewis flax. As such, ‘Appar’ should not be used as a native. My objective in this study was to compare the performance of populations of native Lewis flax to that of ‘Appar’ In: Roundy, Bruce A.; McArthur, E. Durant; Haley, Jennifer S.; Mann, David K., comps. 1995. Proceedings: wildland shrub and arid land restoration symposium; 1993 October 19-21; Las Vegas, NV. Gen. Tech. Rep. INTGTR-315. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. Stanley G. Kitchen is a Botanist at the U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Shrub Sciences Laboratory, Provo, UT 84606. Common Gardens Common gardens were established with transplants in the spring of 1989 at the Orchard and Nephi sites. Transplants were grown using seed collected from 14 native 321 populations and from ‘Appar’ seed (table 1). Replicated plots of 24 plants each were randomly arranged within three blocks at each site. Each plot consisted of four rows (120 cm spacing between rows) of six plants each (60 cm spacing within rows). Spacing between plots was 150 cm. Individual plants were rated yearly from 1989 to 1991 at the Orchard site and from 1989 to 1992 at the Nephi site. All living plants were scored for vigor on a scale from 1 (low) to 5 (high). The abundance of leaves and stems and their color, flower production, and overall succulence were considered in determining individual plant vigor scores. In 1990 plants at the Nephi site were also scored for rust infection on a scale from 0 (no visible evidence of rust) to 5 (rust infection on more than 90 percent of the plant). All plants at the Nephi site were treated with the systemic fungicide, Plantvax, in May 1990. (maximum possible seeds per fruit is 10). After seeds (fruits) had matured and capsules had begun opening, bulk samples were collected from more than 20 plants of each accession. Seed weight was determined with four replications of 100 seeds. Estimates of the total number of seeds and the weight of seeds per plant were extrapolated from these data. Greenhouse Seedling Emergence In November 1991, I evaluated seedling emergence in controlled greenhouse conditions (25 °C during day, 15 °C during night) for 19 Lewis flax accessions and ‘Appar’ (table 1). Fifty seeds of each accession were planted in three replicated rows at a depth of 3.2 cm in well-drained planter boxes. The planting medium was a loamy sand. Row length was 70 cm; spacing between rows was 6 cm. Boxes were watered periodically so soil moisture would not be limiting. Emerged seedlings were counted weekly for 42 days after planting. In addition, germination trials were conducted on each seedlot to determine total germinable seed percentages. Four replications of 25 seeds were placed in Petri dishes on top of two germination blotters moistened to saturation with tap water. Petri dishes were placed in a cardboard box enclosed in a plastic bag and stored in the greenhouse for 21 days. Germinated seeds were counted and removed weekly. Ungerminated seeds were classified as dormant or dead using a simple cut test. Emergence percentages were adjusted by dividing raw emergence percentages by percent germinable for each accession. Seed Production Seed production was determined for eight plants of 13 Lewis flax accessions (table 1) and for ‘Appar’ at the Nephi site during 1990 and 1991. Vigorous, nonborder plants were selected 2 weeks before flowering began. Reproductive stems were counted on each plant after flowering had ceased. Mean numbers of flowers and fruits per stem were estimated by counting aborted flowers and fruits on a subsample of 20 (1990) or 10 (1991) stems from each study plant. Seeds from 20 fruits from each plant were harvested just before capsules were fully ripe. Seeds were counted and divided by 200 to determine percent fruit fill Table 1—Site information for 19 Lewis flax seed collections. ‘Appar’ perennial blue flax seeds were from a commercial lot. Accessions were used in a combination of studies including common garden (1), seed production (2), greenhouse emergence (3), and field seedings (4). Collection name County State Elevation Mean annual precipitation m cm Confusion Range Potosi Burr Trail Yuba Dam Mona Cove Fort Maple Grove Lava Hot Springs Little Antelope Summit Black Hills Fort Collins Asotin Provo Overlook Blue Springs Hill Hyde Park Richmond Parley’s Summit Panguitch Lake Millard Clark Garfield Juab Juab Millard Millard Bannock White Pine UT NV UT UT UT UT UT ID NV 1,870 1,850 2,030 1,630 1,540 1,760 2,000 1,460 2,270 22 25 25 33 34 34 36 36 36 Custer Larimer Asotin Utah Box Elder Cache Cache Summit Garfield SD CO WA UT UT UT UT UT UT 1,340 1,760 320 1,970 1,570 1,540 1,710 2,060 2,580 36 38 38 43 43 44 47 58 58 Elk Knoll ‘Appar’ Sanpete — UT — 3,160 — 71 — 322 Vegetation type Saltbush-grass Pinyon-juniper Pinyon-juniper Sagebrush-grass Sagebrush-grass Sagebrush-grass Sagebrush-grass Sagebrush-grass Pinyon-juniperMountain brush Ponderosa-mahogany Ponderosa-mahogany Palouse grassland Sagebrush-grass Sagebrush-grass Sagebrush-grass Sagebrush-grass Mountain brush Ponderosabitterbrush Alpine meadow — Studies 1,2,3,4 1,2,3 1,2,3,4 1,3,4 1,2,3 1,2,3 1,2,3 1,2,3,4 3 3 3 1,2,3,4 1,2,3,4 1,2,3 1,2,3 1,2,3 1,2,3 3 3 1,2,3,4 Table 2—Survival and vigor of ‘Appar’ perennial blue flax and 14 Lewis flax collections at the Orchard common garden (planted March 1989). For all collections, 1991 survival was less than 5 percent. Plant vigor is on a scale of 1 (low) to 5 (high). Within columns, means followed by the same letter are not significantly different at the P < 0.05 level (Student-Newman-Keuls multiple range test). Field Seedling Emergence Small plots at the Orchard, Nephi, and Skull Valley sites were seeded in October 1991 using seed from six Lewis flax accessions and ‘Appar’ (table 1). Sites were fallowed to reduce competition from weeds. On each site, sets of four parallel furrows 5 to 10 cm deep and 45 cm apart were made perpendicular to the prevailing winter wind direction using common farm equipment. Seeds of each accession were planted in three 8-foot plots (four furrows each) arranged in a randomized complete block design. Seeding depth was 1 to 2 cm. Seeding rate was 67 seeds per meter of row. Seedling density was determined in late spring using eight 60-cm subplots randomly located along the seeded rows of each plot. Emergence percentages were calculated from seeding rate and seedling densities. Subjective observations for plant size, stand survival and vigor, and flowering intensity were made in the summer of 1993. Transplant survival and seedling emergence percentage data were arcsine transformed before statistical analysis. Results of all experiments were subjected to analysis of variance procedures appropriate to the experimental design used. Within a treatment, significant differences among accessions were determined using the StudentNewman-Keuls (SNK) multiple range test. Regression analysis was used to examine the relationship of mortality and vigor to rust resistance at the Nephi garden and the relationship between emergence percentages in the greenhouse and in the field. Collection Transplant survival 1989 1990 Vigor 1990 - - - - - - -Percent - - - - - ‘Appar’ Confusion Range Potosi Burr Trail Yuba Dam Mona Cove Fort Maple Grove Lava Hot Springs Asotin Provo Overlook Blue Springs Hill Hyde Park Richmond Parley’s Summit 35a 33a 31ab 31ab 10bcde 26abc 2e 10bcde 13bcde 8cde 25abc 19abcd 21abcd 17abcd 7de 35a 31ab 28abc 26abcd 10cdef 24abcd 2f 8def 10cdef 7def 22abcde 15bcdef 9cdef 7def 2f 3.8ab 4.1a 3.4ab 3.4ab 3.0abc 3.4ab 4.0a 3.6ab 2.6abcd 2.9abcd 3.3ab 1.5cd 1.6cd 1.3d 2.0bcd was apparent throughout the garden. Blocks of ‘Appar’ remained free of any visible evidence of infection, while rust resistance in Lewis flax accessions varied considerably among accessions (table 3). Mortality from 1990 to 1992 was significantly correlated with mean rust index (r2 = 0.52, d.f. = 13) even though fungicide application Results Common Gardens Initial transplant survival (5 months after transplanting) at the Orchard common garden was 19.2 percent. Survival percentages for eight Lewis flax accessions were not significantly different than for ‘Appar’ (table 2). Mean survival 1 year after planting was 15.7 percent. Most mortality between years occurred among accessions collected from more mesic sites (see mean annual precipitation, table 1). At this time, survival for five Lewis flax collections was not significantly different than that of ‘Appar’. Survival after 2 years was less than 5 percent for all accessions, including ‘Appar’ (data not shown). Mean vigor of surviving plants at the Orchard common garden in August 1989 ranged from 1.0 to 2.0, with no significant differences among collections (data not shown). In 1990, mean vigor for all accessions was 2.9; mean vigor for 12 Lewis flax accessions was not significantly different from that for ‘Appar’ (table 2). Lower vigor tended to be associated with collections that experienced the highest mortality between August 1989 and April 1990. At the Orchard site, ‘Appar’ and Confusion Range were the only accessions that flowered to any significant extent (Kitchen, field notes). Transplant survival for all collections at the Nephi common garden 1 year after planting was 95.3 percent. Survival varied greatly among accessions from the summer of 1990 to 1992 (fig. 1). Visible evidence of rust infection was first noted in April 1990; by mid-May rust infection Figure 1—Survival of eight representative Lewis flax accessions and ‘Appar’ in a common garden near Nephi, UT. Greenhouse-reared seedlings were transplanted in April 1989. Mortality from 1990 to 1992 was significantly correlated with severity of 1990 rust infection (P < 0.05, r 2 = 0.52). All plants were treated for rust in 1991, minimizing infection. 323 effectively controlled rust in 1991. All plants of the Confusion Range accession died by 1991. This accession showed the greatest susceptibility to rust. Significant differences in the vigor of surviving plants were observed each year among accessions at the Nephi site (table 3). In 1989, nine accessions of Lewis flax had mean vigor values that were not significantly different than that of ‘Appar’. In 1990, five accessions of Lewis flax had significantly higher mean vigor values than ‘Appar’ and one had a significantly lower value. Vigor in 1991 was significantly correlated with rust infection of the pre2 vious year (r = 0.61, d.f. = 12). In 1991, mean vigor for the Maple Grove, Asotin, Hyde Park, and Parley’s Summit accessions was not significantly different from that of ‘Appar’. These accessions all had low rust indices. ‘Appar’ remained rust free. Vigor of surviving plants in 1992 was similar for 11 accessions, including ‘Appar’. While several accessions had relatively high mean vigor values in 2 or 3 of the 4 years, only Maple Grove scored among the highest in each of the 4 years. ‘Appar’ scored among the highest during 3 years. set percentages in 1991 (data not shown). Two-year perplant seed production among Lewis flax accessions was highest for the Mona, Maple Grove, Hyde Park, Cove Fort, and Lava Hot Springs accessions. Seedling Emergence and Growth Seedling emergence under greenhouse conditions ranged from 18 to 89 percent (table 5). Emergence percentages for 12 accessions of Lewis flax were not significantly different than that of ‘Appar’. Seedling emergence in the field varied by site. Mean emergence percentage for the seven accessions at the Orchard site was 18 percent. There were no significant differences among accessions. Mean emergence at the Nephi site was 23 percent. Emergence percentages for the Lava Hot Springs (51 percent) and Provo Overlook (33 percent) accessions were significantly higher than those of the other five collections, including ‘Appar’. Mean seedling emergence at the Skull Valley site was 63 percent, significantly higher than at the other two sites. At that site, only the Burr Trail accession had an emergence percentage significantly lower than that of ‘Appar’. Combined field emergence percentages ranged from 42 percent for ‘Appar’ to 22 percent for Burr Trail and were not significantly correlated with greenhouse emergence percentages. No seedlings at the Orchard and Whiterocks sites survived to the summer of 1993. Observations at the Nephi site revealed vigorous plants of each accession, all of which flowered and produced seed. Seed Production Seed production for ‘Appar’ was clearly higher than for all other accessions (table 4). However, ‘Appar’ seeds are smaller than those of all collections of Lewis flax. The gap in seed production between ‘Appar’ and the Lewis flax accessions is reduced when comparing grams of seeds per plant. Though mean first-year (1990) production was 37 g per plant for the eight most productive accessions of Lewis flax (compared to 53 g for ‘Appar’), second year (1991) production was relatively poor or nonexistent for five of those accessions. Data were not taken for four accessions in 1991 because the study plants died. All accessions, including ‘Appar’, experienced reduced flowering and fruit Discussion Factors other than species adaptability affected successful establishment of transplants. Such factors include Table 3—Mean vigor and rust index of surviving transplants of ‘Appar’ perennial blue flax and 14 collections of Lewis flax at the Nephi common garden. Plant vigor is on a scale from 1 (low) to 5 (high). Rust index values were scored from 0 (no infection) to 5 (more than 90 percent of plant infected). Within columns, means followed by the same letter are not significantly different at the P < 0.05 level (Student-Newman-Keuls multiple range test). Collection 1989 Mean vigor 1990 1991 ‘Appar’ Confusion Range Potosi Burr Trail Yuba Dam Mona Cove Fort Maple Grove Lava Hot Springs Asotin Provo Overlook Blue Springs Hill Hyde Park Richmond Parley’s Summit 2.9a 2.9a 2.6abcd 2.7abcd 2.9a 2.4dce 2.7abc 2.9a 2.5bcde 2.2e 2.8ab 2.6abcd 2.4de 2.9a 2.1e 2.1cd 2.3b 2.2bc 2.0cd 2.1cd 2.4ab 2.0cd 2.6a 2.4ab 1.9de 2.1cd 2.0cd 2.1cd 1.9de 1.7e 324 3.9a — 1.4c 1.6e 1.8e 2.9bcd 2.8cd 3.4abc 2.9cd 3.3abc 2.4d 3.1bc 3.5abc 2.9bcd 3.6ab 1992 2.5abc — 2.5abc 2.2abc 2.2abc 2.5abc 2.4abc 2.6abc 2.1abc 2.9a 2.3abc 1.8bc 2.0bc 1.7c 2.2abc Mean rust index (1990) 0.0g 4.6a 1.0d 1.4c 1.8b 1.2c 1.0d 0.4f 0.6e 0.3f 1.7b 1.0d 0.6e 1.0d 0.1g Table 4—Seed production per plant for ‘Appar’ perennial blue flax and 13 collections of Lewis flax at the Nephi common garden. Within columns, means followed by the same letter are not significantly different at the P < 0.05 level (Student-Newman-Keuls multiple range test). Collection Mean seeds per plant 1990 1991 Seeds per gram Mean grams per plant 1990 1991 - - - - - - - -g - - - - - - - - ‘Appar’ Confusion Range Potosi Burr Trail Mona Cove Fort Maple Grove Lava Hot Springs Asotin Provo Overlook Blue Springs Hill Hyde Park Richmond Parley’s Summit 34,012a 15,114b 9,548b 14,784b 14,466b 18,820b 17,323b 13,788b 7,544b 14,781b 16,865b 10,375b 10,582b 8,988b 23,126a — — — 7,122b 5,626b 8,418b 5,266b 4,031b — 8,046b 9,871b 5,939b 5,805b 641 465 343 428 360 474 452 392 513 365 481 362 381 527 53a 33abc 28bc 35abc 40ab 40ab 38ab 35abc 15c 40ab 35abc 29bc 28bc 17bc 36a — — — 20b 12b 19b 13b 8b — 17b 27ab 16b 11b Table 5—Seedling emergence percentages of ‘Appar’ perennial blue flax and 19 Lewis flax collections in greenhouse and field seeding trials. Within columns, means followed by the same letter are not significantly different at the P < 0.05 level (Student-Newman-Keuls multiple range test). Collection Greenhouse Seedling emergence Field sites Orchard Nephi Skull Valley - - - - - - - - - - - - - - - - - - -Percent - - - - - - - - - - - - - - - - - - - ‘Appar’ Confusion Range Potosi Burr Trail Yuba Dam Mona Cove Fort Maple Grove Lava Hot Springs Little Antelope Summit Black Hills Fort Collins Asotin Provo Overlook Blue Springs Hill Hyde Park Richmond Parley’s Summit Panguitch Lake Elk Knoll 63abcd 43bcde 89a 60abcd 41bcde 67abc 88a 68abc 58abcd 59abcd 21a 19a — 8a 13a — — — 17a — 18c 14c — 14c 12c — — — 51a — 88a 66ab — 43b 59ab — — — 55ab — 19e 18e 22de 75ab 61abcd 56abcd 52abcde 67abc 38bcde 24cde — — 24a 24a — — — — — — — — 21c 33b — — — — — — — — 79ab 52ab — — — — — — differences in the condition of seedlings when they were transplanted, variability in transplant method among planters, and variability in conditions (such as air temperature) during transplanting. For this reason, caution is required in evaluating differences in 1989 survival data among accessions at the Orchard site. Changes in survival from 1989 to 1990 combined with 1990 vigor scores provide a truer indication of adaptation to xeric conditions, such as those encountered there (table 2). Based on these data, the drought tolerance of several accessions of Lewis flax may equal that of ‘Appar’. Interpreting data from the more mesic Nephi site requires similar caution. In general, accessions from drier collection sites were more susceptible to rust infection and shorter lived than those from more mesic sites. The role of rust in among-accession differences in plant longevity is unknown. However, rapid development and early reproductive maturity would be a great adaptive advantage 325 for accessions from sites where drought is more frequent and more severe. Investment of resources into rust resistance or reserves necessary for long life would be less advantageous at such sites (Fenner 1985). Slower maturing accessions (those with low 1990 vigor or lower seed production) had greater rust resistance and were longer lived. Some of these accessions were from sites that receive up to 50 percent more annual precipitation than the Nephi site. Because vigor is a qualitative measure subject to considerable bias, comparisons between sites or years should not be made. For example, the overall mean vigor in 1990 was 2.9 at Orchard and 2.2 at Nephi, even though the Nephi plants were undoubtedly larger and more productive than those at Orchard. However, comparisons for the same year and same site are valid because they were made by a single team and completed within 1 day. Vigor, as we measured it, was affected by several intrinsic factors including resource allocation patterns and resulting growth rate, resource depletion in connection with reproduction, and susceptibility to rust. On the Orchard and Nephi study sites, vigor for most Lewis flax accessions was comparable to that of ‘Appar’, both when the accessions were grown as transplants and when they were seeded, suggesting that they may have the ability to perform as well as ‘Appar’. Accessions from collection sites receiving 34 to 44 cm mean annual precipitation (the middle of the range for collection sites) had the highest 2-year seed production, although the production was considerably lower than that of ‘Appar’ (tables 1 and 4). Further work is needed to determine how seed yield might improve if rust were controlled or other culture conditions were improved. Though seed yields for Lewis flax accessions did not equal those of ‘Appar’, they were probably high enough for profitable commercial production. Successful seedling emergence in the field is affected by factors that vary among sites and years. None of the seven accessions tested was superior to the rest when all three sites are considered. In addition, there is no correlation between the emergence means for all three study sites and greenhouse emergence success. Greenhouse emergence from deep planting is not likely to be a useful test or selection technique for this species. These studies suggest that several accessions of Lewis flax have seedling growth capabilities equal to those of ‘Appar’. However, single-year emergence percentages for Lewis flax accessions are reduced by the portion of viable seed that remains in the soil seed bank. In controlled germination studies, all Lewis flax accessions expressed some combination of traits that would enable them to maintain a seed bank (Meyer and Kitchen 1994). In this study, ‘Appar’ lacked this adaptive feature; virtually all seeds germinated at the first occurrence of favorable temperatures. Field retrieval studies at three contrasting sites confirmed the laboratory results. Care should be taken not to lose the capacity for seed banking in Lewis flax releases through intentional or inadvertent selection in an agronomic setting. It appears that obstacles to the production and use of Lewis flax seed are largely those associated with field cultivation. Some accessions with otherwise favorable traits are quite short lived and susceptible to rust. Other accessions show relatively good rust resistance and longevity but may mature more slowly and produce less seed. The impacts of rust control on seed production are not yet known. There will also be additional costs associated with chemical control of rusts. Hybridization could be used to pool favorable traits found in Lewis flax as a species. Hybrids that combine high levels of seed production, rust resistance, and drought tolerance are possible. We can only speculate concerning possible compromises to be encountered in such an effort. Broadly adapted collections or hybrids that pose minimal difficulty in seed production need to be identified. This work represents a first step. Acknowledgments This research was funded in part by the U.S. Department of the Interior, Bureau of Land Management, through Interagency Agreement ID910-IA-8. Special thanks are extended to Stephen B. Monsen, Karl Soerensen, Susan E. Meyer, and Stephanie Carlson for many hours of careful field and laboratory work that have made this work possible. References Fenner, M. 1985. Seed ecology. New York: Chapman and Hall. 151 p. Howard, C. G.; Jorgensen, K. R. 1980. ‘Appar’ Lewis flax (Linum lewisii, Pursh) description, adaptation, use, culture, management, and seed production. U.S. Department of Agriculture, Soil Conservation Service, Plant Materials Center. Kitchen, S. G. 1994. Perennial forb life-history strategies on semiarid rangelands: implications for revegetation. In: Monsen, S. B.; Kitchen, S. G., comps. Proceedings— ecology, management and restoration of annual rangelands; 1992 May 18-22; Boise, ID. Gen Tech. Rep. INTGTR-313. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 342-346. McArthur, E. D. 1988. New plant development in range management. In: Tueller, P. T., ed. Vegetation science applications for rangeland analysis and management. Boston, MA: Kluwer Academic Publishers. Meyer, S. E.; Kitchen, S. G. 1994. Life history variation in blue flax (Linum perenne: Linaceae): seed germination phenology. American Journal of Botany. 5: 528-535. Pendleton, R. L.; Kitchen, S. G.; McArthur, E. D. 1993. Origin of the flax cultivar ‘Appar’ and its taxonomic relationship to North American and European perennial blue flax. Wildland shrub and arid land restoration symposium. Abstracts. Provo, UT: Shrub Research Consortium: 23 Shaw, N.; Monsen, S. B. 1983. Nonleguminous forbs for rangeland sites. In: Monsen, S. B.; Shaw, N., comps. Managing Intermountain rangelands—improvement of range and wildlife habitats. Gen. Tech. Rep. INT-157. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 123-131. 326