This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. RANGELAND SPECIES GERMINATION THROUGH 25 AND UP TO 40 YEARS OF WAREHOUSE STORAGE Richard Stevens Kent R. Jorgensen ABSTRACT be stored. Germinability of each species will affect the seeding rate. Germination data of some range grasses stored anumber of years have been reported (Hafenrichter and others 1965; Little 1937; Tiedemann and Pond 1967). However, longevity germination studies of wildland shrubs and forbs have been somewhat limited. Springfield (1968, 1970, 1973, 1974) worked with germination and longevity of stored winterfat (Ceratoides lanata) and fourwing saltbush (Atriplex canescens) seed. Longevity reports on germination of stored winterfat (Stevens and others 1977) and antelope bitterbrush CPurshia tridentata) seed (Giunta and others 1978) have been made. Plummer and others (1968) listed the duration of good viability for 44 shrubs, 24 forbs, and five grasses. Plummer and others (1970) also reported germination results after relatively short periods of storage on seed of a number of native shrubs and forbs. Two works from the U.S. Department of Agriculture, Forest Service (1948, 1974) included information on germination and longevity studies for native trees and shrubs. Van Haverbeke (1989) reported on viability of 20-year-old ponderosa pine CPinus ponderosa) seed. Kay and others (1984) reported on germination of seed of 22 Mojave desert shrub species following 9 years of storage. Kay and others (1988) ran germination tests on seed of 115 desert grasses, forbs, and shrubs that had been stored for up to 14 years. Seed of rangeland species in the Intermountain West are commonly stored in open, unheated, and uncooled warehouses and granaries, sometimes for extended periods. Little information is available as to the length of life of many warehouse-stored rangeland seeds. Germination results are presented from seed stored in UT, for 19 shrub, 16 forb, and three grass species for 26 years, and 12 shrub, 20 forb, and 29 grass species for 0 to 40 years. Germination, longevity, and afterripening are discussed for each species and group of species through various periods of storage. Each species exhibited its own unique germination characteristics that varied with source and age of seed. Some .species' viability increased with increasing age while others decreased with age. an open warehouse in Sanpete County, INTRODUCTION In 1981 Stevens and others published seed germination data for· 32 shrub and forb species stored in an open warehouse through 15 years. These data along with 20 and 25 years germination data for seed of the same species and sources and six additional species are included in this report. In addition, germination results are given for 61 shrub, forb, and grass species with 0 to 40 years of storage. A limiting factor to range rehabilitation can be the availability of quality seed of desired species at affordable prices. With most wildland species good seed crops are not produced every season: seed needs to be obtained in good years and stored until needed. It is not uncommon to store commercially produced seed. Seed may have to be stored for a few months to many years. Utah State law requires that marketed grass seed be tested every 18 months, and forb and shrub seed every 9 months. These requirements can differ by State. Rules and procedures for testing seed have been established for many wildland species (Stevens and Meyer 1990). Stored seed is generally not tested on a regular basis or prior to seeding when used by the person or agency storing it. Some species possess considerable afterripening, with maximum germinability not reached for a number of years following collection. Some species do not retain viability well, while others experience little change over long periods of storage. To obtain maximum seeding results and maximum return of dollars spent, it becomes imperative to know germinability over years and how long a species can or should METHODS Study 1: Germination Through 20 Years of the Same Seed Sources-During the fall of 1963, current seed of 19 shrub, 16 forb, and three grass species (table 1) were hand-collected from native stands or purchased from commercial sources. Seed was commercially or handcleaned to 85 percent purity or higher, placed in cotton bags, and deposited in metal file cabinets in an open warehouse in Ephraim, Sanpete County, UT, for the duration of the study. Over 25 years (1963-88), temperature in the warehouse ranged from a low of -29.9 to a high of 38.3 Mean daily temperatures during winter, spring, summer, and fall periods were -3.3, 6.7, 20, and 7.2 °0, respectively. Germination tests on the stored seed were conducted in February of 1965, 1966, 1967, 1968, 1970, 1973, 1978, 1983, and 1988. These dates marked the beginning of the oc. Paper presented at the Symposium on Ecology, Management, and Restoration of Intermountain Annual Rangelands, Boise, m, May 18-22, 1992. Richard Stevena is Project Leader and Research Biologist, Utah Division of Wildlife Resources, Ephraim, UT 84621. Kent R. Jorgensen is Assistant Project Iader and Research Biologist, Utah Division of Wildlife Resources, Ephraim, UT 84627. 257 Table 1-Mean percentage germination of seed from 38 shrub, forb, and grass species after 2, 3, 4, 5, 7, 10, 15, 20, and 25 years in storage in an open, unheated, and uncooled warehouse Species Common name Source 2 3 4 5 Years of storage 7 10 15 20 25 ------------------------PeroentgenninaNod------------------------Shrubs: Amelanchier aln/folia Saskatoonsennoebeny Ame/anchier utahensls Utah sennceberry Artemisia nova Black sagebrush Artemisia tridentata ssp. tridentata Basin big sagebrush Atrlplex canfJSCBns Fourwirig saltbush Ceanothus martini/ Martin ceanothus Ceratoldes lanata Winterfat Cercocarpus ledlfollus Curlleaf mountain-mahogany Cercocarpus montanus True mountain-mahogany Chrysothamnus nauseosus ssp. holo/eucus White rubber rabbltbrush Cowan/a stansburlana Cliffrose Ephedra nevadensls Nevada ephedra Ephedra vir/dis Green ephedra Gray/a brandegel Spineless hopsage Peraphyllum ramoslsslmum Squawapple Pui'Shla g/andulosa Desert bitterbrush Purshla tridentata Antelope bltterbrush Ribas aureum Golden currant Symphor/carpos oreophllus Mountain snowberry Forbs: Ba/samorhlza macrophylla Cutleaf balsamroot Balsamorhlza saglttata Arrowleaf balsamroot Erlogonum herac/eoldes Wyeth erlogonum Hedysarum boreale Utah sweetvetch Herac/eum lanatum Cowparsnlp UgusNcum porter/ Porter ligusticum Unum perenne Lewis flax LomaNum kingii Nuttalllomatium Spring City Canyon, UT Henrieville, UT Manti, UT Ephraim, UT Panaca, NV Manti Canyon, UT Corona, NM Mayfield, UT Ephraim Canyon, UT Richfield, UT American Fork, UT Wah-Wah Valley, UT Manti, UT Escalante, UT Ephraim Canyon, UT Bishop, CA Mt. Den, UT Manti, UT Spanish Fork Canyon, UT Bountiful, UT Paradise Valley, NV Brigham City, UT Orem, UT Pleasant Creek Canyon, UT Ephraim Canyon, UT Ephraim, UT Ephraim Canyon, UT 91 80 91 285- 97 99 99 81- 66 55- 73 82 32 72 76- 96 90- 67- 5 0 34- 5 1 0 0 0 67 70- 24- 0 0 0 47 40 40 50 43 37- 18 11 3 5 5 12 10+ 40 36- 5 6 90 83 74- 18 7 0 0 0 0 67 63 80 76 69 64- 44 28 63 65 61 68- 46- 25- 3 0 0 80- 65- 34- 14 11- 7 0 0 0 89 89 84 89 91- 66 63 •so*+ 8 0.3 90 93 91 85 89 91 85- 79 77 88 92 92 84 80 82 88- 24 2 87 92 86- 57- 13 6 0 0 0 42 42 42 37 39- 21 10 78 86 80 80 69 73 65 61 60 79*+ 86 87 94 88 88 85 84- 74 27- 6 2 0 80- 44- 8 10 8 20- 4 0 0 0 0 0 0 0 48 28- 0 80 64 35 28- 40 42 37- 20 51*+ 87 90- 64- 16- 59 67 58 55- 25 40 16 11 21 2 1 0 0 0 0 0 7 8- 92 17 5 0 41 28 24 36- 13 0 0 0 0 66* 72*+ 85 93 83 70- 25 8 0 69 73 73- 37- 8 2 0 0 (con.) 258 Table 1 (Con.) Species Common name Source 2 3 4 5 Years of storage 7 10 15 20 25 ------------------------Peroentgennlna6o~------------------------- Forbs: (Con.) Luplnus slpestrls Mountain lupine Luplnus ssricsus Silky lupine Penstemon palmer/ Palmer penstemon Medlcago sativa Alfalfa 'Rambler' same + hard seed Sangulsorba minor Smallbumet Sphaerslcsa grossulari'lfolla Gooseberry globemallow Tragopogon porrifolius Vegetable oyster salsify Vlgulsra multiflora Showy goldeneye Ephraim Canyon, UT Ephraim Canyon, UT Ephraim Canyon, UT Canada 58 n 69 60- 26 28 13 6 97 99 100 99- 86 85 92- 75 83 81 79- 65 50 69 76 75 75 70 n 92 95 94 93 91 92 96- 79 82 7 6 9 65 66 17 sa·+ Ephraim, UT Benmore, 7 UT Mt Pleasant, 65 UT Ephraim Canyon. 18 UT 76 0 66 73 67 86- 71 87 88 78 69+ 83 6 7 6 66- 46 31- 13 0 0 11 13 13- 0 0 0 63 13 1 71 2 01'88888: Agropyron lntsrmedlum Intermediate wheatgrass Bromus lnsnnls Smooth brome Ssca/8 C8f'6SI8 Winter rye Washington 95 96 93 94 80 78- Colorado 70 71- 52 39- 15 11 3 1 0 Idaho 89 88 82 75- 56 48 32- 2 0 'Results based on four samples of 100 seeds each at 100 percent purity. Llndlcates significant decrease In germination between adjoining years at CP < 0.05). significant Increase in germination between adjoining years at CP < 0.05) • .,.Indicates significant afterrfpenJng at CP < 0.05). 1+1ndlcates second,tbird,fo~,~,seven~,10~,15~,2oth,and 25~ year of storage following harvest. For each species Study 3: Germination of Various Seed Sources over 1"' Years-Seeds of six shrub, eight forb, and 20 grass species (table 3) were placed in paper bags and stored for 1 to 40 years in the open warehouse described in study 1. Some species had various sources of seed with various years of storage. Seed selection and germination tests were carried out as described in study 1, with the exception that a few sources had sufficient seed for only two 100-seed samples. in each year of sampling, four random samples of 100 seeds each were selected and individually placed between damp papers, wrapped in plastic, and placed in a refrigerator. Temperature in ~e refrigerator ranged between 1.1 and 3.3 °C. Care was taken to ensure that only undamaged seed with good fill was selected and used. Results are based on 98 percent fill and 100 percent purity for all species except fourwing saltbush (52 percent fill) and Martin ceanothus (Ceanothus martinii) (59 percent fill). Seed samples were checked at 14-day intervals. All germinated seeds were removed and the number that had germinated was recorded. Seed samples remained in an unlighted refrigerator for the duration of the study. Germination checks were made on schedule for up to 16 months. Seeds were considered germinated when the hypocotyllength reached 5 mm. Data were subjected to analysis of variance with SAS (SAS 1982). STUDY 1 RESULTS Germination and retention of viability varied between species, sources within a species, and years of storage (table 1). Two species, desert bitterbrush (Purshia glandulosa) and 'Rambler' alfalfa (Medicago sativa), did not significantly (P < 0.05) lose any germinability with 25 years of storage. Working with desert bitterbrush, Kay and others (1988) had little loss of germination in 6 years. Antelope bitterbrush seed germination did not decrease significantly for 20 years and only dropped to 74 percent between 20 and 25 years. Nevada ephedra (Ephedra nevadensis) had a significant loss between 15 and 20 years but retained acceptable germination of 77 percent Study 2: Germination of the Same Seed Source Following Collection and 6 to 16 Years Later-For six shrub, 12 forb, and nine grass species (table 2), two 100-seed samples were taken from freshly collected seed and from seed of the same lots 5 to 15 years later. Seed storage, selection, germination tests, and data analysis were made as described in study 1. 259 Table 2-Percentage germination of the same seed source for shrubs, forbs, and grass seed, comparing freshly collected seed and seed with various years of storage In an open warehouse Species Common name 0 5 6 7 8 Years of storage 10 9 11 12 13 14 15 ···-·--·-·····-·--·······-···-·Pe~ntgerminaUon'-·--·-·-----··------··------···· Shrubs: Berberis repens Oregon-grape Caragana arborescsns Siberian peashrub Lonlcsra tatarlca Honeysuckle Psraphy/lum ramoslsslmum Squawapple Peraphy/lum ramoslsslmum Squawapple Shepherd/a argentea Silver buffaloberry Forbs: Astragalus clcer Cicer milkvetch Astragalus clcer Cicer mllkvetch Astragalus gal9glformis Giant astragalus Coron/1/a varia Crownvetch Osmorhlza occldentalis Sweetanise Penstemon eaton// Eaton penstemon Psnstsmon eatonil Eaton penstemon Penstemon palmeri Palmer penstemon Penstemon platyphy/lus Thlckleaf penstemon Penstemon spectabilis Showy penstemon Vlgulsra multiflora Showy goldeneye Vlgu/sra muH/f/ora Showy goldeneye Gra8888: Agropyron slongatum Tall wheatgrass Agropyron elongatum Tall wheatgrass Agropyron elongatum Tall wheatgrass Bromus lnermus Smooth brome Festuca elaUor Meadow fescue Muhlenb8rgla wrlghtii Splkemuhly Oryzopsls hymenoides Indian ricegrass Oryzopsls hymsnoldes lncfian ricegrass Stlpa comata Needle-and-thread 25 23 88 85 57 ~ 31 67 49 58 58 85 88 51* 4+ 8 89 73 65 88 89 41* 70 + 94 44 71* + 87 63* + 82 89 82 74 68 30* + + + 75 44 39 0 72* 91 + 85 87 85 92 91 94 69* + 14* + 9* + 99 62 49 55 88 83 83 'Results based on two samples of 100 seeds each at 100 percent purity. '-Indicates significant decrease In germination between germination years at {p < 0.05). ,.Indicates significant afterripenlng at {p < 0.05). 4 +1ncfic:ates significant Increase In germination between germination years at {p < 0.05). 260 Table 3-Seed germination of selected species with 0 to 40 years of storage. Seed of the same species may not be from the same source Species Common name 1 2 5 6 7 8 Years of storage 9 10 13 14 15 16 17 1 Shrubs: Atrlplex hymenelytra Desert holly Atrlplex lentiformls Big saltbush Caragana pygmaea Pygmy peashrub Csanothus cuneatus Wedgeleaf ceanothus Csanothus fBndlerl Fendler ceanothus Colutea arborBSC6ns Bladdersenna Forbs: Erodlum clcutar/um AHUeria Mfldicago sativa Alfalfa MeO/otus officina/Is Yellow sweetclover Onobtychls sativa Sainfoln Osmorhlza occ/dentalls Sweetanlse Penstemon eaton// Eaton penstemon Penstemon platyphyl/us Hillside penstemon Pentstemon subglsber Smooth penstemon 01'88888: Agropyron crlstatum Fairway wheatgrass Agropyron d8S8rtorum Crested wheatgrass Agropyron slbiricum Siberian wheatgrass Agropyron spicatum Bluebunch wheatgrass Agropyron trichophorum Pubescent wheatgrass A/opecurus arundlnaCBus Reed foxtail Alopecurus pratensls Meadow foxtail Bromus e1'8Ctus Meadow brome Dactyl/s glomerata Orchardgrass Festuca elatior Meadow fescue FBStuca ovina Sheep fescue FBStuca rubra Red fescue Koe/er/a cristata Prairie junegrass Lolium perenne Perennial ryegrass ---------------------------------Pe~ntgennlna~n --------------------------------- 57 92 67 85 63 27 37 45 (40 years. 90) 86 95 59 84 92 27 52 34A 88 86 68 58 52 95 41 39 96 8 91 38 95 94 86 47 97 86 91 91 96 (30 years. 44) 82 89 73 82 89 63 41 72 40 95 93 (con.) 261 40 Table 3 (Con.) Species Common name 1 2 5 6. 7 8 Years of storage 9 10 13 14 15 16 17 1 ···········---··---···--·--·-----Pe~ntgennlnaUon --·--·--------------------------- Graases: (Con.) Ph/sum pratsnss Timothy Poabulbosa Bulbous bluegrass Psalhyrostachys juncsus Russian wildrye Secale montsnum Mountain rye Sporobo/us alroldss Alkali sacaton Sporobolus columbiana Subalpine needlegrass 98 8 99 82 23 88 50 99 63 1 Results basad on two or four samples of 100 seeds each. Utah sweetvetch and 3 years for Lewis flax and vegetable oyster salsify. Kay and others (1988) reported Palmer penstemon had slightly longer viability. Golden currant CRibes aureum), spineless hopsage (Grayia branclegei), and winterfat germination dropped significantly after 4 years of storage; cowparsnip (Jleracleum lanatum), cutleafbalsamroot (Balsamorhiza macrophylla), and smooth brome (Bromus inermis) after 3 years; black sagebrush (Artemesia nova) (llld white rubber rabbitbrush (Chrysothamnus nauseosus ssp. hololeucus) after only 2 years. Kay and others (1988), working with Mojave desert accessions, had similar results with white rubber rabbitbrush but less retention of viability with winterfat. Stevens and others (1977) reported similar results with winterfat. Springfield, (1968, 1973, 1974), working in New Mexico, found that winterfat would lose viability rapidly unless it was stored in a cold environment. Alfalfa viability is a combination of hard and germinated seed. Hard seed is viable seed that does not absorb water nor germinate during the germination test. 'Rambler' alfalfa seed germination percentage did not significantly · change over 25 years. However, germination percentage of hard seed did significantly decrease between the lOth and 15th year of storage. Five species showed considerable afterripening by demonstrating a significant increase in germination with age. Antelope bitterbrush, cliffrose, small burnet, and Wyeth eriogonum germinability increased to the third year and Lewis flax to the fourth year after harvest. Kay and others (1988) demonstrated a similar increase in cliffrose germination over the first 3 years. with 25 years. Nevada ephedra seed stored in a warehouse in California lost significant germination after the first year, but seed stored at -15 oc did not lose germination with 14 years of storage (Kay and others 1988). Silky lupine (Lupinus sericeus) and small burnet (Sanquisorba minor) germination decreased significantly between 5 and 7 years. However, they had 76 and 83 percent germination, respectively, after 25 years. Cliffrose (Cowania stansburiana), curlleafmountainmahogany (Cercocarpus ledifolius), fourwing saltbush, green ephedra (Ephedra viridis), Nevada ephedra, and gooseberry globemallow (Sphaeralcea grossulariifolia) exhibited a significant loss in germination between 15 and 20 years. In work in California with clifftose and green ephedra, Kay and others (1988) obtained results similar to the above. Springfield (1970), working with fourwing saltbush, had a 50 percent loss of germination over 15 years. Utah serviceberry (Amelanch~r utahensis), Lewis flax (Linum perenne), and intermediate wheatgrass (Agropyron intermedium) retained acceptable germination for 10 to 15 years. Mountain snowberry (Symphoricarpos oreophilus), Saskatoon serviceberry (Amelanch~r alnifolia), true mountain-mahogany (Cercocarpus montanus), squawapple CPeraphyllum ramosissimum), and showy goldeneye (Vi&uiera multiflora) had a significant loss in germination between the seventh and lOth year of storage. Basin big sagebrush (Artemisia tridentata ssp. tridentata), arrowleafbalsamroot (Balsamorhiza sagittata), silky and mountain lupine (Lupinus alpestris), Nuttall lomatium (Lomatium nuttallii), Palmer penstemon (Penstemon palmeri), Porter ligusticum (Ligusticum porteri), small burnet, Utah sweetvetch (lledysarum boreale), vegetable oyster salsify (Tragopogon po"ifolius ), Wyeth eriogonum (Eriogonum heracleoides), and winter rye (Secale cereale) retained consistent germination for 5 to 7 years. Duration of good viability was listed by Plummer and others (1968) as 5 years for arrowleafbalsamroot, Palmer penstemon, Porter ligusticum, Nuttalllomatium, and STUDY 2 RESULTS Germination tests of new and several-year-old seed of 20 species, six of which had more than one accession, produced variable results (table 2). Eight species exhibited considerable afterripening. One source of tall wheatgrass (Agropyron elongatum) had 19 percent more germination 262 with 9-year-old seed than from new seed. However, two other sources had no significant change in germination over 9 years. New seed of meadow fescue CFestuca elatior) exhibited 69 percent germination and 8 years later, 99 percent. Spike muhly CMuhlenbergia, wrightii) germination went from 14 to 62 percent in 7 years. For one source of Indian ricegrass (Oryzopsis hymenoides), 14 years after harvest, germination increase4 from 9 to 49 percent. Another source with only 10 years storage had no significant change in germination. One source of cicer milkvetch (Astragalus cicer) went from 51 to 89 percent germination in 7 years. A second source, however, had no significant change in germination with 8 years of storage. Crownvetch (Coronilla varia) seed showed a significant increase in germination from 41 to 70 percent in 8 years. In two accessions of Eaton penstemon (Penstemon eatonii), germination significantly increased over 13 years. One went from 71 to 87 percent and the second from 63 to 82 percent. Showy penstemon (P. spectabilis) exhibited a significant increase in germination from 30 to 75 percent in lOyears. Germination percentage did not significantly change in accessions of 11 species: Oregon-grape (Berberis repens) (25 to 23 percent in 13 years), Siberian peashrub (Caragana arborescens) (88 to 85 percent in 6 years), one source of squawapple (no change in 7 years), silver bu1faloberry (Shepherdia argentea) (85 to 88 percent in 9 years), cicer milkvetch (72 to 85 percent in 9 years), Astragalus galegiformis (88 to 89 percent in 7 years), Palmer penstemon (89 to 82 percent in 13 years), thickleaf penstemon (J'enstemon platyphyllus) (74 to 68 percent in 14 years), tall wheatgrass from two sources (85 to 87 percent and 85 to 92 percent in 9 years), smooth brome (91 to 94 percent in 5 years), and Indian ricegrass (55 to 63 percent in 10 years). Only five species exhibited significant loss in germination percentage over their indiVidual storage periods: honeysuckle (Lonicera tatarica) (57 to 49 percent in 12 years), squawapple (67 to 49 percent in 8 years), sweetanise (Osmorhiza occidentalis) (94 to 44 percent in 13 years), showy goldeneye from two sources (44 to 1 percent in 15 years, 39 to 0 percent in 13 years), and needle-and-thread (Stipa comata) (88 to 63 percent in 8 years). arborescens) showed little difference in germination with 1-year-old (45 percent) and 10-year-old (38 percent) seed. All of the tested forb seed showed considerable longevity. Forty-year-old seed of alfileria (Erodium cicutarium) exhibited 90 percent germination, alfalfa 86 percent with 15-year-old seed, and yellow sweetclover CMelilotus o/ficinalis) 59 percent with 13-year-old seed. The penstemonsEaton, hillside, and smooth (Penstemon subglaber) had 86, 58, and 52 percent, respectively, with 15-year-old seed. The forbs, sweetanise and sainfoin (Onobr:ychis sativa), expressed 34 and 27 percent germination with 10-year-old seed. A majority of the grass species had good germination after 7 to 9 years. The wheatgrasses (Agropyron ssp.), in general, exhibited high germination for 7 to 8 years. Sixyear-old Reed foxtail (Alopecurus arundinaceus) seed germinated at 86 percent, while meadow foxtail germinated at 91 percent after 13 years. Meadow brome (Bromus erectus) showed 96 percent germination with 2-year-old seed and 91 percent with 9-year-old seed. Orchardgrass Wactylis glomerata) germination with 5-, 7-, and SO-yearold seed was 89, 82, and 44 percent, respectively. Germination of 82 percent after 7 years and 23 percent after 15 years was obtained with Russian wildrye (Psathyrostachys junceas) seed. The fescues-meadow, sheep CFestuca ovina), and red CF. rubra)-all had germination of over 63 percent with 7-year-old seed. Prairiejunegrass (/{oeleria cristata) showed 40 percent germination 10 years after harvest. Perennial ryegrass (Lolium perenne) had 93 percent with 6-year-old seed. Of the grasses tested, timothy (J'hleum pratense) exhibited considerable longevity with 98 percent germination from 10-year-old seed. Bulbous bluegrass (Poa bulbosa) and alkali sacaton (Sporobolus airoides) exhibited identical germination (99 percent) with 7-year-old seed. Nine-year-old subalpine needlegrass (Stipa columbiana) seed germinated at 63 percent. Mountain rye (Secale montanum) had 80 percent germination from new seed and 50 percent with 17-year-old seed. VARIATION IN GERMINATION In these studies, difference in germination between sources of a number of shrub, forb, and grass species was found. Comparing results from more than one seed source per species in study 1 (table 1) and additional sources of the same species in study 2 (table 2) demonstrated variations in germination between sources. Two sources of squawapple, one in study 1 and the other in study 2, both held germination for 7 years. A third source in study 2, however, had significantly less germination in the eighth year. Considerable variation between various shrub species' seed sources, especially sources from differing environmental conditions, has been reported for fourwing saltbush, winterfat, rubber rabbitbrush, big sagebrush, and bitterbrush (Meyer 1989, 1990; Meyer and Monsen 1990; Meyer and others 1987; Meyer and Pendleton 1990; Stevens and others 1977). After 13 years, germination of one Palmer penstemon seed source (study 2) did not change significantly (89 to 82 percent). An additional source of Palmer penstemon STUDY 3 RESULTS Even though the source of seed is not the same for most species, results (table 3) can be used as a general guide in estimating longevity for many species. For a good number of the species in this study, there are no other available extended longevity data. Retention of germinability with years of storage varied considerably with the shrubs. Big saltbush Wriple% lentiformis) expressed more longevity (92 percent in 14 years) than did desert holly saltbush (A. hymenelytra) (57 percent in 2 years). Five-year-old seed of pygmy peashrub (Caragana pygmaea) germinated at 85 percent and 10 year-old seed at 67 percent. Wedgeleaf ceanothus (Ceanothus cuneatus) had considerably more germination (63 percent) with 10-year-old seed than did Fendler ceanothus (C. fendleri) (27 percent) with 14-year-old seed. Bladdersenna (Colutea 263 Hafenrichter, A. L.; Foster, R. B.; Schwendiman, J. L. 1965. Effect of storage at four locations in the West on longevity of forage seeds. Agronomy Journal. 57: 143-147. Kay, B. L.; Graves, W. L.; Pergler, C. C. 1984. Storage of seed of Mojave Desert shrubs. Journal of Seed Technology. 9:20-28. Kay, B. L.; Graves, W. L.; Young, J. A. 1988. Long-term storage of desert seed. Mojave Revegetation Notes 23. Davis, CA: University of California. 22 p. Kitchen, S. C.; Meyer, S. E. 1991. Seed germination of intermountain penstemon influenced by stratification and GAS treatments. Journal of Environmental Horticulture. 9: 51-60. Kitchen, S. C.; Meyer, S. E. 1992. Temperature mediated changes in seed dormancy and light requirements for Penstemon palmeri (scrophulariaceae). Great Basin Naturalist. 52: 53-58. Little, E. L. 1937. Viability of seeds of southern New Mexico range grasses. Res. Note SW-6. Albuquerque, NM: U.S. Department of Agriculture, Forest Service, Southwestern Region. 4 p. Meyer, S. E. 1989. Warm pretreatment effects on antelope bitterbrush (Purshia tridentata) germination response to chilling. Northwest Science. 63(4): 146-153. Meyer, S. E. 1990. Seed source difference in germination under snowpack in northern Utah. In: Proceedings, 1990 Billings symposium on disturbed lands in the West. Publ. 9003. Bozeman, MT: Montana State University and Reclamation Research Unit: 184-191. Meyer, S. E.; Jorgensen, G. L.; McArthur, E. D. 1989. Variation in germination response to temperature in rubber rabbitbrush (Chrysothamnus nauseosus: Asteraceae) and its ecological significance. American Journal of Botany. 76: 981-991. Meyer, S. E.; McArthur, E. D.; Monson, S. B.1987. Intraspecific variation in germination patterns of rangeland shrubs and its relationship to seeding success. In: Ecology of rangeland plants; Proceedings of the symposium. Washington, DC: U.S. Department of Agriculture, Agricultural Research Service: 82-89. Meyer, S. E.; McArthur, E. D.; Monson, S. B. 1990. Germination response ofArtemisia tridentata (Asteraceae) to light and chill: Patterns ofbetween-population variation. Botanical Gazette. 151(2): 176-183. Meyer, S. E.; Monson, S. B. 1990. Seed-source differences in initial establishment for big sagebrush and rubber rabbitbrush. In: McArthur, E. D.; Romney, E. M.; Smith, S. D.; Tueller, P. T., comps. Proceedingssymposium on cheatgrass invasion, shrub die-off, and other aspects of shrub biology and management. Gen. Tech. Rep. INT-276. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 200-208. Meyer, S. E.; Pendleton, R. L. 1990. Seed germination biology of spineless hopsage; between-population differences in dormancy and response to temperature. In: McArthur, E. D.; Romney, E. M.; Smith, S. D.; Tueller, P. T., comps. Proceedings-symposium on cheatgrass invasion, shrub die-off, and other aspects of shrub biology and management. Gen. Tech. Rep. INT-276. Ogden, in study 1 held its germination for only 5 years (83 to 79 percent) and then dropped to 65 percent by the seventh year. Kitchen and Meyer (1991, 1992) reported considerable variation in germination between sources of Palmer penstemon and other Intermountain penstemons. Some variations were found between cicer milkvetch sources. One source in study 2 had 51 percent germination the year of harvest and significantly increased to 89 percent in 7 years. The second source showed a slight decrease (73 to 65 percent) over 8 years. Young and Evans (1982) worked with a number of coolseason range grasses and found variation between varieties and sources within the same species. In our studies we had one source of smooth brome whose germination did not significantly change in 5 years (91 to 94 percent) (study 2), and one that showed 70 percent germination with 2-year-old seed and signifiCantly dropped to 52 percent with 4 years of storage. There were three sources of tall wheatgrass in study 2. One source demonstrated afterripening by increasing its germination between the first and ninth year (72 to 91 percent). Germination increased slightly but not significantly in 9 years (85 to 87 percent and 85 to 92 percent) for the two other sources. Two sources of Indian ricegrass in study 2 demonstrated somewhat differing results. The first source started out with 9 percent germination and 14 years later had increased to 49 percent. The second source started out at 55 percent and increased slightly to 63 percent in 10 years. SUMMARY Each species exhibited its own unique germination characteristics that varied with source and age. Seed of many species can be obtained and stored for various lengths of time and still retain acceptable germinability. Some species' germination increased with age; others' decreased with age. Because percentage germination is a measure of the number of seed per 100 that will germinate and have the potential to produce a seedling, all new seed and seed that has been in storage should be tested by an official seed laboratory (Stevens and Meyer 1990) prior to purchase and seeding. Purchase price and seeding rate should be acijusted to the most recent germination test. ACKNOWLEDGMENTS We gratefully acknowledge A. Perry Plummer, Don R. Christensen, Stephen B. Monsen, James N. Davis, and E. D. McArthur for initiating the study, and assisting in data analysis. Funds were provided through Federal Aid in Wildlife Restoration Project W-82-R (Study 4) and Intermountain Research Station, Forest Service, U.S. Department of Agriculture, Ogden, UT. REFERENCES Giunta, B. C.; Jorgensen, K. R.; Plummer, A. P.; Stevens, R. 1978. Antelope bitterbrush-an important wildland shrub. No. 78-12. Salt Lake City, UT: Utah State Division of Wildlife Resources. 48 p. 264 UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 187-192. Plummer, A P.; Christensen, D. R.; Jorgensen, K. R.; Stevens, R. 1970. Highlights, results, and accomplishments of game range restoration studies, 1970. Publ. 70-3. Salt Lake City, UT: Utah Department ofFish and Game. 94p. Plummer, A P.; Christensen, D. R.; Monsen, S. B. 1968. Restoring big-game range in Utah. Publ. 68-3. Salt Lake City, UT: Utah Department ofFish and Game. 183 p. SAS. 1982. Users guide: statistics. Cary, NC: SAS Institute. 584p. Springfield, H. W. 1968. Cold storage helps winterfat seed retain viability. Journal of Range Management. 21(6): 401-402. Springfield, H. W. 1970. Germination and establishment offourwing saltbush in the Southwest. Res. Pap. RM-55. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 48 p. Springfield, H. W. 1973. Winterfat fruits and seeds retain high viability three years in cold storage. Res. Note RM-233. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 3 p. Springfield, H. W. 1974. Winterfat seeds viable after eight years of refrigerated storage. Journal ofRange Management. 27(1): 78. Stevens, R.; Davis, J. N.; Jorgensen, K. R. 1981. Viability of seed from thirty-two shrub and forb species through fifteen years of warehouse storage. Great Basin Naturalist. 41(3): 274-277. Stevens, R.; Giunta, B. C.; Jorgensen, K. R.; Plummer, A P. 1977. Winterfat (Ceratoides lcuuua). PubL 72-2. Salt Lake City, UT: Utah Division ofWJldlife Resources. 41 p. Stevens, R.; Meyer, S. E. 1990. Seed quality testing for range and wildland species. Rangelands. 12: 341-346. Tiedemann, A. R.; Pond, F. W. 1967. Viability of grass seed after long periods of uncontrolled storage. Journal of Range Management. 20(4): 261-262. U.S. Department of Agriculture, Forest Service. 1948. Woody-plant seed manual. Misc. Publ. 654. Washington, DC: U.S. Department of Agriculture, Forest Service. 416p. U.S. Department of Agriculture, Forest Service. 1974. Seeds ofwoodyplants in the United States. Agric. Handb. 450. Washington, DC: U.S. Department of Agriculture, Forest Service. 883 p. Van Haverbeke, D. F. 1989. Effect of storage temperature on germination of seeds of twenty sources of ponderosa pine. Res. Note RM-490. Fort Collins, CO: U.S. Department of Agriculture, Forest Service. Rocky Mountain Forest and Range Experiment Station. 4 p. Young, J. A; Evans, R. A 1982. Temperature profiles for germination of cool season range grasses. ARR-W-27. Washington, DC: U.S. Department of Agriculture, Agricultural Research Service. 91 p. 265