Great Basin Naturalist

Volume 55 Number 2 Article 7

4-21-1995

Lahontan sagebrush ( arbuscula ssp. longicaulis): a new taxon

Alma H. Winward Range and Watershed Management, Intermountain Region, Forest Service, U.S. Department of Agriculture, Ogden,

E. Durant McArthur Sciences Laboratory, Intermountain Research Station, Forest Service, U.S. Department of Agriculture, Provo, UT

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Recommended Citation Winward, Alma H. and McArthur, E. Durant (1995) "Lahontan sagebrush (Artemisia arbuscula ssp. longicaulis): a new taxon," Great Basin Naturalist: Vol. 55 : No. 2 , Article 7. Available at: https://scholarsarchive.byu.edu/gbn/vol55/iss2/7

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Great Basin Naturalist 55(2), © 1995, pp. 151-157

LAHONTAN SAGEBRUSH (ARTEMISIAARBUSCULA SSP. LONGICAULIS): A NEW TAXON

Alma H. "'inwardl and E. Durant McArthur2

ABSTRACT.-A new subspecies of Artemisia arbuscula is described, A arhuscula ssp. longicaulis Winward & McArthur, ssp. nov. This taxon is a landscape dominant in portions ofnorthwestern Nevada and adjacent and at elevations from 1050 to 2000 m on shallow or argillic (clayey) soils. It differs from A arbuscula ssp. arhuscula in its long floral stalks and large leaves. Morphological, chemical, ecological, and cytological data suggest that it is of hybrid origin. It is hexaploid (6x). We hypothesize that 2x A. arhuscula ssp. arbuscula and 4x A tridentata ssp. wyomin­ gensis are its parents.

Key wonts: Nevada, , chemotaxonomy, allopolyploid, hybrid, Tridentatae.

In preliminary reports we (Wim.vard et al. sagebrush (A. nova) was best treated as a sub­ 1986, 1991) provided a brief description of a species of A. arbuscula, but Beetle (1960) new taxon ofArtemisia found in northwestern restored it to Nelsons (1900) original species Nevada and adjacent California and Oregon. status. Beetle (1960) recognized two sub­ We suggested referencing it by the common species of A. arbuscula, arbuscula and ther~ name Lahontan sagebrush pending a formal mopala. His treatment has been generally description. This paper provides that formal accepted (Winward and Tisdale 1977, McArthur description and details concerning its taxono­ et al. 1981, Shultz 1986), although Winward my, distribution, general ecology, and origin. (1980) has observed an unusual variant of A. arbuscula in eastern Oregon that reaches a TAXONOMY height of 1 m. He suggested that further taxo­ nomic treatment of A. arbuscula would be The new taxon is a member of subgenus appropriate. Tridentatae of Artemisia, the true sagebrushes During the past few decades fieldworkers (Beetle 1960, McArtbur et al. 1981, Shultz in western Nevada have observed a sagebrush 1986). We suggested (Winward et al. 1986) that that does not fit the existing Artemisia taxo­ this taxon may have originated as a hybrid with nomic keys. Brunner (1972) termed this sage­ parental lines consisting of low and big sage­ brush (A. arbuscula and A. tridentata). Because brush "wide-lobe" with the comment, "Dr. ofits general morphology and ecology, we sug­ Beetle feels this may be an ecotype ofA. triden­ gested that it be considered a subspecies ofA. tata ssp. wyomingensis. I concur." Others have ar"buscula. Further studies indicate that this ne"v referred to it as "wonder sagebrush," "junk taxon is in fact best treated as a subspecies of sagebrush," or "N" sagebrush (Winward et al. A. arbuscula. 1986). Accessions of two populations (Trough The type specimen of A. arbuscula came Springs, Humboldt County, NY, cultures Ul from a collection along the arid plains of the and U58 and Leonard Creek, Humboldt Lewis (now known as the Snake) River (Nuttall County, NY, culture U55) of this taxon were 1841). Subsequent workers bave submerged the established in common gardens of the Forest species as a subspecies of big sagebrush, A. Service's Shrubland Biology and Restoration tridentata ssp. arbuscula (Hall and Clements Research vVork Unit at several locations 1923), or, in contrast, recognized a number of around central Utah; there they were treated races and subspecies within the species (Ward as an ecotype ofA. tridentata ssp. wyorningensis 1953, Beetle 1960). Ward proposed that black following Beetle and Brunner (Brunner 1972;

JRange and Watershed Management, Intermountain Region, Forest Service, u.s. Department ofAgriculture. Ogden, UT 84401­ 2Shrub Sciences LaboratOlY, Intermountain Research Station, Forest Service, U.S, Department ofAgliculture, Provo, UT 84506.

151 152 GREAT BASIN NATURALIST [Volume 55 e.g., McArthur and Plummer 1978, Welch and • Nevada, Pershing Co., 6.4 km west of McArthur 1979, 1981, 1986, McArthur et al. Toulon, McArthur & McArthur 1683, 1981, 1985, McArthur and Welch 1982, Welch (SSLP, two sheets); et al. 1986, 1987). The new taxon is a landscape • Nevada, Washoe Co., Mustang, dominant over much ofits range (Winward et al. McArthur & McArthur 1684, (SSLP); 1986), and both domestic and wild animals • Oregon, Lake Co., 32 km east ofAdell, feed extensively on it (Brunner 1972, Welch and Sanderson & McArthur 1590, (SSLP); McArthur 1986, Winward et aJ. 1986, Welch • Oregon, Malheur Co., near McDermitt, et al. 1987). Nevada, Winward, s.n. 31 October 1986, (OGDF, two sheets, SSLP). Description Artemi.sia arbuscula ssp. longicaulis Win­ DISTRIBUTION AND ECOLOGY ward & McArthur ssp. nov. Similis A. arbu.scula Artemisia arbuscula ssp. longicaulis occurs ssp. arbuscula sed ramis f10ralibus multo lon­ on several hundred thousand hectares in gioribus et foliis maguioribus differt (Similar northwestern Nevada and in adjacent areas of to A, arbuscula ssp. arbuscula except flower California and Oregon at elevations from stalks are much longer and leaves are larger). about 1050 to 2000 m (Fig. 1). It often occurs The longer flower stalks and larger leaves in pure stands. It may also share dominance also differentiate ssp. longicaulis from ssp. with other sagebrush taxa such as big sage­ thermopola, which differs from ssp. arbuscula brush (A. tridentata ssp. tridentata and wyo­ and langicaulis by having deeply trifid leaves mingensis), low sagebrush (A. arbuscula ssp. (Beetle 1959). arbuscula), and black sagebrush (A. nova). At We chose the common name Lahontan sage­ lower elevations it is interspersed with salt brush because the old shorelines of Pleisto­ desert shrub species such as shadscale cene Lake Lahontan are one of the centers of (Atriplex confertifolia), Bailey greasewood (Sar­ its current distribution and may have provided cobatus baileyi), Mormon tea (Ephedra spp.), the ecological setting for the taxon's origin and budsage (Artemisia spinescens), Shockley's development (Winward et aJ. 1986, 1991). desert thorn (Lycium shockleyi), and horse­ Type: Toulon, Persbing County, Nevada, brush (Tetradymia spp.). Except for Artemisia, USA, 1053 m, S. C. Sanderson and E. D. our taxonomy follows Welsh et al. (1993) and McArthur 1593, 21 August 1986. Holotype: Mozingo (1987). The most common grass BRY. Isotypes: OGDF, RENO, SSLp, and understory species at upper-elevation Lahontan UTe. Other specimens examined: sagebrush sites is bluebunch wheatgrass • Nevada, Douglas Co., Topaz Lake, (Elymus spicatus). At lower elevations Thurber Sanderson & McArthur 1594, (SSLP, and desert needlegrasses (Stipa thurberiana four sheets); and S. speciosa), and Indian ricegrass (Stipa • Nevada, Humboldt Co., Golconda, hymenoides), bottlebrnsh squirreltail (Elymu$ Plummer s.n., 1985, (SSLP); elymoides), and Sandberg bluegrass (Poa • Nevada, Humboldt Co., Leonard secunda) are more common. Areas supporting Creek, Plummer & McArthur, s.n., 3 A. arbuscula ssp. longicaulis receive between October 1975, culture U55, (SSLP); 175 and 350 mm ofprecipitation annually with • Nevada, Humboldt Co., Trough most as winter precipitation. The frost-free Springs, Jackson Mountains, Plummer, season ranges from 90 to 110 days. Lahontan Brunner, & McArthur, s.n., 3 October sagebrush grows most commonly on Aridisols, 1975, culture Ul, (SSLP); but at upper elevations it also occurs on • Nevada, Humboldt Co., Trougb Mollisols. Soil Conservation Service, U.S. De­ Springs, Jackson Mountains, McArthur partment ofAgriculture, personnel have located 1532, culture Ul, (SSLP); A. arbuscula ssp. longicaulis on at least 17 soil • Nevada, Humboldt Co., Trout Creek series. Generally, these soils have low available Basin, Jackson Mountains, McArthur water-holding capacities and a shallow depth 1501, (SSLP, two sheets); to an argillic horizon and/or bedrock. These • Nevada, Lyon Co., Dayton, Sanderson soils are similar to those of low sagebrnsh (A. & McArthur 1595, (SSLP, two sheets); arbuscula ssp. arbuscula) communities 1995] LAHONTAN SAGEBRUSH, A NEW TAXON 153

OREGON McDermitt

• :! Alturas ~ i? ~ .4 Winnemucca

•Susanville

Austin• ,

NEVADA

Fig L Extent ofthe known distribution of Artemisia arbu.scula ssp. longicaulis. 154 GREAT BASIN NATURAliST [Volume 55

(Fosberg and Hironaka 1964, Zamora and McArthur et al. 1981, 1988). The follOwing Tueller 1973, G. K. Brackley and C. A. taxa are thought to have originated as hybrids: Plummer personal communication). • A. argillosa (A. cana ssp. viscidula X A. General distributions of the three sub­ longiloba-Beetle 1959), species of A. arbuscula are as follows: ssp. • A. tddenlata ssp. spicifo1'Tnis (A. arbuscu.la~ western and eastern tridentala ssp. vaseyana X A. cana ssp. Utah to eastern and northeastern viscidula-Beetle 1959, Goodrich et al. California; ssp. themwpoln, western Wyoming 1985, McArthur and Goodrich 1986), and adjacent and northern Utah to • A. tridentata ssp. xericensis (A. tridentata northern Nevada and eastern Oregon; ssp. ssp. tridentata X A. tddentata ssp. longicaulis, western Nevada, extending into vaseyana-Winward 1970, Rosentreter adjacent California and Oregon. and Kelsey 1991), • A. tddentata ssp. wyomingeosis (A. SUPPORTING DATA AND DISCUSSION tddenlata ssp. !ridentala X A. tddenlata ssp. vaseyana with perhaps some Morphological, chemical, and cytological involvement with A. nova as well­ data are consistent with the hypothesis that A. Beetle and Young 1965, Winward 1975, arbu.scuW ssp. longicatw. is of hybrid origin, McArthur 1983). with A. arbu.scula ssp. arbu.scula as one parent Figure 2 shows a polygonal representation of and A. tridentala ssp. wyomingensis as the morphological features of the new subspecies other parent. Hybridization and introgression and its putative parents, including permanent are thought to have been important in the evo­ leaf width, length, and length/width ratio and lutionary development and diJIerentiation of flower and vegetative stalk lengths. Values are Tridentatae taxa (Ward 1953, Beetle 1960, shown in Table 1. Morphological differences

A. arbuscwa ssp. longicaulU

A. arbMculo. ssp. arbuscula A A. tridentata ssp. wyomingensu . ... - . -. - ... .

••

\ ' , \ , \ " \ " \ ' , \ , \ '. ' ..... \ .. ' .... \ .-.':' .... B , -."" \. " '\ "-.;: - , ~'. \ , \ \ \ E - c -----

D

Fig. 2. Polygonal KIUPh comparing m01-phological features ofArtemWia arbuscula ssp.longk;atdi.s and its putative par­ ents, A. arbuscula ssp. arbwcuJ.a and A. tridentata ssp. wyomingen.sis. Data from Table 1: A = flower stalk length. B = vegetative stalk length; C = leafwidth; D = leafllw ratio; E = leaflength. 1995J LAHONTAN SAGEBRUSH, A NEW TAXON 155

TABLE 1. Morphological measurements (means + SD) ofA. arbuscula ssp. arhuscula (ARARa), A. arbuscula ssp. longi­ caulis (ARARl), and A. tridentata ssp. wyomingen.sis (ARTRw)a, ARARa ARARI ARTRw Flower stalk length (em) 16.4+ 1.7 24.5+3A 10.4 + 4.2 Vegetative stalk length (em) 3.9 + 0.8 5.7+ 1.2 4.7+ 2,4 Leaflength (mm) 5.9+ 1.1 7.3 + 1.0 8.3 + 2.4 Leafwidth (mm) 3.0 + 0.4 3A + 0.4 2.7 + 0.8 LeaflIw ratio 2.0 2.2 3.1 aOata for ARAR" are from the following collections: California, Lassen Co., Shaffer Mountain, Srulderson & McArthur 1591 (SSLP); Nevada, Eureka Co., T\lSCarOnl Mountains, 40 km north of Cmtin, Frischknecht 210, (SSLP); NeV'dda, Humboldt OJ., 13 km northwest ofPm...dise Valley Town, Holmgren 128, (SSLP); Nevada, Lander Co" Bade Creek, Thiyabe National Forest, Goodrich 8868, (OGDF); Nevada, Lander Co., Toiyabe Range, Toiyahe Natiooal Forest, Goodrich 9966, (OGDF); Nevada, Nye OJ., Toiyabe Range, Toiyabe National Forest, Goodrich 12201, (OGDF); Nevada, Pershing Co., 40 km northeast ofWinnemucca, Winward s.n., 25 Cktober 1984, (OGDF); Nevad

TABLE 2. Percent frequency ofshared phenolics for A. nova (ARNO), A. tridentata ssp. tridentata (ARTRt), A. tridenta­ ta ssp. VlIscyana (AHTRv), A. tridentata ssp. wyomingensis (ARTRw), A. arbu.scula ssp. arhuscula (ARARa), and A. arbua­ cula ssp.longicaulis (ARABl); data from Wilt et al. (1992). ARNO ARTRt ARTW ARTRw ARARa ARABI ARNO 52 49 57 59 52 ARTRt 52 76 48 52 ARTRv 48 58 53 ARTRw 52 54 ARAR" 54 wngicaulis is an a11opolyploid derivative from FOSBERG, M. A., AND M. HIHONAKA. 1964. Soil properties diploid, n = 9, A. arbuscula ssp. arbuscula, and afhding the distribution of big and low sagebrush communities in southern Idaho. Forage physi~ tetraploid, n = 18, A. tridentata ssp. wyomin­ ology and soil-vegetation relationships. Special gensis. A. m-huscula ssp. arbuscula occurs in Publication of the American Society of Agronomy 5: both diploid and tetraploid popnlations in the 230-236. norfuwestenl Nevada area. Tetraploid A. triden­ GOODRICH, S., E. 0. McAnTHUR, AND A. H. WINWARD. tata ssp. wyorningensis also occurs there 1985. A new combination and a new variety in Artem.­ isia tridentata. Creat Basin Naturalist 45: 99-104. (McArthur and Sanderson in review). Popula­ HALL, H. M., AND F. E. CLEMENTS. 1923. The phylogenet~ tion dynamics of sageblush populations migrat­ ic method in taxonomy, the North American species ing in response to climatic change during the of Artemisia, Chrysothamnus, and At:riplex. Publica­ various phases of ancient Lake Lahontan and tions of the Carnegie Institute of Washington 326: other pluvial lakes of the Lahontan basin 1-3,55. HANKS, 0. L, E. D. McARTHUR, R. STEVENS, AND A. P. (Morrison 1965) could have provided the PLUMMEH. 1973. Chromatographic characteristics opportunity for the origin and establishment and phylogenetic relationships of Artemisia, section ofA. arbuscula ssp. longicaulis. Morphological, Tridentatae. USDA Forest Service Research Paper cytological, and chemical data are consistent INT~141. U.S. Department of Agriculture, Forest with the hybrid origin hypothesis. Service, Intermountain Forest and Range Ex.peri­ ment Station, Ogden, UT. 24 pp. McARTHUR, E. 0. 1983. Taxonomy, origin, and distrihu~ ACKNOWLEDGMENTS tion ofbig sagebmsh (Artemisia tri.dentata) and allies (subgenus Tridentatae). Pages 3-13 in K. L. Johnson, We thank Gary Brackley, Jim Brunner, Chu editor, Proceedings of the first Utah shrub ecology Ge-lin, Shere] Goodrich, Craig Plummer, workshop. Utah State University, Logan. 50 pp. McARTHUR, E. D., AND S. GOODHICH. 1986. Artemisia tri­ Stewart Sanderson, Marty Wilt, and the late dentata ssp. spicifonnis: distribution and taxonomic Perry Plummer for their help in various phas­ placement. Pages 55--57 in E. 0. McArthur and R L. es of this study. We thank field personnel of Welch, compilers, Proceedings-symposium of the the Soil Conservation Service, U.S. Department biology of Artem.isia and Chrysothamnus. USDA of Agriculture, Nevada offices, who provided Forest Service General Technical Report INT~200. Intermountain Research Station, Ogden, ur. 398 much of the information for Figure 1. We also pp. appreciate assistance provided by the McARTHUR, E. D., AND A. P. PLUMMER. 1978. Biogeography Pittman-Robertson W82R Wildlife Habitat and management of important western : a Project and u.s. Department of Agriculture case study, section Iridentate of Artemisia. Great Cooperative State Research Service Grant 91­ Basin Naturalist Memoirs 2: 229-243. McARTHUR, E. D., AND S. C. SANDERSON. In review. 38300-6157. Additional chromosome counts in subgenus Tridenta­ tae ofArtemisia with implications. LITERATURE CITED McARTI-WU, E. D., AND B. L. WELCH. 1982. Growth rate differences among big sagebrush (Artemi8ia tridenta­ BEETLE, A. A. 1959. New names within the section tal subspecies. Journal of Range Management 35: Tridentatae ofArtemisia. Rhodora 61: 82-85. 396--4Q1. __,.-' 1960. A study of sagebrush, the section Tridentar McARTHUR, E. D., C. L. POPE, AND D. C. FREEMAN. 1981. tae of Artemisia. Wyoming Agricultural Experiment Chromosomal studies of subgenus Tridentatae of Station Bulletin 368: 1--83. Artemisia: evidence for autopolyploidy. American BEETLE, A. A., AND A. YOUNG. 1965. A third subspecies in Journal ofBotany 68: 589-605. the Artemisia tridentata complex. Rhodora 67: McARTHUR, E. D., B. L. WELCH, AND D. L. NELSON. ~(15~~(16. 1985. Developing improved cultivars of sagebrushes BllUNNER, J. R. 1972. Observations on Artemisia in Nevada, and other composite shrubs. Pages 188-196 in J. R. JOllrnal ofRange Management 25: 205-208. Carlson and E. 0. McArthur, editors, Symposium: 1995] LAHONTAN SAGEBRUSH, A NEW TAXON 157

range plant improvement, proceedings-selected a superior selection of low-elevation mountain big papers presented at the 38th annual meeting. Society sagebrush. USDA Forest Service Research Paper for Range Management, Denver, CO. 220 pp. INT-370. Intermountain Research Station, Ogden, McARTHUR, E. D., B. L WELCH, AND S. C. SANDERSON. UT. 10 pp. 1988. Natural and artificial hybridization between WELCH, B. L., E. D. McARTHUR, AND R. L. RODRIGUEZ. big sagebrush (Artemisia tridentata) subspecies. 1987. Variation in utilization ofbig sagebrush acces­ Joumal ofHeredity 79: 268-276. sions by wintering sheep. Joumal of Range Manage­ MORRISON, R. B. 1965. Quaternary geology of the Great ment 40: 113-115. Basin. Pages 265-285 in H. E. Wright, Jr., and D. G. WELSH, S. L., N. D. ATWOOD, S. GOODRICH, AND L. C. Frey, editors, The Quaternary of the . HIGGINS. 1993. A Utah flora. 2nd edition, revised. Princeton University Press, Princeton, NJ. 922 pp. Brigham Young University Print Services, Provo, MOZINGO, H. N. 1987. Shrubs of the Great Basin. Univer­ UT.986pp. sity ofNevada Press, Reno. 342 pp. WILT, F. M., J. D. GEDDES, R. V TAM:\1A, G. C. MILLER, NELSON, A. 1900. New from Wyoming-XII. Bul­ AND R. L. EVERErr. 1992. Interspecific variation of letin ofthe Ton'ey Botanical Club 27: 258-274. phenolic concentrations in persistent leaves among NELSON, D. L., AND R. G. KREBILL. 1981. A sagebrush wilt six taxa from subgenus Tridentatae of Artemisia disease ofunknown cause. Great Basin Naturalist 41: (). Biochemical Systematics and Ecology 184-191. 20,41-52. NLTTALL, T. 1841. Descriptions ofnew species and genera WINWARD, A. H. 1970. Taxonomic and ecological relation­ of plants in the natural order of the Compositae. ships of the big sagebrush complex in Idaho. Unpub­ Transactions of the American Philosophical Society lished dissertation, University of Idaho, Moscow. 80 11,7,357-453. pp. ROSENTRETER, R., AND R. G. KELSEY. 1991. Xeric big __~. 1975. Evolutionary development ofArtemisia tri­ sagebrush, a new subspecies in the Artemisia triden­ dentata taxa. Page 163 in H. C. Stutz, editor, tata complex. Journal of Range Management 44: Proceedings of the symposia and workshop, wild­ 330-335. land shrubs. Brigham Young University, Provo, UT. SHULTZ, L. M. 1986. Taxonomic and geographic limits of 168 pp. Artemisia subgenus Tridentatae (Beetle) McArthur __,;e' 1980. Taxonomy and ecology of sagebrush in (Asteraceae: Anthemideae). Pages 20-28 in E. D. Oregon. Oregon Agricultural Experiment Station McArthur and B. L. Welch, compilers, Proceed­ Bulletin 642: 1-15. ings-symposium of the biology of Artemisia and \VINWARD, A. H., AND E. W. TISDALE. 1977. Taxonomy of Chrysothamnus. USDA Forest Service General Tech­ the Artemisia tridentata complex in Idaho. Univer­ nical Report INT-200. Intermountain Research sity of Idaho College of Agriculture. Idaho Agricul­ Station, Ogden, UT. 398 pp. tural Experiment Station Bulletin 19; 1-15. 'WARD, G. H. 1953. Artemisia, section Seriphidium, in WINWARD, A. H., E. D. McARTHUR, AND G. K. BRACKLEY. North America, a cytotaxonomic study. Contributions 1991. Lahontan sagebrush-another sagebrush in ofthe Dudley Herbarium 4: 155-205. Nevada. Page 13 in Abstracts, 44th annual meeting, WELCH, B. L., AND E. D. MCARTHUR. 1979. Variations in Society for Range Management, Denver, CO. 45 pp. winter levels of crude protein among Arlemisia tri­ WINWARD, A. H., E. D. McARTHUR, D. A. KAFFER, C. A. dentata subspecies grown in a unifonn garden. Journal PLUMMER, AND G. K. BRACKLEY. 1986. Another sage­ ofRange Management 32: 467-4.69. brush in Nevada. Technical Note: Range Nevada 44. ___. 1981. Variation of monoterpenoid content among USDA Soil Conservation SeIVice, Reno, NV 2 pp. subspecies and accessions of Artemisia tridentata WOOLF, C. M. 1969. Principles of biometry. D. Van grown in a unifonn garden. Journal ofRange Manage­ Nostrand Co., Inc., Princeton, NJ. 359 pp. ment 34: 380--384. ZAMORA, B., AND P. T. TUELLER. 1973. Artemisia arbuscula, __~. 1986. Wintering mule deer preference for 21 acces­ A. longiloba, and A. nova habitat types in northern sions of big sagebrush. Great Basin Naturalist 46: Nevada. Great Basin Naturalist 33: 225-232. 281-286. WELCH, B. L., E. D. McARTHUR, D. L. NELSON, J. C. Received 1 March 1994 PEDERSON, AND J. N. DAVIS. 1986. 'Hobble Creek'- Accepted 11 August 1994