Ecotones Between Artemisia nova and A. tridentata Plant Communities in the Buckskin Mountains of Western Nevada James A. Young Charlie C. Clements Abstract—The habitat restrictions between the larger woody sagebrush (Artemisia) species and the lower growing species have long intrigued ecologists in the Great Basin. In the Buckskin Mountains of west-central Nevada, ecotones between big sagebrush (A. tridentata) and black sagebrush (A. nova) communities are extremely abrupt and distinct, with no intermixing of the two types. These differences are conditioned by different soils and are reversible with changing edaphic development through erosion or deposition. The relative distribution of the taller, woody sagebrush (Artemisia) species in comparison to the shorter or dwarf woody species has long intrigued plant ecologists. As typified by big sagebrush (A. tridentata), the taller woody species of sagebrush are often landscape characterizing in their distribution. The shorter or dwarf woody species usually have more limited distribution and are often associated with specific edaphic situations (Zamora and Tueller 1973). It was proposed by the soil scientist Harry Summerfield, that the dwarf sagebrush species low (A. arbuscula) and black sagebrush (A. nova) often occupied sites where the surface soil horizons had been removed by erosion on a recent geologic time scale (Young and others 1996). The woody sagebrush species occupy such a variety of habitats over such a large and climatically diverse geographical area, there are many exceptions to this hypothesis, but it provides an innovative approach for considering the ecotones between black and big sagebrush communities. Our purpose was to investigate ecotones between big and black sagebrush to provide insight into the distribution of the two species in the Buckskin Mountains of west-central Nevada. Study Area _____________________ The Buckskin Mountains are located 100 km southeast of Reno, Nevada in the rain shadow of the Sierra Nevada and the Pine Nut Mountains (Morrison 1964). This is the portion of the Carson Desert in which Billings (1945) suggested that shadscale (Atriplex confertifolia)-dominated salt desert shrub communities occurred because of atmospheric In: McArthur, E. Durant; Ostler, W. Kent; Wambolt, Carl L., comps. 1999. Proceedings: shrubland ecotones; 1998 August 12–14; Ephraim, UT. Proc. RMRS-P-11. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. James A. Young is a range scientist and Charlie D. Clements a wildlife biologist for U.S. Department of Agriculture, Agricultural Research ServIce, 920 Valley Road, Reno, NV 89512. USDA Forest Service Proceedings RMRS-P-11. 1999 drought rather than the concentration of soluble salts in the soil. The Buckskin Mountains are a very structurally complex fault-block mountain range typical of the Great Basin (Hudson and Oriel 1979). The bulk of the range is composed of repeated flows of volcanic tephra. The incandescent volcanic ash flows moved across the landscape like water as they were deposited. The heat from each new flow converted the surface of previous landscapes to an indurate tuff (Proffett and Proffett 1976). These hardened strata tend to form a series of cliffs above eroding talus slopes. The landscape of the Buckskin Mountains has existed for a sufficient period of time for considerable soil development to have occurred (Blank and others 1996). On north facing slopes, clay-textured soils occur at the soil surface. Apparently, previously existing surface horizons have eroded over geologic time. This same relationship is apparent on many older alluvial fans that spill from the mountain slopes and canyons. The areas with clay-textured surface soils support black sagebrush plant communities (Young and Palmquist 1992). South facing slopes have soils with no horizonation or only color changes indicating weak horizonation. Shadscalehorsebrush (Tetradymia glabrata)/desert needlegrass (Achnatherum speciosum) plant communities form the vegetation on the south slopes. Big sagebrush communities do not occur on the residual soils of the Buckskin Mountains. Erosion and Deposition __________ Degrading System On the north end of the Buckskin Mountains, a fairly substantial drainage has eroded into the stony clay-textured soils of an alluvial fan complex. Under current climatic conditions, the drainage would only have running surface flow for brief periods immediately after exceptional high intensity short-duration summer thunderstorms. Sometime in the recent geologic past, there was sufficient flow to erode a flat bottomed arroyo 3 m deep into the clay. The width of the arroyo averages 15 m. The erosion surface is partially filled with irregularly stratified erosional products reflecting the mixed volcanic make up of the watershed. The fill ranges in texture from sand to substantially sized boulders. There is no evidence of soil development. The ecotone between the black sagebrush communities of the clay textured fans and the big sagebrush community of the arroyo fill is very striking because of the difference in stature of the two sagebrush species. The ecotone between 29 the two communities is absolutely abrupt, reflecting the soils change. Most of the shrub species in the arroyo community are at least as tall as big sagebrush. Antelope bitterbrush (Purshia tridentata), gray rabbitbrush (Chrysothamnus nauseosus), and true sage (Salvia dorrii) occur with big sagebrush in the arroyo. It is obvious from the ecotone between the two communities that black sagebrush in the Buckskin Mountains is exclusively adapted to the clay-textured soils where it is the only woody species except for a diminutive form of green rabbitbrush (Chrysothamnus viscidiflorus), occasional plants of Nevada ephedra (Ephedra nevadennsis), and horsebrush (Young and Palmquist 1992). Extensive excavation, with mechanical equipment to the bottom of the arroyo fill, failed to provide any evidence of prolonged subsurface flow of moisture in the fill. The sandtextured nature of the fine material in the fill certainly offers contrasting moisture retention characteristics compared to the predominantly clay-textured soils of the older fans. Mutual Exclusion of Artemisia _____ The bulk of the Buckskin Mountain range is tilting to the west (Hudson and Oriel 1979). The mouths of the canyons on the east side of the range (long axis running north and south) spill alluvium on the top of old fans rather than eroding incised arroyos into the old surfaces. The broad basin on the eastern flank of the range, known as Lincoln Flat, supports both big and black sagebrush plant communities. The black sagebrush communities occur on the erosional surfaces with clay-textured surface horizons. The big sagebrush communities, without antelope bitterbrush or true sage, occur on the recent alluvial surfaces. Fulstone Canyon spills on to Lincoln Flat along the northeastern portion of the Buckskin Range. It has a very large watershed, made larger by the capture of several adjoining canyons along a fault strike that parallels the main axis of the range. At the mouth of the canyon, there is an area of several hectares where coarse-textured recent alluvium has buried older erosional surfaces with black sagebrush communities. There is a succession of big sagebrush communities from seedling dominance to maturing plants on this deposition area. There are no black sagebrush plants, seedling or adult, in these recent deposition areas despite the clay-textured horizon being present under moderately or thinly bedded recent erosional products. Big sagebrush plants thrive on the clay-textured soils as long as there is a surface horizon of coarser-textured material. This suggests that seedbed quality may be the factor excluding big sagebrush from the black sagebrush communities. In the Buckskin Mountains we know very little about the seedling ecology of black sagebrush plants, because seedlings are very rare in natural communities (Young and Palmquist 1992). The black sagebrush plants grow on mounds, which is a common feature for many shrubs in the Great Basin (Charley and West 1975) . Young and Palmquist (1992) suggested, based on a very small sample, that in the Buckskin Mountains black sagebrush seedlings establish in the interspace between mounds and new mounds build around the seedlings. Obviously, because of litter deposition and nutrient cycling, the old mounds would appear to be the most desirable seedbed in the black sagebrush communities, but when black sagebrush plants become senescent and die the mound becomes densely colonized by the native perennial grass squirreltail (Elymus elymoides) and by the exotic annual cheatgrass (Bromus tectorum). These herbaceous species preempt the environmental potential released by the death of the shrub, making it very difficult for black sagebrush to become established on the mound. Eventually the herbaceous species exhaust the nitrogen-enriched soils of the old mound and the mound erodes. Considering the great linear extent of the ecotones between big and black sagebrush it would seem logical that occasionally plants of one species would establish in the community dominated by the other species. This seems never to happen in the Buckskin Mountains. Big sagebrush plants would be highly visible in black sagebrush communities. This near total lack of the dominant species crossing of the soils ecotones suggests the seedbed potential of the contrasting soils and the inherent potential of the seeds of the species must be mutually exclusive. The environmental parameters that control establishment and persistence of woody sagebrush species on an extensive array of environmental potentials is an intriguing subject for further study in the Intermountain Area. In the case of the sagebrush communities in the Buckskin Mountains, the hypothesis that erosion of surface soil horizons is related to dominance of dwarf species of sagebrush may have merit and deserves investigation in other areas of the Great Basin. Lack of Hybridization ____________ References _____________________ Artemisia and Chrysothamnus are the two woody genera with species common to both communities across the abrupt ecotonal boundary. They occur exclusively as distinct species with no evidence of hybrid swarms. Perhaps the environmental gradient is so abrupt and occurs over such a short distance there is no environmental potential suitable for intermediate forms even if the genetic make up and breeding system of the species permitted hybridization. Billings, W. D. 1945. The plant associations of the Carson desert region, western Nevada, BuIter Univer. Botanical Studies. 7: 89-123. Blank, R. R.; Young, J. A.; Lugaski, T. 1996. Podogenesis on talus slopes, the Buckskin Range, Nevada, USA. Eeoderma. 71: 121-142. Charley, J. L.; West, N. E. 1975. Plant-induced soil chemical patterns in some shrub-dominated semi-desert ecosystems of Utah. J. Plant Ecology. 63: 945-964. Agrading System 30 USDA Forest Service Proceedings RMRS-P-11. 1999 Hudson, D. M.; Oriel, W. M. 1979. Geologic map of the Buckskin Mountain Range, Nevada. Map 64. Nevada Bureau of Mines, Reno,NV. Morrison, R. B. 1964. Lake Lahonton: Geology of southern Carson desert, Nevada. Prof. Paper 401. Geological Survey, Washington, DC. 477 p. Proffett, J. M., Jr.; Proffett, B. H. 1976. Stratigraphy of the Tertiary ash flow tuffs in the Yerington district, Nevada. Report 27. Nevada Bureau of Mines, Reno, NV. 64 p. USDA Forest Service Proceedings RMRS-P-11. 1999 Young, J. A.; Palmquist, D. E. 1992. Plant age/size distribution in black sagebrush (Artemisia nova): effects on community structure. Great Basin Naturalist. 52:313-320. Young, J. A., Longland, W. S.; Blank, R. R. 1996. Management of woody species of Artemisia in western North America. In: Caligari, P. D. S.; Hind, D. J. N. (eds.) Compositae: Biology and Utilization. Proc. International Compositae Conference, Kew. Kew, UK. Royal Botanical Gardens: 277-290. Zamora, B. T.; Tueller, P. T. 1973. Artemisia arbuscula, A. longiloba, and A. nova habitat types in northern Nevada. Great Basin Naturalist. 33: 225-242 31