The Great Basin Naturalist

Published at Provo, Utah, by B righam Young U niversity

ISSN 0017-3614

Volume 52 Ju n e 1992 N o . 2

Great Basin Naturalist52(2), pp, 95-121

RED BUTTE CANYON RESEARCH NATURAL AREA: HISTORY, FLORA, GEOLOGY, CLIMATE, AND ECOLOGY

James R. Ehleringer , Lais A. AnKw, Ted Aroow, Irving B. McNulty , and Norman C. Negus

Abstract — Red Butte Canyon is a protected near pristine canyon entering Salt Lake Valiev. Utah It contains a well-developed riparian zone and a perennial stream; hillside vegetation ranges from grasslands on the lower limits to Douglas-fir and aspen stands at the upper elevations In this paper we describe the history of human impact, natural history aspects of climate, geology, and ecology and fauna! and floral information for key species in the canyon. The role and importance of Research Natu ral Areas is discussed, particularly with respect to the need to protect Red Butte Canyon—one of the few remaining undisturbed riparian ecosystems in the Intermountain West.

Key wordsi grassland, IntvrmoHntain West, oak-maple, plant adaptationr Red Butte Canyon, Research Natural Area, riparian ecology.

Red Butte Canyon, one of many canyons in stantia] value to society. These might include the Wasatch Range of Utah, opens westward unique geological features, endangered plant into Salt Lake Valley, immediately east of the and animal species, or areas of particular value University of Utah (Fig. 1). Like most canyons for scientific research as baseline bench marks along the Wasatch Front, it is a grassland at the of ecosystems that have been largely destroyed lowest elevations, is forested at its upper end, by human impact. In the case of Red Butte and has a perennial stream. What makes this Canyon, the RNA designation was given canyon u nusual is its history. The canyon was the because this canyon is one of the few remaining watershed for Fort Douglas, the U.S. Army post (if not the last) undisturbed watersheds in the built in 1862 that overlooked Salt Lake City. As Great Basin. The U.S. Forest Service report a protected watershed, these lands were, for the proposing that Red Butte Canyon be declared a most part, kept free from grazing, farming, and Research Natural Area described ihe canyon as other human-impact activities. When the U.S. . a living museum and biological librarv of a Army declared these lands surplus in 1969, the size that exists nowhere else in the Great Basin U.S. Forest Service assumed responsibility for . . . an invaluable bench mark in ecological the canvon. Since that time. Red Butte Can von J J time." The Red Butte Canyon RNA is unique has been kept in its protected state and desig­ because it is a relatively undisturbed watershed nated a Research Natural Area (RNA). adjacent to a major metropolitan area (Salt Lake The Research Natural Area designation Valley). To protect this valuable resource, access denotes an area that has been set aside because to the Red Butte Canyon RNA has been largely it contains unusual or unique features of sub­ restricted to scientific investigators. One of the 96 G r ea t Ba sin N a t u r a l ist [V olum e 52

Salt Lake City'////////// Intl. Airport ////////// ' / / > / / / / / / / * / w /s/s/sr/// Pinecrest f/f/ff/SJf/'fS, rs / / / / i rr f f s t / f f f h /////////j' ////V

JsSSS/SJ/SSfSSSA/SSffSSSfffSSS/SSfJ-SjrssSfSsS fftf.

< 8 0 c mile ///////■ kilometers « / y / y / Mitt C^eek Cnyn

Fig. 1, Location of Red Butte Canyon and oHier sites referred to in text,

goals of the RNA Program is to protect and of biotic communities along an elevation gradi­ preserve a representative array of all significant ent from about 1530 m (5020 ft) on the west end natural ecosystems and their inherent processes to more than 2510 in (8235 ft) at the crest. as baseline areas. A second goal is to conduct The purpose o f this paper is to provide a brief research on ecological processes in these areas description of the his tor)-; flora, geology, cli­ to learn more about the functioning of natural mate, and ecology of diis unusual and valuable versus manipulated or disturbed ecosystems. resource. There is increasing interest in Red Research activities in the Red Butte Canyon Butte Canyon, in part by scientific investigators RNA are directed at both of these goals: under* because of its utility as a protected, undisturbed standing basic ecological processes (physiologi­ watershed, anti in part by curious citizens from cal adaptation, drought adaptation, nutrient the nearby Salt Lake Valley. Yet. there Inis not cycling, etc.) and also the impact of humans on been an overall reference available for those our canyons through both airborne (air pollu­ interested in general features of the canyon or tion, acid rain, etc.) and land-related (grazing, past ecological studies within the canyon. Most human traffic, etc.) activities. The latter are of the information on Red Butte Canyon is conducted through comparison of Red Butte scattered. With the closure of Fort Douglas in with other canyons along the Wasatch Range. 1991, many of the historical records will becom e In size, Red Butte Canyon is relatively small more difficult to access. It is hoped that the compared with other drainages along the, synthesis presented in this paper will provide Wasatch Front. The drainage basin covers an area of approximately 20.8 km2 (5140 acres) the necessary background for those interested (Fig. 2). The drainage arises on the east from a in the history and ecology of the Red Butte minor divide between City Creek and Emigra­ Canyon RNA. Irving McNulty first summarizes tion canyons and drains to the west. The canyon the history of the canyon, followed by Ted lias two main forks (Knowltons and Parleys) and Amows description of geology and soils, James many side canyons. Near the canyon base, a Ehleringer contributed the hydrology, climate, reservoir was constructed earlier this century to and plant ecology sections. The section on vas­ provide a more stable water supply to Fort cular flora was prepared by Lois A mow, and Douglas. The diversity of slope and aspect com ­ Norman Negus wrote the mammalian and avian binations of the terrain contributes to a variety fauna sections.

1992] R e d B u t t e C a n y o n R e s e a r c h N a tu ra l A r ea 99

Table I. Description of geological formations in Red protect the water supply of Fort Douglas. This Butte Canyon, law preven ted any sale of land in the canyon or 1 ...... ■ ■ i ■■ ■ I. - ■ ■■ k m . II ■ a. ftllMlI II, II, I i ■■ ...... fcl *j, . Cenozoic- era, Quaternary' system, Holocene series further watershed development. In 1906 the fa Fhod-phin alhmwm. Sand, cobbly to silty, dark gray at U.S. Army built a dam on Red Butte Creek to top; grading downward to medium to light gray, sandy to supply additional water for Fort Douglas. The cobbly gravel; locally bouideiy. fe Engime/'ed fill Selected earth material that has been present dam was constructed between 1928 and emplaced and compacted, 1930, and the reservoir provided water for Fort Cenozoic era, Quaternary and Tertiary systems, Douglas until its closure in 199L Holocene and Pleistocene series There are no grazing records available for fg Alluvial-fan deposits. Bouldery to clayey silt, dark gray to Red Butte Canyon prior to 1909, by which time brown; rocks angular to subrounded. Id landslide deposits. Composition similar to material the U nited States had acquired title to most of up slope. the land in the canyon, Cottam and Evans Mesozoic era, Jurassic system (1945) reported evidence of some gully erosion Jtc Ttuin Creek Limestone. Brownish gray and pale gray to occurring in the canyon prior to 1909 and pale yellowish gray silty limestone, intercalated with assumed it was due to overgrazing. Although we greenish gray shale. lack quantitative data, there are a few isolated Mesozoic era, Jurassic? and Driassic? systems incidents indicating the occurrence of grazing, JTn Nugget Sandstone. Pale pinkish buff, fine- to medium- grained, well-sorted sandstone that weathers orange- i ncluding an 1S54 report of a young man drown­ brown. Massive outcrops form the ridge called Hed Butte. ing in a flash flood in Red Butte Canyon while Mesozoic era, Triassic system herding animals. Over fort)' head of oxen used Tau Ankareh Formation, upper member. Reddish brown, to haul sandstone from the quarry in the iate reddish purple, grayish red, or bright red shale, siltstone, 1800s remained in the canyon during that time. and sandstone. Tag Ankareh Formation, Gattra Grit Member. White to pale In 1869 the War Department appointed a puiple, thick-bedded, crossbedded, pebbly quartzite. herder to control loose cattle grazing on Fort Forms a prominent white ledge for lon$ distances. Douglas and in the canyon. In 1890 three squat­ Tam Ankat'eh Formation, Mahogany Member, Reddish brown, reddish purple, grayish red, or bright red shale, ters had settled into the canyon., and their forty siltstone, and sandstone. head of cattle were grazing in the Parleys Fork Tt Thaunes Formation. Medium to light gray, fossiliferous, area before being evicted. By 1909 the Army locafly nodular limestone., limy siltstone, and sandstone, had built a gate at the mouth of the canyon to Tiv Woodside Shale. Grayish red, grayish purple, or bright red shale and siltstone. control access, thus further protecting the Paleozoic era, Permian system . watershed. Although this did not prevent occa­ Ppc Park City Formation and related strata- Fossiliferous sional animals from wandering into the canyon sandy limestone, calcareous sandstone, and a medial from adjacent canyons, it did reduce both their phosphatic shale tongue. numbers and their length of stay. Consequently, Paleozoic era, Pennsylvanian system most of the canyon has not been grazed by cattle Pw Webei' Quartzite. Pale tan to nearly white, fine- to medium-grained, crossbedded quartzite and medium or sheep through most of this century. gray to pale gray limestone. Portions of the upper reaches of the canyon Pro Round Valley Limestone. Pale gray limestone with pale were timbered In 1848, when a road was built ;ray siltstone partings. Contains pale pinkish chert that along the canyon bottom, it was repotted that forms irregular nodules. there was an abundance of timber suitable for Paleozoic era, Mississippian system Mdo Doughnut Formation. Medium gray, thin-bedded fence poles. Later The Church of Jesus Christ limestone with pods of dark gray to black chert and of Latter-day Saints built a bowery on. Temple abundant brachiopods and bryozoa. Square in downtown Salt Lake City in. the 1850s Mgb Great Blue Forination. Thick-bedded, locally cliff- with wood obtained from Table Mound forming, pale gray, fine-grained limestone. Mh Humbug Formation. Alternating, tan-weathering, limy (between Knowltons Fork and Beaver Canyon). sandstone and limestone or dolomite. In 1863 the Army constructed 34 buildings at Md Deseret Limestone. Thick ledges of dolomite and lime­ Fort Douglas from ‘‘timber hauled from the stone with moderately abundant lenses and pods of dark chert. canyons/’ but there is no indication as to how Paleozoic era much timber came from Red Butte Canyon. P Paleozoic rocks, undifferentiated, However, apparently not many timber-size trees were available in the lower canyon as indicated by a pioneer who built a log cabin in the canyon. He stated he had to travel five miles up the 100 G r ea t B asin N a t u r a list [Volume 52 canyon to obtain enough logs for tlie cabin in the canyon to the general public. In 1987 the canyon early 1860s. was opened to the public in late spring for There are no available records oi fires that several days; tills weekend opening attracted may have occurred in the canyon, In 1988 a fire over 5000 visitors and led to a trampling on from Emigration Canyon spread into the upper vegetation along the main road in the canyon. headwaters of Red Butte Creek before it was This opening was repeated in 1988 and contained. The land was subsequently reseeded attracted 1100 people. Currently the State with native species by the U.S. Forest Service. Arboretum at the University of Utah conducts Land ownership within the canyon changed natural history education classes ( — 10 individu­ several times during the late 1800s and early als per group) in the lower portions of the 1900s. Land occupied by Fort Douglas in 1862 canyon. Limited deer hunting has been permit­ was officially given to the U.S. Army in 1867 ted by the Forest Service each fall, but the when President Johnson withdrew four square impact of the hunts is unknown. A Red Butte miles from public domain for the use of the Steering Committee, consisting of representa­ Army. However, this included only a small por­ tives from the Forest Sendee, the University of tion of the mouth of Red Butte Canyon, The Salt Utah, and other government agencies con­ Lake Rock Company, which quarried most of cerned with preservation of natural areas, is the sandstone in the canyon, owned part of the involved in making decisions pertinent to the canyon, and the Union Pacific Railroad Co. jurisdiction and management of the Red Butte acquired four sections in the lower portions of Canyon Research N atural Area. the canyon in the lS60s, Smaller portions of the The history of Red Butte Canyon., with the canyon were claimed by private individuals exception of the quarrying activity and some under the Homestead Act of 1862. Such claims grazing in the past century, is largely a history of preservation. The U.S. Army at Fort Douglas could be acquired easily under this act, which was concerned with the protection of the water­ was very liberal and required only a small claim fee. Gradually, between 1884 and 1909, through shed and gradually acquired sufficient control to protect it. The U .S. Forest Service declared a combination of acts of Congress, exchanges of the entire canyon a Research. Natural Area and property, and outright purchases, Fort Douglas thus insured its protection for the future as a obtained title to most of the canyon, by 1896 and bench mark of riparian and shrub ecosystems in almost the entire canyon by 1909, Only three the Intermountain West. small parcels of a total oi less than 90 hectares (—200 acres) are still privately owned today, and G e o l o g y these are close to the margins of tlie canyon. In 1969 the U.S. Department of Defense relin­ The rocks underlying Red Butte Canyon quished ownership of Red Butte Canyon. The range in age from recent Holocene deposits of U.S. Forest Service is now responsible for these our time; to Mississippian rocks that are about lands. Tlie Forest Service recognized tlie natu­ 360 million years old. Holocene and Pleistocene ral state of the area had been preserved through deposits are unconsolidated, consisting mostly many years of closure to the public and desig­ of landslides or alluvium deposited by existing nated Red Butte Canyon a Research Natural streams. Their aerial distribution is shown in Area in 1970. By definition such areas are tracts Figure 3, and a description of the deposits is of land that have not been strongly impacted by given in Table J.. . human-related activities such as logging or graz­ The older rocks range in age from ing by domestic livestock. They are permanently Mississippian to Jurassic, a span of about 220 protected from devastation by humans so they million years. They are all consolidated now, but may serve as reference areas for research and originally they were formed as deposits in ed u cation . oceans or inland seas or as sand dunes in an arid Red Butte Canyon has served as a research environment. No rocks representing the site for biologists for over fifty years and will approximately 140 million years between the continue to do so in the future. Public education end of Jurassic time and the Holocene are pres­ about conservation and the need for the public ent in Red Butte Canyon. Either they were to better understand the importance of never deposited or they have been eroded. Research Natural Areas are major concerns. The consolidated rocks in most parts of the Recently the Forest Service briefly opened the

G r ea t Ba sin N a tu r a list [V olum e 52

Township IN. Range IE seel ion

2 kilometers

Fig. 5, Soils map of Red Butte Canyon. See Table 2 for a description of abbreviations. Adapted from Woodward et a], (1974), until flow is collected in the reservoir located flows driven by snowmelt followed by very near the base of the canyon. The stream has much reduced flows derived from groundwater created a narrow-based canyon with sides rising throughout the remainder of the year (Fig. 6). abruptly at an average slope of about 35 degrees Spring melt flow, which is typically an order of to the north and about 40 degrees to the south. magnitude greater than other periods of the Immediately upstream of the reservoir is a U.S. year, peaks in May and persists for 6-8 weeks. Geological Survey Hydrologic Bench Mark Sta­ The average monthly stream flow rate during tion. This gaging station has been maintained by May is 0.416 m'Vsec (14.7 ft'Vsec), By Septem­ the U.S. Geological Survey since October .1963, ber, the lowest average mondily flow rate, Prior to that, the Corps o f Engineers, U.S. Army, stream discharge has decreased to 0.058 m3/sec recorded monthly discharge at this location (2.0 ftVsec). Mean, stream flow rates do not beginning in January 1942. increase during the summer months, although The average monthly discharge (1964-88) is nearly one-fourth of the annual precipitation 0.133 m'Vsec (—4.7 ft3/sec) as it enters the res­ falls during this period. ervoir at 1646 in (5400 ft) elevation (U.S. Geo­ Average monthly stream flow values, how­ logical Survey records). The stream flow ever, hide much of the stream dynamics and exhibits a straightforward annual pattern, char­ resultant impact on riparian vegetation. On a acteristic of this geographic region— high spring daily basis, stream flows can vary tremendously 1.992] R e d B u t t e C a n y o n R e s e a r c h N a tu r a l A r e a 103

Table 2. Description of unifcs on the soils map of Red overland flow was substantial. This was by far Butte Canyon. the greatest discharge rate in recent times, AGG Agassiz- association, very steep. 40-70 percent having eclipsed the previous maximum single slopes; moderately permeable, well drained. Agassiz—35 day rate of 1.70 m 3/sec (60.0 ft "/sec) measured percent, very cobbly silt loam on ridges and convex areas on 18 May 1975 (U.S. Geological Survey of upper slopes. Picayune—-55 percent, noncalcareous variant, gravelly loam in concave areas and in draws. R ecords). Other soils— 10 percent. The unusually high stream discharge rate in BCG Brad very rocky loamy sand, 40 to SO percent May 1983 is of particular significance because slopes. Very permeable, extremely well drained. Very of its impact on stream geomorphology and rocky, cobbly, loamy sand; dark reddish-brown; shallow. adjacent vegetation. Tlie high, flows quickly BEG Braclshaw-Agassfe association, steep. 40-70 per­ scoured the streambed, taking out beaver dams, cent slopes; moderately permeable, well drained. eroding stream banks, knocking down riparian Bradshaw—55 percent, very cobbly siltdoam in slightly trees, and causing massive erosion. Gullies 5-10 concave areas. Agassis.—35 percent, very cobbly sitf-loain in convex areas and ridgetops where soil is shallow. Other m (16-33 ft) deep were cut into permanent soils— 10 percent. streamheds in Knowltons Fork and throughout DGG Deer Creek-Picayune association, steep. 30-60 Red Butte Creek. Sediment flow associated percent slopes; moderately permeable, well drained. with tins record stream discharge was in excess Deer Creek—55 percent; loam; very dark brown; deep of 269 metric tons ( -“593,000 lbs) per day in on very steep, north- and northeast-facing mountain slopes. Picayune—35 percent; gravelly day loam; very mid-May (compared to typical spring melt con­ dark brown, deep, calcareous on west-facing slopes. centrations of 1 metric ton 1—2200 lbs] per day) Other soils— 10 percent. (U.S. Geological Survey Records); this resulted EMG Emigration very cobbly loam, 40 to 70 percent in a delta formation at the mouth of Red Butte slopes. Moderately permeable, well drained. Cobbly Reservoir. Prior to the 1982-83 winter, no delta Joarn; facing south; dar k, grayish brown; shallow; patches of bedrock. had existed. The delta was soon —SO m (~ 100 ft) long. By 1990 the delta had fanned out more HGG Harkers-Wallsburg association, steep. Moder­ ately permeable, well drained. Harkers—55 percent, than 60 m into the reservoir. The heavy winter loam, 6—40 percent slopes, very dark brown, deep in rains of 1982- 83 saturated soils all along the drainageways and concave areas of slope faces. Walls- Wasatch Front, and landslides were common. burg -35 percent, very cobbly loam, 30-70 percent Red Butte Canyon was no exception. Slope slopes, on ridges and convex areas of slopes where bed­ rock is near the surface, very dark grayish brown, shallow. sloughing, which, killed the overlying perennial, Other soils—.10 percent. vegetation, was common throughout tlie canyon, HHF Ilarkere soils, 6 to 40 percent slopes. Moderately No doubt this compounded the stream sedi­ permeable, well drained. Loam and cobbly loam, on ment load during the spring of 1983 and for sloping old alluvial fans and steep mountain slopes. several years thereafter. In 1990 signs of the LSC Lucky Star gravelly loam, 40 to 60 percent 1982—83 slope sloughing were still clearly obvi­ slopes. Moderately permeable, well drained. Very dark ous in Knowltons Fork as well as in the upper grayish brown, deep on northerly slpes, and. lower portions of the main canyon. Natural Mw Mixed alluvial land. Poorly drained, highly stratified revegetation of both riparian and slope vegeta­ mixed alluvium on undulating, gently sloping, and nearly level flood plains. tion types has occurred since these floods. In particular, Acer negundo (boxelder) and Salix exigu-a (willow) have increased in frequency in during snowinelt, depending on air tempera­ the newly deposited alluvium along the stream- sides (Donovan and Ehleringer 1,991). Recov­ tures and snowpack depth, (primarily that of ery of the sloughed slopes, which were for the upper Red Butte Canyon and Knowltons Fork) . most part covered by A. g randidentatum (bigtooth The 19S2—S3 winter was one of unusually high maple) and Quercus gambelii (Gambel oak), has precipitation along the Wasatch Front. Heavy proceeded at a slower rate, with those slopes still snows in mid-May 1983 were followed by dominated by herbaceous species. equally unusual warm temperatures at the end As part of the bench mark analysis, the U.S. of the month. As a consequence, stream flow Geological Survey monitors several major aspects rates peaked at record values. On 28 May 1983, of stream quality in addition to stream discharge, Red Butte Creek crested at a discharge rate including water temperature, suspended sedi­ exceeding 2.97 rn'Vsec (104,9 itVsec) (stream ment, and chemical quality. Included with flow was above the maximum gage height), and chemical quality are specific conductance, pH, 104 G r e a t B a sin N a t u r a l ist [V olum e 52

1 .5 0

CO co 1 .2 5 ' £

0 . 5 0 - p

© 0 . 2 5 - C/5

0 \9 % %% %*%>

Fig. 6 Mean monthly discharge rates of Red Butte Creek just before it enters Red Butte Reservoir, Large and small tick marks indicate end-of-year and inid-year points, respectively. Data are from U.S. Geological Survey records,

dissolved oxygen concentration, coliform bacte­ water sources used by plants for growth (see ria, and ionic and dissolved elemental concen­ discussion below). Hydrogen isotope ratios trations (ammonium, arsenic, beryllium, cadmium, (ratio of D/H of a sample to that of a standard) calcium, carbonate, chloride, chromium, cobalt, are measured relative to an ocean water stan­ copper, fluoride, iron, lead, lithium, magnesium, dard; samples lighter than ocean water have less manganese, mercury; molybdenum, nickel, deuterium and are therefore negative in their nitrate, nitrite, phosphate, potassium, selenium, values. Over the four-year measurement period silver, sodium, sulfate, strontium, vanadium, (1988-91), hydrogen isotope ratios of stream and zinc). The stream itself is strongly alkaline waters have averaged near —122%channel (Bond 1979). values occurring during spring snowmelt. Typi­ Summertime stream flow represents cally the hydrogen isotope ratio of winter storm groundwater discharge, while the spring flows events (snow) is more negative than that of result primarily from snowmelt at higher eleva­ summer storms, The hydrogen isotope ratios of tions. Not all of the groundwater originating wells and springs near Pinecrest (immediately from upper-elevation sources enters the stream east of Red Butte Canyon) are — 132%C', slightly before it leaves the canyon. Tracing the possible more negative than Red Butte Creek (Dawson sources of water into stream, and therefore that and Ehleringer 199.1), and suggest that a frac­ water which is available to plants, is possible by tion of the groundwater originating from the analyzi ng the isotopic composition of that water. upper portions of the canyon may persist as The deuterium :( II or D) to hydrogen (!TI) underflow and does not enter the creek before ratios of stream waters have been measured leaving the watershed. Hely et al. (1971) indi­ since June 1988 at the USGS Bench Mark sta­ cated that substantial fracturing occurs in the tion and at the mouth of Parleys Fork by the bedrock of Red Butte Canyon, which would Stable Isotope Ratio Facility for Environmental have the effect of increasing groundwater loss Research at the University of Utah (Dawson and from the canyon through these layers and not Ehleringer 1991). These naturally occurring via stream discharge. stable isotopes of hydrogen provide long-term Bond (.1977, 1979) investigated nutrient- data that are useful in addressing both long­ con centration patterns of stream flow in Red term regional climatic patterns and the specific Butte Creek. In particular, his studies focused 1992] R e d B u t t e C a n y o n R e s e a r c h N a tu ra l A r e a 105

Table .3. Locations oi weather stations of Red Rutte Canyon, All stations were operated by the U.S. Army between 1942 and 1964, and only precipitation was recorded. The. U.S. Geological Survey has maintained a storage gage at Red Butte #2 since 1964. The Biology Department at the University of Utah lias maintained daily temperature, humidity, and wind speed records at Red Butte #2, Red Butte #4, and Red Butte #6 since 1982. Red Butte #1, while technically outside the canyon, iorrns an integrated part of the weather station complex.

Station Location Latitude Longitude Eievation Period

Red Butte #1 Fort Douglas 40® 46' 110° .50' 1497 m 1942-1964 Relocated to Biology 40° 46' 110°50' 1515 m 1991-present Experimental Carden Red Butte #2 Head of Red Bntte 40"47' n r 48' 1653 m 1942-1964 Reservoir 1982-present Red Butte #3 Along Red Butte Creek 40" 48' n r 47- 1865 m 1942-1952 at Brush .Basin Red Butte #4 Along Red Butte Creek 40° 48' n r 46' 1890 in 1942-1971 100 m west of Beaver 1982-present Canyon Red Butte #5 Parleys Fork 100 m above 40° 47' 111*48' 1753 m 1942-1956 inlet to Red Butte Creek Red Butte #6 U pper end K nowitons F ork; 40* 49’ n r 45' 2195 m 1946-1971 relocated to top of Ells Fork 40° 49f in °4 6 ' 2195 m 1982-present

on relationships between nutrient transport out soonal systems penetrate into northern Utah. of the watershed and stream discharge rates. Mean annual precipitation ranges from about Solute concentration was not necessarily pro­ 500 mm (20 in) at the lower elevation to approx­ portional to stream discharge. Instead, for many imately 900 mtn (35 in) at the higher elevations ions, such as magnesium, sulfate, and chloride, (Hely et al 1971, Bond 1977; Table 3). the relationship was logarithmic. The slopes of Precipitation stations have been monitored these relationships depend on whether stream in Bed Butte Canyon by several groups. The How is increasing (i.e., spring snowmelt) or U.S. Army had six rain gages in operation decreasing, Over the course of the year, a loop between 1942 and 1964 (Table 3). Bond (1977) or directional trajectory was formed by having collected data at several of these stations two different slopes. For most of the major ions, between 1972 and 1974, In addition, the U.S. the trajectory was clockwise; that is, ionic con­ Geological Survey maintained storage gages at centration wa s greater in winter when flow rates Red Butte #2, Red Butte #4, and Red Butte #6 were low than during summer. Plant growth of between 1964 and 1974. Since that time, they the dominant riparian species commences near have maintained a storage gage at Red Butte #2. the end of the snowmelt period, and it is ques­ Within the watershed, daily precipitation as tionable whether riparian species are able to rainfall is collected at each of the weather sta­ utilize the greater nutrient availability during tions: snowfall is not adequately measured by the snowmelt period. After snowmelt, stream the sensors in place. However, these data are discharge is based primarily on groundwater currently collected at Hogle Zoo in Salt Lake input. Nitrate, ammonium, and phosphate con­ City (same elevation as previous Red Butte #1, centrations in Red Butte Creek during ground­ but 4 km south). water discharge are low (Bond 1979). In Variation in annual precipitation within Red contrast, overall concentrations of calcium, Butte Canyon is strongly dependent on eleva­ magnesium, sodium, chloride, and sulfate are tion (Fig. 7). The slope of this relationship is much greater because of parent bedrock char­ similar to that observed for other mountainous acteristics. areas within the Great Basin (Houghton 1969), and precipitation at. the Salt Lake City reporting C l im a t e station ( Salt Lake City International Airport) falls on this relationship. Thus, while lacking Climate within Red Butte Canyon is charac­ continuous precipitation records (or the canyon terized by hot, dry summers and long, cold proper, precipitation records available for Salt winters. Most precipitation occurs in winter and Lake City can be used as a preliminary basis for spring, with the summer rains less predictable estimating mean annual precipitation at differ­ and dependent on the extent to which rnon- ent locations within the canyon. 106 G r e a t Ba s in N a t u r a l ist [V olum e 52

e E aC.

‘SL 0 £ G. 2 n c 1

Elevation, rn

Fig, 7- Relationship between mean Bfinu&i precipitation and ejevation for Red. Butte Canyon storage gages Red Butte #l-#6, Shown also is the mean annua), precipitation for the primary station of Salt Lake City (Salt Lake City International Airport) as the open symbol

30 -

i o pd o L * c a? 20 - P <0 b r 0 t y Pig, 8. Mean monthly air temperature, vapor pressure, and photosynthetically active solar radiation (400-700 nm) 1 £ 10 measured at Red Butte #2 between 1982 and 1990.

18.7 and 31.8 C (66--S9 F) at Red Butte #2, while nighttime minimum temperatures ranged between 5.2 and 16.4 C (41-62 F) {Fig. 9). At the higher-elevalion stations, daytime maximum air temperatures were lower. The difference in maximum temperatures was negatively related to elevation (maximum temperature [°C] - 34.3 Fig. 9. Mean monthly maximum and minimum air tem­ — 0.00494 ■ elevation jm], r = .91) at approxi­ perature at Red Butte #2 {.1653 m elevation), Red Butte #4 mately half the dry adiabatic lapse rate. On the (1890 m elevation), and Red Butte #6 (2195 m elevation) during the growing season between 1982 and 1990. other hand, nighttime minimum temperatures were not related to elevation, because of cool- Air temperatures have been collected trom air drainage effects (Fig. 9), Red Butte #4 is automated weather stations at Red Butte #2, located streamside within the canyon, whereas Red Butte #4, and Red Butte #6 since 1982. the other two stations are above the channel of Mean monthly air temperatures at Red Butte #2 cold air that develops at higher elevations and were below freezing in December and January pours down the canyon at night. As seen in and above 20 C in June, july, and August (Fig. Figure 9, this cold-air drainage effect at Red S). In contrast, mean monthly temperatures at Butte #4 (1890 m [6180 ft] elevation) depressed Red Butte #6 were below freezing only slightly nighttime minimum air temperatures by 4-8 C longer, from November through February, and (7-14 F) below that observed at Red Butte #6 above 20 C in July and August. During the main (2230 m 17292 ft j elevation). growing period {May through September), day­ Photosynthetically active solar radiation time maximum temperatures ranged between (PAR, 400-700 nm), atmospheric vapor pressure, 1992] R e d Bu t t e C a n y o n R e s e a r c h N a tu r a l A r e a 107 and wind speed are also recorded at each of mers as factors producing the variation in the these stations. Between 1982 and 1990, mean vegetative zones of the eastern boundary of the daily total PAR values have exceeded 40 mo) Great Basin, jumper is present in the central m "d“1 (Fig. S), which is typical for mid-latitude Wasatch Range, but only three Utah juniper sites having only moderate cloud cover and little (juniperus osteospeiyna) are known to exist in summer precipitation. This number is quite Red Butte Canyon: a mature tree with a 0.5 m useful not only in estimating the available (1.6 ft) diameter trunk, located on the south photon flux for photosynthesis, but also in pro­ slope of Parleys Fork and nearly obscured by the viding an estimate of the exten t of solar heating more mesophytic vegetation, and two shrublike of the surface, which ultimately affects air tem­ plants 1-1,3 m (3-4 ft) tall growing on the south­ peratures. Elevation has a limited impact on the west divide. PAR values within Red Butte Canyon, since the With few exceptions, notably the naturalized difference in elevation is relatively small. How­ grasses Agrostis stolonifera (redtop bentgrass), ever, we suspect there may be relatively large Bromus tectonim (cheatgrass), and Poa praten- differences in PAR between Red Butte Canyon sis (Kentucky biuegrass), only the most common and Salt Lake City because of increased air indigenous plants that occur in the various plant pollutants within the city that tend to reflect the communities are listed below, primarily because sunlight before it strikes the earths surface. the presence of introduced plants is usually Most notably we would see this as haze or smog dependent on disturbance and tends to fluctu­ within the valley that is lacking once in. the ate accordingly. Some of the more frequently canyon. occurring introduced plants are listed in a sep­ Average monthly atmospheric vapor pres­ arate section. sure at site #2 showed little annual variation, Rip a r ia n c o m m u n it y .— F rom, the point at ranging only about 3 mbar throughout the year which Red Rutte Creek emerges from the (Fig. 8). Other sites exhibited a similar pattern. canyon and throughout the floor of the canyon This parameter is largely affected by large air the streamside vegetation (plants residing in soil mass movements; and since subtropical air kept moist to wet by the stream) consists chiefly masses do not move into this region during the of western water birch (Betula occidentaiis ) and summer, the monthly changes in atmospheric mountain aider (Alnus incana), accompanied, at vapor pressure change little during the course inteivals by usually dense stands of red osier of the year. However, because of the large dogw ood {Cornus sericea) and willow (Salix spp.). annual change in air temperature and the non­ Adjoining the stream along the floor of the linear dependence oi the evaporative gradient canyon below and above the reservoir is an often on temperature, relative humidity levels are densely wooded strip consisting chiefly of substantially lower and evaporative gradients Gambel oak (Quercus garrdmlii), boxelder (A c e r are substantially higher during the summer n e g u n d o ), and bigtooth maple (A c e r g ra n d i- m onths. d e n ta tu m ), many of these trees ranging from 9 to 18 in {30 to 60 ft) or more talL Also included in this plant community are widely scattered Va s c u l a r F l o r a individuals or small populations of cotton woods (.Populus fremontii, P. angustifolia , and P. X From the mouth ol Red Butte Canyon at acuminata), chokecherry (Primus mrginiana ), about 1530 m (5020 ft), its walls rise to their Woods rose (Rosa toaodm), bearberry honey­ highest point—-2510 m (8235 ft)— at the head su ck le (Lonicera involucrata), thimbleberry of Know! tons Fork in the northeast corner of the (Rubus panrifiorus), serviceberry (Amelanchier canyon. Within this modest rise o f980 m (32-15 a ln ifo lia ), western black currant (Hibes hud- ft) occur four distinct plant communities: ripar­ so n ia n u m ), and golden currant (Ribes aureum). ian, grass-forb, oak-maple, and coniferous. Relatively few species of grass and forbs are Pinon-juniper and ponderosa pine communi­ found here, among them* ties, which often occur in this elevationaJ range in Utah (Daubenmire 1943), are not present in Ely mus giaucus blue wildrye Red Butte Canyon. Billings (1951, 1990), in Lonvitium dissectum giant lomatiuiii Mahonia repens discussions of vegetational zonation in the Great (Berberis repens) Oregon gfape Basin, cites a greater incidence of winter Osmorhiza chilerim sweet cicely cyclonic storms and slightly more moist sum­ Poa eompressa Canada biuegrass 108 G r ea t B a sin N a t u r a l ist [V olum e 52

P. pmtensis Kentucky bluegrass (Typha latifolia) covering approximately 0.25 SmUacina stellata wild lily-of-tbe-valky hectare (0.62 acre), only a few scattered clumps S, racemasa false Solomon-seal Solidago oanademis goldenrod remained. According to Forest Sendee person­ nel, these losses would not have been as severe Beaver, once native, were reintroduced into had the beaver dams been active during flood­ Red Butte Canyon in 1928 (Bates 1963) and ing. Species in the following genera are among were active along Red Butte Creek and some of those undoubtedly affected: Eleochmis, Scir- its tributaries for 54 years thereafter. Numerous pusyjuncus, Agrostis, Catahrosa, Desckampsia, marshy areas between elevations of 1645 m Glyceria, Poa, Polypogon, Eqimetum, Angelica, (5400 ft) and 2133 m (7000 ft) were created by Eetxda, Cimta, Heradeurn , Rudbeckia, Soli­ the impoundment of water due to their darn- dago% Barb area, Cardaniine, Nasturtium,. building activities. To prevent the beaver popu­ Rorippa, Lonicera, Cornus, Trifolnim , M en th a, lations from becoming undesirably large, the Nepetaf Lenina, Epilobmm, Habenaria, Pole- Utah Division of Wildlife Resources in 1971 nionmm, Polygonum, Rumex, AconUum, undertook management of the populations. In Ranunculus, Geum, Ribes, Salix, Mirmdus, December 1981 a recommendation was made, V eron ica, and U rtica. based on an analysis of the water supply to Fort The U.S. Forest Service, Salt Lake Ranger Douglas from Red Butte Canyon, that all beaver District, requested the Utah Division of Wild­ be eliminated trom tlie canyon because their life Resources to reintroduce the beaver during feces could contaminate the water with the par­ the summer of 1991. At the time of this publi­ asite Giardia lamblia. Accordingly, in 1982 the cation, beaver had not yet been reintroduced, It colonel in command of Fort Douglas applied for is hoped that with tim e the plant diversity1 typi­ and received from the Utah Division of Wildlife cally associated with beaver dams will be rees­ Resources a permit to remove the beaver from tablish ed . the canyon. Subsequently, all beaver were 'har­ GrasS-fqrr COMMUNITY—According to v ested , ” Stoddart (194.1), the grasslands of northern Bates (1963) studied the impact of beaver on Utah form the southernmost extension of the stream flow in Red Butte Canyon. The vegeta­ Palouse prairie. Of the two communities into tive cover was affected for approximately 91 m which the Palouse prairie is divided, only that (298 ft) on either side of the portion of the dominated by biuebuneh wheatgrass (E lym u s stream in which the beaver were active, and sp ica iu s, originally known as A g ro p tjro n sediment deposited behind the beaver dams in ■S'picatum) occurs in Red Butte Canyon. Rela­ the canyon varied from 0.6 to 2,4 m (2 to 8 ft) in tively large open areas inhabited by grasses and depth. He also noted that the small alluvial forbs, with an occasional big sagebrush {A rtem i­ plains formed by the sediment made it apparent sia tridentata ), squawbush (Rhus trilobata}, and that during periods of high runoff, and perhaps bitterbrush {Purshia tridentata ), are fou n d during normal How, the dams allowed the reten­ chiefly below the 1829 m (6000 ft) contour tion of quantities of suspended materials. Schef- (Kleiner and Harper 1966), although smaller fer (1938), in a report on beaver as upstream grass-forb associations also occur in forest clear­ engineers, ascertained that two beaver dams ings at higher elevations. Some of the more commonly occurring species within the grass- retained 4468 m° (157,786 ft'1) of silt. It is not forb community at lower elevations are; known whether an actual count of the number of beaver dams in Red Butte Canyon was ever Achillea miUifolium milfcil yarrow made; but the environmental change effected Allium acuminatum tapertip onion Ami>msia psilostachya western ragweed by their ultimate displacement during the 1983 Arnbis holboeUii Holboeii rockcress flooding of what had to have been enormous Ai'istida purjmrea quantities oi sediment has been significant. The (A. iongiseta) purple tlireeawn removal of all inactive beaver dams has inevita­ Artemisia ludi)-oiciaiw Louisiana wormwood Astragalus utahensis U tah imlkvetch bly led to the elimination of or significant reduc­ Aster adsceridens everywhere aster tion in the density of some 55 species of typically Bakamorhiza nwtrophylla cutleaf balsaniroot wetland plants from once marshy areas within Balsamorhiza sagittata arrowleaf balsamroot Bromus tectorum. Red Butte Canyon. For example, in 1990 it was cbeatgrass Cirsium uncbdatum gray thistle noted that in an area which once supported a Collomia linearis narrowLeat collomia nearly pure stand of closely spaced cattails Comandra umbellata- bastard toadflax 1992] R e d B u t t e C a n y o n R e s e a r c h N a tu r a l a r e a 109

Crepis acuminata mountain hawks beard Merterisia brvoistyla VV^atch bluebell Cyvwpterus hngip&s long-stalk spfmg-parsley Microseris nu tan s nodding seorzonellu $lymus trachjcmlus Phacelia heterophuJUi varilea) seorpionweed (Agropyron canimtm) s leader wheatgrass Poafendleriam rtiutbongrass Epdobium bmdiycarfmm P. praten#is Kentucky bluegrass (E. ptmculatum) autumn willowherb Senecio i*tfegerrimus Columbia groundsel Erigeron divergem spreading daisy Gatierrezia barothtai broom snakeweed Mountain mahogany (Cercocarpus ledifo - Hedysamm boreal? northern sweetvtjtch Im s) occurs as individual and as scattered, Heliomeris muUiflura mostly small populations, often in association (Viguiem multi-flora) showy goldeneye Lovuitium f ritermtum tern ate lomatium with oak, sagebrush, or other mountain shrubs, Lufnnus (ifgenteus silvery lupine generally on northwest-facing, sparsely vege­ Microstetis gracilti little polecat tated slopes. It can be seen from the main road Phacelia linearis threadleai sco rpionweed through the canyon as small trees against the sky Phlox longijpltd longleaf phlox Poasecunda fP sandbergti) Sandberg bfuegrass along the exposed, rocky, south rim of die Stipa CAimnta net1 die-and- th read canyon, especially toward its western end As Wyethia anqtlexieauUs mulesears low shrubs it occurs sporadically, chiefly on OAK-MAPLE COMMUNITY.— Cainbel oak exposed dry sites above 1980 in (5500 ft). ( Quercus gambdii) is the dominant type of veg­ Big sagebrush {Artemisia tridentata) occurs etation throughout the altitudinal range of the sporadically in drier sites throughout the canyon. It forms what appear to he randomly canyons altitudinal range. Low sagebrush spaced clones throughout much of the area. In (Artemisia arbuscula ) occurs as relatively pure accordance with the moisture regimen, the stands at about 2133 m (7000 ft) along the clones may range from thickets 0.3 in (1 ft) or southeast rim o( the canyon. less in height in dry upland sites to stands of C o n lf er o u s c o m m u n n r.— D o ugl as-fir stately, well-spaced trees in lowland areas. Both (Pseudotsuga menziesii), white fir (Abies con- walls of the canyon support often nearly co lo r), and aspen (Populas tremuioid-es) d o m i­ impenetrable oak in association with bigtoodi nate this community, either in pure or in mixed maple (Acer grandidentatum), die latter grow­ stands, growing chiefly on north- to northeast- ing chiefly in drain age way s. Few species thrive and northwest-facing slopes; the aspen reach as as understory with dense oak cover. The most low as 1706 m (5600 ft) and the firs occur mostly common are Galium aparine (catchweed bed- above 182S m (6000 ft), Achlorophyllous straw) and Mahonia repens (Oregon grape). Corallorhiza spp. (coralroot orchid) arc among Others appearing seasonally under oak are the few plants able to flourish in the shade of Erythronium grandiflorum (dogtooth violet), dense stands of mixed conifers. Many small Clatjtonia hmceolata (lanceleaf spring beauty), trees, shrubs, forbs, and grasses thrive in less HydrophyUtim capitatum (ballhead water leaf), and H. occidentals (western waterleaf), Among dense stands or in openings between stands of plants commonly fringing oak clones are: trees in this community. Among them are: Agoseris plmtca mountain dandelion Acer glahrurn Rocky Mountain maple Apoctjnwn mtdmsawwfolium spreading dogbane Amclanchier alnifom Saskatoon servidebefry Arahis glabra tower mustard Aqutlegia coemlea Colorado columbine Brvmus carmattis mountain hrotne Arnica spp. arnica Com&ndra umhellata bastard toadflax CastilMa spp. Indian paint brush Delphinium nvttaUmnum Nelson larkspur Ceanothus velutinus mountain lilac D&tcurainia pinnatu bine tansy mustard Ehjmus glaucus blue wild rye Eriogonum henideoides. whorled buckwheat Erigenm spedqsus showy fleabane £, racemosum redroot buckwheat Galium spp. bedstraw Ger/mium viseoMssititum sticky geranium Hord&um brachyantiwrum meadow barley Helumthella uniflora one-headed sunflower Lathyrus pmicifloms Utah sweetpea ffefcwnms nrnttiflora Phtisacarjnis malvacetis mallow ninebark (Viguiem mukifbra) hairy goldeneye Poa nervosa Wheeler bluegrass Hydrophylkim spp, waterleaf Pmrrns virginmna chokecherry Koeleria macrantha Ribcs uiscanssimum sticky currant (fi crUtata) Jmiegrass Rubus parviflora thimbleberry Lewopoa kUigii Sambucus spp elderbern/ (Hespervchloa kingii) spike fescue Sorhm scopidina American mountain ash Lomatium dhsectum giant lomatium Sijmphoncarpus i*reopkilu$ mountain snowherry Machaernnfitera canescem hoary aster ThahctTum fendleti Fendler nieadowrue 110 G r e a t B a sin N a tu r a list [V olum e 52

Plants e n d e m ic to Utah.— Only two spe­ least six of them would not ordinarily occur cies occurring in Red Butte Canyon are said to Within the elevagonal limits of the canyon: be endemic to Utah: Angelica wheeleri W ats. Agmsii? semwerticilhta water poJypogon (Mathias and Constance 1944^45! (Wheeler Arasmckw t^ssellata rough fiddleneek angelica) and Erigevon arenatioides (D . C. Angelico pinrmta small-leaved angelica tasselftovwr Eat.) Cray (rock fleabane). Angelica wheehn *Biickdlia grandiflora Castilleja angusUfolw Indian paintbrush lias, however, been collected close to both the Cirsium floatimnH Fiodmaja thistle Idaho and the Nevada boundaries with Utah Cn/ptanifia flavocvlata yellow-eye cryplanth (Albee et al, 1988). Erigeron arenarioides is Deachantjma mespitom tufted hair grass known from Salt Lake. Utah. Tooele* Weber, *Erigeron ghhellus smooth fleabane *Eriogonum ovaltfoUum cushion buckwheat and Box Elder counties (Albee et at. 1968, GHill mustard M, E. Jones - G. striata Taivxactim officinale common dandelion (Lam.) Hitehe. Thlaspi arvense pennycress Juruxis tracyi Rydb. swordleaf rush Tragoppgon duhius goats beard = j. ensifolius Wikst. Veronica anci-gallis- aquaticc water speedwell Taraxacum, laevigatvm common dandelion (Wild.) DC. - T. officinale The incidence of I sat is tinctoria and L in a ria Wiggers d a h n a tica increased greatly between 1970 and Thus, the 5 11 species representing 73 fami­ 1990. lies reported from Red Butte Canyon by Arnow FtORISTIC DIVERSITY.—The following spe­ (1971) can now be placed at 484 species (390 cies were reported from Red Butte Canyon by indigenous and 94 introduced) known to have Cottam and Evans (1945) and by Bates (1963). Not only is the presence of these plants unveri­ fied by herbarium specimens (see Albee et al. *Wr!h ihe assist urictrof rbme and Leita Shts^ curator? of the ieH>aric 1988, which is based on specimens in the herba­ at Brigham Young and Utah State universities, respectively, a herbarium check ria of Brigham Young University, Utah State was nrtiide to be tsrtmn that no Butt*? Caryfm spedinsns cxi^t for those sivdes mvkd with ao astensVthm, iKtxirdiivg to Albee ei (195SI are apt in University, and the University of Utah), but at Butte Ctnyori or its v:c;ntt^r. 1992] R e d B u t t e C a n y o n R e s e a r c h N atural A r ea i l l

upper port ions o f the canyon typically consisting 2200 of umrner-active scrub oak, aspen, and conifer­ ous forest communities (Fig. 10), Composition within each of these communities is not con­ stant, but instead species vary in their impor­ c 2000 o tance within a community type as orientation CC $ and elevation change. These elevation gradients CD 1800 represent a continuum of moisture availability, with high temperatures and low precipitation amounts at lower elevations making conditions more xeric, while slope orientations less south­ 1600 erly in exposure become progressively more mesic within an elevation band. Soil type (Fig. 5) and depth also play a major role in affecting plant distribution by providing variation in the Fig. 10 Distribution, by elevation, of tlie major plant water-holding capacity oi the substrate. The dis­ communities in Red Butte Canyon. tribution of the scrub-oak community to the highest elevations within the canyon is most been present in the canyon at one time or likely related to soil conditions, since at high another. Only two populations present in 1971 elevations scrub oak persists on south-, east-, are definitely known to have been eliminated: and west-facing slopes that would normally be Lactuca biennis (biennial wild lettuce), which expected to be do m inated by aspen if i t were not was introduced into Utah from the north about for the very shallow, rocky soils that typify these 1967 but did not survive; and S olidago elevations within Red Butte Canyon. occidentalis (western golden rod), a single Red Butte Canyon has been largely pro­ st re am side population at the mouth of the tected from grazing since its acquisition by the canyon taken out by the 1983-84 flooding. U.S. Army almost a century ago, The conse­ According to Albee et al. (1988), the 390 quence of this lack of grazing pressure at low'er indigenous species reported from Red Butte elevations is a recovery to near pristine levels, Canyon (Amow 1971) also occur in at least one and this is clearly reflected in the early commu­ other canyon to the south. A mow et al. (1980) nity analyses of Evans (1936) and Cottam and and Albee et al. (1988) i ndicate that roughly 130 Evans (1945). Within the scrub oak and grass­ native plants not found in Red Butte Canyon land communities of Red Butte Canyon and have been collected between an elevation of adjacent Emigration Canyon, a canyon annually 1826 and 2438 m (6000 and SG00 ft) in canyons exposed to sheep grazing, there are large differ­ having a greater altitudinal range in southern ences in plant density (Fig, 1L). Emigration Salt Lake County. This figure indicates that the Canyon was originally described by early pio­ floristic diversity in Red Butte Canyon, while neers as having a dense vegetation at lower greater than that in heavily disturbed Emigra­ elevations. However, grazing not only reduced tion Canyon (Cottam and Evans 1945), is less that cover but also increased the fraction of the than that in canyons farther south. plant cover occupied by ruderal, weedy species Nomenclatural changes since Amow (1971) (Cottam and Evans 1945). While, plant density are listed in the Appendix, in Red Butte Canyon may be greater and weedy species composition lower as a result of reduced Pl a n t E c o lo g y disturbance and grazing, the canyon is not free of these weedy components and historical Vegetation distribution.— A number o f effects (as noted in early sections). Dam con­ studies have focused on describing the vegeta^ struction during the 1920s and other U.S. Army tion distribution within Red Butte Canyon activities within the lower portions of Red Butte (Kleiner and Harper 1966, Swanson, Kleiner, Canyon have resulted in sufficient disturbance and Harper 1966. Kleiner 1967). There is a that many ruderal weedy species, such as strong xeric to mesic elevation gradient, with Grindelia squarrosa (curly gum weed). L a ctu ca lower portions of the canyon dominated by a se rrio la (prickly lettuce), and Polygonum acd- spring-active grassland community and the cu la re (knotweed), are now common. 112 Great Basin Naturalist [Volume 52

Sainuelson (1950) conducted an analysis 30 # B Red Bjtte similar to that ol’Cottam and Evans (1945) on □ Emigration the algal components of the streams in Red Butte and Emigration canyons. He observed that as a result of livestock grazing and human 20 settlement, sediment load and turbidity were ? much greater in Emigration than in Red Butte o Creek. The consequence of this stream-quality difference was the dominance by algal genera in 10 Emigration Creek that are turbidity tolerant, such as OsCillatoria and Phormidium. Con­ versely, in the clear waters of Red Butte Creek filamentous algae, primarily No.stoc. were most 1515 ‘625 1700 2060 common. Overall algal densities were three times greater in Red Butte Creek, owing to the Transect elevation, m greater light penetration into that stream. At the same time, Whitney (1951) compared the dis­ Fig. 11. A nomparison ot the plant cover ill open grass­ tributions of aquatic insects in the two streams. land communities of different elevations in Red Butte and Emigration canyons. Adapted from Cottam and Evans He found that densities of aquatic insects were (1945). greater in Red Butte Creek. Of those insects persisting in Emigration Creek, there was a even though there may be little summer precip­ preponderance of species characterized by gills itation (Dina 1970, Dina and Klikoff 1973). protected from silt, which would better allow Adaptation.—In the nonforested portions them to tolerate the more turbid conditions in of the Intermountain West, plant growth is Emigration Creek, largely restricted to spring and early summer PHENOLOGY.—Plant activity is governed by periods by cold temperatu res du ri ng winter and two parameters; temperature and soil moisture imited water availability during the summer availability. Cold winter temperatures limit (Caldwell 1985, Dobrowolski, Caldwell, and growth activity between November and March Richards 1990. Comstock and Ehleringer 1992). (Caldwell 1985, Comstock and Ehleringer A number of recent reviews have addressed 1992). While a limited number of species, such adaptation characteristics of plants growing in as the early spring ephemeral Ratwnctdus tes- these emironments (Caldwell 19S5, DeLucia ticidatus (bur buttercup), may begin activity and Schlesinger 1990. Smith and Knapp 1990. during warm periods in February; most annuals Smith and Nowak 1990). For the most part, do not begin growth until the warm periods plants within Red Butte Canyon are exposed to a hot, dry environment, with little relief from between snowstorms in early■r March. At lower elevations, a number of herbaceous perennials developing water stress during the summer such as Balsamorhiza macrophtjUa (cutleaf months. The only clear exception to this pattern balsamroot) may begin to leaf out during March, is the series of plants within the riparian com­ but most woody perennials do not leaf out until munities along the canyon bottom. To gain a mid- to late April. The annuals and most herba­ better understanding of this occurrence, many ceous species at lower elevations have com­ of the recent ecological researchers within the pleted growth and reproduction by mid-June Red Butte Canyon RNA have focused on mech­ and then remain dormant until the following anisms by wllich plant species have adapted to autumn or spring (Smedley et al. 1991). In con­ limited water availability. trast, woody species at lowfer elevations remain Among the first ecophvsiological studies was active from April through October, although the that by Dina (1970). who examined water stress vast majority of the growth will occur during the levels of the dominant tree species in the lower spring (Donovan and Ehleringer 1991). At portions ol the canyon: Acer grandidentatum higher elevations, vegetative and reproductive (bigtooth maple), Acer negtmdo (boxelder), growth are delayed until late May or June by Artemisia tridentata (big sagebrush), Purshia cold temperatures. Plants at the higher eleva­ tridentata (bitterbrush), and Quercus gambelii tions will remain active throughout the summer, (Gambel oak). Dina (1970) observed that 1992] Bed Butte Canyon Research Natural Area 113

6 availability are avoidance and tolerance. Avoid­ ance of water stress is accomplished by comple­ o tion of growth and reproductive activities before E o 2 forbs the onset of the summer dr ought, whereas toler­ E ance is associated with the evolution of features E that allow plants to persist through die drought > period. o c Several interesting studies have been con­ Q> ducted in Red Butte Canyon that shed light onto ■ 0mmm 4 — the nature of a plait’s ability to tolerate water 0 stress and persist through time. Treshow and co0) D Harper (1974) examined longevity of herba­ 1 ceous perennials in grass, mountain brush, aspen, and conifer communities throughout the CD 1 canyon. They obseived that life expectancies of dominant herbaceous perennial species, such as 0 Astragalus utahensis (Utah n 11!kvetch), Bcdsa- mofhiza inacropktflla (cutlcaf balsamroot), April May June Hedy santmbo reale (northern sweetvetch), and Wyethw amplexicattlis (mulesears), are rela­ Fie 12. The mean water-use efficiency values for tively short (3-20 years) when compared to the grasses and forbs within the grassland community of Red 'Utte Canyon during main period of tlie growing season. longer-lived (>65 years) grass species, such as Water-use efficiencies were calculated from carbon isotope Agropyron spicatwn (bluebunch wheat grass) discrimination values from Smedley et aJ. (1991) and the and .Stipa eomata (needle-and-thread'. The vapor pressure data in Figure S. inability to persist through successive drought years may be one of the reasons that dicotyle­ midday leaf water potentials of - 30 to — 65 bars donous species have shorter life expectancies develop in perennials occupying slope sites than monocot) !edonous species. Related to this, during late summer, whereas water potentials of Smedley et al (1991) examined the water-use adjacent riparian tree species are maintained efficiency of these and other herbaceous grass­ between —20 and -30 bars during the same land species. Water-use efficiency, the ratio of periods. Water potentials in the range of -10 to photosynthesis to transpiration, serves as a mea­ -15 bars cause many crop species to wilt and sure of how much photosynthetic carbon gain close their stomata, reducing transpirational occurs per unit water loss from tlie leaf. Dicot water loss. Tolerance of water stress levels as low herbaceous perennials had consistently lower as -40 to -60 bars is thought to occur in only water-use efficiencies than their monocot coun­ the mosl drought-adapted aridland species. terparts (Fig. 12). The differences in intrinsic These late-summer water potential values on water-use efficiencv withm this life form mav be slope species are sufficiently low to close sto­ a major contributing factor to the shorter life mata and reduce photosynthesis to near zero expectancy in dicot herbaceous species. Consis­ values. In Dina’s (1970) study photosynthetic tent with this pattern, Smedley et al. (1991) rates of riparian species decreased by 50-80% observed that water-use efficiencv of annual from nonstress values, but riparian trees were species is significantly lower than that of peren­ able to maintain positive net photosynthetic nial species in grasslands along the lower por­ rates throughout the summer. More recently, tions of the canyon.m They _ # also observed that Dawson and Ehleringer (1992) and Donovan perennials which persist longer into the summer and Ehleringer (1991) conducted related stud­ drought period have higher water-use efficien­ ies and again observed that photosynthetic cies than those species that became dormant in carbon gain of slope species is largely limited to late spring. During 1988-90. precipitation was spring and earl}' summer, whereas riparian spe­ unusually low. The effects of the three-year cies are able to maintain photosynthetic rates drought are now seen in Gambet oak and throughout the year, albeit that photosvnthetic bigtooth maple at their lower distribution limits, rates are lower in summer than in spring. especially on shallow soils, where stem dieback Two common responses to limited water has become prev alent. 114 C heat Basin Naturaijst [Voluing 52

10 cm

M a rc h A pril M ay

Fi^. 13. Height of ( i/i•'(< >',t!rnt Ion“i{i

Ehleringer (198S) examined leaf-level peratures are substantially higher than the opti­ adaptations ol plants along the entire elevational mum photosynthetic temperature for the eleva­ transect within Red Butte Canyon. This study tor plant and result in both a decreased focused on determining patterns of leaf angle photosynthetic rate and a decreased water-use and leaf absorptance variation among species efficiency (Werk et al. 1986’. To increase both af within communities exposed to different degrees the rate of photosynthetic carbon gain and of drought stress. Increased leaf angle and water-use efficiency, the pseudoscape elongates decreased leaf absorptance reduce the solar as spring temperatures progressively increase energye v incident on leaves and are viewed as {Fig. I 3). The result is that what was once a mechanisms tor both reducing leal energy loads prostrate canopy is elevated above the warm soil (reducing leaf temperature) and increasing surface and now exposed to cooler air tempera­ water-use efiiciencv. Along a transect from tures abovC the ground surface. Werk et a!. grassland tlirough coniferous forest. \'oiy few (1986) showed that the rate at which the plant species exhibit: any significant changes in psuedoscape elongates is dependent on die rate leaf absorptance, However, leaf angles among of soi 1-surface heating. Plants from protected or species become progressively steeper in drier north-facing sites elongate less than those from habitats. Phis pattern is consistent with the exposed, southerly .sites. notion that as plants are exposed to progres­ Donovan and Ehleringer (1991) examined sively drier environments, the general adaptive relationships between water use and the likeli­ response of species within the community' is to hood of establishment bvJ common shrub and increase leaf angle, thereby reducing incident tree species in the lower portions of Red Butte solar radiation levels. Canyon. They observed that photosynthesis is In the o grasslands on the lower portions r of greater in seedlings than in adults throughout Red Butte Canyon is a most unusual plant sjk1- most of the growing season, but that water stress cies, Ciftnoptenis longipes (long'stalk spring- and water-use efficiency are lower in seedlings. parsley). Sometimes known as the “elevator Seedling mortality in several of the species is plant," C. longipes is a prostrate herbaceous associated with higher water-use efficiencies, perennial with an elongating pseudoscape (a suggesting that mortality selection occurs With scape is a leafless flowering stalk arising from greater frequency in seedlings that are conser­ ground level; the pseudoscape is an elongation vative in their water use before they have estab­ of the leaf-hearing stem in the region between lished sufficiently deep roots to survive the long the roots and existing leaves). Other summer drought periods CyrrkYpterus species also have a pseudoscape, Few studies have addressed ecophysiologi­ but in none of the other species is it as well cal aspects oi riparian ecosystems in the Inter­ developed as in C. lorigipes. In spring, solar mountain West. This is somewhat surprising heating of the ground surface increases soil and since riparian ecosystems are most often among

!eai temperatures and can result in moderately the first to be damaged*5 by J human-related aetiv- warm leaf temperatures (30-35 C). These tem­ ities. from outdoor recreation to water DBH of main tree trunk, cm

Fig, 14, Hydrogen Isotope ratio of stem water: of tliree common streamside ant! itdiacenr nonst reattisdt trve species in PaHeys For); of Bed Bint'.- Canyon as u function ol the- diameter at breast height of die main trunk, Plotted as gray Kirs liiv also the hyilrogcn isotope? ratios of the three possible water sources Tor these plants: loeal precipitation, stream water, and srm.mdwater. ©pen symbols represent streamside plants and closed symbols represent nonstreamside plants, Adapted from Dawson anti ifhJennEer (ISUi).

impoundment to grazing. Pied Butte Canyon, as sources currently used by that plant. Rain, one if the few remaining riparian systems in the groundwaters, and stream waters differ in their Intermonntain West not severely impacted by hydrogen isolope ratios, providing a signal dif­ human activities, is ideal for studies of the adap­ ference that could he detected by stem-water tations of riparian plants and for comparative analyses. Dawson and Ehleringer (1991) studies of species sensitivities to human-related observed that among mature tree species none activities. were directly using stream water (Fig. 14). All In a recent studv Dawson and Ehleringer were using waters from a much greater depth, (1992) examined water sources used by riparian which had a hydrogen isotope ratio more nega­ plants species. In I Iteir study, plants were segre­ tive than either stream water or precipitation. gated according to niicrohabitat and size:

YoungO streamside...... trees utilized stream water, , stream side versus nonstreamside and juvenile but only when small. Young trees at nonstream­ versus adult (based on diameter at breast height). Their results were rather startling and side locations utilized precipitation, having suggest that a new perspective is necessary' access to neither stream water nor deeper when evaluating riparian communities, their groundwater. One possible reason that stream- establishment potentials, and their sensitivity to side trees mav not depend on stream water is disturbance. Dawson and Ehleringer (1991) that this surface water source may occasionally used hydrogen isotope analyses of stem waters dry up during extreme drought years and to determine the extent to Which different cat­ become unavailable to these trees; another is egories of riparian trees utilize stream water, that stream channels occasionally change their recent precipitation, or groundwater. Hydrogen course, and dependence on .surface moisture isotopes are not fractionated by roots during would then result in increased drought stress water uptake; therefore, the hydrogen isotope and likely increased mortality rates. The long­ ratios of stem water will reflect the water term stream discharge rates suggest that stream 116 G re at Basin Naturalist [ Volume 52 water may he less dependable than deeper to 19S2. The loss of active beaver dams in the groundwater sources (Fig. 6). early 1980s has doubtless greatly reduced the Many plants do not contain both male and populations of small mammals that are female reproductive structures in their [lowers, restricted to die mesic-marshy habitats of the but are present as either male or female plants canyon, (dioecy). Freeman et al. (1976, 1980) noted that Nonetheless, based on the altitudinal gradi­ dioeey is a common feature of plants in the ent and vegetational diversity of Red Butte Intermountain West. Furthermore, they Canyon, a total of 51 species of mammals should observed that the two sexes are usually not ran­ hypothetically occur there. Below is a list of the domly distributed across the landscape. Rather 39 species of mammals known to occur in Ked there is a spatial segregation of tlie two sexes Butte Canyon.j such that iemales tend to predominate in less K sECTJVOjS A—SniUCIDAK stressful microsites (wetter, shadier, etc,), Sorex ptfkislfis water shrew whereas males occur with greater frequencies Sorex Vflgmm wandering shrew on more stressful sites (drier, sunnier, saltier, Sorex cineiws masked shrew etc.). In Red Butte Canyon, Freeman et al. Cm kopteha—Vest Ercrn jonadar * Eptesicus fuscus big brown hat (1976) investigated spatial distributions of Acer LaCOMOIU’Ha—LEFOitlDAtf negundo (boxelder, a riparian tree) and Thalia- Lepm towmendi while-tailed jackrabbit trumjendleri (Fendler meadowme, a perennial Sijlvilagus nuttallii Nuttall cottontail herb). In both species, there was a strong spatial Rode ntia—-Sc i u R \ dal; TaflliaschtTlis hiuhonicys red squirrel segregation of the two sexes. Marnwtajlavivfmter yellpwdbellied marmot Dawson mid Ehleringer (1992) have fol­ Spertnophih^ urrmtius Uinta ground squirrel lowed up on the initial observations of spatial Spermophilus van( ^!tu^ rock squirrel segregation iwAcernegwidv (boxelder), seeking Eutamias mittimus 1 east chipmunk Glaucomas salninix& northern %ins; squirrel to determine whether intrinsic physiological Rodent? a—Get \um>AE differences among the sexes may contribute to Thomormjs talpoides northern pocket gopher plant mortality in different microsites. They Thonumim hotiae botta pocket gopher observed that female trees have significantly Rod k ntj a—Gasi t > tuDaf Castor canadensis beaver lower water-use eJficiencies than male irees on RODEVTiA—MUKIDAE both stream side (where female predominate) Reithrodontoimfi nicgdotti western harvest mouse and nonstream side locations (where males pre­ Peromy$fiu$ nwnicidutiis deer mouse dominate}. Male trees exhibit a higher water- Fenytmjscus hoylii brush ftiouse use efficiency in dry sites than in streamside Clethrioni'>mtjS gapperi red-backed vole Ondatra zibethicu# muskrat locations, but female trees exhibit no such Pkenai'omtjs tattirrnedittf heather V&le response across microhabitats. The lack of a Microtus numUmm montane vole change in water-use efficiency by female trees Microtun fon0ciiudti$ Ibng-t.ailed vole on dry, nonstreamside locations may contribute Aroimla risJiardsoni water vote Rudentia—Zafodibae to an increased mortality rate, which then ZGpus princeps western jumping mouse ultimately results in a male-biased sex ratio at R C)D E NT 1A—E UETH1ZO NTT DAE these sites. EfTthizon dnrsaturrt porcupine Ca hktvo ua —Canidae Cattfa l&trans coyote Mammalian Fauna Ca w Nm j \ ia—P ftOCTON r dae Ba&sariscm tistutus ring-tailed eat Pwcyon htoi racoon The mammalian fauna of Red Butte Canyon CaKMEUOKA — M USTtti i UA El is remarkably diverse, due in part to the altitu- Musteki freruita long-tailed weasel dinal gradient and numerous small patches of Musteld ertninca ermine various plant communities indigenous io the Mnstel# vis cm miuk Taxkha taxus Badge* area. A particularly rich small mammal fauna is Mephitis mephitis stuped skunk associated with the patches of riparian habitat Carnivoiu—Frlidae along Ked Butte Creek and its tributaries. Prior Lynx m/us bobcat to die run-off of 1983., riparian habitats were Fells concolor mountain lion A htjoda< rmA—Cervidae much more extensively developed than at pres­ Cerwtf cantubtnsis elk ent. Numerous marshy meadows existed in Otbcuiletis h&mionus mule deer association with large, active beaver dams prior Akes fimericanits rrsoo.se 1992] Red Butte Canyon Research Natural Area 117

Some of the larger species have been G ALLircmM ES—TETKaONIDaE observed only occasionally, such as die bobcat, Dendraeapus obscitrus Blue Crouse Bonasa umbeUus Ruffed Grouse mountain lion, and moose. But others such as Gallifohmes—Ph a sia nidae the mule deer, elk, and coyote are observed widi Lophortyx calif amicus Ca] ifornia Quail high frequency at some seasons. A rather rich Phasiantts colckUtis Ring-necked Pheasant rodent fauna inhabits the canyon, with many of Alectaris graeca Chukar STiUGtFOHMES—STMOIDAE the species preferentially occupying the moist Oto flammeokii Flammukted. Owl riparian communities of grasses, forbs, and Bubo virgtTuanus Great Homed Owl shrubs. Thus, the red-backed vole, heather vole, Asio Qtus Long-eared Owl montane vole, long-tailed vole, water vole, and CORACIIFORMES—Al/CEDINIDAK Megoceryle alcyon Belted Kingfisher jumping mouse are virtually restricted to the FICIFOHM ES—PlClDAE small mesic meadows along Red Butte Creek Calaptes cafer Red-shafter Flicker and its tributaries. Similarly, the three species of Sphyrapicus varias Yellow-bellied Sapsucker shrews in die canyon are distributed almost Dendrocopus inUosus Haii)' Woodpecker Dendrocopus pubescens Downy Woodpecker exclusively in the riparian habitats. PaSSEMFOkM ES—-CO BV1 DAE In some larger meadows, such as along Par­ Cyanocitta stdlen Stellers Jay leys Fork and at Porcupine Gulch, the microtine Aphektcoma coarulesc-ens Scrub Jay rodents are distributed in a strongly zonal pat­ Pica pica Magpie pASSEFUPORMES—PAJUDAE tern. Long-tailed voles are found in the driest Pams atricapHlw Black-capped Chickadee parts of the meadows, montane voles in the Pams £titnbt;h Mountain Chickadee more mesic areas where grasses, sedges, and Psaltiiparus minimus Common Bushtit forbs comprise a diverse community, and water PassErifohmes—SirrtDAE Sitta canadensis Red-breasted Nuthatch voles in the immediate stream Side area, their PASbEKlFOli MES—CERTHJJDAF burrows often entering the bank at the water's Ceithiajamdiaris Brown Creeper edge. Red-backed voles and heather voles are Passerifohmes—ClNCUDAE typically found around the bases of willows in Cinciw itoexicanus Dipper Pass erifor mes—Tt jrdidae the meadows, as well as around the edges of Myadeste$ tmvnsendi Townsend’s Solitaire conifers at higher elevations. Passebifoames—Sylvudae A few species are found only at higher eleva­ Regulus satropa Golden-crowned Kinglet tions in association with Pseudotsuga menziesii Passerifor S1ES—Stukntdae Stumus vulgaris Stalling (Douglas-fir) and Populm tremuloides (aspen). Passeki formes—Ictebidae These include the red squirrel, Uinta ground Stumpila neglect a Western Meadowlark squirrel, yellow-bellied marmot, and least chip­ Passehifohwes—Fringiludae munk. The oak-mountain mahogany zone Carpodacus mexicanus Hou Finch Spimts pirn is Pme Siskin seems to be die preferred habitat of the rock Junco oreganos Oregon [unco squirrel and perhaps the ring-tailed cat as well. Several dissertations dealing with the ec. ology In addition to the species that are permanent and physiological adaptations of shrews, microtine residents in Red Butte Canyon, the following rodents, and jumping mice have utilized study list of summer residents represents species that sites in Red Butte Canyon (Forslund 1972. probably also nest in the canyon: Cranford 1977), AN5ER [FORM ES—ANaTIDaE Anus platyrlupu.kos Mallard Duck Falcon ufohmes—-Aocipit w dae Avian Fauna Buteo farnaicensis Red-tailed Hawk Aquila chrysmtoS Golden Eagle In his study of the birds of Red Butte F ALCOMI TOR M £5—FALCOtf IDAE Canyon, Periy (1973) found that 106 species Falco sporverius Sparrow Hawk occurred in die area during his study. Of these, C HA RADKIIFO HM ES— SOQLOrACUDAE Acffifs mactdarla Spotted Sandpiper 32 species are permanent residents and 44 are COLUMBIFOHMES— COUJMBIDAE summer residents. The remainder (30) are Jenmdura m/icrowa Mourning Dave migrants or winter residents. The permanent Apodjpormes—Trockilidae resident birds include: Archilochus alexandri BJark-chiiined Hummingbird FaLCONIFOHMES—ACCIPITRtDAE Selasphorus plati/cercus Broad-tailed Accipiier gent Uis Goshawk Hummingbird AccipUer stiiatus Sharp siiinucd Hawk P ASSEHIFOH M ES—TY HANNlDAE Accipiter cooperi Cooper's Hawk Empidutuix oherholseri Dusky Flycatcher 118 Great Basin Naturalist [Volume 52

Evipulonfix d ifficilis Western Flycatcher Research Natural Areas provide several spe­ Cma&tyHu sijrcJuhihis Western Wood Pee wee cific advantages to the nations scientific rjji MRS— HmUNptivi l>aE Tachijcim'Xa thuhissina Violet-green Swallow qorn in unity, which are typically not otherwise lmU>prtKTv bicvlor Tree Swallow available. These include potential use ot an area Ripuria riparUi Bank Swidlow that has had minimal human interference and Std^idopta ifx ntfiatllis Rrm^h-winged Swallow lias a reasonable assurance ol long-term exis­ Hmtntlo nislica Bain Swallow Fetrovhvlidon pyrrhonota (.'lifl Swallow tence, and the potential association and interac­ Passehifokmes—T itGCf yfot)yi ida.e tion of scientists from different disciplines Tn*glodytes tnxhm House Wren leading to discoveries unlikely to occur without Salpi nctes ubmlzhix Rock W ren such an association. Conducting research at —Turdhmk common locations is key to developing these Tardus miffiat&rnts Robin UijlocAchla guttata 1 lermit Thrush interactions. Research Natural Areas not only Htfbxdchln ustulata Swainsou s Tlmisli assist in the progress of basic science, but also Sialia ainiwoidex Mountain Bluebird provide federal and state agencies with informa­ PA^SlvKlFOHMJ'S—SVLVJIDAE Fulioptifa (oendea Bhio-gruy Cnuieafclier tion upon which to base management decisions. P ANSKRTFOH —Vt HKON IDA FI The melding of ecosystem preservation and Virett gift™ Warbling Vineo research on basic ecological processes at 1*AA>K >HMES—-PaIIUI J1MK Research Natural Areas provides numerous Venniwm cduta Ortuige-crowwd Warbler valuable options to society. The Red Butte Vcrutivom rrrginiae V irginias Warble* Vendroica petechia Yellow Warblt.T Canyon RNA serves this purpose well. Although DtmdroU-it (mduhofti Audubon $ Warbler initially affected by human activities during the Qpttrorrite (ohuk i MjcGSlEivr^yV W#hler earlyjr settlement of the Salt Lake Valley, -r the VViZscwii^ mtsilfo Wilson s Waihler canyon was soon set aside by the federal govern­ - / P^K SEJU rtfU MKS— ICTK Kt J > A f t Icivrtis buUickii Bullocks Oriofe ment and has now had nearly a century to Molothnts ater Brown-headed Cowbt td recover (though the loss of beaver represents a PASSKKirOHMKS—TflRAUL'IUAE significant impact to the ecology ol the riparian Viraniitt lud<>inciana We?''tern Tar11iger ecosystem). Other canyons in the Wasatch PaSKEJUFOIIMKS—FWNCILUDAE Vhcuticus mdawtmphalus Bluckheiujed Grosbeak Range have not received equivalent protection, As we move into the twenty-first century, Fdsserina mtwerta Lazuli (hintingC 1 v ■ C(ir}w({tn.w$ (gissinii Cos sin’s Finch there w ill be increasing pressure to understand SpiituS tristix American Goldfinch the dynamics of ecological systems and man s Chiomnt ehltmtrri Green-tailed Towliee Fip:L) litythrothubniifi LUilous-sided Tmvhee impact on ecological processes. Maintained as a Paoeadas in'amineua Vesper Sparrow protected watershed, the Red Butte Canyon funci) ainiceps C»ray-headed Juneo RNA provides a unique opportunity for SjiheUe passatfna Chipping Sparrow addressing these important issues to human aiasiiim m<\',(uc 'Kttig Sparrow society and to the preservation of our environ­ : ■ ment. Unprotected, it is an invaluable resource Role of Research Natural Areas lost Forever.

Federal Iand-managenlent agencies have L it e r a t u r e C it e d been developing a national system of Research Natural Areas since 1927. More than 400 areas have received this designation nationally. Since An asterisk (") refers to studies conducted in C ? ^ inception of the RNA Program, there have been Red Butte Canyon, but not specifically cited in two primary purposes for Research Natural this manuscript. Areas: Albf:k. B. J., L, M, Shultz, and S. Go o d iiic h . I9S\ Atlas 1. to preserve a representative array of all oft he vascular plantsoi Utah. Utah Museum ol Natural History, Salt Like City. 670 pp. significant natural ecosystems and their Anonymous, lUo-t. History oj Fort Diiiiglas. Compiled by inherent processes as baseline areas; and Fori Douglas Army Engineers Office* Am now. J* A. 197], Vascular flora of Rixl Bulte Canyon, Salt 2. to obtain, throughv 7 scientific education and research, information about natural system Lake Count). Utah. Masters thesis. University of Utah, Salt Lake City. 383 pp. components, inherent processes, and com­ ____ . 1 MS I, Foci secunda Presl. versus P. sandbar^ii Vascy parisons with representative manipulated fPoaeeae). Systematic Botany 6: 4.I2-42.L. systems. ____ 1987. Cramineae. Pages 684-7SH in S. I „ Welsh, 1992] Red Butte Canyon Research Natural Area 119

N, Ahrood* L. C Higgins, and S. Goodrich. A Utah lished doctoral dissertation, University of Utuh, Salt flora. Brigham Young University Press, Provo. Utah. Lake City. Arnow. L. A,, B. Albee, and A. WVckoff 1980. Flora of Djna., S. J-, and L, G, Klikoff 1973. Carbon dioxide the central Wasatch Front, Utah. Utah Museum of exchange by several stream side and scrub oak commu­ Natural History. Sal* Lake Gty, 563 pp. nity species of Red Butte Canyon, Utah. American AJUUNGTON. L. and T G, ALEXANDER- 1965. The U.S. Midland Naturalist 89: 70—80, Army overlooks Salt Lake Valley, Fort Douglas 1862^- Dobrgwolskl J. P., Ml M Callwrll. and J. M. Rich­ 196-5. Utah Historical Quarterly 33: 327—350. ards. 1990* Basin hydtology and plant root systems, Bates, J. W. 1963. The effects of beaver on stream flow. Job Pages 243-292 in C, B. Osmond* L, F. Pifelka, and Completion Report for Federal Aid Project W-65TL G. M, Hidy, eds., Plant biology of the Basin and Range, Utah State Department of Fish and Game Information Springer Verlag, New York. Bulletin 63-13. 87 pp. Donovan. L, A*f and J. R Ehlf^jnc;e4 1991. Econhysio- Billings. W D 1951. Vegetation zonation in the Great logica! differences among pre^reproductive anrf repro­ Basir: of western North America. In: Les fosses ductive classes of several woody species ^ Oecologia S6: ecologiques de la regeneration de la vegetation des 594-597. ' zones arides, International Union of Biological Sci EhleuiNGEH. J* R. 1983. Changes in leaf characteristics of enoes Series B ColEoqulaS: 101-122, species along elevation a! gradients in the Wasatch _____ * ? 990. The mountain forests of Nordi .America and Front, Utah. American Journal of Botany 75: 690-689* their environments. Pages 47-36 in C. B* Osmond. EvAMb, F. F\. 1936 A comparative study of the vegetation of L. F. Pitelka, and G- M. Hidy, eds., Plant biafogy of the a grazed and ungrazed canyon of the Wasatch Range, Basin and Range. Springer Verlag, New York, Unpublished masters thesis, University of Utah, Salt Bond, 1-1. W, 1977. Nutrient dynamics of undisturbed eco­ Lake City. systems. Unpublished doctor al dissertation, University Fojislund. L. G> 1972. Endocrine adjustments in Microtm of Utah. Salt l^ake City. montanus populations from laboratory and natural _____r. 1979. Nutrient concentration patterns in a stream environments. Unpublished doctoral dissertation, draining a montane ecosystem m Utah. Ecology 60 > Tulane University, New Orleans, Louisiana. 1184-1196. ME&MAK D, C., K« T. and W K- Ostleh 1980- fBu£w$TEB, W. 1951. GcJl wasps producing falls on the Ecology of plant dioecy in the mtennountam region of scrub oak Que*~Gu$ gambelii Nutt. Unpublished western North America and California. Oecologia 44: masters thesis* University of Utah, Salt Lake City. 410-417. Caldwell, M, M. 1QS5. Cold desert. Pages 19S-212 in FKRgjMAN, D. C, L, G. KLTfcOFiv and K, T. Hamper, 1976. B. F. Chabot and IS. A. Mooney, eds.. Physiological Differential resource utilization by the sexes of dioe­ ecology of North American plant communities. Chap cious plants. Science 193: 597-599, man and Hall, London. Hely, A. G., R. W, MpwEft, and C, A. Harr 197], Water COM>~roeK, J, F-, and], R. Eulkkingek, 1992. Plant adap­ resources of Salt Lake County, Utah Utah Department tation in the Great Basin and Colorado Plateau. Great of Natural Resources Technical Publication 3L Barin Naturalist. In press. HlBBAKD, C- G- 1980. Fort Douglas, 1862-1916. Unpub­ Cottam W P.7 and F. Evans. 1945. A comparative study of grazed ted ungrazed canyons of the Wasatch Range* lished doctoral dissertation, University of Utah. Salt Utah. Ecology 26: 171-181. Lake City. Houghton, J_ ir\ CfaMfokd, J. A. 1977. The ecology of the western jumping G. 1969, Characteristics of rainfall the mouse Unpublished doctoral dissertation. University Great Basin. Desert Research Institute, L-niversity of of Utah. Sa^t Like City. Nevada* Reno. 205 pp. •CnoET. A. R.> L, Woodward, and D. A. Anderson 1943. "James. F. K., Jr. 1950. The ants of Red Butte Canyon. Measurements of accelerated erosion on range-water­ Unpublished masters thesis, University of Utah, Salt shed land. Journal of Forestry 41: 112-116. Lake City CRONQD15T, A, j. 1947. Revision of the North American KleIneh. E, F. 1967 A study of the vegeftitiomt) communi­ species of Erigeron, north ol Mexico, Brittonia 6: 121 ties of Red Butte Canyon, Salt Lake County, Utah, 302. ^ Unpublished master? thesis, University of Utah, Salt ____ . 1961. An integrated system ol classification of flow Lake City. 53 pp, ering plants. Columbia University Press, New York. Klf^inek, E. F., and K* T. Hakpek, 1966. An investigation 1262 pp. of association patterns of prevalent grassland specter in Dau BEN MIRE, R. F. 1943. Vegetationa! zonation in the Red Butte Canyon, Salt Lake County, Utah. Utah Rocky Mountains. Botanical Review 9: 325—393. Academy of Science. Arts* and Letters 13: 29-,36. Dawsom, T- E., and J. R. EiiLERFNGBF.. 1991. Streamside La ffekty, K. M. 1949. A preliminary study of the spiders trees that do not use stream water. Nature 350: 335­ of Red Butte Canyon. Unpublished masters thesis, 337. University of Utah, Salt Lake City. ____ . 1992. Gender-specific physiology, carbon isotope Ld-LLtNCEH. D. B 1985. Ferns and fern allies. Smithsonian discrimination* and habitat distribution in boxelder, Institution Press, Washington, D C, 389 pp, Acer negundtt Ecology. In press. Maxell, R. E.. and R. L. Titref/t 196*0. Geologic map of DeLUCIA* E. H,,andw. H. Sculesingek* L990, Ecophys­ Salt Lake Count); Utah Utah Geological and Mlneral- iology of Great Basin and Sierra Nevada vegetation on ogical Survey Reprint Series, R.S. S3. Scale 1:62,500, contrasting soils. Pages 143^17SmC. B. Osmond, L. K Mathias, M. E., and L, Constance. 194^45, Pitelka* and G. M. Hidy, eds,. Plant biology of the Basin Umbelliferae. North American Flora 28 B: 43-297. and Range. Springer Verlag, New York. Negus, N. C.. P. J. Berger, and L. G. F o k slu n d 1977, DtNA± S J. 1970- An evaluation of physiological response to Reproductive strateg)^ of Microtws numtanus. foumal water stress as a factor influencing the distribution of of Mammalogy 5S: 347-^353- six woody species in Red Butte Canyon, Utah- Unpub­ Petiry, M. L, 1973. Species composition and density of the 120 G r e a t B a s in N a t u r a l is t [Volume 52

birch of Red Butte Canyon. Unpublished masters lished masters thesis, University of Utah, Salt Lake thesis, University of Utah, Salt Lake City. City. 'Peterson. B. V. 1953, Taxonomy and biology of the black Woodward, L,, J. L, Harvey, K. M. Donaldson, J. J. flies oi Salt Lake County. Unpublished master's thesis, Shiozaki, G. W. Letshman. and J. H. Broderick. University of U tah, Salt Lake City. 1974. Soil survey of Salt Lake- area, Utah. U.S. Soil SamuELSON. J. A. 1950. A. quantitative comparison of the Conservation Service in Cooperation with Utah Agri­ algal populations in two Wasatch Mountain streams. cultural Experiment Station. 132 pp. Unpublished masters thesis, University of Utah. Salt Lake City. Received 14 November 1991 Sckeffeji. V B. 1938. Management studies c! transplanted Accepted 1 June 199'2 beaver in the Pacific Northwest. North American Wild­ life Transactions 6: 320-326. Sieren, D. 1. 1981. The taxonomy of the genus Enthamiu. APPENDIX Rhodora S3: 557-579. Smedley. M. P., T. E. Dawson. J, P. Comstock, L, a. Nomenclatujral Changes in the Flora, Donovan, D. E. Sherrill. C. S. Cook, and J. R. Ehuerincer 1991. Seasonal carbon isotnpic discrim­ 1971-1990 ination in a grassland community. Oecologia 85: 314­ 320. " The following is a list of nomenclatural and Smith. S. D., and R. S. Nowak. 1990. Ecophysiology of orthographic changes made since publication of plants in the intermountain lowlands. Pages 170-2* 11 the Vascular Flora of Red Butte Canyon, Salt ui C, B. Osmond L. F. Pitelka, and G. M. Hid)-; eds.. Lake County. Utah (Amow 1971), Fam ily Plant biology of the Basin and Range, Springer Verlag, New York. names of flowering plants are changed to accord Smith, W. K., and A. K. JCnapp. 1990. Ecophysiology of high with those used by Cronquist (.1981), A ll other elevation forests. Pages 87-142in C. B. Osmond, L, F. name changes are contained in Welsh et al. Pitelka. and G. M. Hidy, eds., Plant biology of the Basin and Range. Springer Verlag. New York. (1987) unless otherwise specified. Stoddaht. L A. 1941. The Palouse grassland association in amakanthaceae northern Utah. Ecology 22:158-163. Amaranthus praecizans of American authors, not L, = A Swanson, G,, E. Kleiner, and K. T Harper. 1966. A blitoidcs Wats. vegetational study of Red Butte Canyon, Salt Lake AMARYLLIDACEAE = Lit. [ ACE A E County, Utah, Proceedings of the Utah Academy of Brodiaea douglasii Wats. = Trtteleki grandifiora Lindl. Science. Arts, and Letters 43: 159--160. Anacardmceae 'Treshow, M., and K. T. Harper. 1974. Longevity of Bhus radicnns L - Toxicodendron njdbergU (Small) perennial forbs and grasses. Oikos 25: 93-96. Greene Treshow, M., and D. Stewart. 1973. Ozone sensitivity of Berberudaceae plants in natural communities. Environmental Conser­ Bei'betrs repens Lind). - Mahorua repens (Lindl.) G. Don vation 5: 209--214. Boraginaceae Tryon. R. M., and A. F. Tryon. 1982. Fems and allied Cruptantha nana (Eastw.) Pays. - Crtiptantha humilia plants, Springer-Verlag, New York. 857 pp. (Gray) Pays. Van Horn, R., and M. D. Chittenden, J*. 1987. Map Haekelia jessicae (McGregor) Brand - H. micmtit-ha showing surlicial units and bedrock geo logy of the Fort (Eastw.) J, L, Gentry' Douglas Quadrangle and parts ol the Mountain Dell Lapvidn echinnta Gilib. = L. squarrosa (Retz.) Dumort. and Salt Lake City North Quadrangles, Davis, Salt (Weber 1987) Lake, and Morgan Counties, Utah, U.S. Geological Cactaceae Survey Miscellaneous Investigations Series, Map 1­ Opuntw aurea .Baxter, misapplied to O. mctcrorhiza 1762. Scale 1:24,000. “ Engelm. "VfcXEHY. R. K., Jr 1990. Pollination experiments in the Cahyophyllaceae Minvulus cardkialis—M. ieuTSii complex. Great Basin Cerastntm vulgatum L. = C. fontanum Baumg. Naturalist 50: 155-159. Stellaiia jamesiann Torr. - Pseudastellaria jameswui •Waser. N. M., R. K. Vickery, Jr.. and M . V. Phsce 19S2. [Torr.} Weber & Hartman (Weber and Hartman 1979) Patterns of seed dispersai and population differentia­ C EL ASTR ACE AE tion in Mimulus guttatiis. Evolution 36: 753-761. Fachistima - Paxistima Weber. W. A. 1987. Colorado flora: western slope. Colo­ Chenofodiaceae rado Associated University Press, Boulder. 530 pp. Salspla kah L. = Salsola iberim Sennen & Pau Weber W. A., and R. Hartman 1979. A North American COMPOSITAE - ASTEKaCEaE representative of a Eurasian genus. Phytologia 44:313­ Aster chdensis Nees = A. ascendent Lindl. 314. Haplopapptts njdbergU Blake - B. watsonii Gray Welsh, S. L., N. D. Atwood, L. C. Higcins, and S. Lactuca pulchella (Pursh) DC. = L. tatanca (L.) C. A. Goodrich. 1987. A Utah flora. Brigham Young Uni­ Mey versity Press, Provo, Utah. 894 pp. Matricaria ntiftricarioides (Less.) Porter = ChamomilUr Werk, K. S.. ]. R. Ehlehincek, and P. C. Harley. 1986. saaveolew (Pursh) Rydb. Formation of false stems in Cymoptems lotigipes: an Solidago neniortdis Ait. = S. sparsiflora A. Gray uplifting example of growth form change. Oecologia S- occidentals (Nutt.) T. & G. = Euthamia occidentals 69: 466^470. Nutt. (Sieren 1981) Whitney, R. R. 1951. A comparison of the aquatic inverte­ Taraxacum laexAgatum (Wtlld.) DC. = T. officinale brates of Red Butte and Emigration Creeks. Unpub* Wiggei'f! (Weber 1987) 1992] Red Butte Canyon Research Natural Area 121

Vigttiem multifloru (Nutt.) Blake - Hchomeria midtiflom Labiatae = Lamiaceae Nutt Moldaofca parvi flora (Nutt) Britt, - Dracacephalum Coknaceae paroiflomm Nutt Comus stolonifcra \lichx. = Comtis seticea L. LeguminosaE = Fa&aCEaE Ckui:iferae- BrASSICaCKAE M oracle = GannahaCeaK Aralm divarica^Mi A. Nels. = A, ko&oeUU Homem, Hunmlus lupulus L, = H. antericanm Nutt, Rorippa islcmdiva (OecU Boj'h. = K, pahfiffiris (L,) Besser Onaghaceal; ft truncal*i (JepsJ Stuckey = fl. tenemma Greene Epilobnim jiammdatiwx T. &G> - E, hrachtjcaqyum Presl Ojscutaceae watsonii Barbey = E. ciliatum Raf, Cuscttia campestns Yunck. = C pentagon# Engelm. Oenothera Iiookeri T & G, = 0, elata H.B.K. GlPRRA£EAE Zauchmria gamitii A, Nel s. = Z. latifolia (Horjk.) Carex utriculata Boott =- C. rostrata Stokes Greene Gramweae * POACL:AE(Amowl987) OkobanckacI^\e Agropijmn canimtm f L*) IBeauv. = Elymus trachycmdus Orobttnche califomicu Cham. & Sohlecht = O. (Link) Shinners corymbosa fBydb.) Ferris A. dasystachyum (Hook ? Scribn. - Elymns l/m&'olatus Polsmonuceae (Scribn. & Sfri.J Gould Iponwpsis aggregafa (Pvirsli) V Grant ^ Gilia aggregata A. intermedium (Host) Beauv. “ Elymus hispulus (Opiz) (Pursh) Spreng. Meld. PGlypodiaceaf, as it occurs in Red B^ttt? Canyon, [5 novp A. smithii Rydb* := Elymus smiihU (Rydb.) Gould divided into the following families !Trvon and Trvon A sfriccttum (Bursh) Scribn* - Ehjims spicatus (Pursh) 19S2> Gould Dennsh'akdtiaceae. of which the genus Ptetidium is a Agrostis alba L. - A. siotomfem L. meinber A. sorniverticillata (Forsk) C Christ, - PohjfX)gon semi- DmoPTEPJDACkAE, which includes the geriera Cystapteris

iXM'ticdhitus (Forsk;) I lylanderf and Woodsui Aritfida bngiseta Steua. =A purpurea Nutt. Cystupteris frafflis (L.) Bemh. is now known to include Bronvus brizaefonnis Fi^ch. & Mey. - brizifonrus two taxa (Lellinger 1985), of which only C. tenuis B. corrpnti$chrad. - B,ja])onicus (MiehxJ Desv. occurs in Red Butte Canyon. Ghjccria elata (Nash) M. E, Jones = G. striata (Lam,) R AN U CULAC E A E Hitcha Ranuncvlm longirostiis Godron = R aquatHti U HespervchloG kingii (Wats.) R\xib. = Leucopoa kingii R. twticulatus Crantz = Ceratocephahis dtthveerus DC. (Wits.) W A. Weber ' (Weber 1987)

KfK^eria cristata Pers. m ^ macrantha (Ledeb.) Soliiill. S a l i c a c i : a e Onfzopsis hijnumoides (R. fit S.) Ricker = Stipe Salix rigida MnhL - S. lutca Nutt htjmenoides R. & S. S.VXJl^AGACEAE Foa sartdhcrgn Vasey = P. seennda Presl (Amow 1981) Lithophragma hulbifera Rydb, = L glabra Nutt Sitanion jubatum J. G Smith. misapplied to Ehjmtts Sc^opkula 111 ACEAE chjmoides (Rafj Suezey CaxtilUya kortardii Rydlv - C. rhexifolia Rydb. Stipa accidentalis Thurb, = S ndsonii Sen bn Tamaiiicaceae JuNCaCEaE Tamarlx prntandra Pall. - T. ramosissima Ledeb. Jtmcusbalticus Willi = /. arcticwi Willd U MBELLIFFRAE = AFLACEAE / tracyi Hvdb, = J, ensifolius Wflcst CicuUi doaglasH (D G | Coult & Rose - C, maculaia L. Lonwtwm nuttallii (Gray) Macbr. = fcmg?/ (Wats,) Cronq.