Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Kenneth D. Heil, J. Mark Porter, Rich Fleming, and William H. Romme
Technical Report NPS/NAUCARE/NRTR-93/01
National Park Service Cooperative Park Studies Unit
U.S. Department of the Interior at Northern Arizona University National Park Service Cooperative Park Studies Unit Northern Arizona University The National Park Service (NPS) Cooperative Park Studies Unit (CPSU) at Northern Arizona University (NAU) is unique in that it was conceptualized for operation on an ecosystem basis, rather than being restrained by state or NPS boundaries. The CPSU was established to provide research for the 33 NPS units located within the Colorado Plateau, an ecosystem that shares similar resources and their associated management problems. Utilizing the university's physical resources and faculty expertise, the CPSU facilitates multidisciplinary research in NPS units on the Colorado Plateau, which encompasses four states and three NPS regions—Rocky Mountain, Southwest, and Western. The CPSU provides scientific and technical guidance for effective management of natural and cultural resources within those NPS units.
The National Park Service disseminates the results of biological, physical, and social science research through the Colorado Plateau Technical Report Series. Natural resources inventories and monitoring activities, scientific literature reviews, bibliographies, and proceedings of techni cal workshops and conferences are also disseminated through this series.
Unit Staff Charles van Riper, III, Unit Leader Peter G. Rowlands, Research Scientist Henry E. McCutchen, Research Scientist Mark K. Sogge, Ecologist Charles Drost, Zoologist Elena T. Deshler, Biological Technician Paul R. Deshler, Technical Information Specialist Connie C. Cole, Editor Margaret Rasmussen, Administrative Clerk Jennifer Henderson, Secretary
National Park Service Review Documents in this series contain information of a preliminary nature and are prepared primarily for internal use within the National Park Service. This information is not intended for use in open literature. This report has been reviewed and approved for dissemination by the National Park Service. Approval does not signify that the contents necessarily reflect the views and policies of the National Park Service nor does mention of trade names or commercial products constitute endorsement or recommendation for use.
Copies of this report are available from the following. To order from the National Park Service, Technical Information Center, use the reference number on the bottom of the report's inside back cover.
National Park Service National Park Service Cooperative Park Studies Unit Technical Information Center Northern Arizona University Denver Service Center P. O. Box 5614 P. O. Box 25287 Flagstaff, AZ 86011-5614 Denver, CO 80225-0287 (602) 556-7466 (303) 969-2130 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Kenneth D. HeiP, J. Mark Porter2, Rich Fleming1, and William H. Romme3
'San Juan College, Farmington, New Mexico 'Rancho Santa Ana Botanic Garden, Claremont, California 'Fort Lewis College, Durango, Colorado
Technical Report NPS/NAUCARE/NRTR-93/01
September 1993
National Park Service Cooperative Park Studies Unit U.S. Department of the Interior at Northern Arizona University Technical Report NPS/NAUCARE/NRTR-93/01
Capitol Reef National Park was established in 1971 as a 241,000 acre park in the heart of the Colorado Plateau. The law creating the park called for the immediate phase-out of livestock grazing. In 1982, controversy surrounding the required phase-out precipitated the passage of a second law which called for an extension of grazing activities. This law also called for the establishment of a comprehensive grazing research program, initially overseen by the National Academy of Sciences, to determine the effects of the activity on the park's natural and cultural resources. By 1991, with NPS funding, the Division of Resource Management and science at Capitol Reef National Park eventually shepherded to completion 11 separate studies as part of this effort. This document is the first report to be published as part of this grazing research series.
Norman R. Henderson Chief, Resource Management and Science Capitol Reef National Park n Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Table of Contents P List of Figures iv
List of Tables v
Abstract 1
Introduction 1 Previous Work: Floristics 2 Previous Work: Vegetation 3 Location 5 Geology 5 Climate 5 Specific Objectives 6 Methods 6 Inventory of Vascular Plants 6 Inventory of Threatened and Endangered Taxa 7 Habitat Relationships of Rare Plants 7 Field Sampling of Community Composition 7 Community Classification 8 Gradient Analysis of the Vegetation 8 Vegetation Mapping 9
Results 9 Vascular Flora 9 Threatened and Endangered Taxa 9 Habitats of Rare Plant Species 10 Plant Community Classification 10 Gradient Analysis of the Vegetation 14 Elevational Patterns 14 Substrate Patterns 15 Community Patterns 16 Effects of Livestock Grazing 19 Overall Vegetation Patterns in CARE 22
Discussion 26 Vascular Flora 26 Threatened and Endangered Taxa 27 Patterns of Rarity in CARE 27 Critical Areas for Rare Plant Conservation in CARE 28 Patterns of Endemism 29 Interpreting Anomalous Plant Distributions in CARE 32 Plant Communities of Special Concern 33 Vegetation Patterns in Capitol Reef National Park 34 Random and Deterministic Patterns in Community Composition 34 Discrete and Continuous Patterns in Community Composition 35 Distribution of Communities along Environmental Gradients 35 The Role of Natural Disturbances in the Vegetation of CARE 36 Long Term Stability of Vegetation in CARE 37 Effects of Grazing on the Vegetation of CARE 38
Acknowledgements 40
Literature Cited 41
Appendix I: Annotated Checklist of Vascular Plants, Capitol Reef National Park, Utah 47 iii Technical Report NPS/NAUCARE/NRTR-93/01
List of Figures
Figure 1. Detailed map of Capitol Reef National Park with inset of location map of Capitol Reef National Park within the Colorado Plateau Physiographic Region 2
Figure 2. Relative abundance of rare plant taxa north and south of the Bitter Creek Divide in Capitol Reef National Park 11
Figure 3. Principal component ordination (PCA) of stands containing rare plant taxa in relation to geologic substrate in Capitol Reef National Park 13
Figure 4. Compositional patterns in pinyon-juniper woodlands in relation to elevation in Capitol Reef National Park 15
Figure 5. Patterns in minimum total grass cover within grassland stands along an elevational gradient in Capitol Reef National Park 16
Figure 6. Compositional patterns in pinyon-juniper woodlands in relation to geologic substrate in Capitol Reef National Park 17
Figure 7. Patterns in minimum total grass cover in relation to geologic substrate in grassland stands in Capitol Reef National Park 17
Figure 8. Patterns in species richness among plant community types in Capitol Reef National Park. ...18
Figure 9. Patterns in grazing intensity among plant community types in Capitol Reef National Park. .20
Figure 10. Patterns in minimum total grass cover in relation to elevation and grazing intensity in grassland stands in Capitol Reef National Park 21 Figure 11. Patterns in species richness of grassland stands in Capitol Reef National Park 21
Figure 12. Patterns in relative abundance of increaser (grazing tolerant) and decreaser (grazing intolerant) species in relation to grazing intensity in grassland stands in Capitol Reef National Park 22
Figure 13. Patterns in presence or absence of two especially sensitive grazing intolerant species (decreasers) in relation to grazing intensity in grassland stands in Capitol Reef National Park 24
Figure 14. Patterns in relative abundance of increaser (grazing tolerant) and decreaser (grazing intoler ant) species in relation to grazing intensity in waterpocket communities in Capitol Reef National Park 24
Figure 15. Relative abundance (% of species in a stand) of exotic (non-native) species in relation to elevation in grassland stands in Capitol Reef National Park 25
Figure 16. Overall patterns in distribution of plant community types in Capitol Reef National Park in relation to elevation and soil type 26
iv Vascular Flora and Vegetation of Capitol Reef National Park, Utah
List of Tables
Table 1. List of plant species which appear on federal lists of threatened, endangered, candidate species, and species no longer candidates for listing from Capitol Reef National Park 4
Table 2. Key locations of endangered, threatened and other plants of concern in Capitol Reef National Park 11
Table 3. Geologic substrates associated with species which appear on federal lists of threatened endangered, candidate species and species no longer candidates for listing 12
Table 4. Matrix of correlations among geologic formations in relation to the presence of rare plants at CARE 13
Table 5. Summary of community types and phases in Capitol Reef National Park 14
Table 6. Summary of individual species responses to livestock grazing in southern Utah 23
Table 7. Plant taxa found in CARE that are considered by Cronquist et al. (1972), by Welsh et al. (1975), and by ourselves to be endemic to the Canyon Lands Section 30
v Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Abstract
Capitol Reef National Park (CARE) is a shrub, grassland, and badlands communities; 98000-ha (242000-ac) reserve located in sandstones at low elevations and a variety of southern Utah, USA. It contains outstanding substrates at middle elevations support examples of the rugged topography, flora, several kinds of pinyon-juniper communities; fauna, and vegetation of this portion of the and cool, moist sites at high elevations are Colorado Plateau. In this study, we collected covered by woodland communities domi and documented the vascular flora, with nated by conifers or aspen. Riparian areas at all special attention on taxa of concern (endan elevations support woodlands and wetlands. gered or threatened or candidates for listing), Unusual and rare plant communities in CARE classified the plant communities in the park, include the bristiecone pine-cushion plant and described the distribution and patterns of community and the dogwood-spruce riparian plant communities in relation to gradients of woodland at high elevations; and hanging elevation, geologic substrate, and grazing gardens, waterpockets, and the hornbeam- intensity. boxelder-oak woodland at low elevations. We documented 759 vascular plant Past grazing by livestock has altered the species in CARE, representing 352 genera and composition and structure of many grassland 86 families. CARE supports at least 36 taxa and riparian communities in CARE. It may that are listed as endangered or threatened. require many decades of grazing protection This is one of the greatest concentrations in and possibly active intervention to restore the region of plant taxa of special concern. The these communities to their presettlement high plant diversity in CARE reflects the great condition. Recovery of community structure range of habitats present and the geographic probably will be more rapid in riparian areas location at the intersection of several bio- than in more arid grasslands, but restoration geographic regions. of original species composition may be slow We identified 34 plant community types in both kinds of areas. Establishment during within the park. Distribution of communities the 20th century of exotic plants, e.g., tamarix is controlled primarily by gradients in eleva and cheatgrass, has permanently changed the tion and geologic substrate. Dry, hot areas at composition of many plant communities in the lowest elevations support various upland CARE.
Introduction
Capitol Reef National Park (CARE) is a encompasses geologic strata representing a 98,000-ha (242,000-acre) reserve located in vast geologic time period from the Paleozoic southern Utah, USA (Figure 1). Established as Era to the present (Smith et al. 1963). Numer a National Monument in 1937 and expanded ous streams have cut through the fold, as a National Park to its present size in 1971, it exposing the rock layers and creating a contains outstanding examples of the rugged spectacularly rugged landscape. topography, spectacular geology, and unusual Although Capitol Reef's geology has been fauna and flora that characterize the Colorado mapped and described in some detail (Smith Plateau physiographic province (Fenneman et al. 1963, Billingsley et al. 1987), the flora 1931). The major feature of the park is the (plant taxa present) and vegetation (plant Waterpocket Fold, a large monocline that communities) previously have been relatively extends some 160 km (100 miles) from near little studied. The great diversity in geology the Colorado River in the south to the flanks and topography, the gradient in elevation of Thousand Lakes Mountain in the north, and (1180-2670 m [3900-8800 ft]), and the park's
1 Technical Report NPS/NAUCARE/NRTR-93/01
strategic biogeographic location near the ton 1931 [cited in Everitt 1970], Neese 1981), intersection of several major floristic regions, Uinta Basin (Goodrich and Neese 1986), San have resulted in a remarkable diversity of Francisco Peaks (Rominger and Paulik 1983), plant taxa and communities within the park's Natural Bridges National Monument (Welsh borders, including numerous endemic and and Moore 1968), Arches National Park rare taxa (Welsh and Chatterley 1985). The (Harrison et al. 1964), Canyonlands National purpose of this study was to inventory, map, Park (Loope 1977, Welsh 1970), Glen Canyon describe, and assess the flora and vegetation National Recreation Area (Tuhy and MacMa- of Capitol Reef National Park. hon 1988), and Grand Canyon National Park (Phillips et al. 1989). Within Capitol Reef itself, Previous Work: Floristics Susan Meyer conducted a preliminary inven Capitol Reef National Park lies in a tory in 1981-1982 and identified over 500 plant portion of the Colorado Plateau that has been taxa, but acknowledged that her inventory relatively poorly explored botanically. Previ was not exhaustive. ous floristic studies have been conducted in With the passage of the Endangered Species the general region, including the San Rafael Act in 1973, the protection of rare and en Swell (Harris, 1980), Henry Mountains (Stan demic plant species has increasingly become
Figure 1. Detailed map of Capitol Reef National Park with inset of location map of Capitol Reef National Park within the Colorado Plateau Physiographic Region. 2 Vascular Flora and Vegetation of Capitol Reef National Park, Utah a management concern in national parks and Previous Work: Vegetation other kinds of nature reserves. Thus, one No detailed classifications or analyses of important objective of our research was to the vegetation of the park as a whole have conduct a systematic inventory of rare plant been made, although some portions of the taxa in CARE (Bratton and White 1980, White vegetation have been studied. The vegetation and Bratton 1980). Although no inventories of of CARE has been classified and mapped at a rare plants in the park as a whole have previ coarse scale based on aerial photographs, ously been completed, occasional collections with nine vegetation types identified (e.g., and studies of local areas have been made. In woodlands, grasslands, badlands), and this particular, Susan Meyer and Keith Harrison information is contained within the park's provided several records of rare plant species geographical information system (GIS). Barth in CARE. Additionally, Pamela Camp and and McCullough (1988) analyzed the effects of Alice Lindahl conducted a survey of the flora, current and historic livestock grazing on vegetation and rare plant species of the Hall riparian vegetation in CARE, and Welsh Creek drainage (Figure 1), including lands (1988) described the riparian vegetation along both in and out of CARE, from The Post the Fremont River in the context of an assess southward to Lake Powell (Camp 1978). Camp ment of a proposed water development documented 12 rare plant species, 8 of which upstream. Fisher et al. (1991) and Cole (1991) are endemic to the Colorado Plateau but are have reconstructed changes in grassland and not on federal lists of endangered or threat woodland vegetation that occurred during the ened species. last several centuries to millenia. Vegetation Further documentation of rare plant taxa research in progress in CARE includes studies has come from field reports of the U. S. Hsh and of the effects of soil microphytic crusts on Wildlife Service, Endangered Species Office erosiun (West 1990 and personal communica botanist (e.g., Anderson 1985). Prior to the tion) and of plant community dynamics in initiation of this project, 17 taxa were recorded waterpockets (N. Henderson, personal com from CARE (Table 1) that were included in the munication). Review of Plant Taxa for Listing as Endangered Vegetation classifications have been or Threatened Species, by the U. S. Fish and developed for other units of the National Wildlife Service (Federal Register (F.R.) Park system on the Colorado Plateau, in 50[188]:39526-39584). These records usually take cluding Canyonlands National Park (Loope the form of voucher specimens in herbaria (e.g., 1977), Glen Canyon National Recreation BYU) or documentation in the literature (e.g., Area (Tuhy and MacMahon 1988), Grand Welsh and Chatterly 1985). Canyon National Park (Phillips et al. 1989), The U. S. Fish and Wildlife Service (USFWS) and Zion National Park (K. Harper, personal has classified rare plant species according to communication). Everitt (1970) classified and potential vulnerability, utilizing alpha mapped the vegetation of a portion of the numeric codes to denote the status of the Henry Mountains near CARE, and Singh plants. See Table 1 for a summary of the and West (1971) classified the Atriplex- USFWS status codes as used throughout this dominated plant communities near Green report. These status assessments are based on River, Utah. The USDA Soil Conservation information currently available to USFWS, Service (SCS) classified rangeland plant and they are subject to change. communities of a large area in southern Utah, These previous studies indicated the floristic including portions of CARE. Coniferous richness of Capitol Reef and the concentration of forest habitat types and aspen community rare plant taxa within the park, but none were types of southern Utah were identified by comprehensive surveys. Therefore, our study Youngblood and Mauk (1985) and by was initiated in part to provide the baseline Mueggler and Campbell (1986), respectively. information on park flora and vegetation An excellent overview of semi-desert and necessary for park management and future woodland vegetation in this region is pro research. vided by West (1988).
3 Technical Report NPS/NAUCARE/NRTR-93/01
Table 1. List of plant species which appear on federal lists of threatened, endangered, candidate species, and species no longer candidates for listing from Capitol Reef National Park. Federal status is indicated in parentheses, following the scientific name (see footnote for explanation of status codes). New records of plant species of concern are marked with an asterisk (*).
Cydadenia humilis var. jonesii (E) Near Deer Point Echinocereus triglochidiatus var. inermis (E)» Paradise Flats Erigeron maguirei var. maguirei
= Astragalus chamaeluce currently has a 3c status; however var. laccoliticus has not been assessed for potential threats, and because it represents a small fraction of the distribution of Astragalus chamaeluce, it is possible that it may be listed as a candidate.
The U. S. Fish and Wildlife Service (USFWS) has classified rare plant species according to potential vulnerability, utilizing alpha-numeric codes to denote the status of the plants. Those used in this report are defined as follows: E- Taxa listed as endangered; T- Taxa listed as threatened; 1- Taxa for which the USFWS has on file substantial information on biological vulnerability and threats to support the listing of these taxa as endangered or threatened species; 2- Taxa for which information now in possession of the USFWS indicates the possible appropriateness for the listing of these taxa as endangered or threatened species, but for which substantial information on biological vulnerability and threats is not currently available; 3- Taxa are no longer being considered for listing as endangered or threatened species, (3c) because they have proven to be more abundant or widespread than was previously believed and /or are not subject to any identifiable threat. These codes are used throughout this report. It should be noted, however, that these status assessments are based on information currently available to USFWS, and they are subject to change.
4 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Location Geology Capitol Reef National Park lies in Wayne, The geology of Capitol Reef National Park Garfield, Sevier, and Emery Counties, Utah is complex (Smith et al. 1963, Billingsley et al. (Figure 1). The highest elevation (2670 m [8800 1987). The oldest rocks are Permian in age and ft]) is in the northwestern end of the park, and are found only in the deepest canyons along the lowest elevation (1180 m [3900 ft]) is at the the western side of the park. Mesa Verde southern tip. The Bitter Creek Divide appears sandstone of Cretaceous age, the youngest to represent an important biogeographic sedimentary formation in the park aside from boundary: to the south a low desert type of Quaternary deposits, occurs along the eastern vegetation predominates, whereas to the north boundary. Volcanic activity occurred in the pinyon/juniper and high desert plant commu nearby areas of Boulder and Thousand Lake nities are more characteristic. At Fruita are Mountains at approximately 20 million years irrigated orchards and meadows. The Water- ago (Barnes 1978). Basaltic boulders were later pocket Fold contains many deep, stream-cut carried by glacial melt water down the moun canyons and nearly vertical rock walls. Runoff tains and are now scattered throughout much from north of the Bitter Creek Divide eventu of the park. As the Colorado Plateau was ally flows into the Fremont River; south of the gradually uplifted, the Waterpocket Fold was Bitter Creek Divide, most runoff flows into created; this prominent structure forms an Halls Creek and eventually to Lake Powell. eastward tilted bend that runs from Thousand Many portions of CARE are remote and Lake Mountain nearly 160 km (100 mi) to Lake difficult to reach. State Highway 24 crosses Powell (Olson and 01sonl986). CARE from east to west in the northern third Billingsley et al. (1987) identified 34 of the park. A scenic drive provides access to distinct geological units within Capitol Reef areas along the west side of the Waterpocket National Park and adjacent areas. These units Fold south of Fruita. An unimproved dirt road range from bedrock deposits of sandstone, runs through the Cathedral Valley (Middle limestone, shale, and mudstone; to intrusive Desert) region in the north, and another dirt and extrusive igneous rocks; to Quaternary road follows the Waterpocket Fold south to deposits of aeolian, alluvial, and colluvial "The Post" and then exits the park. A four origin. The variety of habitats created by these wheel drive road provides access to the Upper different geologic substrates has a profound Deep Creek region. The remainder of the park effect on the distribution of plant species and is accessible only by foot. Nearly 31 miles of communities, as is described below. trails provide some access to remote portions of the park, but many areas are far from any Climate trails, and the extremely steep and rugged The climate of Capitol Reef National Park topography makes much of the Waterpocket varies seasonally and with elevation, but few Fold inaccessible or nearly so. instrumental weather data are available for The inaccessibility of most of the park area this area. Records taken from 1948-1986 at is a major reason why its flora has not yet been CARE headquarters in Fruita (1670 m [5500 adequately documented. Accessible and well ft]) indicate a mean annual precipitation of collected areas in the park include: Fruita, 18 cm (7.2 in), with 30% falling in July and Fremont River Valley, Upper Deep Creek, August. Annual precipitation is estimated Cathedral Valley, Lower Halls Creek, Hick to be about 12.5 cm (5 in) at the lowest eleva man Bridge Overlook area, Capitol Gorge, tions along Halls Creek, and about 30 cm Chimney Rock area, Deer Point, Cohab/ (12 in) at the highest elevations in upper Frying Pan Canyons, Spring Canyon, Pleasant Deep Creek. An important reason for the Creek, Sulphur Creek, Grand Wash, and The generally low precipitation throughout Hartnet. One major objective in our research CARE is the location of the Waterpocket Fold was to inventory the more remote and poorly within an apparent rain shadow of the high collected portions of the park as thoroughly as Aquarius Plateau lying to the west (Everitt possible. 1970).
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The seasonal distribution of precipitation growing season, indicating severe and sus is bimodal in this region, with peaks in winter tained drought stress for plants. High soil and late summer (Everitt 1970). Much of the temperatures also create severe conditions for rainfall during July and August comes as plant growth: Wein and West (1972) measured sudden cloudbursts which can cause devastat soil surface temperatures >60 degrees C in ing floods along desert washes (Woolley July at a semi-desert site near Cisco, Utah. 1947). Winter precipitation generally is more Such temperatures exceed the recognized gentle and prolonged; a larger fraction of this upper limit of tolerance for most plant tissues. water percolates into the soil and supports Early spring probably is the only time when plant growth the following summer. moisture and temperature conditions are At the lower elevations, maximum consistently favorable for plant growth at low temperatures in the summer frequently exceed elevations in CARE. The climate diagrams 38 degrees C (100 degrees F) (Smith et al. indicate that higher elevations also have a 1963). Summer temperatures at higher eleva pronounced precipitation deficit in early tions in the park are more moderate (Park summer, but late summer "monsoon" showers Service data). Winter temperatures through may provide sufficient moisture to match out the park are typically cool to cold, with evaporative demands, at least in some years. freezing temperatures sometimes lasting several days. Freezing can occur as late as Specific Objectives May or as early as September. Park records 1. To complete an annotated list of indicate an average January minimum at vascular plant species in Capitol Reef National Fruita of -8 C (18 F) and an average July Park, identifying special status, general maximum of 33 C (92 F). location, abundance, habitat, geology, special West (1983a, 1983b; Charley and West use information, and elevation. 1975) has constructed climate diagrams for 2. To develop a plant community classifi several stations in southern Utah which have cation system by use of multivariate analysis. generally similar climatic conditions as those 3. To describe major vegetation patterns of CARE. These analyses show that in the by means of direct gradient analysis, and to lower elevations, potential evapotranspiration assess some of the effects of livestock grazing exceeds precipitation throughout most of the on grassland and riparian plant communities.
Methods
Inventory of Vascular Plants work we located several sampling points in To develop a complete vascular plant flora areas between the transects to sample the full for the park, it was necessary to systematically range of environmental gradients within the inventory all of the kinds of habitats present at park. various times during the growing season. This We walked each of these transects at least was accomplished by establishing a series of once early and late in the growing season, transects extending from east to west across the sampled the plant communities at each place park. The transects were located at intervals of where substrate, elevation, or topography one to several kilometers, depending on accessi changed substantially, and collected voucher bility and variability of habitats within specific specimens of the vascular plants. In this way we areas. Additional transects were established to inventoried the vascular flora over the entire reach special habitats not included in the east- range of elevations, geologic substrates, and west transects, and in the final year of field topographic features represented in the park, 6 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
throughout the growing season (April - October) examined in two ways. First, we constructed a during three years (1986,1987,1988). Voucher matrix of correlations between presence/ specimens were deposited in the herbaria at absence of a given taxon and geologic sub Brigham Young University (BYU), San Juan strate, based on our field inventory data. (At College (SJNM), and Capitol Reef National Park. each site where we found a population of rare plants, we recorded the geologic substrate and Inventory of Threatened other habitat information.) Secondly, we and Endangered Taxa ordinated the same data by means of prindpal Prior to field investigations, we reviewed component analysis (Gauch 1981). Ordination the taxa listed by the USFWS as threatened, is a multivariate technique of indirect gradient endangered and/or sensitive plant species (F.R. analysis that reveals the relative similarity and 50[188]:39526-39584) to compile a list of those dissimilarity among sampling units, in this taxa which were known from or near the park case geologic formations. (Welsh and Chatterly 1985, Welsh 1978, Welsh et al. 1975, Welsh and Thorn 1978). This initial Field Sampling of Community Composition list induded spedes which were documented Because of the large study area, the from the park as well as spedes reported near rugged and often barely accessible terrain, and enough that there was a potential that they may the diversity of species and habitats present, occur in the park. We also used the herbarium at we adopted a rapid field sampling technique Brigham Young University to locate previously emphasizing semi-quantitative, releve infor documented populations of spedes on the mation (Mueller-Dombois and Ellenberg USFWS list which were known to be in or near 1974). With this approach we sacrificed some the park. The herbarium also provided informa precision and accuracy in our measurements, tion regarding blooming and fruiting periods of but it was the only feasible way to sample the each spedes. vegetation of the entire park within a reason We then visited known localities of threat able time frame. It did provide adequate data ened, endangered and/or sensitive plant spedes for construction of a broad-scale classification during the respective blooming periods to and analysis of plant communities over the become familiar with both reproductive and entire park. Future studies of finer-scale vegetative characteristics of the plants. During patterns within some of the community types these visits, we took photographs and character that we have identified may be more appro ized the habitat (elevation, substrate, topo priately conducted using more intensive graphic position, slope, aspect, associated quadrat or transect sampling techniques. spedes). This habitat information helped us to We sampled a total of 344 stands during recognize similar areas along our sampling the field seasons (April-October) of 1986,1987, transects (described in the previous section) and 1988. Stands were located along the where previously unreported populations of sampling transects that we established to rare plants might be located. inventory the vascular flora (described above). We searched for rare plants along our Before going into the field, we selected the sampling transects (described above) through general locations where we wished to sample out the three field seasons of 1986,1987, and by studying USGS topographical maps and 1988. Wherever threatened, endangered and/ geology maps (Billingsley et al. 1987) and or sensitive plant spedes were encountered, identifying areas that would represent the we recorded the location on USGS topo entire range of variation in elevation and graphic maps, habitat characteristics, number geologic substrate within CARE. When we of plants, any apparent or potential threats, reached one of these general areas in the field, and areal extent of the population. we then selected a representative portion of it, established a sample stand, and recorded its Habitat Relationships of Rare Plants exact position on a 15-minute topographic The possible affinity of each rare plant map. Each stand was an area of approximately taxon for a particular geologic substrate(s) was 100 square meters (usually about 10m x 10m)
7 Technical Report NPS/NAUCARE/NRTR-93/01
within which the environment and the vegeta intent was not to produce a purely objective tion appeared more or less homogeneous. community classification, but to create a Stand borders were not staked or measured practical classification composed of a reason precisely, but were selected visually before able number of types that are easily recog sampling began. nizable in the field and can be used by manag In each stand we first recorded all vascu ers and future researchers. The final result was lar plant species present. Then, we visually our classification system containing 34 com estimated the proportion of total plant bio- munity types (see Results section). mass contributed by each species using five categories: 1-5%, 6-25%, 26-50%, 51-75%, and Gradient Analysis of the Vegetation 76-100%. (The releve method commonly To unravel the effects of environmental employs estimates of percent ground cover of variation and disturbance on the composition each species, but we based our sampling on of the vegetation, we employed direct gradi percent of total biomass present in order to ent analysis. Because our community compo better capture the dominance of the large trees sition data assigned each plant species to an and shrubs in sites where these growth forms abundance class (i.e., 1-5% or 6-25%, etc.) we were present.) We also noted species growing could not directly compare the actual abun in the general area but not present within the dance of each species or group of species borders of the stand. Finally, we recorded among different stands. Instead, we used slope, aspect, elevation, geological substrate, three modified gradient analysis techniques topographic position, and other general for comparison. information for each stand. Recent grazing First, for some analyses, e.g., total percent intensity was estimated visually on the basis cover of grasses in grasslands along gradients of livestock scat and tracks, and categorized as of elevation, soil type, and grazing intensity, none, light, moderate, or heavy. Any other we calculated a minimum total grass cover for kinds of disturbance, e.g., recent fire, also each stand. This was accomplished by taking were noted. the minimum value of the abundance class for each grass species, summing these values, and Community Classification then determining into which abundance class The species abundance data were sub the sum fit. For example, if a stand contained jected to cluster analysis using TWINSPAN two species with abundance class 2 plus two (Cornell Ecology Programs, Gauch 1981) on species with abundance class 3, the sum of the VAX computer at Fort Lewis College. This minimum values would be 1+1+6+6=14. A is a hierarchical, polythetic, divisive method of value of 14 falls within abundance class 3, so classification. The program produced 30 the minimum total cover of grass species clusters after 7 divisions. Next we evaluated within this stand would be abundance class 3. the floristic composition and environmental Although this approach was conservative and characteristics of the individual stands within insensitive to subtle differences among stands, each cluster. Most of the clusters represented it did reveal several large differences in more or less distinctive and easily recogniz vegetation composition that could be ex able species assemblages, and these were then plained in terms of environmental gradients regarded as community types in our clasifica- and /or disturbance history. tion. We combined some clusters to produce a Our second technique of direct gradient single community type because of their analysis was to determine the proportion apparent physiognomic and floristic similar of sampled stands that fell into each of ity. We also divided some clusters into more several categories of interest. For example, than one community type because of differ we calculated the percent of pinyon-juniper ences in dominant species, growth forms, or stands in which pinyon had a higher abun environmental conditions that were conspicu dance rank than juniper, in which juniper ous in the field even though they did not have had a higher rank than pinyon, and in which a strong effect in the clustering process. Our their abundance ranks were equal. Again, 8 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
this method was insensitive to subtle differ The gradient analysis of the vegetation ences, but it did reveal major trends. revealed strong correlations between Finally, we plotted locations of stands community types and combinations of eleva representing each community type along tion, geologic substrate, slope, aspect, and axes of elevation and relative soil moisture topographic position. Therefore, Boolean (a function of substrate) to depict the distribu analysis program was used to identify all tion of plant communities along the two locations within the park where each particu major controlling gradients in the park, viz., lar combination of environmental conditions elevation and substrate. occur; these areas then were classified as the appropriate community type within a Vegetation Mapping new map of vegetation. See Romme et al. A vegetation map was produced using the (1993) for details on producing the vegetation park's geographical information system (GIS). map.
Results
Vascular Flora currently listed with the USFWS. Although To date, 759 species in 352 genera and 86 this is a sizable number of taxa, it represents families are known from Capitol Reef Na only about 4.6% of the park's total vascular tional Park, based on our inventory as well as flora. previous studies. The total number of specific Four of these taxa have been designated as and infraspecific taxa is 806. We found 316 endangered species. Two of the endangered previously unreported taxa in the park. The taxa, Echinocereus triglochidiatus var. inermis largest plant families were the Gramineae and and Cycladenia humilis var. jonesii, are known Compositae, with 105 and 155 taxa respec from single localities in CARE; however, they tively. In contrast, several families were are also known from a few populations quite represented by only one species (e.g., Se- distant from the park in Utah and Colorado. laginellaceae, Cannabinaceae, Lentibu- The remaining two, Sclervcactus wrightiae, and lariaceae, and Commelinaceae. Erigeron maguirei var. maguirei, while local and An annotated checklist of vascular plants restricted in distribution within the park, are is provided in Appendix I. This list includes represented by numerous occurrences within information on general distribution, eleva- CARE. tional range, and plant community(s) for each One species, Townsendia apnea, is classified taxon listed. Nomenclature follows Welsh et as threatened. We found it to be somewhat al. (1987) for the most part, though in some more common than prior accounts indicated, instances Cronquist et al. (1972,1977,1984) with populations scattered from Upper was followed. A total of 432 new plant speci Cathedral Valley to the Hartnet, at Upper mens were deposited in CARE's herbarium. Deep Creek, and overlooking the Fremont River near Miners Mountain. Aside from these Threatened and Endangered Taxa populations in CARE, the only other known New distributional records produced by locations are north of the park in the San our inventory indicate that an additional 19 Rafael Swell. taxa included in the Review of Plant Taxa for In addition to the officially listed endan Listing as Endangered or Threatened Species, gered and threatened taxa, ten other taxa in by the U. S. Fish and Wildlife Service (Federal CARE are federal candidates, of which four Register 50[ 1881:39526-39584; F. R. 55[35]:6184- are classified as Category 1 taxa and should be 6229) are found in CARE (Table 1). This brings officially listed as either endangered or to 36 the total number of species in CARE threatened within the near future. These four 9 Technical Report NPS/NAUCARE/NRTR-93/01
include Gilia caespitosa, Pediocactus winkleri, components, which represent approximately Spiranthes diluvklis, and Schoencranbe barnebyi. 50% of the variance in the data. liabitats Finally, 21 additional taxa are classified as 3C. underlain by the Kayenta, Wingate, Navajo, Kaibab, and Cutler Formations — all of which Habitats of Rare Plant Species produce relatively coarse textured soils — We found several key locations in CARE support similar taxa of rare plants. Habitats (Figure 1) that contained large concentrations underlain by the finer textured Mancos, of rare plant taxa, including upper Deep Creek Curtis, Entrada, Summerville, and Moenkopi (8 taxa), the Hartnet area (6 taxa), the Hickman Formations support a different group of rare Bridge area (6 taxa), Deep Creek in the Water- plant taxa. The Quaternary geologic substrates pocket Fold (5 taxa), and Miners Mountain (5 were isolated in the ordination space, indicat taxa). The southern portion of the park, south ing that they were floristically dissimilar from of Bitter Creek Divide, apparently supports either the coarse-textured or the fine-textured fewer taxa of concern than an equivalent area groups. This probably is because several rare in the northern part of the park (Table 2). plant taxa are found exclusively on Quater To further examine this pattern in rare nary alluvial deposits, and also because plant distribution, we examined two arbi several species from both of the other major trarily selected sets of seven areas on the map, groups are found on Quaternary alluvium as each area comprising two square miles. All of well. The rare plants from the Chinle, Morri the areas in one set were located north of the son, and Carmel Formations were not strongly Bitter Creek Divide, and all in the other set associated with those of any other formations. were located south of the Divide. We found a significant difference (alpha < 0.05) between Plant Community Classification the number of rare plants within comparable We identified 34 plant communty types areas north and south of Bitter Creek Divide within the park, plus several phases within (Figure 2). some community types (Table 5). The vegeta Of the 36 taxa of concern, only 14 are tion in CARE appears to be a continuum apparently restricted to a single geological rather than a series of discrete units, except formation (Table 3). Seven of these 14 are where underlying physical factors, such as soil associated with Quaternary formations, either texture or moisture, change abruptly. Continu alluvial, colluvial, pediment, or eolian depos ously varying vegetation still can be classified, its. However, certain groups of petrologically but it must be recognized that some of our similar geologic formations tend to support distinctions are more or less arbitrary. similar assemblages of rare plants. Table 4 We define a community as a more or less shows the correlations of geologic formations unique and repeated assemblage of plant based on the occurrence of rare plants on these species occupying sites characterized by a formations. High correlations occur between more or less unique combination of elevation Navajo, Kayenta and Wingate formations. and soil texture conditions. A phase is a This indicates that those taxa of concern recognizable subunit of a community type, associated with any one of these predomi characterized by a somewhat unusual assem nantly sandstone formations (e.g., Navajo) are blage of species and occupying a portion of likely to be found on the other two as well the usual habitat for that community type (e.g., Kayenta and Wingate). Significant with no recognizably unique environmental correlations also exist between the Summer conditions. Phases may reflect subtle, unrecog ville and Entrada Formations, which contain nized patterns in underlying environmental mostly clays, mudstones, and siltstones in gradients; or unique historical events of CARE. dispersal, establishment, or disturbance; or These relationships among geologic bom. They cannot be mapped reliably, but substrates also were demonstrated in the they contribute to the park's biological diver principle component analysis (PCA) ordina sity, and so are included in this classification. tion. Figure 3 shows the first three principle Romme et al. (1993) provide a dichotomous
10 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Table 2. Key locations of endangered, threatened and other plants of concern in Capitol Reef National Park (see Table 1 for a list of these species and a summary of their status according to the U.S. Fish and Wildlife Service). Each location represents an area of approximately two square miles and the number of rare species found therein. Many individual species are found in more than one location. The division of the park into northern (N) and southern (S) regions coincides with a natural divide (Bitter Creek Divide) but is otherwise arbitrary.
Location Number of Species North or South
Miners Mountain 5 N Upper Deep Creek 8 N Deep Creek on the Fold 5 N The Hartnett 6 N Cathedral Valley 3 N Paradise Flats N Hickman Bridge N 4 mi. S of Bitter Creek Divide S Halls Narrows S Below Burr Trail S The Post S Red Slide S Wagon Box Mesa s Top of Burr Trail s
Figure 2. Relative abundance of rare plant taxa north and south of the Bitter Creek Divide in Capitol Reef National Park Vertical lines are ±1 standard deviation.
11 Technical Report NPS/NAUCARE/NRTR-93/01
Table 3. Geologic substrates associated with species which appear on federal lists of threatened, endangered, candidate species and species no longer candidates for listing. The asterisk indicates the geologic formation on which the species is most often associated (if any), when it is found on multiple substrates.
Cycladenia humilis var. jonesii (E) Chinle Echinocereus triglochidiatus var. inermis (E) Qal Erigeron maguirei var. maguirei (E) Navajo, Cutler Sclerocactus wrightiae (E) Entrada, Mancos*, Summerville, Qal Townsendia aprica CD Cutler, Entrada, Moenkopi, Summerville Gilia caespitosa (l) Kayenta, Navajo*, Carmel Pediocactus ivinkleri (i) Summerville*, Entrada, Carmel Schoencrambe barnebyi (l) Moenkopi*, Chinle, Cutler Spiranthes diluvialis (i) Qal Cymopterus beckii (2) Navajo, Kayenta, Wingate, Cutler Dalea flavescens var. epica (2) Qal Erigeron maguirei var. harrisonii (2) Kayenta, Navajo Habenaria zothecina (2) Qal Hymenoxys depressa (2) Entrada, Summerville, Curtis, Mancos Thelesperma subnuda var. alpina (2) Carmel Asclepias ruthiae (3c) Entrada, Qal Astragalus barnebyi (3c) Morrison Astragalus chamaeluce var. laccoliticus O Mancos*, Qal Astragalus consobrinus (3c) Kayenta, Entrada, Moenkopi, Summerville, Qal Astragalus harrisonii (3c) Kayenta, Navajo, Wingate, Qal Astragalus malacoides (3c) Mancos, Moenkopi, Qal Astragalus pardalinus (3c) Entrada*, Qal Astragalus woodruffii (3c) Entrada, Summerville, Qal Castilleja scabrida (3c) Kayenta, Navajo, Wingate, Qal Chamaechenactis scaposa (3c) Entrada, Moenkopi Cryptantha johnstonii (3c) Carmel Erigeron abajoensis (3c) Navajo*, Cutler Eriogonum corymbosum revealianum (3c) Carmel, Qal Eriogonutn tumulosum (3c) Moenkopi, SummervUle Geranium marginale (3c) Qal Lomatium junceum (3c) Moenkopi Noma retorsum (3c) Qal Parthenium ligulatum (3c) Entrada, Moenkopi, Summerville Peteria thompsonii (3c) Entrada, Qal Physaria acutifolia var. purpurea (3c) Carmel Xylorhiza confertifolia (3c) Chinle
12 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Table 4. Matrix of correlations among geologic formations in relation to the presence of rare plants (those on federal lists; see Table 1) at CARE.
Morri Curtis Kaibab Chinlc Win- Mancos Cutler Carmel Navajo Moen• Summer - En- Qal son gate enta kopi ville trad.•
Curtis -0.028 1.000 Kaibab -0.040 -0.040 1.000 Chink •0.050 -0.050 -0.071 1.000 Wingate -0.050 -0.050 0.367 •0.088 1.000 Mancos -0.058 0.497 -0.083 •0.103 O.103 1.000 Cutler -0.066 -0.066 0.605 0.172 0.172 -0.138 1.000 Carmel -0.073 -0.073 -0.105 0.131 0.131 •0.153 0.174 1.000 Kayenta -0.073 -0.073 0219 •0.13 1 0.675 -0.153 0.041 0.005 1.000 Navajo -0.081 -0.081 0.495 • 0.143 0.615 -0.168 0.415 -0.025 0.724 1.000 Moenkopi -0.088 -0.088 -0.126 0.085 0.156 0.029 0.176 -0231 O.053 -0254 1.000 Summerville -0.088 0.317 -0.126 •0.15 6 •0.15 6 0240 O.016 -0.053 O.053 -0254 0.362 1.000 Entrada -0.108 0.256 •0.155 -0.193 •0.19 3 0.154 0.084 -0.126 0.126 -0.314 0233 0.664 1.000 Qal -0.154 -0.154 •0220 -0274 0.124 0203 • 0.364 -0258 0.111 -0.168 -0.352 0221 • O.006 1.000
Figure 3. Principal component ordination (PCA) of stands containing rare plant taxa in relation to geologic substrate in Capitol Reef National Park. Factor 1 represents 22.7% of the variance, factor 2 represents 14.8%, and factor 3 represents 13.0% of the variance. Substrates producing coarse textured soils are depicted with a triangle; those with fine textured soils are depicted with a square. Substrates depicted with an "x" or a circle produce a variety of soil textures, and are largely uncorrelated with the three axes.
13 Technical Report NPS/NAUCARE/NRTR-93/01
Table 5. Summary of community types and phases in Capitol Reef National Park. See text for definitions of community types and phases, and Appendix I for more detailed descriptions. A. Badlands Communities D3a. Cushion Plant Phase Al. Bentonite Badlands Community D3b. Galleta Phase A2. Hopsage Badlands Community D3c. Pygmy Sagebrush Phase A3. Mat Saltbush Community D4. Pinyon-Juniper-Tall Shrub Community A3a. Shrub Phase D5. Dwarf Mountain-Mahogany Slickrock A3b. Grassland Phase Community A3c. Buckwheat Phase A4. Gypsum Badlands Community E. Upland Forest and Woodland Communities El. Bristfecone Pine-Cushion Plant Community B. Grassland Communities E2. Ponderosa Pine-Bitterbrush Woodland Bl Shadscale-Grassland Community Community B2. Eolian Grassland Community E2a. Mountain-Mahogany Phase B3. Mixed Grassland Community E3. Ponderosa Pine-Manzanita Woodland Community C. Upland Shrub Communities E4. Mesic Montane Woodland Community CI. Eolian Shrub Community E5. Aspen Woodland Community CI a. Blackbrush-Mormon Tea Phase Clb. Sand Sagebrush Phase F. Riparian and Wetland Communities C2. Blackbrush-Rabbitbrush Community Fl. Cottonwood-Rabbitbrush Woodland C3. Intermittent Riparian Shrub Community Community C3a. Greasewood-Rabbitbrush Phase F2. Waterpocket Community C3b. Fourwing Saltbush-Rabbitbrush F3. Cottonwood-Willow Riparian Woodland Phase Community C3c. Big Sagebrush Phase F3a. Reed Phase C4. Low Shrub Community F3b. Baltic Rush Phase C5. Dry Canyonbottom Shrub Community F4. Alder-Birch Riparian Community C6. Mesic Canyonbottom Shrub Community F5. Dogwood-Spruce Riparian Woodland C7. Sagebrush-Bitterbrush Community Community F6. Hanging Garden Community D. Pinyon-Juniper Woodland Communities F7. Hornbeam-Box Elder-Oak Woodland Dl. Pinyon-Juniper-Grass Community Community D2. Juniper-Mormon Tea Community F8. Cultivated Orchards and Settlements D3. Pinyon-Juniper-Low Shrub Community F9. Perennial Wetland Community
key for identification of plant communities in distinct assemblages of species that we had not CARE, plus a detailed description of each previously recognized in the field, but that were community type and phase, in terms of species quite apparent when we looked for them. composition, associated environmental condi tions, and major responses to disturbances. Gradient Analysis of the Vegetation The community classification that we The major abiotic variables controlling the produced is a synthesis of the objective results distribution and composition of plant commu of the cluster analysis and our more intuitive nities in CARE are elevation and geologic field experience with the park's vegetation. In substrate. Disturbance by livestock grazing, general, the clusters produced by the objective both past and present, also modifies the analysis coincided nicely with the vegetation vegetation. We analyzed each of these factors units that we perceived in the field. Several separately, and then looked for interactions clusters that were very distinct in the objective among them, as presented below. analysis appeared far less distinct in the field, and so they were combined with other, similar- Elevational Patterns appearing groups in the final classification. The We examined vegetational changes along cluster analysis also drew attention to a few an elevational gradient within our sample 14 Vascular Flora and Vegetation of Capitol Reef National Park, Utah stands classified as pinyon-juniper woodland variables, e.g., substrate (see below). These and as grassland communities. All commu results are consistent with those reported nity-types varied with elevation, but these were elsewhere in the southwestern USA (Woodin the communities for which we had an adequate and Lindsey 1954, Tueller et al. 1979, Pieper sample size to uncover the details of the pat and Lymbery 1987). terns. We aggregated all six of the pinyon- The minimum total grass cover in grass juniper types (i.e., pinyon-juniper-grass, juniper- land communities generally increased with Mormon tea, juniper-sal tbush, pinyon-juniper- elevation (Figure 5). For this analysis we low shrub, pinyon-juniper-tall shrub, and dwarf examined only ungrazed and lightly grazed mountain mahogany slickrock communities) for stands, since grazing also influences grass this analysis; and similarly aggregated all three cover (see below). The progressively greater grassland types (i.e., shadscale grassland, eolian cover of graminoid species with increasing grassland, and mixed grassland communities). elevation is probably due to the increasing The relative abundance of Pinus edulis and quantity and reliablility of precipitation at Juniperus osteosperma varied with elevation higher elevations. (Figure 4). Generally, juniper was dominant in stands at lower elevations, especially those Substrate Patterns below 1700 m (5600 feet), and pinyon was Geologic substrate had an important dominant in stands at the highest elevations, influence on the relative abundance of Pinus especially above 2120 m (7000 feet). Between edulis and Juniperus osteosperma in pinyon- 1700-2120 m, there was a mixture of domi juniper woodlands located between 1700-2120 nance patterns, reflecting the influence of other m (Figure 6). Generally, juniper was more
Figure 4. Compositional patterns in pinyon-juniper woodlands in relation to elevation in Capitol Reef National Park. "]" indicates that juniper (Juniperus osteosperma) is the only one of the two species present; "]>V" indicates that juniper is more abundant than pinyon (Pinus edulis); "]=?" indicates that pinyon and juniper are roughly equal in abundance; "P>J" indicates that pinyon is more abundant than juniper; and "P" indicates that pinyon is the only one of the two species present. The numbers in parentheses at the top represent the number of sampled stands in each elevational class.
15 Technical Report NPS/NAUCARE/NRTR-93/01
Figure 5. Patterns in minimum total grass cover within grassland stands along an elevational gradient in Capitol Reef National Park. "3" depicts plant cover class 3 (26-50%) in our releve samples; "4" depicts cover class 4 (51-75%); and "5" depicts cover class 5 (76-100%). The numbers in parentheses at the top are the number of sampled stands in each elevation class. This analysis was restricted to ungrazed and lightly grazed stands only. abundant than pinyon on fine-textured ties (Figure 7). This analysis was done only substrates (clays, siltstones, mudstones), with ungrazed and lightly grazed stands to whereas the two species were roughly equal eliminate grazing interactions. Relatively or pinyon was dominant on coarse-textured sandy soils developing in eolian and alluvial substrates (sandstones, colluvium). This substrates generally supported a higher cover pattern is consistent with the elevational of graminoids than finer textured soils devel partem reported above, in that drought stress oping in terrace gravel deposits and Entrada is generally greater both at lower elevations sandstone. This may be because sandy soils and in finer textured soils. These results most effectively retain the moisture from indicate that juniper has greater drought intermittent summer rain showers within the tolerance than pinyon. Everitt (197051) re upper portion of the soil profile where most of ported that the lower elevational limit of the roots of the grasses are located. pinyon-juniper vegetation in the Henry Moun tains several km east of CARE was dependant Community Patterns on geological substrate: the lower limit was We examined patterns in species richness approximately 1515 m (5000 ft) on coarse- by calculating the mean, standard deviation, textured soils developing from indurated and range in number of species listed for each sandstones, approximately 1820 m (6000 ft) stand classified within a particular community in shallow gravels overlying porphyry, and type or phase. There was a surprising similar approximately 2120 m (7000 ft) on fine- ity in richness among many community types; textured soils developing from Mancos shales. mean number of species per stand was Geologic substrate also influenced mini between 20 and 30 in 21 of the 36 community mum total grass cover in grassland communi types and phases (Figure 8). The richest
16 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Figure 6. Compositional patterns in pinyon-juniper woodlands in relation to geologic substrate in Capitol Reef National Park. "]" indicates that juniper (Juniperus osteosperma) is the only one of the two species present; "J>P" indicates that juniper is more abundant than pinyon (Pmus edulis); "]=V" indicates that pin yon and juniper are roughly equal in abundance; "P>J" indicates that pinyon is more abundant than juniper; and "P" indicates that pinyon is the only one of the two species present. The numbers in parentheses at the top represent the number of sampled stands in each elevational class. This analysis was restricted to stands at 6000-7000 feet (1820-2120 m), which is the elevational zone where pinyon and juniper are roughly equal in abundance.
Figure 7. Patterns in minimum total grass cover in relation to geologic substrate in grassland stands in Capitol Reef National Park "3" depicts plant cover class 3 (26-50%) in our releve samples; "4" depicts cover class 4 (51-75%); and "5" depicts cover class 5 (76-100%). The numbers in parentheses at the top are the number of sampled stands in each elevation class. This analysis was restricted to ungrazed and lightly grazed stands only. Qed=Quatemary eolian deposits; Qat=Quaternary terrace gravel deposits; Je=Entrada formation; Qal=Quaternary alluvium.
17 Technical Report NPS/NAUCARE/NRTR-93/01
Number of Species/Stand
Sagebrush-Bitterbrush Ponderosa Pine-Manzanita Cottonwood-Willow Ponderosa Pine-Bitterbrush Waterpocket Cotton wood-Rabbirbrush Mesic Canyon-Bottom Shrub Mesic Montane Woodland Perennial Wetland Eolian Shrub Pinyon-Juniper Grass Dry Canyon-Bottom Shrub Mixed Grassland P/J-Low Shrub Galleta Phase Pinyon-Juniper Low Shrub Intermittent Riparian Shrub Int. Riparian Shrub Big Sage Phase Blackbrush-Rabbitbrush P/J -Low Shrub Cushion Plant Phase Aspen Pinyon-Juniper Tall Shrub Eolian Grassland Shadscale Grassland Mat Saltbush Grass Phase
Hophornbeam-Boxelder-Oak Gypsum Badlands Dwarf Mt-Mah. Slickrock
Juniper-Saltbush Mat Saltbush Bristlecone Pine Cushion Plant Hanging Gardens Bentonite Badlands Mat Saltbush Shrub Phase Dogwood-Spruce Mat Saltbush Buckwheat Phase Hopsage Badlands
Figure 8. Patterns in species richness among plant community types in Capitol Reef National Park. Horizontal lines represent the mean and range, and shaded areas represent one standard deviation, of the number of vascular plant species recorded in our releve plots of approximately 100 square meters area. Numbers in parentheses indicate the number of sampled stands within each community type.
18 Vascular Flora and Vegetation of Capitol Reef National Park, Utah communities included the ponderosa pine Effects of Livestock Grazing types, various riparian types, and pinyon- The influence of gradients in elevation juniper woodland types. Generally, richness and geologic substrate, described above, was higher in those communities subjected to interacts with the effects of livestock grazing less drought stress, i.e., those at higher eleva to produce some complex patterns in the tions or in well watered habitats at low vegetation of CARE. For example, minimum elevations. Lowest richness generally was total grass cover generally increased with found in the various badlands community elevation in ungrazed and lightly grazed types which are subjected to regular and grasslands (Figure 5), but the opposite pattern severe drought stress, sometimes in combina was seen in moderate to heavily grazed stands tion with high soil temperatures and salinity. (Figure 10). The explanation for this shift is There were some apparent exceptions to unknown (in fact, it may be merely an artifact the generalizations in the previous paragraph. of small sample size). Hanging gardens and the dogwood-spruce Richness, i.e., total number of species per community—both located in habitats with a stand, did not show any consistent pattern perennial water supply—were among the with either elevation or grazing intensity in least rich communities; and the highest mean grassland communities (Figure 11). One richness was in the xeric-appearing sagebrush- would expect heavy and prolonged grazing to bitterbrush community. These seemingly eliminate the most palatable species in a disparate results may be simply an artifact of community, but they may be replaced by small sample size. Alternatively, they may other species tolerant of grazing, such that the reflect other unrecognized processes that total number of species remains unchanged. influence richness and community structure. Figure 12 shows the relative abundance of The relatively high richness in the pinyon- grazing intolerant species ("decreasers") and juniper types was somewhat surprising, since grazing tolerant species (native "increasers" these communities occupy only an intermediate plus exotic "invaders") in the grassland position in the elevational moisture gradient. communities. Grazing tolerance was deter However, pinyon-juniper woodlands usually mined from the literature (Table 6). The are found in habitats characterized by deep, minimum total cover of grazing tolerant coarse textured soils or shallow soils over frac species was generally greater than that of tured bedrock, e.g., in sandstones and colluvium. intolerant species even in lightly grazed These kinds of soils provide deep soil moisture stands, but the dominance of "increasers" and which can be reached by the roots of woody "invaders" became even greater with increas shrubs and trees, thus ameliorating the adverse ing grazing intensity (Figure 12). effects of relatively low total precipitation We also examined the presence or absence (Everitt 1970, Loope 1977). Thus, pinyon-juniper of two preferred forage species that are very woodlands in CARE actually can be viewed as sensitive to grazing pressure, the shrub occupying comparatively mesic habitats. Ceratoides lanata and the grass Stipa comata, in The various community types and phases relation to grazing intensity in grassland differed greatiy in the degree to which they have communities. Approximately half of even the been affected by past and present livestock lightly and moderately grazed stands, and grazing (Figure 9). Generally, the most affected three fourths of the heavily grazed stands, communities were those characterized by gentle lacked both species (Figure 13). topography, easy accessibility, and lower Grazing has modified the structure of elevations. The most heavily grazed community lower-elevation riparian plant communities as types include the various grassland communi well as grasslands in CARE (also see Barth ties, and both intermittent and perennial and McCullough 1988). Figure 14 shows the riparian communities at low elevations. The relative abundance of grazing tolerant vs communities least affected by grazing were intolerant species in waterpocket communities those characterized by steep and rocky habitats, subject to either light or heavy grazing pres lack of accessibility, or very little palatable sure. Grazing intolerant species were equal or vegetation. greater in abundance than tolerant species in 19 Technical Report NPS/NAUCARE/NRTR-93/01
Percent of Sampled Stands
Dry Canyon-Bottom Shrub Intermittent Riparian Shrub Cottonwood-Rabbitbrush Mixed Grassland Mat Saltbush Shrub Phase Eolian Grassland Waterpocket Int. Riparian Shrub Big Sage Phase Mat Saltbrush Grass Phase Mat Saltbush Pinyon-Juniper Grass Shadscale Grassland Cottonwood-Willow P/J-Low Shrub Galleta Phase Pinyon-Juniper Low Shrub Juniper-Saltbush Bentonite Badlands Aspen Sagebrush-Bitterbrush Blackbrush-Rabbitbrush Eolian Shrub Dogwood-Spruce Hanging Gardens Ponderosa Pine-Bitterbrush Hopsage Badlands Mesic Canyon-Bottom Shrub Gypsum Badlands Pinyon-Juniper Tall Shrub Mat Saltbush Buckwheat Phase Mesic Montane Woodland Hophornbeam-Boxelder-Oak P/J -Low Shrub Cushion Plant Phase Dwarf Mt.-Mah. Slickrock Bristlecone Pine Cushion Plant Ponderosa Pine-Manzanita Perennial Wetland
Figure 9. Patterns in grazing intensity among plant community types in Capitol Reef National Park. "H" depicts heavy grazing; "M" depicts moderate grazing; and "L" depicts light or absence of grazing. These are subjective estimates of grazing intensity, made at each sampling site. The numbers at the top indicate the number of sampled stands in each community type.
20 Vascular Flora and Vegetation of Capitol Reef National Park, Utah Figure 10. Patterns in minimum total grass cover in relation to elevation and grazing intensity in grassland stands in Capitol Reef National park. "3" depicts plant cover class 3 (26-50%) in our releve samples; "4" depicts cover class 4 (51-75%); and "5" depicts cover class 5 (76-100%). The numbers in parentheses at the top are the number of sampled stands in each eleva tion/grazing class.
Figure 11. Patterns in species richness of grassland stands in Capitol Reef National Park. Horizontal lines represent the mean and range, and shaded areas represent one standard deviation, of the number of vascular plant species recorded in our releve plots of approximately 100 square meters area. "H" depicts heavy grazing; "M" depicts moderate grazing; and "L" depicts light or absence of grazing. These are subjective estimates of grazing intensity, made at each sampling site. Numbers in parentheses indicate the number of sampled stands within each elevation/grazing class. 21 Technical Report NPS/NAUCARE/NRTR-93/01
Figure 12. Patterns in relative abundance of increaser (grazing tolerant) and decreaser (grazing intolerant) species (see Table 6) in relation to grazing intensity in grassland stands in Capitol Reef National Park. "D>I" indicates that decreaser species are more abundant (greater total cover) than increaser species; "D=I" indicates that decreasers and increasers are roughly equal in abundance; "I>D" indicates that increasers are more abundant than decreasers; and "I»D" indicates that increasers are far more abundant than decreasers. Numbers in parentheses at the top indicate the number of sampled stands in each grazing class. Grazing classes are based on subjective estimates of grazing intensity at each sampling site.
all five of the lightly grazed stands; but in the variables are elevation and soil characteristics. two heavily grazed stands, tolerant species Elevation is a complex gradient incorporating were more abundant than intolerant in one changes in both temperature and precipita stand, and the two classes were equal in the tion. Generally, temperatures decrease and other. Interestingly, we found no moderately precipitation increases with increasing eleva grazed waterpockets; all were either lightly tion. grazed (perhaps ungrazed) or heavily grazed. The soil gradient in Figure 16 represents This probably is because the waterpockets are primarily a gradient in water availability. The such favorable habitats for livestock that they most mesic soils are those with a perennial graze heavily in all waterpockets that they can water supply. The next most mesic soils are reach. either coarse-textured (e.g., colluvium) or Grazing and other disturbances have overlie deeply fractured bedrock (e.g., some created numerous opportunities in CARE for sandstones); both of these conditions provide exotic species to become established. Many of deep water supplies that can be tapped by these are undesirable weedy species (Table 6). woody species having deep roots (Everitt Exotic species comprised an average of 7% of 1970, Loope 1977). The most xeric soils are the flora in grassland stands located below either very shallow and developing over 5000 ft in CARE, but were smaller components impenetrable bedrock (e.g., some sandstones), of the flora at higher elevations (Figure 15). or of very fine texture such that precipitation tends to run off rather than infiltrate (e.g., Overall Vegetation Patterns in CARE clays and silts). Figure 16 summarizes the major patterns It is evident from Figure 16 that several in the distribution of plant communities in community types in CARE are largely re CARE. The two most important controlling stricted to a single soil or rock type, even 22 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Table 6. Summary of individual species responses to livestock grazing in southern Utah. Numbers in parentheses indicate the references which are listed at the bottom of the table. Species with an asterisk (*) have been classified as both increasers and decreasers, depending upon specific site and grazing conditions. Invaders are non-native species (mostly of Eurasian origin). Decreasers under Grazing Pressure Shrubs: Grasses: Artemisia biglovii (1) Bouteloua eriopoda (1) Artemisia spinescens (1,2) Hilaria jamesii (1) * Atriplex canescens (1,4) Muhlenbergia porteri (1) Atriplex cuneata (1,4) * Oryzopsis hymenoides (1,4) Ceratoides lanata (1,4) Poa fendleriana (1) Ephedra torreyana (1) Sitanion hystrix (1) Eriogonum microthecum (1) Sporobolus airoides (1) Kochia americana (1,2) Sporobolus cryptandrus (1) Salix spp. (1) Stipa cotnata (1,4) Forbs: Sphaeralcea spp. (1) Increasers under Grazing Pressure Shrubs: Trees: Artemisia filifolia (1,4) Juniperus osteosperma (1) Artemisia tridentata (1,2) Pi'nus edu/is (1) Atriplex confertifolia (1,2) Forbs: Atriplex corrugata (2) Astragalus spp. (1) Atriplex cuneata (2) * Caesalpinia repens (1) Atriplex falcata (2) Coldenia hispidissima (1) Atriplex gardneri (2) Cordylanthus spp. (1) Chrysothamnus nauseosus (1,4) Cryptantha flava (1) Chrysothamnus viscidiflorus (1,4) Eriogonum inflatum (1,4) Coleogyne ramosissima (1,4) Lupinus pusillus (1) Ephedra cutleri (1) Machaeranthera canescens (4) Ephedra viridis var. viscida (4) Opuntia spp. (1,4) Eriogonum corymbosum (1) Sphaeralcea spp. (4) * Gutierrezia sarothrae (1,4) VVyef/iw scabra (1) Quercus gambelii (1) Grasses: Sarcobatus vermiculatus (1,4) Aristida purpurea (1) Tetradymia spp. (3) Bouteloua gracilis (1,4) Vanckuefl stylosa (1,4) Hilaria jamesii (1,4) * Yucca spp. (1) Muhlenbergia pungens (1,4) Sitanion hystrix (3) *
Invader Species Forbs: Atriplex heterosperma (2) Halogeton glomeratus (1,2) Atriplex hortensis (2) Helianthus annuus (1,4) Atriplex patula (2) Lepidium perfoliatum (2) Atriplex rosea (2) Malcomia africana (2) Ceratocephalus testiculatus (2) Sfl/so/fl to/i (1,4) Chenopodium album (2) Grasses: Chenopodium botrys (2) Bromus rubens (2) Bromus tectorum (1,2) References: (1) Jarman, T., and H. Swenson. 1984. Ecological Site Descriptions, SCS-BLM Utah, unpublished draft report. (2) West (1988), p. 221. (3) West (1983c), pp. 341-343. (4) Tuny and MacMahon (1988).
23 Technical Report NPS/NAUCARE/NRTR-93/01
Figure 13. Patterns in presence or absence of two especially sensitive grazing intolerant species (decreasers) in relation to grazing intensity in grassland stands in Capitol Reef National Park. "A" indicates that both winterfat (Ceratoides lanata) and needle-and-thread grass (Sripa comata) are absent from habitats that appear suitable for them. "P" indicates that one or both species are present. Numbers in parentheses at the top indicate the number of sampled stands in each grazing class. Grazing classes are based on subjective estimates of grazing intensity at each sampling site.
Figure 14. Patterns in relative abundance of increaser (grazing tolerant) and decreaser (grazing intolerant) species (see Table 6) in relation to grazing intensity in waterpocket communities in Capitol Reef National Park. "D>I" indicates that decreaser species are more abundant (greater total cover) than increaser species; "D=I" indicates that decreasers and increasers are roughly equal in abundance; and "I>D" indicates that increasers are more abundant than decreasers. Numbers in parentheses at the top indicate the number of sampled stands in each grazing class. Grazing classes are based on subjective estimates of grazing intensity at each sampling site. 24 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Figure 15. Relative abundance (% of species in a stand) of exotic (non-native) species in relation to elevation in grassland stands in Capitol Reef National Park. Horizontal lines represent the mean and range, and blocked areas represent one standard deviation, of the percent of non-native species among all of the vascular plant species recorded in our releve plots of approximately 100 square meters area. though they extend over a broad range of This powerful control by the geologic elevation. Moreover, these soil types are substrate may be a general characteristic of closely correlated with geologic substrate. desert vegetation, where soils are poorly Topographic position, aspect, and slope developed and water is the overwhelmingly generally had far less influence on community limiting resource. By contrast, in regions type and structure in CARE than did substrate where precipitation is generally greater, and elevation. Everitt (1970) and Loope (1977) community types appear to occur over a also reported that geologic substrate exerted wider range of rock and soil types, and their a powerful control on vegetation patterns in distribution is controlled more by elevation the Henry Mountains and in Canyonlands and topographic position than by geologic National Park, respectively. substrate (e.g., Whittaker 1956, Peet 1981).
25 Technical Report NPS/NAUCARE/NRTR-93/01
Figure 16. Overall patterns in distribution of plant community types in Capitol Reef National Park in relation to elevation and soil type. The solid vertical and horizontal lines indicate that a particular community type rarely or never extends across the boundary depicted by the line. The community types in parentheses are restricted to the areas designated by the broken horizontal lines, but the community types without parentheses may extend across the broken horizontal lines. In portion of the figure where more than one type of community is indicated, the local distribution of communities is controlled by other, more subtle factors not depicted here. "H-B-O" designates the hophornbeam-boxelder-oak riparian woodland community.
Discussion Vascular Flora regions. CARE's flora includes representatives The florao f Capitol Reef National Park of the Rocky Mountain, Sierra Madrean, and represents one of the largest documented Great Plains floras, as well as several species floras of any national park on the Colorado endemic to southern Utah or the Colorado Plateau. This park also contains a greater Plateau, and cosmopolitan species distributed number of threatened, endangered, and rare widely over North America and Eurasia (Bar taxa than any other park unit in the continen bour and Billings 1988). These biogeographic tal United States. patterns are discussed further below under Two reasons for the large number of plant threatened and endangered taxa. Habitat species in CARE are its great variety of diversity in CARE results from the wide variety habitats and its strategic location near the of geologic substrates and topographic condi intersection of several major biogeographic tions, as well as the broad range in elevation.
26 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Many plant species in CARE are strongly Waterpocket Fold, south through the North associated with specific geologic formations, Blue Hills region to Notom. and are secondarily influenced by the pres By contrast, Erigeron maguirei var. maguirei ence of alluvium, colluvium, and soils having is known only within CARE and from two distinctive characteristics of texture and very small populations outside the park. We salinity (Welsh et al. 1987, Weber 1987). Some found it scattered across the Waterpocket Fold examples of highly restricted taxa include on Navajo Sandstone slickrock, from Upper Maidenhair fern (Adiantum capillus-veneris), Deep Creek southeast to the Fremont River. found on Navajo sandstone in hanging The only other known occurrences of Erigeron gardens; Harrison milkvetch (Astragalus maguirei var. maguirei are in the San Rafael harrisonii), Beck cymopterus (Cymopterus Swell just north of CARE. In 1986, before we beckii), Maguire daisy (Erigeron maguirei), and began our survey, the only known population the Rabbit Valley gilia (Gilia caespitosa), all on of Erigeron maguirei var. maguirei in the world Navajo sandstone; and Jones cycladenia had less than 30 individuals, and the only (Cycladenia humilis var. jonesii), found only on record from CARE was of Erigeron maguirei Chinle formation. Many of the species having var. harrisonii, an endemic variety apparently these kinds of restricted distributions are restricted to one canyon in the park. Our field endemic to the Colorado Plateau and the state inventories have now revealed that var. of Utah (Welsh and Chatterley 1985). maguirei is more abundant in CARE, and that var. harrisonii apparently represents pheno- Threatened and Endangered Taxa typic response (i.e., plasticity) of individuals Although our study was the most thor growing in shaded, moist canyons. ough inventory of rare plants yet conducted in CARE's population of Echinocereus tri- CARE, the distributional records that we glochidiatus var. inermis shows the greatest report here probably are still incomplete. The geographic disjunction from other known recounted distribution of rare plants in the populations. It is found in the park at a single park is obviously limited by the areas where station, on Paradise Rats. The remaining transects were done. There were many specific populations are in the Paradox Valley of locations in the park that we did not sample western Colorado, along the Gunnison and because of time constraints and the general Colorado Rivers in Colorado and in the La Sal vegetational and physiographic similarity of Mountains of San Juan County, Utah. Simi these areas to those places that we did sample. larly, Cycladenia humilis var. jonesii is found in This included areas in which rare taxa were CARE at a single location in the southern expected to occur. For example, we have no portion of the park, which fits the general doubt that there exist additional undescribed pattern of geographically disjunct populations locations of Erigeron maguirei var. maguirei, throughout the entire range of this taxon. Gilia caespitosa, Cymopterus beckii and Townsen- Two of the Federal Candidate Category 1 dia aprica in CARE. species, Gilia caespitosa and Pediocactus winkleri, were found to be more widespread in the park Patterns of Rarity in CARE than previous records would indicate. Indeed, The rare plants in CARE represent a the CARE populations of Gilia caespitosa variety of patterns of rarity. For example, represent the greatest concentrations of this Sclerocactus wrightiae is regionally restricted to species so far discovered, the only other the Colorado Plateau but occurs at numerous occurrence being near Teasdale, Utah (Wilken locations both in and out of CARE. We found 1979). Gilia caespitosa is scattered on the it at several locations throughout Cathedral Navajo Sandstone formation from Golden Valley, as well as along the Hartnet and in the Throne to Deep Creek (though the largest South Desert. Populations also are known populations are north of the Fremont River). from the Notom area just outside the park, Pediocactus winkleri, previously known in the and we expect that it occurs in scattered, small park from a single location on the Hartnet, populations from the Hartnet, east of the was found scattered from the Hartnet,
27 Technical Report NPS/NAUCARE/NRTR-93/01
through parts of the South Desert, south to the Bitter Creek Divide; Figure 2) probably results Golden Throne area. from the higher elevation, the difference in Two additional Federal Candidate Cat orientation of the Waterpocket Fold, and egory 1 species, Spiranthes diluvialis and greater exposure of many of the formations in Schoencrambe barnebyi, represent significant the north. Several of the locations with excep discoveries in the park. One previously was tionally high numbers of rare taxa, e.g., Upper known only from a single historical record Deep Creek, Miners Mountain, The Hartnet and the other was unknown in CARE. Spiran and above Hickman Bridge (Table 2), are at thes diluvialis is a polyploid orchid species the higher elevations in the park. known from a very few scattered localities This local variation in the abundance of across the Rocky Mountains and Great Basin. rare taxa throughout the park suggests that Although there is a collection of this species there are critical regions within CARE where from the park, in the Fremont River Canyon disturbance could adversely affect a large west of the Visitor Center, the location could number of plants of critical concern. The not be relocated. Indeed, the habitat described most notable is the Upper Deep Creek area on the voucher specimen, Tamarix grove, (Table 2, Figure 1). This high-elevation expo could not be found at the location cited on the sure of Navajo Sandstone and Carmel specimen. On the opposite side of the park, Formation supports one endangered, one two populations (one very small and the other threatened, two federal candidate and four more sizable) were located. Schoencrambe 3C plant species. This concentration of rare barnebyi was previously known only from two plants is likely due in part to the great diversity populations of less than 50 individuals in the of habitats, which range from open sagebrush to southern San Rafael Swell (a single record in aspen stands and even brisfiecone pine associa field notes referred to another location along tions. Moisture regimes vary from dry, sandy Pleasant Creek in CARE, which we were bench-tops to perennial streams with associ unable to relocate). We found this species in ated wetlands and seeps. This, coupled with the park along the Fremont River and Sulphur nearly 450 m (1500 ft) of relief, provides an Creek. The five populations in CARE now incredible diversity of potential habitats that represent the largest populations of this rare species may inhabit. Upper Deep Creek species discovered to date. The Sulphur Creek also supports numerous high-elevation plant East population possessed over 200 individu species that are not endemic or federally als in 1986-1988. listed, but that occur in the park only in this area. Examples include Habenaria sparsiflora, Critical Areas for Rare Plant Conservation in Fritillaria atropurpurea, Pinus longaeva, and CARE Zigadenus elegans. The distribution of rare and endemic taxa Other areas supporting high concentra throughout the park is not uniform. This is not tions of rare plant taxa include the Miners surprising, considering the great number of Mountain/Fremont Canyon area, the Hartnet, geologic formations, and the variation in the Hickman Bridge region, and perhaps a elevation, drainage patterns and moisture large portion of the Waterpocket Fold north of regimes. These various sources of complexity the Fremont River (Table 2, Figure 1). Because result in considerable edaphic variation, and of the large number of endemics and rare these edaphic factors may be responsible for plants in these areas, it is recommended that the nonrandom distribution of rare taxa, i.e., these areas be given special management edaphic endemism (Mason 1946a, 1946b; consideration to prevent unnecessary human Raven 1964; Fiedler 1985). One such feature, disturbances. Unfortunately, Upper Deep geologic substrate, is often used as a proxy for Creek, the Hartnet, and portions of the Water more specific edaphic parameters (see, for pocket Fold still have grazing periodically example, Welsh and Thorn 1978). through the year. In these areas, trampling The greater abundance of rare taxa in the may have adversely affected some of the northern portion of the park (i.e., north of federally protected plants that are present.
28 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Patterns of Endemism Section and the Canyon Lands Section, while Capitol Reef National Park lies within the the southern portion of the park extends deep Great Basin Floristic Province (Gleason and into the the Canyon Lands Section. Moreover, Cronquist 1964). This province represents the according to Welsh et al. (1975), the park's arid region of western North America, com southern boundary is close to the Colorado posed of plateaus and lesser mountain ranges River endemic region. Based on its location, that lie between the two major mountain then, it would be expected that CARE's chains, the Rocky Mountain Range and the endemic and rare plants would have their Cascade-Sierra Nevada Ranges, and it corre floristic origins largely in the Utah Plateaus sponds to the Intermountain Floristic Province and the Canyon Lands Sections. This indeed is of McLaughlin (1992). Holmgren, in Cronquist the case: by far the greatest influence on et al. (1972), partitioned much of this floristic endemic plant species in CARE is from the province into sixteen sections which represent Canyon Lands Section (Table 7). "natural areas that can be recognized by A more localized center of endemism is in vegetative, floristic and/or physiographic the San Raphael Swell, located just north of features" (Cronquist et al. 1972:78). These CARE in the northwest corner of the Canyon "floristic sections" are at a much finer scale Lands Section. Harris (1983) recognized eight than the "floristic areas" described by taxa which he considered endemic to this area. McLaughlin (1986,1989). These sections are Four of these taxa are also found at CARE: further grouped into four major divisions, one Cryptantha johnstonii, Erigeron maguirei, Sclerocac- of which is the Colorado Plateau Division, tus xvrightiae, and Schoencrambe barnebyi. The which is nearly identical to McLaughlin's occurrence of the first of these species in CARE (1992) Northern Colorado Plateau Floristic is not surprising, since the Cathedral Valley District and principally conforms to the portion of the park is rightfully considered as northwestern portion of Fenneman's (1931) part of the San Rafael Swell. However, the last Colorado Plateau physiographic province. It is three spedes were found farther south, well out within this division that Capitol Reef National of the San Rafael Swell. Park is positioned (Figure 1). One of Harris' San Raphael Swell endem Because the boundaries of Holmgren's ics, Schoencrambe barnebyi, is known from two floristic province sections are "in part influ sites in the San Rafael Swell (S/s Butte and enced by local regions of species endemism" about three miles east at Bullberry Spring), (Cronquist et al. 1972:78), a feature not in and from our newly found locations along cluded in the multivariate analyses of floristic Sulphur Creek and the Fremont River in areas based on species lists from published CARE. The geologic substrates in the Sulphur local floras (McLaughlin 1986,1989), they may Creek/Fremont River area are very similar to be helpful in evaluating patterns of endemism those where the populations are found in the within the park. According to Cronquist et al. southern San Rafael Swell. This geologic (1972), the three sections, or regions of ende setting can be described as extensive areas of mism, which make up the Colorado Plateau Moenkopi Formation assodated with steep Division are the Canyon Lands Section, the slopes that expose the Chinle, Wingate, Utah Plateaus Section, and the Uintah Basin Kayenta, and Navajo Formations as well as Section. In a review of endemic plants of Utah canyons that cut through the Kaibab and by Welsh et al. (1975), four regions of ende Coconino/Cutler Formations. Furthermore, mism on the Colorado Plateau are recognized. this type of habitat is not uncommon in the San This classification differs only in the recogni Rafael Swell, but Schoencrambe barnebyi is found tion of the Colorado River and adjacent areas only at the farthest south point of such a geo as a separate region of endemism, in addition logic setting in that region. While the popula to the three recognized by Holmgren. tions at CARE are considerably larger, the occur The geographic location of CARE is rence of habitat, by comparison, is much less. significant in that the northern portions of the These disjunctions between the San Rafael park are at the interface of the Utah Plateaus Swell and Capitol Reef are important for
29 Technical Report NPS/NAUCARE/NRTR-93/01
Table 7. Plant taxa found in CARE that are considered by Cronquist et al. (1972), by Welsh et al. (1975), and by ourselves to be endemic to the Canyon Lands Section. Those marked with an asterisk are on the U. S. Fish and Wildlife Service threatened and endangered plant species list. Taxa marked by (=) were considered by the compilers to be rare enough to warrant protection whether or not they were formally listed as endangered or threatened. A. Endemic taxa compiled by Cronquist et al. (1972): Aquilegia micrantha Eremocrinum albomarginatum Asclepias ruthiae* Erigeron abajoensis* Astragalus coltonii var. coltonii Erigeron maguirei* (vars. maguirei and harrisonii) Astragalus desperatus* (variety conspectus=A. Eriogonum bicolor barnebyi) Eriogonum leptocladon var. leptocladon Astragalus episcopus Gilia caespilosa* Astragalus flavus var. argillosus Gilia (lpomopsis) roseata Astragalus harrisonii* Gilia subnuda Astragalus malacoides* Mimulus eastwoodiae Astragalus moencoppensis Camissonia eastwoodiae Astragalus musiniensis Penstemon cyanocaulis Astragalus pardalinus* Penstemon marcusii Astragalus woodruffii* Psoralea juncea Atriplex corrugata Sclerocactus parviflorus Cycladenia humilis var. jonesii* Sclerocactus wrightiae* Ephedra viridis var. cutleri Vanclevea stylosa
B. Endemics compiled by Welch et al. (1975): Asclepias labriformis Hymenoxys depressa*= Castilleja scabrida* Lomatium junceum* Cryptantha johnstonii*- Nama retorsum Dalea flavescens var. epica*= Phacelia rafaelensis Eriogonum corymbosum var. revealianum*= Xylorhiza confertifolia*= Gaillardia spathulata
C. Additional endemic taxa from the Canyon Lands Section that have been described or reassessed taxonomically since 1975:
Astragalus barneby? Habenaria zothecina* Astragalus chamaeleuce var. laccoliticus* Pediocactus winkleri* Cymopterus becki? Schoencrambe barnebyi*
several reasons. First, given that CARE and centers farther south. Finally, these disjunc the San Rafael Swell are both part of the tions suggest that other San Rafael Swell western extent of the Colorado Plateaus endemics are likely to be found in similar Section, these disjunctions are indicative of an habitats in CARE. overall floristic similarity between the two Another plant that may fit this category is areas. Secondly, some of the rare species the newly described Gilia tenuis. We searched known from very few localities in the Swell for it in CARE, but did not find it; however, (i.e., Erigeron maguirei var. maguirei, Toumsen- we think there is a high likelihood of its occur dia apnea, and Schoencrambe barnebyi) are now rence on white, limy substrates associated known to be more abundant in CARE than in with the Curtis Formation in the Cathedral the San Raphael Swell, suggesting that the Valley, Hartnet, and South Desert of CARE, populations in the San Rafael Swell may repre or on light colored sandy substrates associated sent dispersals from the actual population with the Dakota Formation. These substrates 30 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
are similar to the Cedar Mountain Shale on found on nearby Boulder Mountain at eleva which one population occurs in the San tions of 2240-2485 m (7400-8200 ft) and on Raphael Swell. Thousand Lakes Mountain at 2210-2730 m Another important region of endemism in (7300-9000 ft). The Utah Plateau endemics that close proximity to CARE is the Utah Plateaus occur in CARE are found at similarly high Region (Cronquist et al. 1972). This region elevations on Miners Mountain (ca. 2210 m comprises the fault-block plateaus which cut [7300 ft]) and at Upper Deep Creek (2180- across the middle of the state, including the 2665 m [7200-8800 ft]), as well as in protected, Fish Lake Plateau (Thousand Lakes Moun wet canyons nearby. In fact, these high tain), Awapa Plateau and the Aquarius elevation areas of the park may be better Plateau (Boulder Mountain). A portion of this considered as the easternmost extent of the region lies only a few kilometers west of Utah Plateaus Section, rather than as a part of CARE. Holmgren (in Cronquist et al. 1972) the Canyon Lands Section. considers this region second only to the Two species found in CARE, Eriogonum Canyon Lands Section in the number of tumulosum and Parthenium ligulatum, were endemic taxa on the Colorado Plateau. previously considered endemics of the Uintah Holmgren recognizes 39 endemics to this Basin (Cronquist et al. 1972). It is now clear region. Of these, four are also found at CARE: that these species are not truly endemic to that Astragalus consobrinus, Geranium marginale, region. More recent distributional information Penstemon leiophyllus, and Toumsendia aprica. Inindicate s that Eriogonum tumulosum occurs not addition, the newly described Erigeron awapen- only at CARE, but it is scattered both north of sia, an endemic of the Awapa Plateau, extends the Uinta Mountains in Sweetwater County, down the Fremont Valley into CARE (Welsh Wyoming, and south of the Tavaputs Plateau, 1983a). cutting across the higher elevations of the San Toumsendia aprica was mentioned above as Rafael Swell (part of the Canyon Lands a species that occurred in both CARE and the Section, as previously noted) to the foot of the San Rafael Swell; it also was considered by Aquarius and Fish Lake Plateaus. It also has Holmgren as an endemic to the Utah Plateaus been recognized that, in spite of the consider region because of the few records of this able barriers posed by the Uinta Mountains species available in 1972. Since that time and the Tavaputs Plateau (Welsh et al. 1976, additional records indicate that T. aprica is in Reveal 1979), some taxa endemic to the fact endemic to the eastern Fish Lake Plateau Colorado Plateau cross the Tavaputs Plateau and adjacent San Rafael Swell. A recently to share the Uinta Basin and the Canyon described endemic taxon of the Fish Lake Lands regions of endemism. Among these (Thousand Lakes Mountain) and Aquarius species are Chamaechenactis scaposa and Sclero- (Boulder Mountain) Plateaus is Thelesperma cactus parviflorus, which are both found in the subnuda var. alpinum (Welsh 1983b). This park. species was found in CARE in the upper Deep A more recently described taxon, Physaria Creek area. acutifolia var. purpurea, was previously known The occurrence in CARE of Utah Plateau only from the southern Uinta Basin. We endemics is to be expected given the high discovered a small population at Upper Deep elevation of portions of the park, including Creek, south of Billings Pass, which represents areas over 2120 m (7000 ft), and the general a considerable disjunction. Rather than a similarity of geologic substrates in the two disjunction, however, it is possible that the areas. Although most soils on the plateaus are purple (or rightly, maroon) flowered Physaria derived from either tufaceous sedimentary at Upper Deep Creek is in fact an exceptional rocks and volcanics or quaternary colluvial form of the more common Physaria acutifolia and boulder deposits, there are some regions var. acutifolia. Regardless of which interpreta with outcrops of Navajo, Kayenta, Wingate, tion is correct, this morphologically distinctive and Entrada Sandstones, similar to what is population represents a unique gene pool, found in CARE. These substrates also can be worthy of protection.
31 Technical Report NPS/NAUCARE/NRTR-93/01
In addition to the regions of endemism recognized regions of endemism are discussed discussed above, Welsh et al. (1976) recog below. nized the Colorado River Drainage as a region The first, Cycladenia humilis var. jonesii, is of endemism; however, they did not cite found in isolated populations scattered across specific examples to support this claim. There the Canyon Lands Section: in the northern La are only a few species in CARE that could be Sal Mountains, near Green River, in the San regarded as members of this region, primarily Rafael Swell, in CARE, in Glen Canyon because there is very little area in the park at National Recreation Area southwest of CARE, low enough elevation to be considered as part and at least one historical record in Arizona, in of the river system (in the narrow sense). The the Dixie Corridor Section of the Colorado most likely candidate is Dalea flavescens var. Plateau. The closest relative of the var. jonesii epica, whose type locality is in the low eleva is presumably the polymorphic C. humilis, of tion area along the Colorado River in Glen the Coast Ranges in California. One hypoth Canyon. It is possible as well that Sphaeralcea esis to explain the present distribution of this grossulariifolia var. moorei, an endemic of low taxon is that var. jonesii may be Cycladenia elevation Colorado and San Juan Rivers, in humilis of early origin, possibly even Madro- the Glen Canyon area, could be considered as tertiary (Axelrod 1958), and its present an endemic of this region. Although not a distribution represents the remnants of a federal candidate, the Moore Globemallow formerly broader distribution across the is quite rare in the park, being restricted to a western U.S. (i.e., the present distribution is a few waterpocket areas in the south end of relictual distribution). If this hypothesis is CARE. correct, then speciation would have to have occurred prior to the period when the various Interpreting Anomalous Plant Distributions in populations became isolated from each other. CARE The second species with an anomalous Regions of endemism are often thought distribution, Pteria thompsonii, is known from to represent locations where populations of Utah, Arizona, Nevada and Idaho, on the what had been previously a more widespread Colorado Plateau and in the Great Basin. The species have reproduced in isolation from the present distribution suggests that this species remaining populations of the "species" from either originated in the southern Great Basin which they were descended. The internment and later migrated along the Colorado River in the isolated areas presumably resulted into the Canyon Lands Section, or evolved in eventually in divergence and allopatric specia the Canyon Lands region and later migrated tion. If such a scenario is correct, then the through the Dixie Corridor into the southern present distributions of these taxa at CARE Great Basin (the Idaho disjunction would have may represent either (1) range extensions to be explained by a dispersal event). Both of from the areas where speciation occurred or these patterns of migration have been sug (2) the actual region in which speciation gested for other taxa of the Colorado Plateau occurred. For many of CARE's rare plant (Reveal 1979). Other close relatives of P. species, it is not difficult to interpret the thompsoniae are P. scoparia (southern New current patterns of distribution as represent Mexico, southwest Texas and Chihuahua), P. ing in situ speciation, migration, or a combina pintorum (Sonora), and P. glandulosum (Mexi tion of the two. However, there are several can Highlands). It is evident that the general taxa whose present distributions present pattern of distribution is from the Mexican difficulties in inferring either in situ speciation Highlands northward; however, the present or later migration. These difficult taxa are not distribution is not particularly helpful in necessarily restricted to a single region of the determining if P. thompsoniae diverged in the Colorado Plateau, but instead may even southern Great Basin or on the Colorado extend beyond the Colorado Plateau. Three Plateau. representative taxa whose distributions are The final example of an anomalous distri more or less anomalous in relation to currently bution pattern is Spiranthes diluvialis. This 32 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
species is known from scattered locations from adverse effect on the natural values of this the east slope of the Rocky Mountains near area. The most important step for preserving Golden, Colorado, to near Salt Lake City, this community is to keep it remote and Utah, in the Bonneville Basin Section of the inaccessible, as it is presently. New trails, Great Basin, and from near Panaca in Lincoln roads, or campgrounds should not be built in County, Nevada, in the eastern Tonopah the vicinity, and backpacking in the area Section of the Great Basin, south to Garfield should not be encouraged. County, Utah in the Canyon Lands Section of The second community type of special the Colorado Plateau. Although the isolated concern is the waterpocket community (see and widespread nature of the populations is Romme et al. [1993] for details). This commu confusing, it is nevertheless possible to infer nity is rare in the park and the region, and is the origin of S. diluvialis. Based on cytological vulnerable to damage by livestock grazing evidence (Sheviak 1984) it is believed that S. and recreational use. Livestock effects on diluvialis (2n=74) is the amphiploid derivative waterpockets are discussed below. Recre of a hybridization between S. magnicanporum ational damage mainly involves pollution of (2n=30) and S. romanzoffiana (2n=30). Sheviak the water with human wastes and garbage, (1984) postulated that the hybridization and disturbance of wildlife dependent on occurred during the Pleistocene, when these these rare sources of permanent water. People two parental taxa potentially could have will want to visit these areas, since they are a coexisted. If this scenario is correct, the current unique and fascinating component of CARE, pattern of distribution is likely the result of but camping, swimming, and extended stays habitat fragmentation, the scarcity of low should be discouraged, especially at remote, elevation mesic habitats, and /or multiple, backcountry waterpockets. independent hybridization events. The third community type of special concern is the hanging garden community, Plant Communities of Special Concern which is widely distributed in canyons of the Many of the plant communities in CARE Colorado Plateau, but is everywhere rare and are common and widespread on the Colorado fragmented in its distribution (see Romme et Plateau, and require no special management al. [1993] for details). The major threat to actions to ensure that they persist into the hanging gardens is invasion by exotic weeds, future. However, four of the community types such as cheatgrass (Bromus tectorum), which that we recognized in CARE are rare or could displace the native mesophytic species. vulnerable or both. Managers should be We found this weed in several hanging garden especially careful to protect these communities sites in CARE. There is no sure control for from unnecessary disturbance. spread of these exotic weeds, but monitoring of The first plant community of special the hanging gardens would be useful to remain concern is the bristlecone pine - cushion plant aware of the magnitude of the problem. community, which is restricted to a very small The final community type of special area (ca. 1 ha) on a flat ridgetop formed by the concern is the hornbeam-boxelder-oak wood Carmel Formation in the upper Deep Creek land community, which is restricted to a few area (see Romme et al. [1993] for more details). small, sheltered sites in Halls Creek Narrows It is significant both for its rarity and for the (see Romme et al. [1993] for details). The major extreme age of the trees. Major threats include threats to this community are related to wildfire, which could kill the trees and recreational use and damage to the trees from consume the ancient dead wood lying on the woodcutting and campfire activities. This ground, and recreation, which could involve problem can be minimized by discouraging cutting and burning the ancient wood. Fire people from camping within the Narrows, may be a minor threat, since fuels are sparse though ordinary hiking through the area and discontinous on the ridgetop inhabited by should pose no threat. Livestock grazing also the old bristlecone pine, but uninformed could degrade this community, but the known recreational activities could have a serious and stands are largely inaccessible to livestock.
33 Technical Report NPS/NAUCARE/NRTR-93/01
Vegetation Patterns component of grasses, whose shallow fibrous in Capitol Reef National Park root systems can effectively absorb the mois The vegetation of CARE consists of a ture available in surface soil layers imme complex mosaic of species and communities diately after a rain. Soils of intermediate distributed along two principal environmental texture commonly support a mix of these two gradients. The first major gradient is elevation. contrasting plant forms. As elevation increases, temperatures generally The chemical composition of the soil also decrease and precipitation increases, resulting influences the kinds of plants that can grow in in generally more mesic conditions at higher that area. For example, the high sodium elevations. The park's vegetation strongly content and alkalinity of some soils derived reflects this moisture and temperature gradi from Mancos shale inhibit plant growth ent, changing from semi-desert grasslands and (Potter et al. 1985a, b). Both Everitt (1970) in shrublands at low elevations to woodlands the Henry Mountains and Loope (1977) in and forests in the highest portions of the park. Canyonlands National Park reported vegeta At any given elevation, temperature and tion patterns related to patterns of soil texture potential evapotranspiration (hence soil and chemical composition that were similar to moisture conditions) also are influenced by the patterns we report here for CARE. aspect and topographic position, e.g., north- facing exposures generally are more mesic Random and Deterministic Patterns in than south-facing exposures. Aspect and Community Composition topographic position appeared to have a Although it was possible on the basis of relatively small influence on the composition our field observations to predict the general of upland vegetation in CARE, especially at type of plant community present on a site the lower elevations where soil texture (see given its elevation and geologic substrate, it below) had an overwhelming influence. Loope frequently was not possible to predict exactly (1977) made a similar observation in Canyon- the composition and relative abundance of lands National Park, although Everitt (1970) shrub species making up the community on that stated that slope and aspect were very impor site. This was true especially of sites in the tant in controlling vegetation patterns in the various pinyon-juniper community types. We Henry Mountains. observed enormous variation in relative abun The second major gradient is soil texture, dance of the shrub species on sites that appeared which influences plant distribution and to be very similar in terms of habitat conditions. growth primarily through its effects on Three explanations for this observation moisture availability. Moisture-holding are possible. First, the distribution of shrub capacity generally decreases from deep sandy species within the broad zone of pinyon and soils, through rocky loams, to shallow fine- juniper dominated communities may be textured clays and silts. Soil texture is controlled in a deterministic fashion by other, determined primarily by geologic substrate. possibly subtle, environmental gradients that Sandy soils result from the weathering of we did not recognize in this study. Further coarse-textured sandstones, such as the research into soil chemistry, micrometeorol- Navajo, and occur also in alluvial and eolian ogy, local herbivore densities, etc., would be deposits. Fine-textured soils are commonly required to rule out this hypothesis. Secondly, associated with shales and mudstones, such as many of the stands that we sampled may have the Mancos and Morrison Formations. been disturbed at various times in the past, Coarse-textured soils generally support and are all converging deterministically into communities dominated by woody shrubs or some "climax" community composition. trees, since the deep roots of these species can Additional research on disturbance history reach deep sources of water during the long and succession would be required to address intervals between precipitation events that this hypothesis. characterize this semi-arid environment. Fine- The third hypothesis, and the one that we textured soils generally support a larger favor, is that there is a large random element 34 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
to the distribution of shrub species in the locations where substrates change abruptly, pinyon - juniper community type. There may and here one finds abrupt changes in the be numerous species capable of occupying a vegetation as well. For example, sharply tilted limited number of micro-sites where growing sandstone outcroppings at the lower eleva conditions are suitable for a shrub growth tions generally support a pinyon-juniper form. Whichever species' propagules reach a woodland, whereas adjacent shales and suitable site first will be the species occupying mudstones support a mat saltbush or mixed that site. Once an individual shrub becomes grassland community. Eolian deposits are established on a micro-site, it persists there for characterized by a distinctive plant commu a long time, probably decades or centuries. nity with usually sharp boundaries that Thus, there are limited opportunities for correspond exactly to the boundaries of the replacement in a successional sense, and underlying sand deposit. Numerous addi species composition of a stand reflects both tional examples could be elaborated in which the deterministic abiotic characteristics of the the vegetation changes abruptly with an site and the more-or-less random events of abrupt change in geologic substrate. individual species establishment on that site. There also are many locations in CARE Table 8 summarizes the woody species where environmental conditions change associated with relatively mesic and relatively gradually, and here the vegetation varies xeric sites within the pinyon-juniper wood gradually as well, supporting the continuum land type. The composition of any particular concept (Whittaker 1975). For example, the stand appears to represent a more-or-less east slope of Miners Mountain in the central random sample of the many species capable of part of CARE represents an elevational living in that particular habitat. gradient of several hundred meters over a Investigators in other kinds of communi more or less uniform geologic substrate. The ties have suggested this kind of stochastic entire slope is covered by a pinyon-juniper element of community composition. Malanson community, but pinyon becomes progressively (1980,1982; Malanson and Kay 1980) found more abundant and juniper less abundant as that hanging garden communities in Zion one moves to higher elevations. On the top of National Park showed low floristic similarity the mountain, ponderosa pine begins to appear. despite high habitat similarity. (We observed These continuous changes in plant community the same pattern in CARE's hanging gardens.) composition reflect an underlying continuous He concluded that any particular site could change in temperature, precipitation, and support a wide variety of assemblages of potential evapotranspiration. mesophytic species, and that species-specific dispersal capabilities and the frequency of Distribution of Communities along disturbance by flooding interacted to control Environmental Gradients community composition at any particular time Although we have described many and place within the potential habitat of this different types of plant communities in CARE, community type. A similar conclusion was it must be emphasized that they all overlap to reached by McCune and Allen (1985) who some extent in their characteristic composition found different species compositions in and habitat. We have taken several natural similar-aged forests occupying essentially patterns of continuous variation and broken identical habitats in a series of deep, isolated them into discrete units for purposes of canyons in the Bitterroot Range of Montana. description and classification. For example, canyon bottom habitats in Discrete and Continuous Patterns in CARE represent a gradient in moisture and Community Composition temperature conditions ranging from rela The vegetation in CARE is an interesting tively dry and hot in shallow canyons at low mix of stands showing both discrete and elevations to relatively moist and cool in deep continuous compositional patterns. Given the canyons at high elevations. The three types of complex geology of this area, there are many canyon bottom communities that we have
35 Technical Report NPS/NAUCARE/NRTR-93/01
identified represent more or less arbitrary community. As coarse soil particles begin to divisions of the continuous variation in largely replace the fine particles altogether, the vegetation associated with this gradient in plant community becomes dominated more habitat conditions. The dry canyon bottom strongly by woody plants, and grasses become shrub community occupies the hot, dry end of a minor component of the community. These the gradient; the mesic canyon bottom shrub kinds of sites may support a low shrub or community is found in the middle portion; pinyon - juniper - low shrub community. and the mesic montane woodland community occupies the cool, wet end of the gradient. The Role of Natural Disturbances in the Similar patterns can be seen across the Vegetation of CARE wide range of soil conditions in CARE. The vegetation of any landscape consists Among sites characterized by coarse textured of a mosaic of individual stands, some with soils, those with a shallow, poorly developed more or less sharp boundaries, others with soil overlying a relatively impermeable very gradual boundaries. The mosaic results bedrock generally are associated with the from underlying patterns in environmental blackbrush-rabbitbrush or low shrub commu conditions, e.g., soil type and elevation, as well nity. At the other end of the spectrum are as historical events such as disturbance and deeper, though still coarse textured, soils or successional processes (Romme and Knight shallow soils overlying heavily fractured 1982, Forman and Godron 1986). This present bedrock. These kinds of sites allow deeper study was focused primarily on environmental root growth and support the more mesophytic gradients and their influence on the vegetation species of the pinyon - juniper - tall shrub patterns in CARE. An important direction for community. On intermediate sites one finds future vegetation research would be to identify the pinyon - juniper - low shrub community, the important kinds of natural disturbances that which contains a mix of species from both of have shaped the various plant communities in the other kinds of communities occupying the park, and to clarify the role of disturbance relatively more xeric and more mesic positions and succession in creating and maintaining the on the soil gradient just described. striking diversity of plant species and communi Soils having a texture suitable for growth ties in this area. of grasses nevertheless vary continuously Our observations suggest that fire has across the landscape, and support a range of been an important form of natural disturbance grass - dominated communities. Deep, sandy in the high elevation ecosystems of the upper soils appear to provide the best growing Deep Creek portion of the park. We saw conditions for grasses, and are associated with lightning scars and fire scars on several large eolian grassland, mixed grassland, or pinyon - ponderosa pine and Douglas fir trees in this juniper grassland communities. It is not clear area. One large stump contained five exter exactly what determines which of these three nally visible fire scars. Reconstructions of types of communities will be present on a pre-settlement fire history in ponderosa pine given site; it may involve historical events forests elsewhere on the Colorado Plateau (e.g., grazing intensity) as well as environ have revealed that these kinds of forests mental factors. As one moves gradually into burned at intervals ranging from several years soils containing more silts and clays, species to a few decades prior to the onset of fire composition also changes and one finds a suppression activities in the twentieth century grassland phase of the mat saltbush commu (Cooper 1960,1961; Dieterich 1980, Madany nity or a mixed grassland community having a and West 1983, White 1985). Periodic, low- different mix of species than the mixed intensity fires tended to maintain an open grassland developing on sandy soils. Where forest structure composed of large, widely the fine soil particles are mixed with larger spaced canopy trees and a well developed pebbles and cobbles, the plant community ground layer vegetation of grasses and forbs. becomes a shadscale grassland or the galleta With the greatly reduced frequency of fire phase of the pinyon - juniper - low shrub during the last century, the canopies of these 36 Vascular Flora and Vegetation of Capitol Reef National Park, Utah kinds of forests generally have become more a flood. In these places much of the former dense, and the ground layer vegetation has plant biomass had been removed, and new become sparse because of shading by the individuals were colonizing the exposed canopy. Many of the large ponderosa pine and ground. As we noted in our discussion of Douglas fir trees in upper Deep Creek were plant communities in CARE, the waterpockets cut many years ago, but the spatial pattern of and riparian communities in general are the stumps suggests that the presettlement characterized by extremely variable species forest was more open and contained larger composition from site to site. A part of this trees than the forest present in this area today. variability may be due to frequent disturbance The forest now consists of trees that appear to by floods and subsequent recolonization, such be mostly less than 100 years of age. In places that a stable or climax community never we saw dying bitterbrush plants in the shade develops. Severe windstorms are a type of of young pine; the shrubs apparently were disturbance that is at least locally important. being killed by competition from the thicken We found an area of about half a hectare in ing forest canopy. upper Deep Creek where all of the large trees Accumulation of dead woody material in had snapped and were lying with their boles the absence of fire also has resulted in fuel pointed in the same direction (northeast). conditions in which a fire today is likely to burn more intensely than the typically low- Long Term Stability of Vegetation in CARE intensity fires of the nineteenth century Paleo-ecological research has revealed (Dodge 1972). Thus, fire suppression may that the vegetation of the Colorado Plateau have significantly modified the structure and has changed during at least the last 25,000 dynamics of high-elevation forests in CARE. years in response to fluctuating climatic Fire probably has not been important in conditions (Cole 1982,1990; Betancourt 1990). most of the low elevation communities The greatest changes occurred at the end of because fuels generally are not abundant or the Pleistocene some 12,000 -14,000 years ago. continuous enough to carry a fire. Grassland Two important paleo-ecological studies communities may be an exception. In the have been conducted in CARE. One was by absence of heavy grazing, grassland fuels are Fisher et al. (1991), who examined opal abundant and continuous enough to carry a phytoliths in grassland soils, and determined low-intensity, rapidly moving fire. Such fires that grasslands in CARE today are different in rarely kill the grasses or forbs, though they some respects than they were 200-800 years may exact heavy mortality in woody plants. ago. Grasslands today appear to contain fewer Some investigators have suggested that forbs and/or shrubs and fewer cool-season pinyon and juniper are encroaching on grasses. Stipa spp. appear to have decreased in southwestern grasslands as a consequence of abundance, whereas Oryzopsis hymenoides and fire suppression in this century (e.g., Johnsen Hilaria jamesii appear to have increased. These 1962, Arnold et al. 1964; also see Tausch et al. changes probably are due mainly to climatic 1981). However, we are aware of no recent changes since the end of the Little Ice Age studies addressing this question in the vicinity some 100-150 years ago. Heavy livestock of CARE. grazing also may have been involved (see Other kinds of important natural distur below). bances in CARE may include periodic floods A second important paleo-ecological and landslides. Little research has been done study was Cole's (1991) analysis of packrat in this area to investigate the possible effects middens in CARE. He reconstructed vegeta and significance of these disturbances, but it tion over the last 5400 years in the Harnet area seems likely that they influence the structure of the park (Figure 1), and concluded that the and dynamics of riparian communities and greatest changes in vegetation composition communities on steep, unstable slopes. In the throughout that time were related to over course of our fieldwork, we observed water- grazing by livestock during the last 100 years. pockets that had been recently scoured out by Plant species that are relatively intolerant of
37 Technical Report NPS/NAUCARE/NRTR-93/01
grazing, such as Ceratoides lanata and Oryzopsis Further research into the interactions hymenoides, were more abundant in the older between environmental gradients and various packrat deposits; whereas species that increase kinds of disturbances that shape CARE's in the presence of heavy sustained grazing, vegetation would aid park management and such as Chrysothamnus zriscidiflorus and Gutier- interpretation not only by clarifying the causes rezia sarothrae, were more abundant in recent of present vegetation patterns but also by deposits. Cole (1991) also found some packrat helping us to anticipate the potentially devas middens in the Hall's Narrows area (Figure 1) tating effects of global climate change over the dating from the Pleistocene. These contained next century. plant species that today are restricted to higher elevations, such as Pseudotsuga men- Effects of Grazing on the Vegetation of CARE ziesii and Juniperus scopulorum. These two The role of natural disturbances — e.g., paleo-ecological studies demonstrate that fire, floods, landslides, and climatic variation substantial changes have occurred in the — in shaping the vegetation of CARE has been vegetation of this area over at least the last discussed above. In addition to these natural 25,000 years, in response both to broad-scale disturbances, livestock grazing during the last climatic change and to more localized effects century has profoundly influenced the compo of heavy livestock grazing. sition and dynamics of some types of plant As a result of anthropogenic increases in communities in CARE. Our research indicates carbon dioxide and other greenhouse gases, that the most significant grazing effects have another major climatic change is expected to occurred in accessible grassland and riparian occur over the next century. This change may communities, though nearly all communities be comparable in magnitude to that at the end of have been affected to some extent (Figure 9). Pleistocene, but the rate of change will be far Sparks et al. (1990) assessed vegetational greater than previous events (Bolin et al. 1986). changes during the last century in two town Current projections based on global circulation ships in northern Utah, and reported the models (e.g., Hansen et al. 1988, Schlesinger and greatest livestock-related changes in mid- Zhao 1988, Wetherald and Manabe 1988) elevation bench, foothill, and bajada sites — suggest an average rise in global temperature of similar to the patterns we found in CARE. 15-4.5 degrees C (Dickinson 1986). Reduced Livestock grazing in CARE's grasslands precipitation and soil moisture at middle evidently has had little effect on total cover or latitudes also are predicted, although regional number of species, except in very heavily and differences in temperature and precipitation continuously grazed stands where both of changes are likely, and currently beyond the these parameters are reduced (Figure 11). resolving powers of the climate models. However, our results suggest that even light The impending global climate change has grazing has produced substantial changes in far-reaching ramifications for natural areas species composition of grassland communi (Peters and Darling 1985, Tangley 1988, Davis ties. The relative abundance of more palatable 1989, Graham et al. 1990, Romme and Turner species (e.g., bunchgrasses) decreases along a 1991). Potential ranges for species may change gradient of increasing grazing intensity, more rapidly than populations can migrate to whereas the abundance of unpalatable species new areas or adapt to new local conditions. (e.g., Gutierrezk sarothrae) shows the opposite Disturbance regimes also are likely to be pattern (Figure 12). Some of the most pre altered, e.g., fires may become more frequent ferred plant species, e.g. Ceratoides lanata and (Sandenburgh et al. 1987). Managers need to Stipa comata, may have been locally extirpated understand the present complex of biotic and by grazing (Figure 13). abiotic factors, including disturbance, that In addition to changes in species composi control the distribution of species and plant tion, heavy and prolonged livestock grazing communities if they are to anticipate and deal appears to have altered the overall structure of with the substantial changes likely to occur in some grasslands in CARE. For purposes of the coming century (Neilson et al. 1989). comparison, we visited the top of the South 38 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Caineville Butte, approximately 10 km east of cannot reach (Figure 14). Ungrazed stands CARE, which supports a relatively xeric were characterized by relatively high cover of grassland community comparable to xeric palatable bunchgrasses (Oryzopsis hymenoides, grasslands in CARE. However, the grassland Poa fendleriana, Sporobolus cryptandrus, and on South Caineville Butte has not been grazed Stipa comata). These same species were uncom for many years (since the 1950s according to mon or absent in the nearby grazed stands. one local landowner; since the 1930s according Because all of the waterpockets that we to another). It probably never was heavily sampled in this area were within 100 m of grazed since there are no reliable sources of each other and shared very similar habitats, water. One of the most striking features of the the differences in vegetation that we detected grassland on South Caineville Butte is the were probably due primarily to differences in large quantity of biomass and litter. There are livestock use. Interestingly, both grazed and enormous clumps of Oryzopsis hymenoides, for ungrazed stands contained Bromus tectorum example, with heights of 1 m and diameters of and other species associated with disturbed 0.5 m containing a mass of dead leaves and sites, apparently because of frequent distur culms accumulated over many years or bance by flooding even in the absence of decades of growth. By contrast, comparable grazing. grassland sites in CARE support only small Barth and McCullough (1988) documented clumps of Oryzopsis hymenoides with minimal similar differences between heavily and light accumulation of litter as a result of repeated ly grazed streamside communities in CARE. grazing. Their data also suggested that prolonged These kinds of differences in structure heavy livestock use of riparian areas may probably are associated with important prevent reproduction of Fremont cotton wood. differences in energy flow and nutrient An examination of historic photographs cycling in grazed and ungrazed grasslands. further suggested that many of the riparian Net primary productivity may be higher in areas in CARE were severely damaged by lightly grazed stands because removal of dead the uncontrolled grazing that occurred prior leaves and culms improves light penetration to the Taylor Act and the beginnings of to photosynthetically active tissues. Net grazing regulation in the late 1930s (Barth primary productivity probably is reduced in and McCullogh 1988). heavily grazed stands, however. Wildlife Both grassland and riparian areas are habitat also appears to be quite different in vulnerable to damage by excessive livestock heavily grazed and ungrazed stands. Dave use, but these two kinds of communities Willey (personal communication, 1989) probably differ in two important ways in their reported a greater abundance and diversity of sensitivity and response to heavy grazing. breeding songbirds on South Caineville Butte First, grazing effects are likely to be more than in heavily grazed but otherwise compa widespread in accessible riparian areas, rable sites in CARE. He attributed the because the presence of water and succulent differences in part to the nesting habitat vegetation makes these areas especially provided by the large thick clumps of bunch- attractive to animals. We noted, for example, grasses in the ungrazed area. that the waterpockets we sampled fell into Regarding the effects of livestock grazing only two grazing categories: heavily grazed in riparian areas, our data suggest similar or ungrazed. The latter category occurred shifts in species composition from dominance only where steep or rocky terrain made it by relatively palatable species to dominance physically impossible for animals to reach the by less palatable species. For example, in the area. series of waterpockets comprising the Willow By contrast, we found a variety of grass and Cottonwood Tanks in the southern part of land stands ranging from heavily grazed to CARE, those waterpockets that are accessible lightly grazed, even in areas where the terrain to livestock supported a measureably different did not prevent animal access. In grasslands, flora than the waterpockets that livestock livestock tend to concentrate around sources
39 Technical Report NPS/NAUCARE/NRTR-93/01
of water; areas remote from reliable water sites may have come into equilibrium with the may be lightly used even though livestock abiotic environment and thus represent new, numbers are high. Thus, the intensity of stable forms of vegetation. livestock grazing appears to be more variable Cheatgrass establishment in many por in grassland than in riparian communities. tions of the northern Great Basin also has led The second important difference between to a profound change in the fire regime. grasslands and riparian areas relates to the Whereas fires formerly occurred infrequently rate and probability of recovery from exces and burned in a patchy fashion because of the sive grazing once the grazing pressure is discontinuity of the fuels in native sagebrush alleviated. Riparian areas appear to recover steppe, the continuous and flammable fuels relatively quicky, probably because of the created by the cheatgrass permit very exten favorable growing conditions that promote re- sive fires to occur now as frequently as every establishment of native plants that had been few years in places (Whisenant 1990). This has reduced or even extirpated by grazers. A led to almost complete destruction of the comparison of photographs from the 1930s native perennial shrubs and grasses and their and 1940s with the same sites today (Barth replacement with cheatgrass and other and McCullogh 1988) suggests that over annuals. Such an extreme vegetational conver grazed streamsides in CARE have largely sion does not appear to have occurred in recovered from the over-grazing of the early CARE, though it could potentially occur in part of this century. some areas where cheatgrass is presently very By contrast, reestablishment of native abundant. plants in the semi-arid environments that It appears, therefore, that even with characterize CARE's grasslands probably is far elimination of livestock grazing, some of slower and even problematical. Suitable CARE's grassland areas may never return to climatic conditions for establishment of new their pre-grazing structure and composition bunchgrasses, for example, may occur only without some form of active intervention. every several decades. Moreover, seed sources Some arid, over-grazed sites may now be in a may have been eliminated in many local areas, new equilibrium with the abiotic environ and the well-established exotic species that ment, and further degradative changes may dominate some sites (e.g., cheatgrass) may potentially occur, judging from experiences competitively exclude seedlings of the native elsewhere. Additional research in this area is grasses. The depauperate plant communities needed if the goal is to restore the vegetation on some of these arid, heavily over-grazed of CARE to its presettlement condition.
Acknowledgements
This research was supported by the and Kay Throne assisted with identification of University of Wyoming - National Park plant specimens. Mary Laspe and Jennifer Service Research Center (Contract #PX 1200-8- Walker helped with multivariate analysis. 8039). We thank Norman Henderson, director John Spence provided helpful comments on of resource management in Capitol Reef fitting our classification to the Brown-Lowe- National Park, for encouragement and logisti Pase system. Ross Worley, David Wilson, Tim cal assistance throughout the study. Field Mietty, and Connie Cole produced the figures. assistance was provided by Wayne Mietty, Kimball Harper, Stan Welsh, and Kay Throne Bob Melton, and Suzette Durall. Stan Welsh reviewed earlier drafts of the manuscript.
40 Vascular Flora and Vegetation of Capitol Reef National Park, Utah Literature Cited
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Mason, H. L. 1946a. The edaphic factor in Peet, R. K. 1981. Forest vegetation of the narrow endemism. I. The nature of Colorado Front Range: composition and environmental influences. Madrono dynamics. Vegetatio 37:65-78. 8:209-240. Peters, R. L., and J. D. S. Darling. 1985. The Mason, H. L. 1946b. The edaphic factor in greenhouse effect and nature reserves. narrow endemism. II. The geographic BioScience 35:707-735. occurrence of plants of highly restricted patterns of distribution. Madrono 8:241- Phillips, B. G., A. M. Phillips, HI, and M. A. S. 257. Bernzott. 1989. Annotated checklist of vascular plants of Grand Canyon National McCune, B., and T. F. H. Allen. 1985. Will Park. Grand Canyon Natural History similar forests develop on similar sites? Association Monograph No. 7, Flagstaff, Canadian Journal of Botany 63:367-376. Arizona.
McLaughlin, S. P. 1992. Are floristic areas Pieper, R. D., and G. A. Lymbery. 1987. hierarchically arranged? Journal of Influence of topographic features on Biogeography. Volume 19 (in press). pinyon-juniper vegetation in south-central New Mexico. Pages 53-57 in Everett, R. L. McLaughlin, S. P. 1989. Natural floristic areas (compiler), Proceedings — Pinyon-juniper of the western United States. Journal of conference. USDA Forest Service General Biogeography 16:239-248. Technical Report INT-215.
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Potter, L. D., R. C. Reynolds, Jr., and E. T. Sheviak,CJ. 1984. Spiranthes diluvialis Louderbough. 1985b. Mancos shale and (Orchidaceae), a new species from the plant community relationships: analysis western United States. Brittonia 36:8-14. of shale, soil, and vegetation transects. Journal of Arid Environments 9:147-165. Singh, T., and N. E. West. 1971. Comparison of some multivariate analyses of perennial Raven, P. H. 1964. Catastrophic selection and Atriplex vegetation in southeastern Utah. edaphic endemism. Evolution 18:336-338. Vegetatio 23:289-313.
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White, A. S. 1985. Presettlement regeneration Wilken, D. H. 1979. The taxonomic status of patterns in a southwestern USA ponde- Gilia caespitosa (Polemoniaceae). Madrono rosa pine (Pinus ponderosa) stand. Ecology 26:91-95. 66:589-594. Woodin, H. E., and A. A. Lindsey. 1954. White, P. S. and S. P. Bratton. 1980. After Juniper-pinyon east of the Continental preservation: philosophical and practical Divide as analyzed by the line-strip problems of change. Biological Conserva method. Ecology 35:473-489. tion 18:241-255. Woolley,R. R. 1947. Utilization of surface- Whittaker, R. H. 1956. Vegetation of the Great water resources of Sevier Lake Basin, Smoky Mountains. Ecological Mono Utah. United States Geological Survey. graphs 26:1-80. Water Supply Paper 920.
Whittaker, R. H. 1975. Communities and Youngblood, A. P., and R. L. Mauk. 1985. Ecosystems. Second edition. Macmillan, Coniferous forest habitat types of central New York. and southern Utah. USDA Forest Service General Technical Report INT-187.
46 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
APPENDIX I ANNOTATED CHECKLIST OF VASCULAR PLANTS CAPITOL REEF NATIONAL PARK, UTAH
Distributional categories used in the checklist are: (W)-widespread throughout Capitol Reef National Park; (Q-common throughout much of the park or to a particular part of the park; (O)occasional, widely scattered in the park; (U)-uncommon, of limited distribution; (R) rare, of very limited distribution and only known from one or two locations in the park. The elevational range is given for each taxon in feet above sea level.
FERN ALLIES Notholaena limitanea Maxon R; 5000; canyon bottoms; pinyon-juniper SELAGINELLACEAE woodland, rock crevices; Muley Twist Canyon. Selaginelk mutica D. C. Eaton U; 5500 to 7500; on rocks in shady sites; slickrock Woodsia pregana D.C. Eaton communities; near Hickman Bridge, Water- U; 8500; mountains; mixed conifer woodland; pocket Fold, Miners Mountain, Cohab and Upper Deep Creek. Frying Pan Canyons.
EQUISETACEAE GYMNOSPERMS
Equisetum arvense L. CUPRESSACEAE O; 4000-4500; bottomland; riparian; oxbow of the Fremont River and Halls Creek. ]uniperus communis L. U; 8500; mountains; mixed conifer woodland; Equisetum hyemaleL. Upper Deep Creek. O; 5500-6500; bottomland; riparian; oxbow of the Fremont River, Pleasant Creek, and Spring Juniperus osteosperma (Ton.) Little Canyon. W; 3900-8000; desert shrub to mixed conifer woodland; widespread throughout the park. Equisetum laevigatum A. Br. R; 4000; bottomland; riparian; Halls Creek. Juniperus scopulorum Sarg. U; 7500-8000; mountains; mixed conifer wood land; Upper Deep Creek. FERNS EPHEDRACEAE POLYPODIACEAE Ephedra nevadensis Wats. Adiantum capillus-veneris L. R; 4000; canyon bottoms; desert shrub; Halls U; 3900-4000; moist slickrock sandstone walls; Narrows. hanging garden community; lower Halls Creek. Ephedra torreyana Wats. W; 3900-7000; upland, bottomland, hills, mesas, Cheihmthes fed Moore benches, washes, and canyons; desert shrub to O; 5000-7000; bottomlands, canyons, and pinyon-juniper woodland; widespread through benches; sagebrush and pinyon-juniper wood out the parlc land; Fremont River Valley, Water Canyon, Sulphur Creek and Miners Mountain. Ephedra viridis Cov. W; 3900-7500; upland, bottomland, hills, mesas, Cystopteris fragilis (L.) Bemh. in Schrader benches, washes, and canyons; desert shrub to R; 8500; mountains; mixed conifer woodland; pinyon-juniper woodland; widespread through Upper Deep Creek. out the park.
47 Technical Report NPS/NAUCARE/NRTR-93/01
PINACEAE ANACARDIACEAE
Picea pungens Engelm. Rhus aromatica Ait. var. simpliafolia (Greene) U; 8000; mountains; riparian; Upper Deep Cronq. Creek. O; 5000-6000; canyons, canyon bottoms, and mesas; pinyon-juniper woodland; Wagon Box Pinus edulis Engelm. in Wisliz. Mesa and Hickman Bridge. W; 5000-7500; upland, mesas, hills and benches; pinyon-juniper woodland; widespread through Rhus aromatica Ait. var. trilobata (Nutt.) Gray out the park. W; 4000-7000; upland, bottomland, mesas, canyons, benches, and canyon bottoms; Pinus longaeva D.K. Bailey riparian, and desert shrub to pinyon-juniper U; 8500; mountain benches; bristlecone pine woodland; widespread throughout the park. community; Upper Deep Creek. Toxicodendron rydbergii (Small) Greene Pinus ponderosa Dougl. ex P. & C. Lawson O; 4000-5500; bottomland; riparian; oxbow of the C; 6500-8800; upland, mountains, and canyons; Fremont River and Halls Creek. mixed conifer woodland; Upper Deep Creek, Longleaf Flats, Pleasant Creek, Grand Wash, APOCYNACEAE and Spring Canyon. Amsonia jonesii Woodson Pseudotsuga menziesii (Mirb.) Franco O; 4000-4500; canyon bottoms; desert shrub and O; 5000-8800; canyons and mountains; moist blackbrush; Halls Creek sites and mixed conifer woodland; Pleasant Creek, Muley Twist Canyon, Spring Canyon, Apocynum cannabinum L. and Upper Deep Creek O; 4500-5500; bottomlands and wet places; riparian; Fruita, Halls Creek and The Post.
FLOWERING PLANTS Cycladenia humttis Benth. var. jonesii (Eastw.) DICOTYLEDONS Welsh & Atwood R; 6000; hillsides; desert shrub-juniper commu ACERACEAE nity; Deer Point.
Acerglabrum Torr. ASCLEPIADACEAE O; 7000-8500; mountains and upland canyons; riparian; Spring Canyon and Upper Deep Creek. Asclepias asperula (Decne.) Woodson R; 6000; canyons; pinyon-juniper woodland; Acer negundo L. Frying Pan Canyon. O; 4000-6500; upland canyons, seeps, springs and waterpockets; riparian; Fremont Valley, Asclepias cryptoceras Wats. Capitol Wash, Pleasant Creek, Halls Creek O; 5500-6500; mesas, benches and hillsides; Upper Muley Twist, Upper Deep Creek Cohab- desert shrub to pinyon-juniper woodland; Frying Pan, Sulphur Creek, Spring Canyon and Lower Deep Creek, Sulphur Creek, Burr Trail, Grand Wash. and The Hartnet.
AMARANTHACEAE Asclepias labriformis Jones O; 5000-6000; washes; desert shrub to pinyon- Anmranthus graecizans L. juniper woodland; Capitol Gorge, Middle Desert O; 5500; bottomland; disturbed sites; Fremont Wash, Spring Canyon and Pleasant Creek. River Valley. Asclepias latifolia (Torr.) Raf. Amaranthus powellii Wats. R; 4000-5500; hillsides and canyon bottoms; O; 5500; bottomland; orchards along the desert shrub and pinyon-juniper woodland; Fremont River. Fruita and Halls Narrows.
48 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Asclepias maarosperma Eastw. Cryptantha capitata (Eastw.) Johnst. R; 5000; hillsides; desert shrub; The Post. U; 6000-7000; flats, meadows, and rocky sites; sagebrush communities and pinyon-juniper Asclepias ruthiae Maguire & Woodson woodland; Paradise Flats and Hickman Bridge. U; 5500-6500; bottomland, hills, and benches; Cryptantha cinerea (Torr.) Cronq. desert shrub and pinyon-juniper woodland; W; 5500-8000; bottomland, mesas, benches, and Middle Desert Wash, Cathedral Valley and washes; desert shrub to mixed conifer wood Longleaf Flat. land; Fremont River Valley, Cathedral Valley, South Desert, Upper and Middle Deep Creek, Asclepias speciosa Torr. and Pleasant Creek. O; 5500-6000; moist sites; riparian; Fruita. Cryptantha circumscissa (H. & A.) Johnst. Asclepias subvertkellata (Gray) Vail O; 3900-5500; canyon bottoms; desert shrub; R; 4000; canyon bottoms; desert shrub; Halls Halls Creek and Spring Canyon. Narrows. Cryptantha confertiflora (Greene) Payson BERBERIDACEAE C; 4000-6000; hillsides, benches, and canyon bottoms; desert shrub to pinyon-juniper Berberis fremontii Torr. woodland; Halls Narrows, Burr Trail, and South C; 4000-5500; canyons, benches, and mesas; Desert. pinyon-juniper woodland; Halls Creek and Circle Cliffs. Cryptantha crassisepala (T. & G.) Greene O; 5000-6500; bottomland, mesas, and benches; Berberis repens Lindl. desert shrub to pinyon-juniper woodland; The O; 6000-8000; mountains and canyon bottoms; Post and The Hartnet. mixed conifer woodland; Cohab Canyon and Upper Deep Creek. Cryptantha fendleri (Gray) Greene U; 5000-7000; hillsides and canyons; desert Berberis vulgaris L. shrub to pinyon-juniper woodland; Middle Deep Creek, Big Thomson Mesa, and Pleasant U; 5500; bottomland; orchards; Fruita. Creek.
BETULACEAE Cryptantha flava (A. Nels.) Payson W; 4000-6500; bottomland, upland, hills, and Alnus tenuifolia Nutt. mesas; desert shrub to pinyon-juniper wood R; 7500-8500; canyon bottoms; mixed conifer land; widespread throughout the park. woodland; Upper Deep Creek. Betula occidenlalis Hook. Cryptantha flavoculata (A. Nels.) Payson O; 5500-8000; uplands and canyon bottoms; C; 5000-6500; upland, hillsides and benches; riparian; Pleasant Creek, Water Canyon, South desert shrub to pinyon-juniper woodland; Burr Desert Overlook, and Upper Deep Creek Trail, Sulphur Creek, Water Canyon, and South Desert Overlook. Ostrya knowltonii Cov. R; 4000; canyon bottoms; riparian; Halls Cryptantha fulvocanescens (Gray) Payson Narrows. W; 5000-6000; bottomland, upland, hills, mesas, and benches; desert shrub to pinyon-juniper BORAGINACEAE woodland; Fremont River Valley, Chimney Rock, Sulphur Creek, and Hickman Bridge. Cryptantha abata Johnst. C; 5000-8000; mesas, benches, mountains, and Cryptantha gracilis Osterh. canyon bottoms; desert shrub to mixed conifer C; 4000-6500; upland, hillsides, and benches; woodland; Sulphur Spring, Burr Trail, Upper desert shrub to pinyon-juniper woodland; Red Deep Creek and Miners Mountain. Slide, Pleasant Creek, Hickman Bridge, and Lower Muley.
49 Technical Report NPS/NAUCARE/NRTR-93/01
Cryptantha humilis (Gray) Payson Tiauilia latior (Johnst.) A. Richards. O; 8000-8800; mountains; mixed conifer wood R; hills; desert shrub; Cathedral Valley. land; Upper Deep Creek.
Cryptantha johnstonii Higgins CACTACEAE R; 7500-8500; mountains and washes; mixed conifer woodland (moist sites); jMiddle and Coryphantha vivipara (Nutt.) Britt. & Rose Upper Deep Creek. R; 5000-6000; hillsides and benches; pinyon- juniper woodland; Burr Trail. Cryptantha mensana (Jones) Payson O; 5000-6000; upland, hillsides, mesas, and EcMnocereus triglochidiatus Engelm. var. inermis benches; desert shrub to pinyon-juniper (K. Schum.) Rowl woodland; Burr Trail, Fruita, and Deer Point. R; 7000; hillsides; pinyon-juniper woodland; Paradise Draw. Cryptantha micrantha (Torr.) Johnst. R; 4000-4500; hillsides and benches; desert EcMnocereus triglochidiatus Engelm. var. melana- shrub; Hall Divide. canthus (Engelm.) L. Benson W; 4500-7000; upland, mesas, hills, benches, and Cryptantha pterocarya (Torr.) Greene canyons; desert shrub to pinyon-juniper U; 4000-6000; bottomland and canyon bottoms; woodland; widespread throughout the park. desert shrub to pinyon-juniper woodland; Bun- Trail and Halls Creek Opuntia basuaris Engelm. & Bigel. var. heilii Welsh & Neese Cryptantha pustulosa (Rydb.) Payson R; 5500; benches; desert shrub; Visitor Center. R; 5000-5500; hillsides; desert shrub; The Post. Opuntia fragUis (Nutt.) Haw. Cryptantha recurvata Cov. C; 6500-8500; upland, mesas, and mountains; C; 5000-6000; hills and benches; desert shrub to pinyon-juniper woodland to mixed conifer pinyon-juniper woodland; Burr Trail, Cedar woodland; Upper Deep Creek and Pleasant Mesa, Hickman Bridge, and Spring Canyon. Creek
Cryptantha rollinsii Johnst. Opuntia phaeacantha Engelm. O; 5000-6000; mesas and benches; desert shrub C; 3900-6500; upland, hills, mesas, benches, and to pinyon-juniper woodland; The Hartnet. canyons; desert shrub to pinyon-juniper woodland; Fremont River Valley, Pleasant Cryptantha setosissima (Gray) Payson Creek, Hickman Bridge, Halls Creek and Spring U; 8000; mountains; mixed conifer woodland; Canyon. Upper Deep Creek. Opuntia polyacantha Haw. Cryptantha tenuis (Eastw.) Payson W; 3900-7500; upland, hills, mesas, benches, and R; 5000-6000; hills; desert shrub to pinyon- canyons; desert shrub to pinyon-juniper juniper woodland; Burr Trail. woodland; widespread throughout the park.
Lappula ocridentalis (Wats.) Greene Pediocactus simpsonii (Engelm.) L. Benson C; 4000-5500; bottomland; disturbed sites; Fruita, O; 8000; mountains; mixed conifer woodland; Oyster Shell Reef, Spring Canyon, and Halls Upper Deep Creek Creek Pediocactus winkkri Heil Lithospermum incisum Lehm. R; 5000-5500; hills and benches; desert shrub to C; 4000-8500; canyon bottoms, mountains, and pinyon-juniper woodland; Jailhouse Rock and benches; desert shrub to mixed conifer wood Ackland Spring. land; Upper Deep Creek, Halls Creek, and Hickman Bridge. Sclerocactus parviflorus Clover & Jotter O; 4000-7000; upland, hills, mesas, benches, and Lithospermum multiflorum T. & G. canyons; desert shrub to pinyon-juniper U; 5500-8500; upland and mountains; pinyon- woodland; Fremont River Valley, Cedar Mesa, juniper woodland to mixed conifer woodland; Hickman Bridge, Cohab/Frying Pan Canyons, Upper Deep Creek and Sheets Gulch. Muley Twist Canyon, and Burr Trail. 50 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Sclerocactus wrightiae L. Benson CARYOPHYLLACEAE U; 5000-6000; hills and benches; desert shrub to pinyon-juniper woodland; Ackland Spring, Arenaria fendleri Gray var. eastwoodiae (Rydb.) Cathedral Valley, and South Desert. Welsh U; 6000; mesa; pinyon-juniper woodland; The Hartnet. CAMPANULACEAE Arenaria fendleri Gray var. glabrescens Wats. Campanula parryi Cray O; 7200-8200; hillsides and benches; mixed R; bottomland; saline soils (riparian); Fremont conifer woodland and bristlecone pine commu River Valley and Oak Creek. nity; Miners Mountain and Upper Deep Creek.
Lobelia cardinalis L. Cerastium nutans Raf. R; 3900; canyon bottoms; riparian; HallsNar- R; 8500; mountains; mixed conifer woodland; rows. Upper Deep Creek
Lychnis drumtnondii (Hook.) Wats. CANNABINACEAE R; 8800; mountains; meadows; Upper Deep- Creek Humulus americanus Nutt. U; 5000-5500; bottomland; riparian; Fremont Paronychia sessiliflora Nutt. River Valley. O; 7000-8000; mountains and benches; pinyon- juniper woodland and mixed conifer woodland; CAPPARIDACEAE Upper Deep Creek and Miners Mountain.
Cleome lutea Hook. Saponaria officinalis L. O; 4000-6000; bottomland and canyon bottoms; O; 5500; bottomland; riparian; Fremont River desert shrub to mixed conifer; widespread Valley. throughout the park Silene antirrhina L. Cleome serrulata Pursh U; 4000-5000; canyon bottoms; disturbed sites; R; 5000-6000; bottomland; riparian and road Willow Tanks and Halls Creek sides; Fremont River Valley and Cathedral Valley. Stellaria longipes Goldie R; 8500; mountains; mixed conifer woodland; Cleomella palmerana Jones Upper Deep Creek R; 4000; benches; desert shrub; Halls Creek CELESTRACEAE CAPRIFOLIACEAE Pachystima myrsinites (Pursh) Raf. Lonicera utahensis Wats. U; 8000-8800; mountains; mixed conifer wood O; 8500; mountains; mixed conifer woodland; land; Upper Deep Creek. Upper Deep Creek Sambucus caerulea Raf. O; 8500; mountains; mixed conifer woodland; CHENOPODIACEAE Upper Deep Creek Allenrolfea occidentalis (Wats.) Kuntze Symphoricarpos longiflorus Gray R; 5700; saline bottomland; riparian; Lower O; 4500-7000; upland, mesas, benches, and Deep Creek canyons; desert shrub to pinyon-juniper woodland; occasional throughout the park Atriplex argentea Nutt. R; 5500; benches; desert shrub; Chimney Rock. Symphoricarpos oreophilus Gray O; 5000-8000; upland, bottomland, and moun Atriplex canescens (Pursh) Nutt. tains; pinyon-juniper woodland to mixed conifer W; 4000-6500; bottomlands, hills, and mesas; woodland; Upper Deep Creek South Desert desert shrub to pinyon-juniper woodland; Overlook, and the Fremont River Valley. widespread throughout the park 51 Technical Report NPS/NAUCARE/NRTR-93/01
Atriplex confertifolia (Ton. &. Frem.) Wats. Chenopodium fremontii Wats. W; 4000-6500; hillsides, mesas, bottomland, and W; 4000-6500; bottomland; desert shrub, benches; desert shrub to pinyon-juniper sagebrush, and pinyon-juniper woodland; Halls woodland; widespread throughout the park. Creek, Fremont River Valley, and Muley Twist Canyon. Atriplex corrugate Wats. U; 5000-5500; bottomland and hillsides; desert Chenopodium glaucum L. shrub; The Post. O; 5500; canyon bottoms; riparian; Fremont River Valley. Atriplex gardneri (Moq.) D. Dietr. var. omenta (A. Nels.) Welsh Chenopodium leptophyllum Nutt. in Wats. W; 4000-6500; bottomland, hills, and benches; O; 5000-7000; bottomland, benches, and upland; desert shrub to pinyon-juniper riparian and pinyon-juniper woodland; Sulphur woodland; widespread throughout the park. Creek, Fremont River Valley, and The Hartnet.
Atriplex graciliflora Jones Corispermum villosum Rydb. R; 4000; canyon bottoms; riparian; Halls U; 5000-5500; bottomland; disturbed sites; Narrows. Fremont River Valley.
Atriplex patulaL. Grayia spinosa (Hook.) Moq. R; 5500; bottomland; saline soils (riparian) R; 3900; hills, benches, and canyons; desert- Fremont River Valley. shrub; Halls Creek
Atriplex powellii Wats. Halogeton glomeratus C.A. Mey. in Ledeb. U; 5000-5500; bottomland and hillsides; desert O; 5000-6000; bottomland; desert shrub; Fremont shrub; The Post. River Valley and Middle Desert Wash.
Atriplex rosea L. Kochia americana Wats. R; 5500; bottomland; saline soils (riparian); O; 5000-5500; canyon bottoms; desert shrub to oxbow of the Fremont River. pinyon-juniper woodland; Muley Twist Canyon and Chimney Rock. Atriplex saccaria Wats. O; 5000-6000; bottomland, washes, and canyon Kochia scoparia (L.) Schrader bottoms; desert shrub; Burr Trail, Grand Wash, C; 5000-6500; bottomland; disturbed sites; Ackland Spring. Fremont River Valley, Spring Canyon, and Pleasant Creek. Bassia hyssopifolia (Pallas) Kuntze R; bottomland; disturbed sites; Bitter Spring Monolepis nuttalliana (Schult.) Greene Creek area. U; 5000-6500; bottomland; disturbed sites; Muley Twist Canyon, Burr Trail, and Longleaf Ceratoides lanata (Pursh) J.T. Howell Flat. W; 5000-6000; upland, mesas, and benches; desert shrub, sagebrush, and pinyon-juniper Salsola iberica Sennen & Pau woodland; Burr Trail, Grand Wash, and W; 4000-7000; bottomland, washes, and canyon Cathedral Valley. bottoms; desert shrub to pinyon-juniper woodland (disturbed sites); widespread Chenopodium album L. throughout the park R; 8500; mountains; disturbed sites in riparian- Upper Deep Creek Sarcobatus vermiculatus (Hook.) Torr. in Emory C; 5000-6000; bottomland; desert shrub; Fremont Chenopodium botrys L. River Valley, Cedar Mesa, Cathedral Valley, and R; 6000; waste places; riparian; Oak Creek. Spring Canyon.
52 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Suaeda calceoliformis (Hook.) Moq. Artemisia campestris L. U; 5500; bottomland; riparian; oxbow of the O; 5500-8000; canyon bottoms, benches, dis Fremont River. turbed sites, mountains; desert shrub, pinyon-juniper woodland, and mixed conifer; Suaeda torreyana Wats. Cathedral Valley, Upper Deep Creek, and C; 4000-6500; bottomland; riparian; Pleasant Middle Desert Wash. Creek, Sulphur Creek, Brimhall Arch, Halls Creek, and Burr Trail. Artemisia dracuncutus L. W; 5500-8000; canyon bottoms and washes; Zuckia brandegei (Gray) Welsh & Stutz var. pinyon-juniper to mixed conifer; widespread arizonica (Standley) Welsh throughout the park. R; 5000; hillsides; desert shrub; Jailhouse Rock Artemisia filifolia Torr. Zuckia brandegei (Gray) Welsh & Stutz var. O; 3900-6000; benches, washes and canyonbot- brandegei toms (sandy sites); desert shrub to 0^000-6500; upland, mesas, benches, and pinyon-juniper woodland; widespread through canyons; desert shrub to pinyon-juniper out the park. woodland; The Hartnet, Cohab Canyon, Fremont River Valley, and Chimney Rock. Artemisia frigida Willd. R; 8500; canyons; mixed conifer and sagebrush CELASTRACEAE flats; Upper Deep Creek. Forsellsia meionandra (Koehne) Heller O; 5000-8000; bottomland, mountains; upland, Artemisia ludoviciana Nutt. hills, mesas, and benches; desert shrub to mixed W; 4000-8500; benches, canyon bottoms, washes, conifer woodland; Fremont River Valley, Middle mesas, mountains; desert shrub to mixed and Lower Deep Creek, and Jailhouse Rock conifer; widespread throughout the park.
COMPOSITATE Artemisia nova A. Nels. O; 8000-8800; mountains; sagebrush communi Achillea millefolium L. ties; Upper Deep Creek. R; 8800; mountains; Aspen community; Upper Deep Creek. Artemisia pygmaea Gray R; 6000-6500; mesas and uplands; sagebrush Ambrosia acanthicarpa Hook. communities and pinyon-juniper woodland; top W; 5000-6000; bottomland; disturbed sites; of the Burr Trail and South Desert Overlook. Fruita, Cathedral Valley, Chimney Rock. Artemisia spinescens D.C Eaton in Wats. Antennaria parvifolia Nutt. R; 4000; benches; desert shrub; Lower Halls O; 8000-8800; mountains; mixed conifer forest; Creek. Upper Deep Creek. Artemisia tridentata Nutt. var. tridentata Antennaria rosea Rydb. W; 5000-6000; benches; sagebrush communities; R; 8500; mountains; mixed conifer forest; Upper widespread throughout the park. Deep Creek. Artemisia tridentata Nutt. var. vaseyana (Rydb.) B. Antennaria rosukta Rydb. Boi. R; 8500; mountains; mixed conifer forest; Upper O; 7500-8500; benches and mountains; sage Deep Creek. brush communities; Upper Deep Creek.
Arctium minus (Hill) Bernh. Aster brachyactis Blake in Tidestr. O; 5500; bottomland; waste places; Fremont R; 5500; bottomland; riparian communities; River Trail. Fremont River Valley
Artemisia bigebvii Gray Aster chUensis Nees W; 4500-6500; benches and mesas; pinyon- W; 5000-6000; bottomland and canyon bottoms; juniper woodland and sagebrush flats; riparian; Fremont River Valley, Oak Creek, and widespread throughout the park. Pleasant Creek.
53 Technical Report NPS/NAUCARE/NRTR-93/01
Aster foliaceus Lindl. in DC. Brickellia microphylla (Nutt.) Gray var. scabra O; 5000-6000; bottomland and canyon bottoms; Gray moist sites; Cathedral Valley, Fremont River O; 5000-6500; mesas and benches; slickrock Valley and Pleasant Creek. communities; widespread throughout the park.
Aster frondosus (Nutt.) T. & G. Brickellia oblongifolia Nutt. R; 6000; canyon bottoms; riparian; Oak Creek. O; 5000-6500; hillsides and benches; desert shrub; Burr Trail, Muley Twist Canyon and Deer Aster glaucodes Blake Point. R; 7000-8500; mountains; mixed conifer wood land; Upper and Middle Deep Creek. Chaenactis douglasii (Hook.) H. & A. O; 7500-8800; mountains; mixed conifer wood Aster occidentalis (Nutt.) T. & G. land; Upper Deep Creek. O; 5500; bottomland; orchards; Fruita. Chaenactis steviodes H. & A. Aster pauciflorus Nutt. O; 4000-6000; bottomland; desert shrub; Halls C; 5500-6000; bottomland and canyon bottoms; Creek and Burr Trail. salt marsh and riparian; Fremont River Valley, Pleasant Creek, and Sulphur Creek Chamaechaenactis scaposa (Eastw.) Rydb. R; 6000; benches and hillsides; pinyon-juniper Baccharis emoryi Gray in Ton. woodland; South Desert Overlook. O; 4000; washes and canyon bottoms; riparian; Halls Narrows. Chrysothamnus depressus Nutt. O; 5500-6500; hillsides and benches; pinyon- Baccharis salicina T. & G. juniper woodland; Burr Trail and Muley Twist R; 5000; washes and waterpockets; riparian; Canyon. Cottonwood Tanks. Chrysothamnus linifolius Greene Bahia dissecta (Gray) Britt. O; 5000-6000; washes; riparian; Sulphur Creek U; 6000; hillsides and canyons; pinyon-juniper and Cathedral Valley. woodland; Pleasant Creek Chrysothamnus nauseosus (Pallas) Britt. var. Bidens cernua L. abbreviata (Jones) Welsh O; 5500; bottomland; saline soils (riparian); O; 6000-7000; mesas, benches, and canyons; oxbow of the Fremont River. desert shrub to pinyon-juniper woodland; Frying Pan Canyon, Hickman Bridge, and Brickellia atracty bides Gray Wagon Box Mesa. R; 4500-5000; steep hill sides; desert shrub; slopes leading to Halls Creek. Chrysothamnus nauseosus (Pallas) Britt. var. albicaulis (Nutt.) Rydb. Brickellia califomica (T. & G.) Gray O; 6000-8000; uplands and mountains; pinyon- C; 5500-7500; canyon bottoms; riparian and juniper woodland and mixed conifer woodland; desert shrub; Fruita, Paradise Draw, Spring Pleasant Creek and Upper Deep Creek Canyon, and middle Deep Creek. Chrysothamnus nauseosus (Pallas) Britt. var. Brickellia grandiflora (Hook.) Nutt. biglovii (Gray) Hall O; 5500; canyon bottoms; pinyon-juniper U; 6500; ledges; slickrock communities; Pleasant woodland and mixed conifer woodland; Creek. Fremont River Valley. Chrysothamnus nauseosus (Pallas) Britt. var. Brickellia longifolia Wats. consimilis (Greene) Hall O; 4000-5000; washes; hanging gardens; Lower O; 5500; bottomland; desert shrub; Cathedral Muley Twist Canyon and Halls Creek. Valley.
54 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Chrysothamnus nauseosus (Pallas) Britt. var. Cirsium calcareum (Jones) Woot. & Standi, var. graveolens (Nutt.) Piper bipinnatum (Eastw.) Welsh W; 3900-7000; bottomland, benches and mesas; R; 6000; benches; pinyon-juniper woodland; Rim desert shurb to pinyon-juniper woodland; Overlook Trail. widespread throughout the park. Cirsium calcareum (Jones) Woot. & Standi, var. Chrysothamnus nauseosus (Pallas) Britt. var. calcareum junceus (Greene) Hall W; 6000-8000; washes and canyon bottoms; O; 4000-6000; bottomland, mesas, and benches; riparian; wide spread throughout the park. desert shrub to pinyon-juniper; Cathedral Valley, Wagon Box Mesa, and Hall Divide. Cirsium calcareum 0ones) Woot. & Standi, var. pulcheUum (Greene) Welsh Chrysothamnus nauseosus (Pallas) Britt. ssp. O; 5000-6000; benches and canyon bottoms; latisquameus (Gray) Hall & Clem. riparian; Burr Trail and Pleasant Creek. O; 5000-5500; bottomland; disturbed sites; Fruita. Cirsium neomexicanum Gray var. neomexicanum O; 5000-6000; bottomland and canyon bottoms; Chrysothamnus nauseosus (Pallas) Britt. var. desert shrub to pinyon-juniper woodland; Burr leiospermus (Gray) Hall Trail and Sheets Wash. O; 5500-6500; mesas and benches; pinyon- juniper woodland; Meeks Mesa and Hickman Cirsium undulatum (Nutt.) Spreng Bridge. O; 5500; bottomland and benches; riparian; Fremont River Valley. Chrysothamnus nauseosus (Pallas) Britt. var. nitidus L.C. Anderson Cirsium vulgare (Savi) Ten R; 6000-6500; mesas; pinyon-juniper woodland; O; 5500; bottomland; riparian (waste places); Wagon Box Mesa. Fruita.
Chrysothamnus parryi (Gray) Greene var. Conyza canadensis (L.) Cronq. attenuatus (Jones) Kittell in Tidestr. & Kittell O; 4000-5500; bottomland and canyon bottoms; O; 7500-8500; mountains; mixed conifer wood disturbed sites; Halls Creek and Fremont River land; Upper Deep Creek. Valley.
Chrysothamnus viscidiflorus (Hook.) Nutt. var. Crepis runcinata T. & G. lanceolatus (Nutt.) Greene O; 4000-6000; bottomland and canyon bottoms; O; 7500-8500; mountains; mixed conifer wood riparian; Halls Creek, Fremont River Valley, and land; Upper Deep Creek. Pleasant Creek.
Chrysothamnus viscidiflorus (Hook.) Nutt. var. Dicoria brandegei Gray puberulus (D.C. Eaton) Jeps. R; 4000; canyon bottoms; desert shrub; Halls O; 6000-8500; canyon bottoms and mountains; Creek. pinyon-juniper woodland and mixed conifer woodland; Oak Creek and Upper Deep Creek. Dyssoidia papposa (Vent.) A.S. Hitchc. U; 5000-5500; bottomland; disturbed sites; Chrysothamnus viscidiflorus (Hook.) Nutt. var. Fremont River Valley. stenophyllus (Gray) Hall W; 5000-6500; bottomland, mesas, and benches; Dyssodia pentachaeta (DC.) Robins desert shrub to pinyon-juniper woodland; O; 3900; canyons and hillsides; desert shrub; Frying Pan Canyon, Fremont River Valley, Halls Creek. Wagon Box Mesa, and Brimhall Arch. Eceliopsis nudicaulis (Gray) A. Nels. Chrysothamnus viscidiflorus (Hook.) Nutt. var. R; 5000-5500; hillsides; desert shrub; Fremont viscidiflorus River Valley and Chimney Rock. O; 5500; bottomland; riparian; Sulphur Creek and Fremont River Valley. Erigeron abajoensis Cronq. R; 7000-8000; canyon bottoms and benches; pinyon-juniper and slickrock communities; Upper Deep Creek and Miners Mountain. 55 Technical Report NPS/NAUCARE/NRTR-93/01
Erigeron aphanadis (Gray) Greene Erigeron speaosus (Lindl.) DC. R; 5000-5500; hillsides and canyons; desert R; 8500; mountains; mixed conifer woodland; shrub; Sulphur Creek and Chimney Rock. Upper Deep Creek.
Erigeron awapensis Welsh Erigeron utahensis Gray var. utahensis O; 5000-8000; upland, hills, mountains, and W; 5000-8000; upland, bottomland, hills, mesas, canyon bottoms; pinyon-juniper woodland to benches, and mountains; desert shrub to mixed mixed conifer woodland; Fremont River Valley conifer woodland; Fruita, Fremont River Valley, and Upper Deep Creek. Burr Trail, Capitol Gorge, Hickman Bridge, Muley Canyon, and Upper Deep Creek. Erigeron bellidiastrum Nutt. R; 3900; canyon bottoms; desert shrub; Halls Erigeron utahensis Gray var. sparsifolius (Eastw.) Creek. Cronq. R; 4000; canyon bottoms; slickrock communities; Erigeron compadus Blake Halls Narrows. O; 5000-7000; upland, hills, and benches; desert shrub to pinyon-juniper woodland; South Desert GaUlardia pinnatifida Torr. and South Desert Overlook. O; 4000-5000; benches; desert shrub; The Post, Burr Trail, and Halls Creek. Erigeron divergens T. & G. W; 4000-6000; canyons, upland, hills, and mesas; GaUlaridia spathulata Gray riparian, desert shrub, and pinyon-juniper W; 5000-6500; bottomland, mesas, benches, and woodland; Halls Creek, Hickman Bridge, The washes; desert shrub to pinyon-juniper wood Post, and Burr Trail. land; widespread throughout the park.
Erigeron eatonii Gray Glyptopleura setulosa Gray R; 7000-8000; mountains, mesas, and benches; R; 4000; bottomland; desert shrub; Halls Creek. pinyon-juniper woodland to mixed conifer woodland; Upper Deep Creek and Miners Grindelia stfuarrosa (Pursh) Dun. in DC. Mountain. O; 5000-6500; bottomland; disturbed sites; Fremont River Valley and Pleasant Creek. Erigeron flagellars Gray O; 5500-8000; mountains and bottomland; Gutierrezia microcephala (DC.) Gray riparian and mixed conifer woodland; Fruita O; 5000-6000; upland, hills, mesas, benches, and and Upper Deep Creek. washes; desert shrub to pinyon-juniper wood land; Wagon Box Mesa, Fremont River Valley, Erigeron lonchophyllus Hook. and Grand Wash. O; 5500-6500; bottomland; riparian; Pleasant Creek and oxbow of the Fremont River. Gutierrezia sarothrae (Pursh) Britt. & Rusby W; 3900-7500; upland, bottomland, hills, mesas, Erigeron maguirei Cronq. benches, washes, and canyons; desert shrub to U; 5500-8000; canyon walls; slickrock communi pinyon-juniper woodland; widespread through ties; Fremont River Valley, Pine Canyon, Upper out the park. Deep Creek, Longleaf Flat, and Meeks Mesa. Haplopappus acaulis (Nutt.) Gray Erigeron pulcherrimus Heller R; 5000-5500; canyon bottoms; slickrock O; 5000-8000; mountains, upland, hills, and communities; Sheets Gulch benches; pinyon-juniper woodland to mixed conifer woodland; Upper Deep Creek, Grand Haplopappus armerioides (Nutt.) Gray Wash, and Chimney Rock. O; 5000-6000; upland, hills, mesas, benches, and canyons; desert shrub to pinyon-juniper Erigeron pumilus Nutt. woodland; Spring Canyon, Capitol Gorge, W; 5000-8000; upland, hills, mesas, mountains, Muley Canyon, and Cohab Canyon. and benches; desert shrub to mixed conifer woodland; Upper Deep Creek, Fruita, Cotton Haplopappus drummondii (T. & G.) Blake wood Tanks, Deer Point, Sheets Gulch, and R; 4000; desert shrub; along uranium road above Miner Mountain. the Halls Narrows and Cottonwood Tanks.
56 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Helianthella microcephala (Gray) Gray Hymenoxys depressa (T. & G.) Welsh & Reveal R; 5000-7500; Desert shrub to pinyon-juniper O; 5000-7000; upland, hillsides, and mesas; woodland; Wagon Box Mesa and top of the Bun- pinyon-juniper woodland; South Desert, Trail. Fremont River Valley, near Jailhouse Rock, and Miners Mountain. Helianthella uniflora (Nutt.) T. & G. O; 5500; bottomland; riparian; Fruita. Hymenoxys richardsonii (Hook.) Cockerell W; 5000-8800; upland, hills, mesas, mountains, Helianthus annuus L. benches, and canyons; pinyon-juniper woodland O; 5500-6500; bottomland; disturbed sites; to mixed conifer woodland; widespread Fremont River Valley and Pleasant Creek. throughout the park.
Helianthus anomalus Blake Iva axillaris Pursh R; 5000; upland; desert shrub; The Post. O; 5000-5500; bottomland; riparian; Fremont River Valley. Helianthus petiolaris Nutt. O; 4000-5500; bottomland; disturbed sites; Iva xanthifolia Nutt. Fremont River Valley and Halls R; 5500; bottomland; riparian; Fremont River Creek. Valley.
Heliomeris multiflora Nutt. Lactuca canadensis L. R; 8000-6800; mountains; mixed conifer wood R; 4000; canyon bottoms; riparian; Halls Creek. land; Upper Deep Creek. Lactuca serriola L. Heterotheca villosa (Pursh) Shinners C; 5000-6500; bottomland; disturbed sites; Fruita W; 4000-8000; upland, bottomland, hills, mesas, and Pleasant Creek. mountains, benches, washes, and canyons; desert shrub to mixed conifer woodland; Leucelene ericoides (Tore.) Greene widespread throughout the park. C; 5000-6000; upland, hills, benches, and mesas; desert shrub to pinyon-juniper woodland; Hymenopappus filifolius Hook. Capitol Gorge, Hickman Bridge, Cohab/Frying W; 4000-8000; upland, bottomland, hills, mesas, Pan Canyons, and Muley Twist Canyon. mountains, benches, washes, and canyons; desert shrub to mixed conifer woodland; Lygodesmia grandiflora (Nutt.) T. & G. var. widespread throughout the park. arizonica (Tomb) Welsh O; 5000-6000; upland, mesas, and benches; Hymenoxys acaulis (Pursh) Parker var. arizonica desert shrub; Middle Desert Wash and (Greene) Parker Cathedral Valley. W; 4000-6500; upland, hills, mesas, benches, washes, and canyons; desert shrub to pinyon- Lygodesmia grandiflora (Nutt.) T. & G. var. juniper woodland; Longleaf Flat, Chimney Rock, grandiflora The Hartnet, Halls Creek, and Cathedral Valley. O; 5000-6500; benches and canyons; sagebrush communities and pinyon-juniper woodland; Hymenoxys acaulis (Pursh) Parker var. caespitosa Fruita, Sheets Gulch, Cedar Mesa, and Pleasant (A. Nels.) Parker Creek. R; 8500; mountains; bristlecone pine community; Upper Deep Creek. Machaeranthera canescens (Pursh) Gray var. aristata (Eastw.) Turner Hymenoxys acaulis (Pursh) Parker var. ivesiana O; 5500-6000; bottomland and canyons; desert (Greene) Parker shurb to pinyon-juniper woodland; Fruita and O; 4000-6500; canyons, upland, hills, mesas, and Frying Pan Canyon. washes; desert shrub to pinyon-juniper wood land; Halls Creek, Pleasant Creek, Capitol Machaeranthera canescens (Pursh) Gray var. Gorge, and Grand Wash. canescens C; 5000-8000; canyons, canyon bottoms, and mountains; desert shrub to mixed conifer woodland; Meeks Mesa, Upper Deep Creek, Sulphur Creek, and Fremont River Valley. 57 Technical Report NPS/NAUCARE/NRTR-93/01
Machaeranthera canescens (Pursh) Gray var. Senecio multUobatus T. & G. ex Gray latifolia (A. Nels.) Welsh C; 4000-7000; upland, bottomland, hills, mesas, U; 8500; mountains; mixed conifer woodland; and washes; desert shrub to pinyon-juniper Upper Deep Creek. woodland; Danish Hill, Middle Deep Creek, Sulphur Creek, Fremont River Valley, Pleasant Machaeranthera grindelioides (Nutt.) Shinners Creek, Halls Creek, and Muley Twist Canyon. O; 5000-6500; benches and canyon bottoms; desert shrub to pinyon-juniper woodland; Burr Senecio spartoides T. & G. var. spartoides Trail, Muley Twist Canyon, and Pleasant Creek. O; 5500-8000; benches and canyon bottoms; desert shrub to mixed conifer woodland; Upper Machaeranthera tanacetifolia (H.B.K.) Nees Deep Creek, Oak Creek, Jailhouse Rock, and C; 4000-6000; benches, canyons, and canyon Longleaf Hat. bottoms; desert shrub to pinyon-juniper woodland; Fremont River Valley, Sulphur Senecio spartoides T. & G. var. multicapitatus Creek, Halls Creek, Cottonwood Tanks, and (Greenman) Welsh Capitol Gorge. U; 6500; upland; pinyon-juniper woodland; Pleasant Creek Malacothrix glabrata Gray R; 3900; canyon bottoms; desert shrub; Halls Senecio streptanthifolius Greene Creek. R; 6000; washes and waterpockets; sagebrush communities; near Burr Trail. Malacothrix sonchoides (Nutt.) T. & G. O; 5000-5500; canyon bottoms and benches; Solidago canadensis L. desert shrub; Middle Desert Wash and The Post. C; 5000-6500; bottomland and canyon bottoms; riparian; Fremont River Valley, Pleasant Creek, Oxytenia acerosa Nutt. and Spring Canyon. O; 4000-5500; bottomland; riparian; Fremont River Valley, Halls Creek, and Spring Canyon. Solidago missouriensis Nutt. R; 4000; canyon bottoms; riparian; Halls Parthenium ligulatum (Jones) Barneby Narrows. R; 6000; mesas and benches; slickrock communi ties; South Desert Overlook. Solidago occidentalis (Nutt.) T. & G. R; 4000; canyon bottoms; riparian; Halls Petradoria pumila (Nutt.) Greene Narrows. U; 5000; mesas and hills; desert shrub; Muley Twist Canyon. Solidago sparsiflora Gray O; 4000-5000; canyon bottoms; riparian, hanging Platyschkuhria integrifolia (Gray) Rydb. var. gardens; Halls Narrows and Fountain Tanks. desertorum (Jones) Ellison O; 4000-6500; upland, canyons, hills, mesas, and Solidago spathulata DC. benches; desert shrub to pinyon-juniper O; 8500; mountains; mixed conifer woodland; woodland; Brimhall Arch, Deer Point, The Upper Deep Creek Hartnet, Halls Creek, and Chimney Rock. Sonchus asper (L.) Hill. Psilostrophe sparsiflora (Gray) A. Nels. O; 4000-5500; canyon bottoms and bottomland; W; 5000-6500; upland, canyons, hills, mesas, and disturbed sites; Halls Creek and Fremont River benches; desert shrub to pinyon-juniper Valley. woodland; widespread throughout the park. Stephanomeria exigua Nutt. Senecio douglasii DC. var. longUobus (Benth.) L. C; 4000-6500; canyon bottoms and hillsides; Benson desert shrub to pinyon-juniper woodland; U; 5000-6500; benches, washes, and canyon Pleasant Creek, Halls Creek, Fremont River bottoms; desert shrub to pinyon-juniper Valley and Muley Twist Canyon. woodland; Pleasant Creek, Cathedral Valley, and Muley Twist Canyon. Stephanomeria runcinata Nutt. R; 5500; hills; desert shrub; Chimney Rock.
58 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Stephanomeria spinosa (Nutt.) Tomb Townsendia incana Nutt. U; 6000-8000; mesas and mountains; pinyon- C; 4000-6500; upland, hills, mesas, and benches; juniper woodland to mixed conifer woodland; desert shrub to pinyon-juniper woodland; Upper Deep Creek and Wagon Box Mesa. Fremont River Valley, Capitol Gorge, Pleasant Creek, Hickman Bridge, Muley Twist Canyon, Stephanomeria tenuifolia (Torr.) Hall Sulphur Creek, Grand Wash, The Hartnet, and C; 4000-6500; upland, bottomland, hills, mesas, Halls Creek. and benches; desert shrub to pinyon-juniper woodland; Cohab/Frying Pan Canyons, Tragopogon dubius Scop. Longleaf Rat, Fremont River Valley, Cedar O; 5000-6500; bottomland; disturbed sites; Mesa, Pleasant Creek, Muley Twist Canyon, and Fremont River Valley and Pleasant Creek Halls Creek. Vanclevea stylosa (Eastw.) Greene Tetradymia canescens DC. U; 3900-5000; hills and benches; desert shrub; O; 5500-8000; upland, hills, benches, and The Post and Halls Creek mountains; pinyon-juniper woodland to mixed conifer woodland; Upper Deep Creek, Fremont Vernonia marginata (Torr.) Raf. River Valley, and Longleaf Flat. R; 5500; bottomland; disturbed sites; Fremont River Valley. Tetradymia glabrata Gray O; 5500-6500; upland, bottomland, mesas, and Viguiera multiflora (Nutt.) Blake canyons; sagebrush communities and U; 8000-8500; mountains; mixed conifer and pinyon-juniper woodland; Fremont River aspen woodlands; Upper Deep Creek. Valley, Longleaf Flat, and Cohab/Frying Pan Canyons. Wyethia scabra Hook. C, 4000-5500; hills, benches, canyons, and mesas; Tetradymia nuttallii T. & G. desert shrub to pinyon-juniper woodland; U; 8000-8500; mountains; mixed conifer wood Fremont River Valley, Capitol Gorge, Chimney land; Upper Deep Creek. Rock, Muley Twist Canyon, Grand Wash, and Halls Creek. Tetradymia spinosa H. & A. R; 6000; hills and washes; pinyon-juniper Xanthium strumarium L. woodland; The Hartnet. O; 5500-6500; bottomland; disturbed sites; Fruita and Pleasant Creek. Thelesperma subnudum Gray var. subnudum W; 4000-6500; upland, hills, mesas, benches, and Xylorhiza confertifolia (Cronq.) TJ. Watson canyons; desert shrub to pinyon-juniper R; 5000-6000; hills and benches; desert shrub; woodland; widespread throughout the park. Muley Twist Canyon and Sheets Gulch.
Thelesperma subnudum Gray var. alpinum Welsh Xylorhiza tortifolia (T. & G.) Greene R; 8500; benches; bristlecone pine communities; R; 5500-6000; hills, benches, and canyons; desert Upper Deep Creek shrub to pinyon-juniper woodland; Deer Point and Burr Trail. Townsendia annua Beaman U; 4000-5500; hills and benches; desert shrub; Xylorhiza venusta (Jones) Heller Brimhall Arch, Cathedral Valley, and The Post. R; 5000; hills and benches; desert shrub; The Post. Townsendia aprica Welsh & Reveal U; 6000-8500; mesas and benches; pinyon- CONVOLVULACEAE juniper woodland and bristlecone pine communities; South Desert Overlook, Miners Convolvulus arvensis L. Mountain, and Upper Deep Creek O; 5000-6000; bottomland; disturbed sites; Fruita and Pleasant Creek. Townsendia exscapa (Richards.) T.C. Porter O; 5000-6000; hills, upland, mesas, and benches; Evolvulus nuttallianus Schult. in R. & S. pinyon-juniper woodland; O; 5000-6000; bottomland, upland, hills, and Danish Hill and Fremont River Valley. mesas; slickrock communities; Fremont River Valley and Hickman Bridge. 59 Technical Report NPS/NAUCARE/NRTR-93/01
CORNACEAE Capsella bursa-pastoris (L.) Medicus O; 5000-6000; bottomland; riparian; Fremont Corttus stolonifera Michx. River Valley and Pleasant Creek. O; 5500-8000; bottomland; riparian; Fremont River Valley, Pleasant Creek and Upper Deep Cardaria draba (L.) Desv. R; 5500; bottomland; riparian; Fremont River Creek. Valley.
CRASSULACEAE Caulanthus crassicaulis (Torr.) Wats. R; 6500; benches; desert shrub; Deer Point. Sedum lanceolatum Torr. U; 8500; mountains; mixed conifer woodland; Chorispora tenella (Pallas) DC. O; 5500-6000; bottomland; disturbed sites; Upper Deep Creek. Fremont River Valley and Pleasant Creek.
CRUCIFERAE Descurainia californica (Gray) Schultz R; 8500; mountains; mixed conifer woodland; Alyssum minus Upper Deep Creek. R; 5500; bottomlands; disturbed sites; Fruita. Arabis demissa Greene Descurainia pinnata (Walt.) Britt. var. filipes O; 8000-8500; mountains; sagebrush and mixed (Gray) Peck conifer woodland; Upper Deep Creek. W; 5500; bottomland, washes and canyon Arabis drummondii Gray bottoms; riparian; Fremont River Valley. R; 8500; mountains; sagebrush; Upper Deep Creek. Descurainia pinnata (Walt.) Britt. var. osmiarum (Cockerell) Shinners Arabis hirsuta (L.) Bernh. O; 4000-4500; washes and canyon bottoms; O; 8000-8500; mountains; pinyon-juniper, disturbed sites; Halls Creek. sagebrush and mixed conifer; Upper Deep Creek. Descurainia richardsonii (Sweet) Schulz var. sonnei (Robins.) C.L. Arabis holboellii Hornem. Hitchc. R; 8500; mountains; shaded coniferous forest; O; 8500; mountains; mixed conifer woodland Upper Deep Creek. and sagebrush communities; Upper Deep Creek.
Arabis microphylUi Nutt. ex T. & G. Descurainia sophia (L.) Webb in Engler & Prantl O; 6000-8500; canyon bottoms and mountains; W; 5500; bottomland; riparian; Fremont River pinyon-juniper, sagebrush, and mixed conifer; Valley. Pleasant Creek and Upper Deep Creek. Dithyrea wislizenii Engelm. in Wisliz. Arabis perennans Wats. O; 4000-4500; canyon bottoms; desert shrub; O; 5000-8000; hillsides, mesas, mountains and Halls Creek. benches; pinyon-juniper to mixed conifer; Brimhall Arch, The Post, and Upper Deep Draba cuneifolia Nutt. ex T. & G. Creek. O; 5000-5500; hills, mesas, benches, and canyons; desert shrub to pinyon-juniper woodland; Arabis pulchra Jones Sulphur Creek and Big Thomson Mesa. R; 5000; uplands and benches; desert shrub; Big Thomson Mesa. Draba stenobba Ledeb. R; 8500; mountains; mixed conifer woodland; Arabis selbyi Rydb. Upper Deep Creek W; 4000-8500; hillsides, mesas, mountains and benches; desert shrub, pinyon-juniper to mixed Erysimum asperum (Nutt.) DC. conifer woodland; widespread throughout the R; 6500; benches; riparian; Pleasant Creek. park.
60 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Erysimum inamspicuum (Wats.) Macmillan Malcomia africana (L.) R. Br. in Ait. O; 5500-8500; mountains and hills; sagebrush U; 5000; roadside; disturbed sites; The Post. communities and mixed conifer woodland; Upper Deep Creek and Cathedral Valley. Nasturtium officinale R. Br. in Ait. R; 5500; moist sites; riparian; oxbow of the Hutchinsia procumbens (L.) Desv. Fremont River. R; 4000; canyon bottoms; riparian; Halls Narrows. Physaria acutifolia Rydb. var. acutifolia C; 4000-6500; upland, mesas, benches, and Lepidium densiflorum Schrad. var. densiflorum canyons; desert shrub to pinyon-junipr wood O; 4000-6000; upland, hiUs, mesas, benches, land; Muley Twist Canyon, Burr Trail, Halls washes, and canyons; desert shrub to pinyort- Creek, Cedar Mesa, Chimney Rock, Pleasant juniper woodland; Burr Trail, Spring Canyon, Creek, and Sulphur Creek. and Halls Creek. Physaria acutifolia Rydb. var. purpurea Welsh and Lepidium densiflorum Schrad. var. ramosum (A. Reveal Nels.) Thell. R; 8500; mountain benches; bristlecone pine O; 6000-8000; mesas and mountains; pinyon- community; Upper Deep Creek. juniper woodland to mixed conifer woodland; The Hartnet and Upper Deep Creek Physaria neivberryi Gray in Ives U; 5000; mesas; desert shrub; Big Thomson Lepidium montanum Nutt. in T. & G. var. jonesii Mesa. (Rydb.) C.L. Hitchc. C; 4000-6000; upland, hills, mesas, benches, Rorippa islandica (Oed.) Borbas washes, and canyons; desert shrub to pinyon- O; 6500-8000; bottomland; riparian; Pleasant juniper woodland; Halls Creek, The Hartnet, Creek and Upper Deep Creek. Fremont River Valley, Capitol Gorge, Cedar Mesa, Chimney Rock, Muley Twist Canyon, Rorippa tenerrima Greene Hickman Bridge, and Grand Wash. O; 5000-5500; bottomland; riparian; Fremont River Valley. Lepidium perfoliatum L. R; 4000; canyon rims; desert shrub; Big Thomson Schoencrambe bamebyi (Welsh & Atwood) Rollins Mesa. U; 5500-7000; canyons (talus); desert shrub; Fremont River Valley, Sulphur Creek, and Lesauerella intermedia (Wats.) Heller Miners Mountain. W; 5000-8500; upland, hills, mesas, mountains, benches, and canyons; desert shrub to mixed Schoencrambe linifolia (Nutt.) Greene conifer woodland; widespread throughout the R; 6000-7000; mesas and benches; pinyon- park. juniper woodland; The Hartnet and Miners Mountain. Lesauerella ludoviciana (Nutt.) Wats. O; 4000-8500; upland, mesas, mountains, Sisymbrium altissimum L. benches, and canyons; desert shrub to mixed C; 5000-5500; bottomland; disturbed sites; conifer woodland; Burr Trail, Upper Deep Fremont River Valley. Creek, Pleasant Creek, and Halls Creek. Staneleya pinnata (Pursh) Britt. Lesauerella rectipes Woot. & Standi. W; 4000-7500; upland, mesas, and benches; W; 4000-6500; upland, mesas, and canyon desert shrub to pinyon-juniper woodland; bottoms; desert shrub to pinyon-juniper widespread throughout the park woodland; Halls Narrows; Fremont River Valley, Cedar Mesa, Pleasant Creek, Muley Staneleya viridiflora Nutt. in T. & G. Twist Canyon, and Sulphur Creek. C; 5000-6000; upland, hills, mesas, and benches; desert shrub to pinyon-juniper woodland; Lesauerella wardii Wats. Chimney Rock, Grand Wash, Fremont River R; 6500; hills; pinyon-juniper woodland; Valley, Sulphur Creek, and South Desert Pleasant Creek. Overlook.
61 Technical Report NPS/NAUCARE/NRTR-93/01
Streptanthella longirostris (Wats.) Rydb. Euphorbia ghjptosperma Engelm. in Torr. W; 4000-7000; upland, bottomland, hills, mesas, W; 4000-6000; bottomland and canyon bottoms; canyons, and benches; desert shrub to pinyon- waste places; Halls Narrows; Hickman Bridge, juniper woodland; widespread throughout the and Fruita. park. Euphorbia parryi Engelm. Streptanthus cordatus Nutt. ex T. & G. R; 4000; canyon bottoms; desert shrub; Halls U; 5000-6500; upland, hills, mesas, and benches; Narrows. sagebrush communities and pinyon-juniper woodland; Fremont River Valley, Pleasant Euphorbia robusta (Engelm.) Small ex Britt. & Creek, and Cathedral Valley. Brown O; 5500-6500; bottomland and hills; desert shrub Thehjpodiopsis divaricate (Rollins) Welsh & to pinyon-juniper woodland. Atwood O; 4000-6000; hills, benches, and canyons; desert FAGACEAE shrub to pinyon-juniper woodland; Chimney Rock, Halls Creek, Deer Point, and Sulphur Quercus eastwoodiae Rydb. Creek R; 4000; narrow canyons; riparian; Halls Creek.
Thelypodium integrifolium (Nutt.) Endl. in Walp. Quercus gambelii Nutt. O; 4000-6500; bottomland; riparian; Pleasant C; 4000-8000; canyons, mountains, mesas, and Creek, Halls Creek, and Fremont River Valley. benches; desert shrub to pinyon-juniper woodland; Pleasant Creek, Muley Twist ELAEAGNACEAE Canyon, Upper and Middle Deep Creek, Halls Creek, Oak Creek, and Cottonwood Tanks. Elaeagnus angustifolia L. O; 5000-6000; bottomland; riparian; Fruita and Quercus undulata Torr. Pleasant Creek. O; 5000-6000; canyons; desert shrub to pinyon- juniper woodland; Burr Trail. Shepherdia canadensis (L.) Nutt. O; 8000-8500; mountains; mixed conifer wood FUMARIACEAE land; Upper Deep Creek. Corydalis aurea Willd. Shepherdia rotundifolia Parry O; 5000-6000; bottomland and canyon bottoms; W; 4000-7000; upland, mesas, benches, and riparian; Paradise Draw and Fremont River canyons; desert shrub to pinyon-juniper Valley. woodland; widespread throughout the park. GENTIANACEAE ERICACEAE Centaurium exaltatum (Griseb.) Wight ex Piper Arctostaphybs patula Greene R; 5000-6000; bottomlands; saline soils (riparian); O; 5000-8000; mesas and mountains; pinyon- Oxbow of the Fremont River and Pleasant juniper and mixed conifer woodland; Burr Trail Creek. and Upper Deep Creek. Frasera albomarginata Wats. EUPHORBIACEAE O; 5500-6500; upland, hills, mesas, and benches; desert shrub to pinyon-juniper woodland; Burr Euphorbia bradrycera Engelm. in Emory Trail and Pleasant Creek. R; 5500; bottomland; riparian; Fremont River Valley. Frasera speciosa Dougl. ex Griseb. in Hook. R; 8500; mountains; mixed conifer woodland; Euphorbia fendleri T. & G. Upper Deep Creek. W; 5000-6500; bottomland, mesas, benches, and washes; desert shrub to pinyon-juniper wood Gentiana affinis Griseb. in Hook land; Fremont River Valley, Longleaf Flat, and R; 8000; mountains; mixed conifer woodland; Muley Twist. Upper Deep Creek.
62 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Gentianella heterosepala (Engelm.) Holub Lycopus asper Greene R; 8000; mountains; mixed conifer woodland; R; 5500; bottomland; riparian; oxbow of the Upper Deep Creek. Fremont River.
GERANIACEAE Marrubium xmlgare L. U; 5000-5500; bottomland and canyon bottoms; Erodium ticutarium (L.) L'Her in Ait. disturbed sites; Fruita and Sulphur Creek. W; 4500-6500; bottomland, washes, and canyon bottoms; disturbed sites; The Post, Fremont Mentha arvensis L. River Valley, and Spring Canyon. O; 5000-5500; bottomland; riparian; Fremont River Valley. Geranium caespitosum James ex Gray O; 6500-8000; upland and mountains; pinyon- Mentha spicata L. juniper woodland to mixed conifer woodland; O; 5500; bottomland; orchards; Fruita. Pleasant Creek and Upper Deep Creek. Monardella odoratissima Benth. HIPPURIDACEAE R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Hippuris vulgaris L. R; 5500; pond; aquatic; oxbow of the Fremont Nepeta cataria L. River. U; 5000-6000; bottomland and canyon bottoms; riparian; Cohab Canyon and Fremont HYDROPHYLLACEAE River Valley.
Nama retorsum J.T. Howell Poliomintha incana (Torr.) Gray U; 5500-6000; upland and hills; pinyon-juniper O; 4500-6000; upland, hills, mesas, benches, and woodland; Hickman Bridge and Longleaf Flat. canyons; desert shrub to pinyon-juniper woodland; Burr Trail, Red Slide, Grand Wash, Phacelia crenulata Torr. in Wats. Sulphur Creek, and Hickman Bridge C; 4000-6000; hills, mesas, benches, and canyons; desert shrub to pinyon-juniper woodland; Salvia reflexa Hornem. Spring Canyon, Sulphur Creek, Halls Creek, and O; 5500; bottomland; riparian; Fruita. Burr Trail. Scutellaria galericulata L. Phacelia demissa Gray R; 5500; canyon bottoms; riparian; Fremont C; 5000; hills and benches of Mancos clay; desert River Valley. shrub; Oyster Shell Reef and The Post. Stachys palustris L. Phacelia ivesiana Torr. in Ives. R; 5000-5500; bottomland; riparian; Fremont C; 4000-6000; canyon bottoms, mesas, and River Valley. benches; desert shrub to pinyon-juniper woodland; Brimhall Arch, The Hartnet, Halls LECUMINOSAE Creek, Fremont River Valley, Capitol Gorge, and Spring Canyon. Astragalus agrestis Dougl. ex G. Don R; 8500; mountains; mixed conifer woodland; Phacelia rafaelensis Atwood Upper Deep Creek. O; 5000-6000; hills, benches, and canyons; desert shrub; Fremont River Valley and Chimney Rock. Astragalus amphioxys Gray U; 4000-6000; benches and canyon bottoms; LABIATAE desert shrub; Halls Creek and Middle Desert Wash. Hedeoma drummondii Benth. O; 5000-5500; bottomland; riparian; Fremont Astragalus asclepiadoides Jones River Valley. U; 5000-6000; hillsides; desert shrub; Middle Desert Wash and Oyster Shell Reef.
63 Technical Report NPS/NAUCARE/NRTR-93/01
Astragalus barnebyi Welsh & Atwood Astragalus harrisonii Barneby R; 5000; benches and mesas; desert shrub; Big O; 5500-7500; canyons (talus); pinyon-juniper Thomson Mesa. and desert shrub; Fruita, Hickman Bridge, Spring Canyon, and Middle Deep Creek. Astragalus brandegei T.C. Porter in Port. & Coult. O; 5000-6000; bottomland and hillsides; salt Astragalus kentrophyta Gray marsh and desert shrub; Fremont River Valley C; 5500-8500; benches and hillsides; pinyon- and Upper Cathedral Valley. juniper woodland and bristlecone pine communities; Fremont River Valley and Astragalus cah/cosus Torr. Upper Deep Creek. R; 6500; hilltops; pinyon-juniper woodland; Pleasant Creek. Astragalus lentiginosus Dougl. ex Hook. O; 4000-5500; benches; desert shrub and pinyon- Astragalus ceramicus Sheld. juniper woodland; Brimhall Arch and Sulphur C; 5500-7500; bottomland to mountains; desert Creek shrub to mixed coniferous woodland; Fremont River Valley, Longleaf Flat and Upper Deep Astragalus lonchocarpus Torr. Creek. C; 5500-8000; hillsides, benches and canyon bottoms; pinyon-juniper woodland to mixed Astragalus chamaeleuce Gray in Ives conifer woodland; Upper and Middle Deep R; 5000-6700; hillsides; pinyon-juniper wood Creek, Sulphur Creek, and Sheets Gulch. land; Burr Trail area. Astragalus malaarides Barneby Astragalus coltonii Jones U; 4000-4500; bottomland and canyon bottoms; O; 5500-7500; mountains, mesas, and canyons; desert shrub; Halls Creek and near Brimhall mixed conifer woodlands; Upper Deep Creek, Arch. South Desert Overlook, and Pleasant Creek. Astragalus miser Dougl. ex Hook. Astragalus consobrinus (Barneby) Welsh R; 8500; mountains; mixed conifer woodland; U; 5000-6500; upland, benches and canyons; Upper Deep Creek. pinyon-juniper woodland; Danish Hill, Fruita, and Water Canyon. Astragalus moencoppensis Jones U; 4500-5500; benches; desert shrub; Brimhall Astragalus convallarius Greene Arch. R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Astragalus mollissimus Torr. var. thompsonae (Wats.) Bameby Astragalus desperatus Jones var. petrophilus Jones C; 4500-6000; benches, hillsides and canyon R; 5600; hillside; desert shrub; Ackland Spring. bottoms; desert shrub and pinyon-juniper woodland; Fremont River Valley, Fruita, Bun- Astragalus episcopus Wats. Trail and Halls Creek. O; 5500-6000; uplands; desert shrub; Middle Desert Wash, Cathedral Valley, and South Astragalus musiniensis Jones Desert. U; 6500; hillsides and talus slopes; pinyon- juniper woodland; Burr Trail and Muley Twist Astragalus flavus Nutt. in T. & G. var. argillosus Canyon. (Jones) Barneby R; 5800; hillsides; desert shrub; Jailhouse Rock. Astragalus nuttallianus Nutt. R; 4000; benches and mesas; desert shrub; Halls Astragalus flavus Nutt. in T. & G. var. candiams Creek Gray U; 5500-6500; hillsides, washes and bottomland; Astragalus pardalinus (Rydb.) Barneby pinyon-juniper woodland and riparian; Fremont U; 5500-6000; bottomland; desert shrub; Upper River Valley and The Hartnet. Cathedral Valley and South Desert.
64 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Astragalus praelongus Sheld. Lathyrus brachycalyx Rydb. var. zionis (C.L. O; 5000-6000; bottomland and hillsides; desert Hitchc.) Welsh shrub; Burr Trail and Circle Cliffs. R; 4000; canyon bottoms; riparian; Halls Narrows. Astragalus sabulonum Gray U; 5000-6000; hills and benches; desert shrub; Lathyrus pauciflorus Fern. Cathedral Valley. R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Astragalus sesquiflorus Wats. O; 5500-6500; benches; pinyon-juniper wood Lupinus argenteus Pursh var. argenteus land; Pleasant Creek, Sheets Gulch and Sulphur O; 5000-5500; upland, canyons, and mesas; Creek. desert shrub to pinyon-juniper woodland; Cathedral Valley, Cedar Mesa, and Pleasant Astragalus tenellus Pursh Creek. O; 7000-8000; mountains; mixed conifer wood land; Upper and Middle Deep Creek. Lupinus argenteus Pursh var. rubricaulis U; 5500-8000; upland and mountains; pinyon- Astragalus wardii Gray juniper woodland to mixed conifer woodland; R; 8500; mountains; mixed conifer woodland; The Hartnet and Upper Deep Creek. Upper Deep Creek. Lupinus brevicaulis Wats. Astragalus woodruffii Jones U; 4000-4500; washes and canyon bottoms; R; 5500; hillsides; desert shrub; Fremont River desert shrub; Red Slide and Brimhall Valley. Arch.
Astragalus zionis Jones Lupinus pusillus Pursh U; 4000-4500; canyons and canyon bottoms; C; 4000-6500; upland, hills, mesas, benches, desert shrub; Halls Creek. washes, and canyons; desert shrub to pinyon- juniper woodland; Pleasant Creek, Cottonwood Dalea flavescens (Wats.) Welsh var. flavescens Tanks, Willow Tanks, Cathedral Valley, and O; 4000-5500; hillsides; desert shrub; Redslide Halls Creek. and Burr Trail. Lupinus sericeus Pursh Dalea flavescens (Wats.) Welsh var. epica (Welsh) O; 8500; mountains; mixed conifer woodland; Welsh & Chatterley Upper Deep Creek. R; 5000; hillsides; desert shrub; The Post. Medicago lupulina L. Dalea oligophylla (Ton.) Shinn. U; 5500; bottomland; riparian; oxbow of the O; 4000-5500; hills, canyon bottoms, and washes; Fremont River. desert shrub; Red Slide and Cathedral Valley. Medicago sativa L. C; 5000-6500; bottomland; riparian; Fruita and Glycyrrhiza lapidota Pursh Pleasant Creek. O; 5000-5500; bottomland; riparian; Fremont River Valley. Melilotus albus Medicus C; 5000-6500; bottomland; disturbed sites; Hedysarum Occident ale Greene Fremont River Valley, Pleasant Creek, and O; 5000-6000; canyon bottoms; desert shrub to Grand Wash. pinyon-juniper woodland; Cathedral Valley, Sulphur Creek, and Upper Muley. Melilotus officinalis (L.) Pallas C; 4000-6500; bottomland and canyon bottoms; Lathyrus brachycalyx Rydb. var. brachycalyx disturbed sites; Fremont River Valley, Pleasant R; 5000; hills; pinyon-juniper woodland; Muley Creek, and Halls Creek. Twist Canyon.
65 Technical Report NPS/NAUCARE/NRTR-93/01
Oxytropis deflexa (Pallas) DC Vicia americana Muhl. ex Willd. R; 8500; mountains; mixed conifer woodland; U; 4000-8000; mountains and canyon bottoms Upper Deep Creek. (shady sites); hanging garden communities and mixed conifer woodland; Upper Deep Creek, Oxytropis lambertii Pursh Halls Narrows, and Pleasant Creek. O; 5000-6000; hills and benches; desert shrub to pinyon-juniper woodland; Middle Desert Wash, Ackland Spring, and Cathedral Valley. LENTIBULARIACEAE
Oxytropis oreophua Gray var. juniperina Welsh Utricularia vulgaris L. U; 7000-8000; mountains and benches; pinyon- C; 5500; pond; aquatic; oxbow of the Fremont juniper woodland to mixed conifer woodland; River. Upper Deep Creek and Miners Mountain.
Oxytropis viscida Nutt. UNACEAE R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Linum aristatum Engelm. in Wisliz. O; 4000-6500; upland, bottomland, mesas, Peteria thompsonae Wats. benches, and canyons; desert shrub to R; 5000; margins of washes; desert shrub; pinyon-juniper woodland; Middle Desert, The Cathedral Valley. Post, Fremont River Valley, Halls Creek, Red Slide, and Pleasant Creek Psoralen juncea Eastw. R; 4000; hills and canyons; desert shrub; Halls Linum perenne L. Divide. O; 5000-8000; canyons, mesas, and mountains; pinyon-juniper woodland to mixed conifer Psoralea lanceolate Pursh var. lanceolata woodland; Upper Deep Creek, Burr Trail, C; 4500-5500; bottomland and canyon bottoms; Pleasant Creek, and Muley Twist Canyon. desert shrub to pinyon-juniper woodland; Fruita, Red Slide, and Sulphur Creek Linum puberulum (Engelm.) Heller U; 5000-6000; hills and upland; desert shrub to Psoralea lanceolata Pursh var. stenophylla (Rydb.) pinyon-juniper woodland; Middle Desert Wash Toft and Welsh and Fruita. R; 4000; canyons and canyon bottoms; desert shrub; Halls Creek. LOASACEAE Psoreothamnus fremontii (Torr.) Barneby O; 4000-5000; bottomland; desert shrub; Halls Metttzelia aMcaulis Dougl. in Hook. Creek and Spring Canyon. W; 4000-6500; bottomland, hills, benches, mesas, and canyons; desert shrub to pinyon-juniper Thermopsis montana Nutt. in T. & G. woodland; widespread throughout the park. O; 5500-8000; bottomland and canyon bottoms; riparian; Fremont River Valley and Upper Deep Mentzelia cronauistii Thompson & Prigge Creek. O; 4000-5000; hillsides and roadsides; disturbed sites; Halls Narrows and The Post. Trifolium fragiferum L. R; 5000-5500; bottomland; riparian; Fremont Mentzelia multiflora (Nutt.) Gray River Valley. O; 5000-6000; bottomland, hills, benches, and washes; desert shrub; Fremont River Valley, Trifolium pratense L. Cathedral Valley, and Chimney Rock. U; 5500; bottomland; riparian; Fremont River Valley. Mentzelia pterosperma Eastw. O; 4000-5500; upland, hills, mesas, benches, and Trifolium repens L. canyon bottoms; desert shrub; -Middle Desert O; 5000-6500; bottomland; riparian; Fremont Wash, Burr Trail, and Halls Creek. River Valley and Pleasant Creek.
66 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
MALVACEAE Mirabilis oxybaphoides Gray in Torr. O; 5000-6000; canyons; pinyon-juniper wood Sphaeralcea coccinea (Nutt.) Rydb. land and riparian; Fremont River Valley, Fruita, W; 4000-6500; upland, hills, mesas, benches, and Hickman Bridge, Lower Muley, Grand Wash, canyons; desert shrub to pinyon-juniper Cohab and Frying Pan Canyons. woodland; widespread throughout the park. OLEACEAE Sphaeralcea grossulariifolia (H. & A.) Rydb. var. grossulariifolia Fraxinus anomala Torr. in Wats. O; 5000-5500; benches and hills; desert shrub; W; 3900-7000; upland, hills, mesas, benches, Cathedral Valley and Spring Canyon. washes, and canyon bottoms; desert shrub to pinyon-juniper woodland; widespread through Sphaeralcea grossulariifolia (H. & A.) Rydb. var. out the park. moorei Welsh R; 4000; canyon bottoms; riparian; Halls Creek ONAGRACEAE
Sphaeralcea parvifolia A. Nels. Calylophus lavandulifolius (T. & G.) Raven W; 4000-6500; upland, hills, mesas, benches, and O; 5500-7000; benches and hillsides; pinyon- canyons; desert shrub to pinyon-juniper juniper woodland; South Desert Overlook and woodland; widespread throughout the park. Miners Mountain.
NAJADACEAE Camissonia eastwoodiae (Munz) Raven O; 4000-5000; bottomland; desert shrub; Halls Najas guadalupensis (Spreng.) Morong Creek and base of Swap Mesa. R; 6500; slow moving streams; aquatic; Pleasant Creek. Camissonia scapoidea (T. & G.) Raven W; 5000-6000; benches and canyon bottoms; NYCTAGINACEAE desert shrub; Fremont River Valley and Jail- house Rock Abronia fragrans Nutt. ex Hook W; 3900 to 8250; washes, canyon bottoms and Camissonia walkeri (A. Nels.) Raven hillsides; pinyon-juniper and desert shrub; O; 5000-6000; hillsides; desert shrub to pinyon- widespread throughout the park. juniper woodland; Burr Trail and Sulphur Creek Abronia nana Wats. R; 5600; hilltops and mesas; pinyon-juniper Epilobium andenocaulon Hausskn. woodland; Danish Hill. O; 5500-8000; canyon bottoms and bottomland; riparian; Upper Deep Creek, Pleasant Creek Allionia incarnata L. and oxbow of the Fremont River. O; 4000-5500; Uplands, washes and canyon bottoms; desert shrub; Lower Halls, Hickman Epilobium angustifolium L. Bridge area, Cohab/Frying Pan and the Fremont O; 8000-8500; mountains; mixed conifer wood River Valley. land, aspen, and sagebrush flats; Upper Deep Creek Mirabilis linearis (Pursh) Heimerl var. decrpiens (Standley) Welsh Gaura parviflora Dougl. ex Hook. R; 8500; mountains; sagebrush community; O; 5500; bottomland; riparian; Fruita. Upper Deep Creek Gayophytum ramosissimum Nutt. in T. & G. Mirabilis linearis (Pursh) Heimerl var. linearis O; 8000-8500; mountains; mixed conifer wood W; 5000-7000; benches, bottomland, and canyon land; Upper Deep Creek. bottoms; desert shrub to pinyon-juniper woodland; widespread throughout the park. Oenothera albicaulis Pursh O; 5000-6000; upland, bottomland, hills, and Mirabilis multifbra (Torr.) Gray in Torr. mesas; desert shrub to pinyon-juniper wood W; 4500-7000; mesas, washes, and canyon land; oxbow of the Fremont River and The Post. bottoms; pinyon-juniper woodland; widepsread throughout the park 67 Technical Report NPS/NAUCARE/NRTR-93/01
Oenothera caespitosa Nutt. var. marginata (Nutt.) Plantago paUigonica Jacq. Munz C; 5000-6000; upland, hills, mesas, and benches; W; 4000-8500; upland, hills, mesas, benches, desert shrub to pinyon-juniper woodland; Burr washes, and mountains; desert shrub to mixed Trail, Capitol Gorge, Hickman Bridge, Sulphur conifer woodland; widespread throughout the Creek, and Cohab/Frying Canyons. park. POLEMON1ACEAE Oenothera caespitosa Nutt. var. macroglottis (Rydb.) Cronq. Eriastrum diffusum (Gray) Mason U; 7500; flats,meadows , and hillsides; pinyon- R; 4000-4500; bottomland; desert shrub; south of juniper woodland; Paradise Flats. Brimhall Arch near the exclosure.
Oenothera coronopifolia T. & G. Cilia aggregata (Pursh) Spreng. R; 8500; mountains; mixed conifer woodland; W; 5000-8000; upland, mesas, mountains, and Upper Deep Creek. benches; desert shrub to mixed conifer wood land; widespread throughout the park. Oenothera hookeri T. & G. O; 4000-6500; bottomland and canyon bottoms; Cilia caespitosa Gray riparian; Fremont River Valley, Pleasant Creek R; 6000-7000; sides of washes and canyons; and Halls Creek. slickrock communities; Pine Canyon, Middle Deep Creek, and Longleaf Hat. Oenothera howardii Jones R; 5000-6000; hills and benches; pinyon-juniper Gilia congesta Hook. woodland; Burr Trail and Deer Point. O; 6000-6500; canyons; slickrock communities and pinyon-juniper woodland; Burr Trail and Oenothera longissima Rydb. Pleasant Creek R; 5500; upland and washes; pinyon-juniper woodland (shady sites); Grand Wash. Gilia gunnisonii T. & G. O; 4000-4500; bottomland; desert shrub; Halls Oenothera pallida Lindl. Creek. W; 5000-7000; benches, hills, upland, mesas, and canyons; desert shrub to pinyon-juniper Gilia inconspicua (J.E. Sm.) Sweet woodland; widespread throughout the park. W; 4500-6000; hills, benches, and washes; desert shrub; Brimhall Arch, The Post, Big Thomson ORBANCHACEAE Mesa, and Burr Trail.
Orobanche fasciculata Nutt. Gilia latifolia Wats. O; 5000-5500; upland, hills, mesas, and benches; R; 5500; bottomland; riparian; near the oxbow of desert shrub; Middle Desert Wash. the Fremont River.
Orobanche ludoviciana Nutt. Gilia leptomeria Gray O; 5000-6500; bottomland, upland, hills, mesas, W; 4000-6500; upland, hills, mesas, benches, and and washes; desert shrub to pinyon-juniper washes; desert shrub to pinyon-juniper wood woodland; Middle Desert Wash, Fremont River land; widespread throughout the park. Valley, and Pleasant Creek. Gilia longiflora (Torr.) G. Don PLANTAGINACEAE O; 5000-6000; washes and canyon bottoms; desert shrub to pinyon-juniper woodland; Plantago lanceolata L. Sulphur Creek, Pleasant Creek, Hickman Bridge, O; 5000-5500; bottomland; disturbed sites; and Cathedral Valley. Fruita. Gilia polycladon Torr. in Emory Plantago major L. O; 4500^6500; upland, hills, mesas, and benches; O; 5500; bottomland; riparian; oxbow of the desert shrub to pinyon-juniper woodland; Fremont River. Brimhall Arch, The Hartnet, Sulphur Creek, and Hickman Bridge.
68 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Gilia pumila Nutt. POLYGONACEAE O; 5000-6500; upland, hills, mesas, benches, and canyons; desert shrub to pinyon-juniper Eriogonum alatum Torr. in Sitgr. woodland; Sulphur Creek, Burr Trail, and The W; 3900-8000; canyon bottoms, hills, and Hartnet. benches; desert shrub to mixed conifer wood land; Upper Deep Creek, Cathedral Valley, and Gilia roseata Rydb. Halls Creek. O; 5500-6500; upland and benches; slickrock communities; Pine Canyon, Hickman Bridge, Eriogonum bicolor Jones and Spring Canyon. U; 5000-6500; hillsides, upland, and benches; desert shrub to pinyon-juniper woodland; The Gilia subnuda Torr. ex Gray Hartnet, Sheets Gulch, and Cedar Mesa. W; 4000-6500; canyon bottoms, upland, hills, mesas, and benches; desert shrub to pinyon-juniper Eriogonum cernuum Nutt. woodland; widespread throughout the park. W; 5000-8500; upland, mountains, hills, mesas, and benches; pinyon-juniper woodland to leptodactylon pungens (Torr.) Nutt. mixed conifer woodland; widespread through O; 4000-6000; upland and canyon bottoms; out the park. slickrock communities; Burr Trail, Muley Twist Canyon, and Halls Creek. Eriogonum corymbosum Benth. var. aureum (Jones) Reveal Leptodactylon watsonii (Gray) Rydb. R; 5500; hills and benches; desert shrub; R; 5500-8000; canyon walls; slickrock communi Chimney Rock and Burr Trail. ties; Longleaf Flat, Burr Trail; Upper and Middle Deep Creek. Eriogonum corymbosum Benth. var. corymbosum W; 5000-7000; upland, hills, benches, and Phlox austromontana Cov. var. austromontana canyons; desert shrub to pinyon-juniper O; 5000-8000; mesas and mountains; pinyon- woodland; widespread throughout the park. juniper woodland to mixed conifer woodland; Upper Deep Creek, Cedar Mesa, and Muley Eriogonum corymbosum Benth. var. orbiculatum Twist Canyon. (Stokes) Reveal & Brotherson O; 5000-6500; upland, benches, and canyons; Phlox austromontana Cov. var. lutescens Welsh slickrock communities; Muley Twist Canyon, R; 7000; hills and benches; blackbrush commu Burr Trail, and Hickman Bridge. nity; Deer Point. Eriogonum corymbosum Benth. var. revealianum Phlox hoodii Richards (Welsh) Reveal O; 5000-6500; canyons; pinyon-juniper wood R; 8500; benches; bristlecone pine communities; land; Fremont River Valley, Pleasant Creek, and Upper Deep Creek. Spring Canyon. Eriogonum deflexum Torr. in Ives Phlox longifolia Nutt. U; 3900-5500; bottomland, washes, and canyon U;.5000-6500; upland and mesas; pinyon-juniper bottoms; desert shrub; oxbow of the Fremont woodland; Cedar Mesa and Cathedral Valley. River and Halls Narrows.
Phlox muscoides Nutt. Eriogonum flexum Jones U; 6000-7000; canyons; sagebrush and slickrock C; 4000-5500; upland, hills, benches, washes, and communities, and pinyon-juniper woodland; canyons; desert shrub to pinyon-juniper near Jailhouse Rock (in the fold) and South woodland; Brimhall Arch, Grand Wash, and Desert Overlook. Halls Narrows.
POLYGALACEAE Eriogonum gordonii Benth. in DC. U; 6000-7000; hillsides and canyon bottoms; Polygala subspinosa Wats. desert shrub to pinyon-juniper woodland; O; 5000-6000; upland, hills, mesas, benches, and Meeks Mesa and Middle Deep Creek. canyons; desert shrub to pinyon-juniper woodland; Burr Trail, Red Slide, Grand Wash, Sulphur Creek, and Hickman Bridge. 69 Technical Report NPS/NAUCARE/NRTR-93/01
Eriogonum hookeri Wats. Eriogonum umbellatum Torr. C; 5000-6500; bottomland, hills, benches, and U; 6500-7500; canyons and benches; pinyon- washes; desert shrub to pinyon-juniper wood juniper woodland; Oak Creek, Miners land; oxbow of the Fremont River, Hickman Mountain, and Pleasant Creek. Bridge, Chimney Rock, and Pleasant Creek. Eriogonum wetherillii Eastw. Eriogonum inflatum T. & G. O; 4000-6000; bottomland, washes, and canyon O; 5000-6500; upland, hills, and benches; desert bottoms; desert shrub to pinyon-juniper shrub; Fremont River Valley, Pleasant Creek, woodland; Halls Creek, Muley Twist Canyon, and The Post. and Cohab/Frying Pan Canyons.
Eriogonum leptocladon T. & G. var. leptocladon Polygonum aviculare L. U; 5000; upland and hills; desert shrub; Cathe O; 5500; bottomland; disturbed sites; oxbow of dral Valley. the Fremont River and Fruita.
Eriogonum leptocladon T. & G. var. ramosissimum Polygonum douglasii Greene (Eastw.) Reveal O; 6000-8500; upland and mountains; pinyon- W; 5000-6000; upland, hills, mesas, benches, and juniper woodland to mixed conifer woodland; canyons; desert shrub to pinyon-juniper Oak Creek and Upper Deep Creek. woodland; widespread throughout the park. Polygonum kelloggii Greene Eriogonum microthecum Nutt. var. foliosum (T. & R; 8500; mountains; mixed conifer woodland; G.) Reveal Upper Deep Creek. W; 4000-6500; upland, hills, mesas, benches, and canyons; desert shrub to pinyon-juniper Rumex crispus L. woodland; widespread throughout the park. C; 5000-5500; bottomland; riparian; Fremont River Valley. Eriogonum microthecum Nutt. var. laxiflorum Hook. Rumex hymenosepalus Torr. R; 6000; benhces; pinyon-juniper woodland; Oak O; 4000-4500; washes and canyon bottoms; Creek. desert shrub; Halls Creek and the Red Slide.
Erigonum ovalifolium Nutt. Rumex salicifolius Weinm. O; 5000-6500; hills, mesas, and benches; pinyon- U; 5500; bottomland; riparian; Fremont River juniper woodland and sagebrush communities; Valley. Burr Trail, Capitol Gorge, and Pleasant Creek. PORTULACEAE Eriogonum palmerianum Reveal in Munz R; 3900; hillsides; desert shrub; Halls Creek. Portulaca oleracea L. O; 5500; bottomland; disturbed sites; Fruita. Eriogonum racemosum Nutt. U; 8000-8500; mountains; mixed conifer wood Talinum brevifolium Torr. land; Upper Deep Creek. R; 6000; upland; pinyon-juniper woodland; Longleaf Flat. Eriogonum shockleyi Wats. O; 5000-6000; upland, hills, mesas, and benches; PRIMULACEAE desert shrub to pinyon-juniper woodland; Middle Desert Wash, Meeks Mesa, Muley Twist Androsace septentrionalis L. Canyon, and Burr Trail. U; 8000-3500; mountains; aspen communities and mixed conifer woodland; Upper Deep Eriogonum subreniforme Wats. Creek. U; 4000-5000; bottomland and washes; desert shrub; The Post and Halls Creek. Glaux maritima L. U; 5500; bottomland; riparian; oxbow of the Eriogonum tumulosum (Barneby) Reveal Fremont River. R; 6500; hills and benches; pinyon-juniper woodland; South Desert Overlook.
70 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
PYROLACEAE Rhamnus betulifolia Greene O; 4000-5000; canyon bottoms; riparian and Pterospora andromeda Nutt. hanging garden communities; Muley Twist U; 8500; mountains; mixed conifer woodland; Canyon and Halls Narrows. Upper Deep Creek. ROSACEAE RANUNCULACEAE Amelanchier alnifolia (Nutt.) Nutt. Aamitum columbianum Nutt. in T. &. G. O; 7000-8500; mountains; mixed conifer wood R; 8800; mountains; shaded coniferous forest; land; Upper Deep Creek. Upper Deep Creek. Amelanchier utahensis Koehne Aauilegia caerulea James in Long W; 5000-7500; uplands; pinyon-juniper wood R; 8800; mountains; shaded coniferous forest; land; widespread throughout the park. Upper Deep Creek. Cercocarpus intricatus Wats. Aquilegia mkrantha Eastw. W; 4000-6500; uplands, benches, rimrock cliffs R; 4000; canyon walls; hanging gardens; Lower and slopes (slickrock); widespread throughout Halls Creek. the park.
Clemantis Columbiana (Nutt.) T. & G. Cercocarpus ledifolius Nutt. O; 7500-8500; mountains; mixed conifer wood O; 8000-8500; mountains; mixed conifer wood land; Upper Deep Creek. land; Upper Deep Creek.
Oemantis ligusticifolia Nutt. in T. & G. Cercocarpus montanus Raf. O; 4000-7500; bottomland, washes, canyon O; 7500-8500; mountains; mixed conifer wood bottoms; desert shrub to mixed conifer land; Upper Deep Creek. woodland; widespread throughout the park. Coleogyne ramosissima Torr. Delphinium andersonii Gray var. scaposum O; 4000-5000; uplands; black brush communi (Greene) Welsh ties; Burr Trail and Halls Creek. 5000-6000; upland, mesas, benches, and hill sides; desert shrub to pinyon-juniper woodland; Fallugia paradoxa (D. Don) Endl. Pleasant Creek, Cathedral Valley, and Spring W; 4000-6500; canyons and washes; desert shrub Canyon. to pinyon-juniper woodland; widespread throughout the park. Ranunculus cymbalaria Pursh O; 5000-6500; bottomland; riparian; oxbow of the Fragaria xrirginiana Duchesne Fremont River, Pleasant Creek, and a spring at O; 8000; mountains; mixed conifer woodland; South Desert Overlook. Upper Deep Creek.
Ranunculus scleratus L. Holodiscus dumosus (Nutt.) Heller O; 5000-5500; bottomland; riparian;Fremon t O; 5000-8000; canyons, mountains, mesas, River Valley. benches, and hillsides; pinyon-juniper woodland to mixed conifer woodland; Upper Ranunculus testiculatus Crantz Deep Creek, Water Canyon, Fruita, Fremont R; 5500; bottomland; disturbed sites; Burr Trail. River Valley, Capitol Gorge and Pleasant Creek.
Thalictrum fendleri Engelm. in Gray Petrophytum caespitosum (Nutt.) Rydb. O; 8000-8800; mountains; mixed conifer wood O; 5500-7500; upland and canyons; Navajo land; Upper Deep Creek. Sanstone rock crevices in the pinyon-juniper woodland; in the fold near Jailhouse Rock and RHAMNACEAE Upper and Middle Deep Creek.
Ceanothus fendleri Gray Physocarpus ailerons (Jones) J.T. Howell O; 7000-8000; mountains; mixed conifer wood U; 5000-8000; canyons; desert shrub to mixed land; Upper Deep Creek. conifer woodland; Muley Twist Canyon and Upper Deep Creek. 71 Technical Report NPS/NAUCARE/NRTR-93/01
Potentilla concinna Richards SAUCACEAE O; 8000-8500; mountains; sagebrush communi ties and mixed conifer woodland; Upper Deep Populus acuminata Rydb. Creek. O; 5000-6500; bottomland; riparian; Water Canyon and Pleasant Creek. Potentilla crinita Gray O; 8000-8500; mountains; sagebrush communi Populus angustifolia James ex Torr. ties; Upper Deep Creek. O; 5000-6500; bottomland and canyons; riparian; Cathedral Valley, Pleasant Creek, and Spring Potentilla diversifolia Lehm. Canyon. R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Populus fremontii Wats. W; 3900-7500; bottomland and canyon bottoms; Potentilla mppiana Lehm. riparian; widespread throughout the park. O; 5500-8000; bottomland and mountains; riparian and mixed conifer woodland; Populus tremuloides Michx. Fremont River Valley and Upper Deep Creek. O; 8000-8800; mountains; mixed conifer wood land and aspen communities; Upper Deep Potentilla pensylvanica L. Creek R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Salix amygdaloides Anderss. O; 5000-6500; bottomland; riparian; Fremont Purshia mexicana (D. Don) Welsh River Valley, Pleasant Creek, and Oak Creek. W; 5000-7000; mesas, benches, canyon bottoms, and mountains; pinyon-juniper woodland to Salix exigua Nutt. mixed conifer woodland; Sulphur Creek, Bun- C; 4000-6500; bottomland and canyon bottoms; Trail, Upper Deep Creek, and Miners Mountain. riparian; Halls Creek, Fremont River Valley, Pleasant Creek, and Muley Twist Canyon. Purshia tridentata (Pursh) DC. C; 7500-8000; mountains; mixed conifer wood Salix geyeriana Anderss. land and sagebrush communities; Upper Deep R; 5500; canyon bottoms; riparian; near Jailhouse Creek Rock in the fold.
Rosa woodsii Lindl. Salix gooddingii Ball C; 5000-8000; canyon bottoms and mountains; U; 4000-5000; bottomland and canyon bottoms; riparian; Fremont River Valley, Pleasant Creek, riparian; Cottonwood Tanks and Halls Creek. and Upper Deep Creek. Salix lasiandra Benth. RUBIACEAE R; 5000; bottomland; riparian; Spring Canyon.
Galium aparine L. Salix lasiolepis Benth. O; 4000-8000; mountains and canyon bottoms; U; 5000-5500; canyon bottoms; riparian; Fremont hanging gardens and other moist sites; Halls River Valley, Grand Wash, and Spring Canyon. Creek and Upper Deep Creek. Salix lutea Nutt. Galium multiflorum Kellogg O; 4000-8000; bottomland and canyon bottoms; O; 4000-6000; canyons; desert shrub to pinyon- riparian; Upper Deep Creek, Hickman Bridge, juniper woodland; Jailhouse Rock, Lower Muley and Halls Creek. Twist Canyon, Chimney Rock Canyon, and Pleasant Creek. SANTALACEAE
Galium trifidum L. Comandra umbellata (L.) Nutt. O; 5500; bottomland; riparian; oxbow of the C; 5000-6000; bottomland, mesas, benches, and Fremont River. canyon bottoms; desert shrub to pinyon-juniper woodland; Fremont River Valley and Cathedral Valley.
72 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
SAXIFRAGACEAE Mimulus glabratus H.B.K. R; 5500; bottomland; riparian; near the oxbow of Heuchera parvifolia Nutt. in T. & G. the Fremont River. R; 8000; mountains; mixed conifer woodland; Upper Deep Creek. Mimulus guttatus Fisch. in DC. O; 4000-6500; canyons and bottomland; riparian; Pamassia Palustris L. Fremont River Valley, Pleasant Creek, and Halls R; 6500-8000; bottomland; riparian; Reasant Creek. Creek and Upper Deep Creek. Pedicularis centranthera Gray in Torr. Philadelpfms microphyllus Gray O; 6000-7000; canyons and hills; pinyon-juniper U; 5000-5500; bottomland; riparian; Fremont woodland; Sheets Gulch, Upper Muley, and River Valley. Miners Mountain.
Ribes aureum Pursh Penstemon ambiguus Torr. U; 5000-5500; bottomland; riparian; Fremont R; 3900; hills and benches; desert shrub; Halls River Valley. Creek.
Ribes cereum Dougl. Penstemon angustifolius Nutt. O; 5000-8000; canyon bottoms and mountains; O; 4000-6000; bottomland, canyons, hills, mesas, riparian and mixed conifer woodland; Water and benches; desert shrub to pinyon-juniper Canyon, Pleasant Creek, and Upper Deep Creek. woodland; Muley Twist Canyon, Halls Creek, and Burr Trail. Ribes inerme Rydb. O; 8000-8500; mountains; mixed conifer wood Penstemon barbatus (Cav.) Roth land; Upper Deep Creek. O; 6000-8000; canyons, benches, and mountains; pinyon-juniper woodland to mixed conifer SCROPHULARIACEAE woodland; Upper Deep Creek, Pleasant Creek, and Upper Muley. Castilleja chromosa A. Nels. W; 4000-6500; uplands, mesas, benches; desert Penstemon caespitosus Nutt. ex Gray var. perbrevis shrub to pinyon-juniper woodland; Burr Trail, Pennell Cedar Mesa, and Halls Creek. R; 7000; upland, hills, and benches; pinyon- juniper woodland; Miners Mountain. Castilleja exilis A. Nels. O; 5000-6000; bottom land; riparian; Fremont Penstemon carnosus Pennell in Graham River Valley and Pleasant Creek. O; 5000-5500; hills, mesas, and benches; desert shrub to pinyon-juniper woodland; Burr Trail, Castilleja linariifolia Benth. Chimney Rock, and Fruita. W; 4000-8800; upland, mesas, and mountains; desert shrub to mixed conifer woodland; Halls Penstemon comarrhenus Gray Creek, Bitter Spring Creek and Upper Deep O; 6500-8000; upland, mesas, mountains, and Creek. benches; pinyon-juniper woodland to mixed conifer woodland; Upper Deep Creek, Pleasant Castilleja scabrida Eastw. Creek, and Miners Mountain. O; 4000-6000; hillsides and benches; slickrock communities; Hickman Bridge, Oak Creek, Penstemon eatonii Gray Cottonwood Tanks, and Halls Creek. W; 3900-7000; canyon bottoms, upland, hills, mesas, and benches; desert shrub to pinyon- Cordylanthus parviflorus (Ferris) Wiggins juniper woodland; widespread throughout the O; 4000-6500; mesas and canyon bottoms; desert park. shrub to pinyon-juniper woodland; Halls Creek, Wagon Box Mesa, and Pleasant Creek. Penstemon leiophyllus Pennell U; 8000-8500; mountains; mixed conifer wood Mimulus eastwoodiae Rydb. land; Upper Deep Creek. R; 4000; canyons; hanging gardens; Halls Narrows.
73 Technical Report NPS/NAUCARE/NRTR-93/01
Penstemon moffattii Eastw. Nicotiana attenuata Torr. ex Wats. U; 5000; canyon rims; desert shrub; Big Thom O; 5500-7000; bottomland, washes, and mesas; son Mesa. disturbed sites; Fremont River Valley and The Hartnet. Penstemon ophianthus Pennell O; 5000-7000; hills, mesas, and benches; desert Physalis longifolia Nutt. shrub to pinyon-juniper woodland; South Desert C; 5000-5500; bottomland; riparian; Fremont Overlook, Cedar Mesa, and Cathedral Valley. River Valley.
Penstemon pachyphylhis Gray ex Rydb. Solarium elaeagnifolium Cav. U; 6000-8000; mesas, hills, and benches; pinyon- U; 5500; bottomland; disturbed sites; Hickman juniper woodland to mixed conifer woodland; Bridge parking lot. South Desert Overlook and Upper Deep Creek. Solanum sarrachoides Sendt. in Mart. Penstemon palmeri Gray var. eglandulosus (Keck) U; 5500; bottomland; disturbed sites; Fremont N. Holmgren River Valley. O; 5000-6500; upland, hills, mesas, benches, and canyons; desert shrub to pinyon-juniper Solanum triflorum Nutt. woodland; Burr Trail, Capitol Gorge, and R; 5500; bottomland; disturbed sites; Fremont Pleasant Creek. River Valley.
Penstemon rostriflorus Kellogg TAMARICACEAE R; 5000-6500; canyons, benches, and mesas; desert shrub to pinyon-juniper woodland; Tamarix ramosissima Ledeb. Pleasant Creek and Muley Twist Canyon. W; 4000-7000; bottomland and canyon bottoms; riparian; widespread throughout the park. Penstemon strictiformis Rydb. U; 8000-8500; hillsides; mixed conifer woodland; ULMACEAE Upper Deep Creek. Celtis occidentalis L. Penstemon strictus Benth. in DC. O; 5000-5500; bottomland; orchards; Fruita. R; 8500; mountains; mixed conifer woodland; Upper Deep Creek. Celtis reticulata Torr. O; 4000-5500; canyon bottoms; sagebrush, desert Penstemon utahensis Eastw. shrub and pinyon-juniper woodlands; Halls C; 5000-6500; upland, hills, mesas, benches, and Creek, Cottonwood Tanks, and Grand Wash. canyons; desert shrub to pinyon-juniper woodland; Muley Twist Canyon, Burr Trail, UMBELUFERAE Deer Point, Cedar Mesa, Chimney Rock, and Spring Canyon. Anethum graveolens L. O; 5500; canyon bottoms; orchards along the Veronica americana Schwein. in DC. Fremont River. U; 5500-6500; bottomland; riparian; near the oxbow of the Fremont River and Pleasant Creek. Cicuta maculata L. R; 5500; benches; riparian; Settling pond along Veronica anagallis-aauatica L. the Fremont River. O; 5000-8000; bottomland; riparian; Upper Deep Creek, Water Canyon, oxbow of the Fremont Cymopterus becldi Welsh & Goodrich River, and Pleasant Creek. U; 5500-6000; bottomland and canyon bottoms; pinyon-juniper woodland; Fremont River SOLANACEAE Valley, Cohab Canyon, Spring Canyon, and Pine Canyon. Datura meteloides Dunal in DC. O; 4500-6000; washes; desert shrub; Fremont Cymopterus buWosus A. Nels. River Valley and Halls Creek. U; 5000-5500; hillsides; desert shrub; Burr Trail.
74 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Cymopterus fendleri Gray VERBENACEAE U; 5000-8000; hills and mountains; desert shrub to mixed conifer woodland; Fremont River Verbena bracteata Lag. & Rodr. Valley and Upper Deep Creek. O; 5000-6500; bottomland; disturbed sites; Fremont River Valley and Pleasant Creek. Cymopterus lemmonii (Coult. & Rose) Cronq. R; 8000-8500; mountains; mixed conifer wood VIOLACEAE land; Upper Deep Creek. Viola canadensis L. Cymopterus purpurascens (Gray) Jones R; 8800; mountains; mixed conifer woodland; O; 5000-6000; mesas and canyon bottoms; desert Upper Deep Creek shrub to pinyon-juniper woodland; Spring Canyon, Brimhall Arch, and Cedar Mesa. Viola nephrophyUa Greene R; 5500; bottomland; riparian; oxbow of the Cymopterus purpureus Wats. Fremont River. O; 4000-6000; hills, mesas, benches, and canyons; desert shrub to pinyon-juniper woodland; VISCACEAE Muley Twist Canyon, Sulphur Creek, Cedar Mesa and Halls Creek. Arceuthobium campylopodium Engelm. O; 6000-7000; mesas, upland and benches; Ligusticum porteri Coult. & Rose parasite on pinyon pines; Wagon Box Mesa and R; 8800; mountains; mixed conifer woodland; Pleasant Creek Upper Deep Creek. Arceuthobium douglasii Engelm. ex Wheeler Lomatium junceum Barneby and Holmgren R; 8500; parasite on douglas-fir; mixed conifer U; 5000-5500; upland, benches, and washes; woodland; Upper Deep Creek desert shrub to pinyon-juniper woodland; Meeks Mesa, Chimney Rock and Sheets Gulch. Arceuthobium vaginatum (H.B.K.) Eichler R; 8500; parasite on ponderosa pine; mixed Lomatium parryi (Wats.) Macbr. conifer woodland; Upper Deep Creek. R; 4000; canyon bottoms; desert shrub; Halls Creek. Phoradendron juniperinum Engelm. O; 5000-6500; parasite on junipers; pinyon- Lomatium scabrum (Coult. & Rose) Mathias juniper woodland; Hickman Bridge and R; 4000-4500; canyon bottoms; desert shrub; Cedar Mesa. Halls Creek and Muley Twist Canyon. VITACEAE Osmorhiza chilensis H. & A. R; 8800; mountains; mixed conifer woodland; Parthenocissus vitacea (Knerr) A.S. Hitchc. Upper Deep Creek. R; 4000-5500; bottomlands and canyon bottoms; riparian communities and hanging gardens; Sium suave Walt. Fremont River Valley and Halls Narrows. Uj 5000-5500; bottomland; riparian; Fremont River Valley. ZYGOPHYLLACEAE
URTICACEAE Tribulus terrestris L. O; 5500; bottomland; disturbed sites; Fruita. Urtica dioica L. U; 5000-5500; bottomland; riparian; Fremont River Valley.
75 Technical Report NPS/NAUCARE/NRTR-93/01
FLOWERING PLANTS- Carex nebrascensis Dewey MONOCOTYLEDONS O; 5500; bottomland; riparian; Fruita.
AGAVACEAE Carex occidentalis Bailey R; 6500-7000; canyon bottoms; riparian; Middle Yucca anustissima Engelm. ex Trel. Deep Creek. O; 4000-5000; upland, hills, mesas, benches, and canyon bottoms; desert shrub to pinyon-juniper Carex rossii F. Boott in Hook. woodland; Halls Creek and Muley Twist O; 6500-8000; mountains and hillsides; mixed Canyon. conifer woodland; Upper Deep Creek and Miners Mountain. Yucca harrimaniae Trel. O; 5000-6000; hills, mesas, benches, and canyons; Carex stenophylla Wahl. desert shrub to pinyon-juniper woodland; R; 8000-8500; benches of mountains; mixed Capitol Gorge, Chimney Rock, Hickman Bridge conifer woodland; Upper Deep Creek. Trail, Cohab/Frying Pan Canyons, and Grand Wash. Carex vallicola Dewey R; 8000-8500; benches of mountains; mixed ALISMACEAE conifer woodland; Upper Deep Creek.
Sagittaria cuneata Sheldon Carex xerantica Bailey R; 5500; bottomland; riparian; Fremont River R; 7000-7500; canyon bottoms; riparian; Middle Valley. Deep Creek.
COMMELINACEAE Cyperus aristatus Rottb. R; 4000; bottomland; riparian; Halls Creek. Tradescantia occidentalis (Britt.) Smyth R; 4000; canyon bottoms; riparian; Halls Creek. Eleocharis acicularis (L.) R. & S. C; 5500-6500; bottomland; riparian; Fremont CYPERACEAE River Valley and Pleasant Creek.
Carexaurea Nutt. Eleocharis palustris (L.) R. & S. W; 4000-8000; bottomland; water pockets W; 4000-6500; bottomland and canyon bottoms; (riparian); Upper Deep Creek, Cottonwood riparian; Fremont River Valley, Pleasant Creek, Tanks; Halls Narrows. Halls Narrows, and Paradise Draw.
Carex douglasii F. Boott in Hook. Eleocharis rostellata (Torr.) Torr. R; bottomland; riparian; oxbow of the Fremont R; 5500; bottomland; riparian; oxbow of the River. Fremont River.
Carex geophila Mack. Scirpus acutus Muhl. ex Bigel. U; mountains; mixed conifer woodland; Upper U; 5500; bottomland; riparian; oxbow of the Deep Creek. Fremont River.
Carex hystricina Muhl. ex Willd. Scirpus maritimus L. R; 4000; canyon bottoms; riparian; HallsNar- R; 5000-5500; bottomland riparian; Fremont rows. River Valley.
Carex lanuginosa Michx. Scirpus pungens Vahl C; 4000-5500; bottomland and canyon bottoms; C; 4000-6500; bottomland and canyon bottoms; riparian; Fruita, oxbow of the Fremont River, riparian; Fremont River Valley, Pleasant Creek, and Halls Narrows. Halls Creek, The Hamet, Muley Twist Canyon, and Sulphur Creek Carex microptera Mack. O; 6000-8000; bottomland and canyon bottoms; Scirpus validus Vahl riparian; Upper Deep Creek, Paradise Draw and R; 5000; bottomland; riparian; Oyster Shell Reef. Water Canyon.
76 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
GRAMINEAE Andropogon glomeratus (Walt.) B.S.P. R; 3900; canyon bottom; riparian; Halls Nar Agoseris glauca (Pursh) Raf. rows. R; 8000-8500; mountains; meadows; Upper Deep Creek. Andropogon hallii Hackel R; 3900; sandy benches; desert shrub; Lower Agropyron cristatum (L.) Gaertn. Halls Creek. O; 5000-6000; bottomlands; disturbed sites; Fremont Valley and Pleasant Creek. Aristida adscensionis L. R; 5500; bottomland; disturbed sites; Sulphur Agropyron dasystachyum (Hook.) Scribn. Creek U; 8500; mountains; montane meadows; Upper Deep Creek. Aristida purpurea Nutt. W; 4000-7000; disturbed sites, benches, canyon Agropyron intermedium (Host) Beauv. bottoms; desert shrub, sagebrush, and pinyon- R; 7000; canyon valley bottom; riparian; Middle juniper woodlands; widespread throughout the Deep Creek. park.
Agropyron repens (L.) Beauv. Avena sativa L. O; 5000-5500; canyon bottoms and bottomlands; R; 5500; bottomland; disturbed sites; Fremont riparian; Fruita. River Valley.
Agropyron smithii Rydb. Bauteloua barbata Lag. O; 5000-8500; canyon bottoms, bottomlands U; 4000-5500; bottomland and canyon bottoms; anddisturbed sites; Fremont Valley and Upper riparian; Halls Narrows and oxbow of the Deep Creek. Fremont River.
Agropyron spicatum (Pursh) Scribn. & Sm. Bouteloua curtipenduk (Michx.) Gray R; 8000-8500; mountains; mixed conifer wood O; 5500-6000; bottomland and canyon bottoms; land; Upper Deep Creek. disturbed sites; Frying Pan Canyon and Fruita.
Agropyron trachycaulum (Link) Make Bouteloua eriopoda (Ton.) Torr. in Whipple C; canyon bottoms and bottomlands; desert O; 4000-7000; mesas and canyon bottoms; desert shrub, sagebrush, riparian; Hickman Bridge shrub and pinyon-juniper woodlands; Lower area, Fremont River Valley, Spring Canyon and Muley Twist Canyon, Halls Narrows, and Pleasant Creek. Longleaf Flat.
Agrostis semivertitilkta (Forsskal) C. Chr. Bouteloua gracilis (H.B.K.) Lag. ex Steud. O; canyon bottoms; riparian; Hickman Bridge W; 4000-8000; mountains, benches, mesas, area, Fremont River Valley, and lower Halls canyon bottoms, and hillsides; desert shrub to Creek. mixed conifer woodland, and grassland; widespread throughout the Park. Agrostis stolonifera L. C; 5000-6000; canyon bottoms; riparian; Sulphur Bouteloua hirsuta Lag. Creek, Fremont River Valley, Muley Twist and R; 5500; bottomlands; orchards; Fruita. Deer Point. Bouteloua trifida Thurb. in Wats. Agrostis tenuis Sibth. O; 5500; bottomlands; disturbed sites; Fruita. R; 5000-6000; canyon bottoms and bottomlands; disturbed site; Spring Canyon. Bromus carinatus H. & A. R; 5500; bottomlands; saline soils (riparian); Alopecurus carolinianus oxbow of the Fremont River. R; 6000; canyon bottoms; riparian; Pleasant Creek. Bromus inermis Leyss. W; 5500-8500; bottomlands and mountains; Andropogon gerardii Vitman pinyon-juniper woodland and mixed conifer O; 5500; bottomlands; orchards and riparian woodland; Fruita and Upper Deep Creek. sites; Fruita. 77 Technical Report NPS/NAUCARE/NRTR-93/01
Bromus rubens L. Elymus junceus Fisch. O; 3900-4500; canyon bottoms; disturbed sites; O; 5500; bottomlands; riparian; Fruita. Halls Creek. Elymus salina Jones Bromus tedorum L. W; 4000-6500; hillsides, mesas, and benches; W; 3900-8000; upland, bottomlands, hillsides, desert shrub to pinyon-juniper woodland; mesas, mountains, benches, washes, canyon Redslide, Spring Canyon, Longleaf Flat, Miners bottoms; desert shrub to mixed conifer wood Mountain, Fremont River Valley, and The land; widespread throughout the park. Hartnet.
Bromus willdenowii Kunth Enneapogon desvauxii Beauv. O; 5500; bottomlands; orchards; Fruita. O; 4000-5000; upland and canyon bottoms; desert shrub; Muley Twist Canyon and Halls Calamagrostis scopulorum Jones Creek O; 5000-6000; bottomlands and canyon bottoms; hanging gardens, seeps and springs; Middle Eragrostis pectinacea (Michx.) Nees. Deep Creek, Fremont River Valley, Spring R; 4000; canyon bottoms; riparian; Halls Canyon and oxbow of the Fremont River. Narrows.
Calamagrostis stricta (Timm) Koeler Erioneuron pilosum (Buckley) Nash in Small R; 5000; canyons; hanging garden community; O; 4000-6000; washes and benches; desert shrub Spring Canyon. to pinyon-juniper woodland; Sulphur Creek, Ackland Spring, Halls Creek, and Grand Wash. Cenchrus longispinus (Hackel) Fern. O; 5500; bottomlands; disturbed sites; Fruita. Erioneuron pulchellum (H.B.K.) Tateoka O; 5000-6000; hills and benches; desert shrub to Dactylis glomerata L. pinyon-juniper woodland; Halls Creek, Fruita, O; 5000-6500; bottomlands; riparian; Fruita, and Fremont River Valley. Fremont River Valley, and Pleasant Creek. Festuca arundinacea Schreb. Dichanthelium lanuginosum (Ell.) Could var. O; 5500; bottomlands; disturbed sites; Fruita. fasciculatum (Torr.) Spellenberg R; 4000; canyons; hanging gardens; Halls Festuca ovina L. Narrows. O; 6500-8000; moist sites and mountains; pinyon-juniper woodland to mixed conifer Dichanthelium lanuginosum (Ell.) Gould var. woodland; Pleasant Creek and Upper Deep sericeum (Schmoll) Spellenberg Creek R; 4000; canyons; hanging gardens; Halls Narrows. Festuca elatior L. O; 5500; bottomlands; riparian; Fremont River Distichlis spicata (L.) Greene Valley. O; 5500-6500; bottomlands, seeps, and springs; riparian; Fruita, The Hartnet, and Pleasant Glyceria striata (Lam.) A.S. Hitchc. Creek. R; 5500; bottomlands; riparian; oxbow of the Fremont River. Echinochloa crus-galli (L.) Beauv. R; 5500; bottomlands; riparian; oxbow of the Hilaria jamesii (Torr.) Benth. Fremont River. W; 3900-7000; upland, hills, benches, mesas, and washes; desert shrub to pinyon-juniper Elymus canadensis L. woodland, and grassland; widespread O; 4000-6500; bottomlands; riparian; Fremont throughout the park. River Valley, Halls Creek, and Pleasant Creek. Hordeum glacum Steud. Elymus glaucus Buckl. O; 5500; bottomlands; disturbed sites; Fruita. O; 6500-8500; bottomlands and canyon bottoms; riparian; Upper Deep Creek and Pleasant Creek
78 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Hordeum jubatum L. Munroa squarrosa (Nutt.) Torr. in Whipple W; 5000-6500; bottomlands, canyons, mesas, and U; 4000-6000; washes; disturbed sites; Halls benches; desert shrub to pinyon-juniper Creek and Hickman Bridge. woodland; Fruita, Sheets Gulch, oxbow of the Fremont River, and Pleasant Creek. Oryzopsis hymenoides (R. & S.) Ricker in Piper W; 3900-7500; upland, bottomlands, hills, mesas, Hordeum pusillum Nutt. benches, and canyons; desert shrub to pinyon- R; 5000; hills and benches; desert shrub; Bun- juniper woodland; widespread throughout the Trail. park.
Koeleria nitida Nutt. Oryzopsis micrantha (Trin. & Rupr.) Thurb. R; 8000; mountains; mixed conifer woodland; U; 5000-6500; bottomlands, benches and canyon Upper Deep Creek. bottoms; pinyon-juniper woodland; Fruita, Pine Canyon, and Paradise Draw. Lolium perenne L. U; 5500-6000; bottomlands and washes; riparian; Paniatm capillare L. Fruita and Hickman Bridge. O; 5500; bottomlands; disturbed sites; Fruita.
Lycurus phleoides H.B.K. Panicum oligosanthes Schultes U; 4000-6500; upland and canyons; slickrock R; 5000; water pockets; riparian; Cottonwood communities; Sheets Gulch, Pleasant Creek, Tanks. Hickman Bridge, and Halls Creek. Panicum virgatum L. Muhlenbergia andtna (Nutt.) A.S. Hitchc. O; 4000; canyon bottoms; riparian; Halls W; 3900-6500; bottomlands and canyon bottoms; Narrows. water pockets (riparian); Fremont River Valley, Pleasant Creek, Cottonwood Tanks, Spring Phalaris arundinaceae L. Canyon, Capitol Gorge, Halls Creek and U; 5500; bottomlands; riparian; Fremont River Sulphur Creek. Valley.
Muhlenbergia asperifolia (Nees. & Mey.) Parodi Phleum pratense L. C; 5000-6500; bottomlands; moist disturbed sites C; 5000-5500; bottomlands; disturbed and moist (riparian); Burr Trail, Fremont River Valley, sites; Fremont River Valley. Spring Canyon, Pleasant Creek, and Sulphur Creek. Phragmites australis (Cav.) Trin. ex Steud. C; 4000-6500; bottomlands; riparian; Fremont Muhlenbergia paucifbra Buckl. River Valley, Halls Creek, and Pleasant Creek. U; 5000-6000; canyons and waterpockets; riparian; Hickman Bridge, Cohab, and Frying Poa bigbveu Vasey & Scribn. in Vasey Pan Canyons. R; 4000; canyon bottoms; riparian; Halls Narrows. Muhlenbergia pungens Thurb. in Gray W; 4000-6000; bottomlands, hills, washes, and Poa compressa L. canyon bottoms; desert shrub to pinyon-juniper O; 5000-6500; bottomlands and water pockets; woodland; Hickman Bridge, Fremont River riparian; Pleasant Creek and Cottonwood Tanks. Valley, Capitol Gorge, Muley Twist Canyon, Cohab Canyon, Grand Wash, and Halls Creek. Poa carta Rydb. R; 5000; bottomlands; riparian; Fremont River Muhlenbergia racemosa (Michx) B.S.P. Gorge. R; 6000; canyon bottoms; riparian; Cohab Canyon. Poa fendleriana (Steud.) Vasey W; 5000-8000; upland, mesas, benches, and Muhlenbergia thurberi Rydb. hillsides; desert shrub to mixed conifer wood U; 5000-6000; washes and canyon bottoms; land; widespread throughout the park. riparian and pinyon-juniper woodland; Hickman Bridge, Capitol Gorge, and Spring Canyon.
79 Technical Report NPS/NAUCARE/NRTR-93/01
Poa palustris L. Spartina gracilis Trin. R; 8000; canyon bottoms; riparian;Uppe r Deep O; bottomlands and canyon bottoms; riparian; Creek. Fremont River Valley, Halls Creek, and Spring Canyon. Poa pratensis L. C; 5000-S000; bottomlands, springs, and Sporobolus airoides (Torr.) Torr. in Parke orchards; riparian and waterpockets; Fremont W; 4000-6000; bottomlands, washes and canyon River Valley, Pleasant Creek, Hickman Bridge, bottoms; desert shrub to pinyon-juniper Willow Tanks, and Upper Deep Creek. woodland; widespread throughout the park.
Poa sandbergii Vasey Sporobolus contractus A.S. Hitchc. U; 5500-6000; upland, mesas, and benches; C; 4000-6000; upland and canyon bottoms; desert shrub; Burr Trail. desert shrub to pinyon-juniper woodland; Halls Narrows; Frying Pan Canyon, Fremont River Polypogon monospeliensis (L.) Desf. Valley, Longleaf Flat, Muley Twist Canyon, and O; 4000-6500; bottomlands; riparian; oxbow of Spring Canyon. the Fremont River, Pleasant Creek, Halls Creek, Spring Creek, and Sulphur Creek. Sporobolus cryptandrus (Torr.) Gray W; 4000-6500; uplands, bottomlands, hills, PuctineUia distorts (L.) Pari. mesas, and canyons; desert shrub to pinyon- U; 5000-6500; bottomlands and canyon bottoms; juniper woodland; widespread throughout the riparian; Cathedral Valley and Pleasant Creek. park.
PuctineUia pauciflora (Presl.) Munz Sporobolus flexuosus (Thurb.) Rydb. U; 6500; canyons; riparian; Pleasant Creek. C; 4000-6500; upland, hills, mesas, benches, and canyon bottoms; desert shrub to pinyon-juniper PuctineUia nuttalliana (Schult.) A.S. Hitchc. in woodland; Jailhouse Rock, Pleasant Creek, Halls Jeps. Narrows, and Muley Twist Canyon. U; 5000-5500; bottomlands; riparian; Fremont River Valley. Sporobolus giganteus Nash U; 4000-5000; bottomlands, hills, and canyon Schedonnardus paniculatus (Nutt.) Trel. in Banner bottoms; desert shrub; Halls Narrows and & Coville Spring Canyon. U; 5500; bottomlands; disturbed sites; Fruita. St ipa arida Jones Schizachne purpurascens (Torr.) Swallen U; 4000-5500; upland, hills, and canyon bottoms; R; 6000; canyon bottoms (moist sites); pinyon- desert shrub to pinyon-juniper woodland; Burr juniper woodland; Oak Creek Canyon. Trail, Capitol Gorge, and Halls Creek.
Schizachyrium scoparium (Michx.) Nash in Small Stipa columbiana Macoun U; 3900-6500; hills, benches, and canyon U; 5500-8000; mesas and mountains; pinyon- bottoms; desert shrub to pinyon-juniper juniper woodland to mixed conifer woodland; woodland^Halls Creek and Pleasant Creek. Frying Pan Canyon and Upper Deep Creek.
Setaria glauca (L.) Beauv. Stipa comata Trin. & Rupr. C; 5000-5500; bottomlands; disturbed sites; W; 4000-7500; mesas, benches, and upland; Fremont River Valley. desert shrub to mixed conifer woodland, and grassland; widespread throughout the park. Setaria viridis (L.) Beauv. C; 5500; bottomlands; disturbed sites; Fruita. Stipa coronata Trin. & Rupr. R; 5000-6500; hills, mesas, benches, and canyon Sitanion hystrix (Nutt.) J.G. Sm. bottoms; desert shrub to pinyon-juniper W; 4000-8000; hills, benches, canyons bottoms; woodland; Deer Point and Capitol Gorge. upland, mountains, bottomlands, and washes; desert shrub to mixed conifer woodland; Stipa neomexicana (Thurb.) Scribn. widespread throughout the park. R; 5000-6000; canyon bottoms, benches, and mesas; desert shrub to pinyon-juniper wood land; Deer Point and Hickman Bridge. 80 Vascular Flora and Vegetation of Capitol Reef National Park, Utah
Stipa pinetorum Jones Junais ensifolius Wikstr. var. brunnescens (Rydb.) U; 6500; hills and mesas; pinyon-juniper Cronq. woodland; The Hartnet. O; 5500-6500; bottomlands and canyon bottoms; riparian; Pleasant Creek, Sheets Gulch, and Stipa scribneri Vasey Spring Canyon. U; 5000-8000; upland, mountains, mesas, benches, and canyons; desert shrub to mixed Juncus ensifolius Wikstr. var. montanus (Engelm.) conifer woodland; Upper Deep Creek, Hickman C.L. Hitchc. Bridge, Pleasant Creek, and Muley Twist O; 5500-6500; bottomlands; riparian; oxbow of Canyon. the Fremont River and Pleasant Creek.
Stipa speciosa Trin. & Rupr. Juncus longistylis Torr. in Emory C; 4000-6000; upland and canyon bottoms; O; 5500-6500; bottomlands and canyon bottoms; desert shrub to pinyon-juniper woodland; riparian; Pleasant Creek and the Fremont River Hickman Bridge, Halls Creek, Fremont River Valley. Valley, Spring Canyon, Sulphur Creek, and Grand Wash. Juncus torreyi Cov. O; 5500-6500; bottomlands and canyon bottoms; Stiporyzopsis bloomeri (Boland) B.L. Johnson riparian; Pleasant Creek and the Fremont River R; 5000-5500; canyon bottoms; pinyon-juniper Valley. woodland; Fremont River Valley. JUNCAGINACEAE Vulpia octoflora (Walt.) Britt. O; 4000-5000; canyon bottoms and upland; Triglochin maritima L. desert shrub; Halls Creek and Muley Twist O; 5000-6000; bottomlands; riparian; oxbow of Canyon. the Fremont River, The Hartnet, and Sulphur Creek. ZeamaysL. ULIEACEAE O; 5500; bottomlands; disturbed sites; Fruita. Allium cernuum Roth IRIDACEAE O; 6000-8800; mountains and canyons; mixed conifer and aspen woodlands; Pleasant Creek, Iris missouriensis Nutt. Oak Creek and Upper Deep Creek. R; 5500; bottomlands; riparian; Fremont River Valley. Allium macropetalum Rydb. Sisyrinchium demissum Greene U; 4000-5000; benches; desert shrub to pinyon- U; 5500-6500; bottomlands (saline); riparian; juniper woodland; Burr Trail and Upper Halls Fremont River Valley and Pleasant Creek. Creek.
JUNCACEAE Androstephium breviflorum Wats. R; 4000; benches; desert shrub; Lower Halls Juncus alpinus Vill. Creek. O; 6000-6500; bottomlands; riparian; Pleasant Creek. Asparagus officinalis L. O; 5500; along irrigation ditiches; orchards; Juncus baltiais Willd. var. mexiamus Kuntze Fruita. W; 4000-8000; bottomlands and canyon bottoms; riparian; Upper Deep Creek, Pleasant Creek, Calochortus aureus Wats. and Halls Narrows. R; 5000-5500; benches; desert shrub; Oyster Shell Reef. ]unais balticus Willd. var. montanus Engelm. R; 6000-6500; canyon bottoms; riparian; Pleasant Calochortus nuttallii T. & G. in Beckwith Creek. O; 6000-8000; mesas and hillsides; pinyon- juniper woodland and mixed conifer woodland; Juncus bufonius L. Upper Deep Creek and Cedar Mesa. O; 5500; bottomlands and canyon bottoms; riparian; Fremont River Valley. 81 Technical Report NPS/NAUCARE/NRTR-93/01
Eremoaynum albomarginatum Habenaria sparsiflora Wats. R; 3900; bottomlands; desert shrub; Halls R; 8000; bottomlands and canyon bottoms; Creek. riparian; Upper Deep Creek.
Fritillaria atropurpurea Nutt. Habenaria zothecina Higgins and Welsh R; 8500-8800; mountains; mixed conifer R; 4000; canyon bottoms; riparian; Halls Narrows. woodland and aspen woodlands; Upper Deep Creek. Spiranthes diluvwlis Shev. R; 5500; bottomlands; riparian; Fremont River Smilacina stellata (L.) Desf. Valley. O; 4000-8000; canyon bottoms; riparian; Fremont River Valley, Upper Deep Creek, and Halls TYPHACEAE Creek. Typha domingensis Pers. Zigadenus elegans Pursh O; 5000-5500; bottomlands; riparian; Fremont U; 8000; mountains, riparian; Upper Deep River Valley. Creek. Typha latifolia L. ORCHIDACEAE C; 4000-6500; bottomlands and canyon bottoms; riparian; Fremont River Valley, Pleasant Creek, Corallorhiza maculate Raf. and Halls Creek. R; 8800; mountains; mixed conifer and aspen woodlands; Upper Deep Creek. ZANNICHELLIACEAE
Empactis gigantea Dougl. ex Hook. Zannichellia palustris L. O; 4000-5500; canyon bottoms; riparian; Halls O; 5500; ponds; aquatic; oxbow of the Fremont Narrows and the Fremont River Valley. River.
82 As the nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural and cultural resources. This includes fostering of our land and water resources, protecting our fish and wildlife, preserving the environmental and cultural values of our national parks and historical places, and providing for enjoyment of life through outdoor recreation. The department assesses our energy and mineral resources and works to ensure that their development is in the best interests of all our people. The department also promotes the goals of the Take Pride in America campaign by encouraging stewardship and citizen responsibility for the public lands and promoting citizen participation in their care. The department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.
NPS D-55 September 1993