The Rocky Mountain Herbarium, Associated Floristic Inventory,

and the Flora of the Rocky Mountains Project

Ronald L. Hartman, Curator

Rocky Mountain Herbarium

Department of Botany

University of Wyoming

Laramie, WY 82071-3165 U.S.A.

ABSTRACT -- The growth of, and improvements to, the Rocky Mountain Herbarium (RM, RMS, USFS) are reviewed with emphasis on the goals of the program. The intensive and systematic floristic inventories carried out in recent years by the RM in Wyoming and adjacent states are discussed, as are plans to complete the survey of the vascular plants of the region. Finally, an update is provided on the Flora of the Rocky Mountains project, including interactions with other similar projects, circumscription of the area covered, contents (particularly of the first volume), and contributions by specialists. To this end, the Rocky Mountain Flora Association is being established to coordinate the inventories and the data basing of specimens and to help with the preparation of the taxonomic treatments.

INTRODUCTION

I was asked to present a talk on the Flora of the Rocky Mountains project, but felt it necessary to expand the topic to include discussions on our recent floristic inventories in Wyoming, Idaho, Colorado, and New Mexico and the significant changes which have occurred at the Rocky Mountain Herbarium (RM) over the past 15 years. Thus a more complete picture emerges of project's origin, resources, and goals. Although the flora of Idaho is touched on but briefly, I feel that the inclusion of the last two topics may provide helpful information to those involved in similar work in that state and elsewhere.

THE ROCKY MOUNTAIN HERBARIUM

What is now called the Rocky Mountain Herbarium (RM) was founded by Aven Nelson in 1893. It was so designated by the Board of Trustees of the University of Wyoming (UW) in 1899, following Nelson's successful botanical exploration of Yellowstone National Park. Since 1960, it has been housed on the third floor of the Aven Nelson Building. This edifice was built in 1923 for the University Library and School of Law following Nelson's tenure as UW president (1917-1922). For a detailed history of the Herbarium, mostly prior to the mid-1950s, the reader is referred to the excellent biography Aven Nelson of Wyoming by Roger L. Williams (1984). Aven Nelson was Curator of the Herbarium for nearly 50 years (1893-1942). His was succeeded in this position by Cedric Lambert Porter (1942-1968), John R. Reeder (1968-1976), and Ronald L. Hartman (1977-present). The following discussion will focus primarily on the past 15 years, the period of my tenure as Curator.

The staff of the RM includes myself (nine-month contract; 30% time as Curator), B. Ernie Nelson, Herbarium Manager since 1974, who does an admirable job of filling a crucial role in the daily operations (full-time), and part-time help. We normally employ three to six students per semester working 10-20 hours per week. They are involved in various aspects of processing specimens including label typing (and data basing), the mounting of specimens (we average about 13,000 per year), accessioning, updating the Wyoming dot maps, filing, and the processing of loans and exchanges.

Over the years, the RM has grown gradually through the efforts of staff and graduate students in obtaining specimens from throughout the Rockies. Major spurts came with the integration of the George E. Osterhout (20,000 sheets in 1937), the Hapeman (30,000, 1951), the Wilhelm G. Solheim Mycological (RMS, 48,000, 1978), and the U. S. Forest Service [National] (USFS, 120,000, 1982) herbaria. In 1978, an intensive and systematic floristic inventory of Wyoming and other Rocky Mountain states was initiated. To date the program has obtained 192,000 numbered collections and possibly an equal number of duplicates for exchange. Acquisition of specimens through the inventory and from the RMS and USFS has more than doubled the size of the Herbarium (Fig. 1).

The RM is the largest facility of its kind between the University of Minnesota (St. Paul), Missouri Botanical Garden (St. Louis), and the University of Texas (Austin) to the east and the University of California (Berkeley), the California Academy of Sciences (San Francisco), and Rancho Santa Ana Botanic Garden (Claremont, CA) to the west. It ranks 18th in size based on number of accessions of over 630 herbaria listed in Holmgren, Holmgren, and Barnett (1990) for the United States; 8th in size for a state university. If one includes either the number of specimens in our backlog (70-80,000) or the number of specimens in the Range Management Herbarium (WYAC; 60,000) of the College of Agriculture, we would rank 16th overall or 7th for a state university. The combined herbaria (RM, RMS, USFS, WYAC) plus the backlog total 750,000 specimens. The WYAC includes the 50,000 accession, A. A. Beetle Grass Collection, with a good representation of graminoids from throughout the World.

In order to house the expanding holdings of the RM, a National Science Foundation facilities grant ($238,859, to R. L. Hartman and Meredith A. Lane, Acting Curator 1985-86) was obtained in 1986 for the purchase of a manual-assist SpaceSaver mobile storage system and 100 new cabinets (since 1977, 26 new cases have been provided by UW and 80 old cases were acquired with the USFS). The system consists of two modules, one with eight movable and three stationary rows (holding 230 standard herbarium cases), the other with five movable and two stationary rows (holding 122 cabinets). This system has increased by 60-70% the storage capacity on the south half of the Herbarium. There are an additional 83 cabinets not in the system, the majority of which are in the north half of the RM. The NSF grant also provided funds to hire a number of undergraduates to facilitate the move, reorganize the collection, and help with the backlog. The reorganization consisted of the following: accessioning and intercalation of the USFS into the RM, replacement of many of the genus covers, segregation of Rocky Mountain area specimens (yellow folders; excluding Wyoming's red folders) from extra-regional sheets (manila folders), and reordering of plant families from the antiquated Engler-Prantl system to one which is alphabetical within major groupings (algae, fungi, mosses/liverworts, lichens, ferns/fern allies, gymnosperms, monocots, and dicots). This reorganization and the processing of new acquisitions has also been facilitated by new funding from the U. S. Forest Service's Rocky Mountain Range and Experiment Station (since 1982) and the College of Arts and Sciences at UW (since 1986).

Despite the growth of the Herbarium, there is still ample work space for the processing of specimens, plant identification, herbarium research (seven work areas each with a dissecting microscope), for the geographic information system (GIS), and for ancillary items (extensive microfiche, reprint, map, and gazetteer collections, library, light table, three microcomputers with printers, literature, type, and Wyoming dot map card files, etc.). A Wyoming reference collection (containing a representative specimen of each taxon of vascular plant known to occur in the state, initially assembled in 1978) is conveniently located to facilitate identification, thus reducing handling of the research material.

Associated with the reference collection are two Wyoming Taxon Checklists (initially compiled in 1979), one of which contains relevant synonymy. Through two cost-share agreements with the national office of the Soil Conservation Service (SCS), a database containing a revised checklist was completed last fall. To each taxon, a literature citation is attached. This has been incorporated into the Plant List of Accepted Nomenclature, Taxonomy, and Symbols (PLANTS) of the SCS. As a consequence of this project, a taxonomic literature database of nearly 3,000 citations pertaining to the flora of Wyoming and the Rockies has been established. The database greatly facilitates plant identification and revisionist work.

A number of steps have been taken to make the processing of new acquisitions as efficient and reliable as possible. The greatest improvement has been in the production of specimen labels, a procedure computerized since 1979 (using the UW mainframe computer). We now use a Zenith 248 with 120 MB hard drive and a Gateway 2000, 486/33C, 8 MB RAM, 500 MB hard drive with a HP LaserJet III printer. We have used three label programs, each of which also captured five categories of data for archiving and eventually data basing. We now use PLabel by Kent D. Perkins (Herbarium, University of Florida) which places all of the data into dBase IV for the Wyoming Specimen Database and produces labels on the LaserJet. A species name dictionary increases both the efficiency and accuracy of data entry. We recently captured data from nearly 14,000 specimens related to a floristic inventory of the west slope of the Wind River Range using this system. We now have a cost-share agreement with the state and national offices of the Bureau of Land Management and the SCS to initiate the capture of specimen label data from the Wyoming sheets in the RM.

Thus far data from more than 80,000 Wyoming collections have been entered into the database. We are also coordinating the project with other groups such as the Specimen Management System for California Herbaria (SMASCH; Tom Duncan), the Intermountain Database Working Group (Mary Barkworth), and the Southeastern Regional Floral Information System (SERFIS; Robert R. Haynes) to establish standards to facilitate the exchange of data.

In 1986, the RM Library was officially recognized as a branch of the UW Libraries. This has greatly facilitated the acquisition of needed taxonomic literature. Also, with the purchase of standard library shelving in 1988, the collection is now consolidated with room for expansion. The RM Library traditionally has been strong in systematic literature for North America. Through the efforts of the previous curator, microfiche editions of many of the important older European works and herbaria were obtained. This microfiche collection continues to grow and includes the herbaria of Candolle, Humboldt, Bonpland, and Kunth, Lamarck, Lindley (orchids), Linnaeus, Michaux, Rousseau-Aublet, and Willdenow; the correspondence of Linnaeus; seven botanical journals; selected works of over ninety important 18th and 19th century plant taxonomists (the above from Inter-Documentation Company, the remainder from Meckler Publishing); the type collections of the New York Botanical Garden, the Academy of Natural Sciences of Philadelphia, and the United States National Herbarium; and the Plant Taxonomic Literature Microfiche Collection with over 5,000 titles, most pre-20th Century.

Associated with the Library, but property of the RM, is an extensive reprint collection. It is now housed (tightly) in seven, 4-drawer file cabinets.

The type specimens have been segregated for several decades. The total now stands at over 5,000 (including holotypes, isotypes, lectotypes, and paratypes, categories originally separated by C. L. Porter). The RM Type Register, developed initially by John and Charlotte Reeder and myself (while a M.S. degree student) between 1969 and 1971, lists the types alphabetically by basionym and contains label information and literature citations. It has now been entered into a dBase IV file so that requests for information can be filled quickly and so it can be included in the proposed national database of types.

The RM exchange program has been expanded considerably in recent years and now includes 55 domestic and 15 international institutions. The average number of specimens sent on and received from exchange annually has increased from 1,200 and 1,400, respectively, for the period 1962 through 1976 to 3,900 and 3,500 for the period 1977 through 1992. The goal of this program is to obtain representative material, in order of priority, from: the Rocky Mountains, western North America, eastern North America, and arctic, alpine, and temperate regions elsewhere in the Northern Hemisphere. Such programs are extremely important in obtaining material for research from throughout the range of a taxon. Relatively few of the estimated 4,300 species in the Rocky Mountains are restricted to the region and many are found throughout much of the Northern Hemisphere. Plant families of special interest (and staff and associates who specialize in each) include the Apiaceae (Hartman), Asteraceae (Hartman and G. K. Brown, Associate Professor of Botany), Bromeliaceae (Brown), Caryophyllaceae (Hartman), Cyperaceae and Poaceae (A. A. Beetle, Professor Emeritus of Range Management), and Salicaceae (R. D. Dorn, RM Associate). Dr. Steven L. Miller (Assistant Professor of Botany; mycology) specializes in Basidiomycetes.

An additional method which helps accomplish the above goal is the interinstitutional loan of specimens. At any one time, 10,000 to 15,000 sheets from the RM are under study at herbaria and museums throughout the World. Likewise, we borrow thousands of sheets annually for our own research.

Another source of specimens has been gifts, either outright (e.g., private herbaria of Osterhout, Hapeman, and Solheim; R. D. Dorn, Cheyenne) or in exchange for identifying a set of duplicates (e.g., Bureau of Land Management; National Park Service; U. S. Fish and Wildlife Service; U. S. Forest Service; Wyoming Game and Fish; E. Evert, Park Ridge, IL, Roger L. Williams, Laramie). It is imperative that all acquisitions be of high quality and accompanied by adequate data. The last major source of specimens for the RM will be covered under the next major heading.

Associated with the RM is a molecular systematics laboratory which was established by Greg Brown and myself. Greg and his students have study isozyme variation and restriction fragment length polymorphisms in chloroplast DNA to address evolution in the Bromeliaceae (funded by the National Science Foundation), the Apiaceae, and the Asteraceae.

Housed at the Herbarium is the Wyoming Natural Diversity Data Base (WYNDD) of The Nature Conservancy (TNC). The five full-time employees include a botanist Walter Fertig; (Hollis Marriott, who held this position from 1985 to 1992, is now with the Wyoming Field Office, TNC), two ecologists George Jones, (replacing Hollis as director), and Gillian Walford, a zoologist Chris Garber with an office  in the Department of Zoology and Physiology, and database manager Mary Neighbours. Our association with this organization has been mutually beneficial in many ways, especially in obtaining cost-share agreements with federal agencies. As the issue of "endangered species" so often cannot be addressed until a careful inventory has been completed, agencies frequently are referred to the Herbarium and the projects are usually done as Master's theses (see below). Data on sensitive taxa collected during a project are then incorporated in WYNDD or similar databases in other states in which we do inventories. We also serve in an advisory roll as to what plant taxa, and rank, should be included in lists of special concern for Wyoming and Colorado.

The Curator chaired a committee on the implementation of a geographic information system for research at UW. Initial funding has enabled the establishment of four nodes (Botany, Geography and Recreation, Geology and Geophysics, and Zoology and Physiology) on the campus ethernet. Housed in the Herbarium is a Sun SPARCstation 2 with 16 MB RAM, 414 MB of storage, a Calcomp 36 X 48" digitizer, ARC/INFO and GRASS software, and access to a 36", eight-pen plotter, a 6.6 GB read/write optical disk jukebox, a tape backup, and a CD reader. Recent additions to the GIS network include five Sun SPARCstations (2 Botany--Ecology, 1 Geography, 2 Wyoming Water Center [WWC]), a Sun file server [WWC], a Howtek Scanner, 1,200 dpi, 12 H 18" (Geography), and a 55 GB optical disk jukebox with two drives [WWC]. Since nearly all of these items were purchased with matching funds from the state, they are available within reason for use by all on the system.

In order to provide instruction in the applications of computers to field and museum research, William J. Gribb (Geography) and I have just received NSF funding ($124,000 including UW match) to establish the Digital Earth Sciences Laboratory (primarily for Botany, Geography and Recreation, Geology and Geophysics, Plant, Soil, and Insect Sciences, Range Management, and Zoology and Physiology). It will have ten PCs (486/66 EISA with 500 MB drives), a 35 mm film color scanner, a color printer, an 8 mm video camcorder, a Global Positioning System community base station (to be housed under cooperative agreement at the Casper BLM office for use by UW and federal agencies in differential GPS throughout Wyoming), a 6-channel field GPS with barcode data logger, three GPS units for recording way stations, a micro-meteorological station with data logger, 2 digital image projectors, an audio-to-digital converter, and software (dBase IV, SPSS-PC, PC/ARC/INFO, MIPS, ERDAS, GRASS, Geo-Link, Delta Classification System, Pankey, Tropicos, Mecca, ImageQuery, etc.). It will also house parts of the GIS research network mentioned above and have access to other hardware and software on the network. This facility will be used in a number of courses and for various purposes. In addition to its use for teaching GIS, GPS, and remote sensing, other applications for undergraduate and graduate courses in plant systematics include the use of available software for generating multiple-access keys, dichotomous keys, parallel descriptions, the data basing of label data for monographic work, etc.

As a member of the Wyoming GIS Steering Committee, the Wyoming GIS Users Group, and the State Mapping Advisory Committee, and through interaction with the Rocky Mountain Mapping Center of the U.S. Geological Survey, Denver, I am interacting with state and federal agencies to assist in the building of a state GIS with a minimum duplication of effort. For example, computerized data in the RM will serve as a layer on species distributions, and the mapping of plant communities in Wyoming and Colorado through remote sensing (William A. Reiners and associates, Botany) will provide other layers. Data from the former will help in the "ground" truthing of the vegetation maps.

RECENT FLORISTIC INVENTORIES

IN THE ROCKY MOUNTAINS

As mentioned above, in 1978 we initiated an intensive and systematic floristic inventory of Wyoming and other Rocky Mountain states with the following goals: 1) document the flora of the Rockies; 2) determine taxa truly in need of protection (many candidates for status as Endangered or Threatened, as well as recent novelties, have proven to be relatively common although often restricted); 3) provide data for the Flora of North America project (the Curator is a regional coordinator), and for regional (the Curator is a member of the Great Plains Flora Association; Hartman 1986a, 1986b, 1986c), state (we share data freely with Dorn for revisions of Vascular Plants of Wyoming) and local floras; 4) provide data for monographic and revisionary studies and for research in plant geography; and 5) computerize data from specimen labels.

During 12 of the past 15 summers, this inventory by the RM staff, graduate students, and associates (Dorn, Evert, Williams, WYNDD, etc.) has amassed 192,000 numbered collections, not including duplicates for exchange. While many have been talking about a National Biological Survey, we have accomplished a great deal towards that end for many areas in the Rockies with relatively little funding. Such inventories provide a wealth of information on the geographical and ecological distribution of native species as well as introductions, which often have great economic impact on agriculture (the Herbarium Manager is coauthor of Weeds of the West, Whitson et al. 1991). Representative material collected in a systematic manner also documents morphological variation or, of equal importance, uniformity throughout this portion of the geographical range of each species. Thus, floristic work is invaluable in obtaining research material for monographers who might otherwise be unable to acquire samples from remote and often road less areas. It also provides excellent taxonomic training at the master's degree level prior to specialization in a doctoral program or employment in areas of natural resource management (e.g., as botanists with Natural Heritage Programs and state and federal agencies).

The various floristic inventories which have been conducted in recent years, and one which is anticipated, through the RM are discussed below (Fig. 2). But first I will cover methodology.

During the first summer of a thesis project, Ernie and I alternate working with the student during three, sometimes four, two-week periods. This insures that our standards, as well as continuity with previous studies, are maintained. During the second field season, each of us may spend one two-week period with the student. Some of the projects have not been associated with Masters theses, but have been done solely by Ernie and myself.

Three things we request when doing an inventory are work space, access to a freezer for ice (made in one-gallon plastic milk jugs), and access to a shower. Because of the scale of the projects, it is imperative that we have a work center for the pressing and drying of specimens. We have been fortunate in nearly all of our studies to have had space and electricity supplied by state or federal agencies (UW Agricultural Extension Service, U. S. Forest Service, Bureau of Land Management, National Park Service, etc.).

In topographic basins, our general mode of operation is to choose an area which has not been surveyed. We drive to the site and collect all taxa of vascular plants encountered in flower and/or fruit, covering as many plant communities as possible, and often hiking several miles. Over the next ten miles, we look for new and diverse habitats, often stopping one to three times, to obtain material of species not collected at the first or subsequent sites. We then repeat the process over the next ten-mile segment of road, or one elsewhere in the basin. In part, the goal is to sample at frequent intervals in order to capture the diversity of taxa in the project area. But more so, we attempt to document morphological variation, or lack there of, and ecological and geographical distribution at a relatively fine scale. The samples are placed in plastic bags by location and habitat, stored on ice, and field data are recorded. Upon return to the work center (often late at night), the worker(s) showers, gets a good nights sleep (usually in personal vehicles), and then spends the next day pressing the material, again often late into the night. Of utmost importance is placing the plants into the ice chest (100 quart capacity) within a reasonable period of time. In montane areas, it is sufficiently cool to keep plant material fresh, stored in a backpack, for most of the day.

In mountainous regions (due to greater diversity, to being on foot with backpack, and to the significant elevational relief), the interval for repeating the collecting cycle is about five miles and often there are many more stops along the way. Following each trip, the route traveled and date are marked on a U. S. Geological Survey 7.5 minute sheet as well as a small-scale map of the area. The latter helps in planning trips as the season progresses.

Collecting tools have varied, but due to the hard and rocky substrates frequently encountered we mostly use a bricklayer's hammer with its chisel end. In the alpine we may employ a sturdy screw driver or a pocket knife.

Our plant driers have become more streamlined through the years but basically consists of a box 6' long, 20" wide, and 24" high built from 1/4" plywood. Two small door hinges, with removable pins, on each inside corner make it collapsible. Vent holes (1-1/4" in diameter) along the bottom on each side allow for the circulation of air. A frame covered with screen (1/4" mesh) at 20" above the floor prevents debris from falling into the box. Five sockets with 150 watt incandescent bulbs spaced along the floor provide the heat. End boards for plant presses are cut from 1/4" plywood; straps from 1/4 or 3/8" cotton sash cord (16.7' long; each with a loop tied at one end). Double-faced corrugates are purchased commercially.

Generally the collecting/drying cycle is 48 hrs. Plants obtained one day are placed on the drier the next and remain there for about 36 hrs. In our arid climate, most species dry within one cycle. What is not dry (e.g., conifers for which cones are stored separately in small paper bags numbered correspondingly; cacti and other succulents) is returned to the press. The dried plants in newspaper are placed in 6-8" stacks between two corrugates and tied with heavy string. They are stacked for eventual transport to the RM. Upon concluding a study, a map showing the collecting locations (e.g., Fig. 3), often a list of sites (by township, range, and section), and a species checklist are prepared and made available to individuals, often speciality collectors, interested in doing further fieldwork.

When working on federal lands, a set of specimens containing one of each taxon, for which we have a duplicate, is given to the agency for a reference collection. These are supplied in newspaper with labels, although we will mount them if funding for supplies and labor is provided. Also, material from a project area is made available through our exchange program to appropriate colleges and universities in the area (e.g., when collecting in Colorado, we exchange with COLO, COCO, CS, and Mesa College, Grand Junction, the last too small [4,000 specimens] to be included in Holmgren et al. 1990).

Except for our first project (Powder River Basin), funding has been low and often from several sources. These include contributions from individuals (the late Louis O. Williams and his wife Terua, J. Vickers Brown, Roger L. Williams, R. L. Hartman, Orval C. Harrison, John F. Freeman, Ron Schreibeis, Jean Oxley, and Brenda Schladweiler), small grants and scholarships (The Nature Conservancy, the Colorado Natural Heritage Program, Colorado Native Plant Society, Wyoming Native Plant Society, Paul Stock Scholarship, Payson Scholarship, Devils Tower Natural History Association, the U. S. Forest Service, and the Bureau of Land Management), and donation of time and resources by graduate students, staff, and associates). We feel fortunate to raise $2,500 to $3,000 for a student per summer to cover gas and food (as mentioned above, we sleep in our own vehicles although the Forest Service occasionally has provided a cabin or house trailer).

Black Hills (Wyoming; Fig. 2, area 1)

The Black Hills have long been of interest to botanists (Dorn 1977b), but much of the work has centered in the South Dakota portion (two-thirds of the area). The Wyoming segment covers 2,500 sq mi and includes the Bear Lodge Mountains (Warren Peaks, 6,600 ft), Devil's Tower, and portions of the Hogback Rim, Red Valley, Minnelusa Foothills, and the Limestone Plateau. The floristic affinities lie largely with the Rocky Mountains and, to a lesser extent, the Great Plains. Mostly during 1983 and 1984, over 12,000 collections (an average of 4.8 specimens/sq mi) were obtained. The documented flora now contains 955 taxa, an increase of 66%. This number includes 14 additions to the vascular flora of Wyoming and 62 new or clarified records for the Black Hills as a whole (Hartman and Marriott 1980, Marriott 1985, 1986).

Powder River Basin (Wyoming; Fig. 2, area 2)

Located in northeastern Wyoming, the Basin is defined as the region underlain by vast coal deposits of Tertiary and Upper Cretaceous age. The areal extent is nearly 18,000 sq mi, equivalent in size to Vermont and Massachusetts. This region, which has undergone rapid energy development, had been little known floristically. Approximately 12,500 collections (0.7/sq mi) were obtain during 1978 and 1979, thanks to funding by the Rocky Mountain Institute of Energy and Environment, UW. Of the 900 taxa of vascular plants documented, 281 were reported as new to the Basin (a 31% increase). Furthermore, eleven species were found to be new to the flora of Wyoming. These data were provided to the Flora of the Great Plains project. The region contains largely a mixture of Great Plains and Great Basin elements with scattered islands of species on buttes (especially the Pumpkin Buttes) and ridges having montane affinity (Dueholm and Hartman 1981, Hartman and Dueholm 1979a, 1979b, Hartman, Dueholm, and Nelson 1985, Hartman et al. 1980).

Southern Powder River Basin/Southeastern Plains (Fig. 2, area 3)

Although the Powder River Basin (PRB) was inventoried in the late 1970s, the southern portion needs additional work. The Rawlins District of the Bureau of Land Management is funding a project covering the northern 60 to 70% of Natrona and Converse counties and all of Niobrara County (PRB) as well as Platte and Goshen counties (southeastern plains). We will also include Laramie County (plains) in this thesis area. Hopefully the fieldwork will be done in 1993 and 1994.

Big Horn Mountains (Fig. 2, area 4)

This crescent-shaped range (3,600/sq mi) has a maximum elevation of 13,175 ft (Cloud Peak). To the northwest it is isolated from the Pryor Mountains (Montana) by the Big Horn Canyon, while to the southwest a depression separates it from the Bridger Mountains. Knowledge of the flora has accumulated over the years through the efforts of numerous collectors. Nevertheless, vast areas of the Big Horns have remained unexplored botanically. During 1979 and 1980, in excess of 8,000 collections (2.2/sq mi) were obtained which, along with earlier accessions at RM, provide a fairly detailed coverage. Over 1,100 taxa are now documented from the Big Horns, an increase of 46% (Nelson and Hartman 1983, 1984). Four species, Aquilegia jonesii Parry, Erigeron allocotus Blake, Penstemon caryi Pennell, and Sullivantia hapemanii (Coult. and Fish.) Coult., previously considered rare were found to be fairly common although often restricted to specific substrates. Of special relevance is limestone and dolomite which constitute a large proportion of the sedimentary strata. Cymopterus williamsii Hartman and Constance (1985), a previously undescribed taxon, was found throughout the southern half of the range.

Bighorn Basin (Fig. 2, area 5)

The Basin covers 7,300 sq mi, ranging in elevation from 3,640 to 7,000 ft (8,123 ft on Heart Mountain). Very little floristic work had been done prior to this decade. Since 1980, more than 9,000 collections (1.2/sq mi) have been obtained. Interestingly, many of the taxa in common with the Powder River Basin bloom up to one month earlier in the Bighorn Basin, presumably due to the early onset of summer drought. The known flora now stands at 687 taxa, an increase of 108%. During this period, the following taxa were described from the area as new to science: Astragalus jejunus Wats. var. articulatus Dorn (1988) and (see discussions below) Antennaria aromatica, Cymopterus evertii, Lomatium attenuatum, and Shoshonea pulvinata. An additional novelty in Cymopterus remains to be described (Hartman 1987). The following sensitive species were also found: Eriogonum brevicaule Nutt. var. canum (Stokes) Dorn, Kelseya uniflora (Wats.) Rydb., Rorippa calycina (Engelm.) Rydb., Stanleya tomentosa Parry, Sullivantia hapemanii, Townsendia nuttallii Dorn, and T. spathulata Nutt. (Nelson and Hartman 1990).

Owl Creek/Bridger Mountains (Fig. 2, area 6)

An inventory began, primarily in 1991 on these east-west trending ranges (1,800 sq mi), approximately half of which is on the Wind River Indian Reservation. The relief varies from 5,000 to 9,684 ft (Phlox Mountain) in the Owl Creeks and to 8,272 ft (Copper Mountain) in the Bridgers. The two are bisected by the Wind River Canyon with a depth of up to 2,240 ft. Being relatively dry and composed largely of limestone and other sedimentary strata, this area is of special interest as a potential refuge of novelties. Furthermore, these ranges may have served as a migratory route between the Absarokas and the Big Horns. About 2,000 specimens have been obtained (by W. Fertig, Hartman, R. Jones, H. Marriott, and Nelson; 1.1/sq mi) but much work remains to be done, especially on the Reservation. Thus far only a small portion of the material has been identified. Species of note include Cryptantha subcapitata, Shoshonea pulvinata, and Townsendia nuttallii (see discussions below).

Wind River Basin (Fig. 2, area 7)

This Basin, excluding the Wind River Indian Reservation, was covered during 1985 and 1986 (Haines 1988), although some work was also done by June Haines in the late 1970s and early 1980s. Specimens from the earlier period are deposited at Central Wyoming College, Riverton. Prominent features of this xeric basin include Beaver Rim, Gas Hills, Rattlesnake Hills, and the Granite Mountains. Except for the last area, much of it is composed of shales, mudstones, sandstones, and derivatives. A total of 6,600 collections have been obtained from this 5,000 sq mi area (1.3/sq mi). Despite severe drought conditions during the thesis study, 885 taxa (including previous holdings in RM) were documented (79% increase). Over 214 first county records were obtained. Further work is needed during years of favorable growth. Recently described taxa from the Basin, some of which were collected in the course of the inventory, include: Cryptantha subcapitata Dorn and Lichvar (1981a), Phlox pungens Dorn (1988), Physaria eburniflora Rollins (1981), P. saximontana Rollins (1984), Trifolium barnebyi (Isely) Dorn and Lichvar (1981b; originally as T. haydenii Porter var. barnebyi Isely 1980), Cirsum sp. (Dorn, unpubl.), and Yermo xanthocephalus Dorn (1991). The last genus was new to science.

Northern Absarokas (Fig. 2, area 8)

The Absarokas were divided into four study areas: the Northern, the Southeastern, the Southwestern, and also the North Fork of the Shoshone River Drainage. The range is composed predominately of volcaniclastics, largely andesite, and is one of the most rugged in North America (Fenneman 1931). Along the eastern margin, sedimentary strata, especially limestone, are exposed.

During 1985 and parts of 1988 and 1989, 6,500 collections were made throughout the Northern Absarokas (900 sq mi; 7.2/sq mi). This study was prompted by a visit in 1984 of the Wyoming Native Plant Society led by E. F. Evert to the calcareous bog at the base of Cathedral Cliffs near Crandall Ranger Station. During a four hour reconnaissance, nine species new to Wyoming were encountered, one of which was new to the conterminous United States (Evert et al. 1986). Some additional records were discovered in subsequent years (Snow, Nelson, and Hartman 1990) and a total of 748 taxa were documented, but the bog remains unique in the state in the number of unreported taxa it harbored. Prominent features include alpine expanses such as Hurricane Mesa (11,010 ft), Trout Peak (12,244 ft), Black Mountain (11,562 ft), Dead Indian Mountain (12,216 ft), and the divide along the eastern border of Yellowstone National Park. As much of the area is road less and remote, at least one more summer will be needed to complete the inventory.

North Fork Shoshone River Drainage (Fig. 2, area 9)

This drainage arises near the east entrance of Yellowstone National Park and separates the Northern from the Southeastern and Southwestern Absarokas. Erwin F. Evert has been seriously inventorying the area since about 1978. It is undoubtedly the most thoroughly studied in the state with more than 12,000 collections from 800 sq mi (an average of 15/sq mi). Approximately 1,070 taxa have been documented representing an increase of over 500% in the known flora (Evert 1982, 1991, Evert and Hartman 1984). Included are more than 24 species new to Wyoming and the following taxa which were new to science: Antennaria aromatica Evert (1984a), Carex luzulina Olney var. atropurpurea Dorn (1988), Lomatium attenuatum Evert (1983), Penstemon absarokensis Evert (1984b), and Shoshonea pulvinata Evert and Constance (1982). The last genus also was new to science.

Southeastern Absarokas (Fig. 2, area 10)

This portion of the highly dissected volcanic plateau ranges in elevation from 7,500 to 13,148 ft (Francs Peak). During 1983 and 1984, over 10,500 collections were obtained in this essentially road less area of 1,700 sq mi (6.2/sq mi). Over 915 taxa were documented representing an increase in the known flora by 231% (Kirkpatrick 1987). Although Cymopterus evertii Hartman and Kirkpatrick (1986) was first encountered on a small sandstone ridge at 5,800 to 6,000 ft in the Bighorn Basin, it was found to be common on pyroclastic andesites at 8,600 to 10,800 ft in the vicinity of on Carter and Phelps mountains. Other novelties encountered include: Ipomopsis spicata (Nutt.) V. Grant ssp. robruthii Wilken and Hartman (named in honor of Rob and Ruth Kirkpatrick; also found in the Northern Absarokas; Wilken and Hartman 1991), a new variety of Silene kingii (Wats.) Bocq. (Hartman and Dorn, unpubl. manuscript), and an undescribed species of Cymopterus (mentioned under Bighorn Basin). Astragalus gilviflorus Sheld. var. purpureus Dorn (1988) was characterized from the extreme southern edge of the area.

Southwestern Absarokas (Fig. 2, area 11)

It is separated from the Southeastern Absarokas by the South Fork of the Shoshone River and Shoshone Pass on the north and the East Fork of DuNoir Creek on the south. Much of the area forms the headwaters of the Yellowstone River; it is road less, and includes the most remote portions of Wyoming (up to a 25 mi hike through prime grizzly bear country to furthermost point). From this study area of 1,500 sq mi, over 5,600 collections were obtained (3.7/sq mi) during the summers of 1987 and 1988. A total of 891 taxa were documented from an area virtually unbotanized previously. Although no novelties nor state records were encountered, first records for the following counties were obtained: Teton, 56; Fremont, 22; and Park, 36 (Snow 1989, 1990, Snow, Nelson, and Hartman 1990). Descurainia torulosa Rollins (1983) had been described from the southern edge of the area. Much additional work is needed in the alpine (Trident, Thorofare, and Buffalo plateaus and Yellow Mountain).

West Slope Wind River Range (Fig. 2, area 12)

During the summers of 1990 and 1991, through a cost-share agreement with the Bridger-Teton National Forest, this area was inventoried. It extends from Togwotee Pass southeast along the Continental Divide to South Pass and ranges in elevation from 7,500 to 13,700 ft. The areal extent is 1,700 sq mi. The northern portion is sedimentary with some volcanics, whereas the southern part is primarily Precambrium in origin. More than 13,800 collections were obtained (8.1/sq mi) and the total flora now stands at 1,038 taxa (58% increase). Thirty-seven sensitive species of vascular plants (WYNDD and U. S. Forest Service lists) were documented. Many of these are restricted to exposed limestone on the mountains adjacent to the Green River Lakes. Four species new to the flora of Wyoming were also discovered (Fertig, Hartman, and Nelson 1991, Fertig 1992a, 1992b, Hartman, Nelson, and Fertig 1991).

The alpine of the eastern slope of the Wind River Range was inventoried by Richard W. Scott during 1963, 1964, and 1965 (Scott 1966). He returned to Wyoming in the mid-1970s to teach at Central Wyoming College (CWC) and continues his inventory of the east flank of the Wind River Range and adjacent areas in preparation for publishing an alpine flora of Wyoming and adjacent areas. The RM maintains an active exchange program with CWC (herbarium not in Holmgren et al. 1990; contains about 22,000 accessions).

Gros Ventre (Fig. 2, area 13)

During the summer of 1977 (prior to my arrival) and to a limited extent in 1978, roughly 1,600 collections were obtained from this area (960 sq mi; 1.7/sq mi). A total of 959 taxa were documented, increasing the known flora by 51% (Lichvar 1979a). Included in this list is Draba borealis DC., a boreal species previously known in the conterminous United States only from Colorado (Lichvar 1979b). Furthermore, 125 additions to the vascular flora of Teton County were discovered in the course of perusing the RM and during the fieldwork (ones not covered by Shaw 1976, Hartman and Lichvar 1980).

Targhee National Forest (Wyoming and Idaho), (Fig. 2, area 14)

Fieldwork was conducted in 1991 and 1992 with funding provided by the Forest Service via The Nature Conservancy. The first season focused on the Snake River Range, the Big Hole Mountains, and the west slope of the Teton Range (1,260 sq mi). The second covered the remainder of the Forest and adjacent areas including: Beaverhead Range, Centennial Range, Henry Lake Mountains, and Island Park area (1,620 sq mi). In 1991, 8,701 specimens (6.9/sq mi) representing 765 taxa were obtained including seven species of special concern to the Forest (Markow 1992). This past summer, a drought year, 5,063 specimens (3.1/sq mi) were collected and currently are being identified.

Wyoming and Salt River Ranges (Fig. 2, area 15)

The Willow Creek drainage (150 sq mi) in the northern Wyoming Range, was inventoried in 1990 through a cost-share agreement with Bridger-Teton National Forest. A total of 1,821 collections were obtained (12/sq mi) and the number of taxa documented was 532 including four of special concern (WYNDD; Hartman, Nelson, and Fertig 1991). Under a similar agreement, the remainder of the northern portion of the ranges was inventoried in 1992 (1,400 sq mi). A total of 7,776 collections were acquired by B. Embry, W. Fertig, Hartman, and Nelson (5.4/sq mi). The southern portion will be inventoried in 1993. The ranges are part of the overthrust belt which extends along the western edge of Wyoming and are composed solely of sedimentary materials, especially limestone. In recent years, several taxa new to science have been described from here including: Astragalus shultziorum Barneby (1981), Physaria dornii Lichvar (1983), P. integrifolia (Rollins) Lichvar var. monticola Lichvar (1984). Species of special concern found to be relatively abundant, often in scattered areas, are Astragalus paysonii (Rydb.) Barneby, A. shultziorum Barneby, Lesquerella paysonii Rollins, and Lomatium bicolor (S. Wats.) Coult. & Rose var. bicolor, whereas Draba borealis was encountered only twice.

Medicine Bow Range (including the Snowys; Fig. 2, area 16)

This area, located about 30 miles west of Laramie, has undergone intensive collecting by A. Nelson, C. L. Porter, and other and has been the site of many ecological studies by W. D. Billings, L. C. Bliss, R. F. Daubenmire, D. H. Knight, H. A. Mooney, and their students. For nearly 50 years, the S. H. Knight Science Camp (UW) served as the base for teaching field taxonomy and ecology courses. During the summer of 1973, B. Ernie Nelson collected about 1,000 numbers to supplement those already in the RM. These served as the basis of his M.S. thesis which contained dichotomous keys to the plants of the area. Historically, fieldwork has been concentrated along the highway traversing the range. In recent years, attempts have been made to inventory other areas including Sheep, Elk, Sheephead, and Jelm mountains, Kennedy Peak, and North Gate Canyon (Dorn, Hammel, Hartman, Nelson). A revised edition of the flora (Nelson 1984) containing 867 species of vascular plants has incorporated much of the new data and the inventory will continue as time permits.

Sierra Madre/Park Ranges (Wyoming and Colorado; Fig. 2, area 17)

This area, covering 2,200 sq mi, was inventoried during 1988, 1989, and 1990. Funding was obtained in part from the Routt National Forest and the Wyoming Native Plant Society. It ranges in elevation from 6,800 to 11,007 ft (Bridger Peak) in the Sierra Madre and to 12,180 ft (Mount Zirkel) in the Park Range. Composed of sedimentary strata along portions of the flanks, both ranges have a large, exposed core of Precambrian gneisses, schists, granites, quartz, etc. Also, volcanic activity has occurred in areas of the Park Range. A total of 5,290 collections (2.4/sq mi) representing 825 taxa were obtained. One new record for Wyoming (Ipomopsis aggregata [Pursh] V. Grant ssp. weberi Grant and Wilken) and 13 sensitive species for Colorado and/or Wyoming were documented (Kastning 1990).

Flat Tops/White River Plateau (Colorado; Fig. 2, area 18)

This inventory, conducted during 1990 and 1991, was funded in part by White River National Forest, The Nature Conservancy, and the Colorado Native Plant Society. The study area (Fig. 3) included all of the Flat Tops (Little, Dunkley, and Beaver; volcanic plateaus), White River Plateau, parts of the Yampa, Williams Fork, White, and Colorado river valleys, and most of the Grand Hogback (roughly 2,500 sq mi). It ranges in elevation from 5,300 ft near Rifle to 12,245 on Sheep Mountain. Approximately 6,500 collections (2.6/sq mi) representing 852 vascular plant taxa were obtained. A total of 27 populations of 11 species of special concern were located; 20 of the populations were newly discovered; seven of the taxa previously had not been reported from the study area. All but one of these taxa occurred in the southern 40% of the range, the portion composed largely of limestone (Vanderhorst 1992).

Philmont National Scout Ranch (New Mexico; Fig. 2, area 19)

During 1968, about 1,290 collections were obtained from this 210 sq mi area in the Sangre de Cristos. Over 760 taxa were documented including 11 new to the state (Hartman 1973) and two new to science: Eriogonum aliquantum Reveal (1976) and Heuchera hallii A. Gray var. novum (Hartman, unpublished). In 1991, Bruce Embury obtained an additional 500 numbers (average now of 9/sq mi) in preparation for a floristic inventory of a larger area in north-central New Mexico (Raton west to Costilla and southwest to Ocate and Taos; western Colfax, eastern Taos, and northwestern Mora counties; 2,500 sq mi).

The next decade or more of fieldwork at RM will focus on completing the inventory in Wyoming, southeastern Idaho, the west slope of Colorado, and the north-central 20% of New Mexico. The last area undoubtedly is the most poorly known botanially. We will coordinate efforts with botanists in these neighboring states to minimize duplication of effort. Because of limited resources, I do not foresee much fieldwork above the 45 parallel (Wyoming's northern border) in the near future. Our hope is to encourage botanists in Montana, Idaho, Alberta, and British Columbia to cover those states and provinces. Utah botanists, especially those associated with Brigham Young University (Stanley Welsh, students, and associates), are doing an admirable job of covering their state. Likewise, workers in the Great Plains, especially H. A. "Steve" Stephens and Ralph Brooks, have made a fairly good effort of inventoring the Black Hills of South Dakota (ca. 10,000 numbers). Unfortunately, the eastern plains of Colorado and Montana desperately are in need of detailed inventories.

What have we and others involved in similar projects in Wyoming accomplished during the past 15 years? A total of 306 new state records have been documented (Table I) and forty-six taxa new to science have been described (at least five more will be published soon)! Forty-one "resurrected" taxa (ones previously considered synonyms of other species but now recognized) have been added. Finally, 32 reports of taxa have been confirmed (with specimens to document the literature reports). The grand total is 425 taxa! Comparing only the number of species new to Wyoming with the number in Dorn (1977a; he did not include infraspecific taxa), there was an increase of 15.8% or one in every six species was new! Of the total number of taxa new to Wyoming, 24% are introduced (Hartman and Nelson unpubl.). Several of these represent potential agricultural pests.

From 1974 through 1990, 142 taxa have been described from the Rocky Mountains (United States) compared with 174 from the Intermountain Region, 163 from California, 121 from the Southwest, 92 from the Southeast, 45 from the Northwest, 39 from the Northeast, and 14 from the Great Plains (Table II; subtotals, therefore, not including named hybrids nor formas). Based on these data (Hartman and Nelson unpubl.), it is apparent that the Rocky Mountains continue to warrant considerable floristic study. Despite this fact, there is a real need to begin the publishing of taxonomic treatments for the region.

Table I. New Wyoming plant records since Dorn (1977a).

	Species	Subspecies	Varieties	TOTAL
New Records	258	4	44	306
New to Science	36	3	7	46
Resurrected Taxa	17	1	23	41
Reports Confirmed	28	0	3	32
New Wyoming Plant Records GRAND TOTAL				425
Introduced taxa	98	0	5	103
from above list = 24% of New Wyoming Records

Table II. Number of taxa by category described from various regions of the conterminous United States and from California during the past 16 years.

The only two regional floras are Coulter and Nelson's New Manual of Botany of the Central Rocky Mountains (Vascular Plants) published in 1909 (covering Colorado, Wyoming, most of Montana, the Black Hills of South Dakota, southern Idaho, the eastern half of Utah, the northern half of New Mexico, and adjacent Arizona) and Rydberg's Flora of the Rocky Mountains and Adjacent Plains: Colorado, Utah, Wyoming, Idaho, Montana, Saskatchewan, Alberta, and Neighboring Parts of Nebraska, South Dakota, North Dakota, and British Columbia published in 1917. The former was a new book, written by Nelson to replace John M. Coulter's earlier work (1885), but the latter was given senior authorship (Williams 1984). It was a relatively conservative treatment for the time and thus quite popular. Rydberg's Flora covered a much lingerer area, but extensive splitting at the species level greatly decreased its utility. Obviously, both are now obsolete except for historical purposes. Weber's Rocky Mountain Flora (1967 and revisions) is a misnomer representing little more than a revision of his useful Handbook of the Plants of the Colorado Front Range (1953 and revisions); it is of limited use beyond the Colorado Rockies.

The current project (Hartman 1990a, 1990b) was initiated with a symposium entitled The Flora of the Rocky Mountain Region (RMR) held at the Southwestern and Rocky Mountain Division of the American Association for the Advancement of Science Meeting in Colorado Springs, May of 1990. Speakers were Jane M. Beiswenger, Paleofloristics (Quaternary) of the RMR; Stanley A. Morain, Origin and evolution of the Rocky Mountain flora; William A. Weber, Phytogeographical affinities between the Rocky Mountains and Asia; William H. Moir, Vegetation zones and plant communities of the RMR; Theodore M. Barkley, The success of the Flora of the Great Plains Project; Stanley L. Welsh, The success of the Utah Flora Project; and R. L. Hartman, Floristic inventories and the Flora of the Rocky Mountain Project. Many of these authors will be contributing chapters to the Introduction for the Flora. Other contributors include Brainerd Mears, Jr., Physiography and geomorphology of the RMR; Roger L. Williams, Botanical exploration of the RMR; S. A. Morain, Plant geography and floristic divisions of the RMR; and R. L. Hartman, Endemism in the RMR.

The Flora of the Rocky Mountains: Vascular Plants of the Rocky Mountains and Adjacent Plains and Basins, North America (FRM) will consist of five or six volumes with a format similar to that of the Intermountain Flora (Cronquist et al. 1972 and subsequent ones) although certain modifications will be adopted from Flora of North America (FNA Editoral Committee 1990). A guide for contributors of taxonomic treatments to the Flora is nearing completion.

The goal is to coordinate the publication of the Flora of the Rocky Mountains with that of the Flora of North America, although the plant families occurring in two volumes of FNA will be covered in one volume of the Flora. The Flora volume will be published approximately two years after the corresponding ones for North America. Consequently, the Flora will be written along the lines of FNA. That is, in the pejorative, it will be a "flora by committee." More correctly, it will be a flora by experts, at least wherever possible. Although this method can lead to considerable hardship for the editor, more often than not, it leads to a greater approximation of reality with regard to the taxa recognized and thus a more enduring work. As a member of the Editorial Committee of the Flora of North America project, I am in a good position to evaluate the work of a specialist prior to inviting the individual to participate in the Flora. Unfortunately, for many authors, the Flora is queued behind The Vascular Plants of Arizona project and likely other floras.

Volume one, like the floras of North America and Intermountain Region, will consist of the introductory chapters (mentioned above) and taxonomic treatments of the ferns, fern allies, and gymnosperms. Illustrations from the Vascular Plants of the Pacific Northwest (Hitchcock, et al. 1955, 1959, 1961, 1964, and 1969) and the Intermountain Flora will be used with permission where appropriate. For the first volume, about 25% of the taxa will need illustrations.

The area covered by the Flora (Fig. 4) includes the Canadian Rockies south of Pine Pass (Hart and Continental ranges and the Purcell and Selkirk mountains; Alberta and British Columbia), all of the Montana, Wyoming, and Colorado, and portions of Washington (Stevens and Pend Oreille counties), Idaho (Bannock, Portneuf, Chesterfield, Bear River, Aspen, Caribou, and Snake River ranges and intervening areas of the southeast portion and that north of the Snake River Plains, excluding the Snake River Canyon and adjacent land below ca. 4,000 ft), Utah (La Sal, Uinta, Wasatch, and Wellsville mountains), South Dakota (Black Hills) and New Mexico (an inverted, roughly triangular area covering the north-central 20% of the state, south to the break in the mountains between Santa Fe and Albuquerque).

Justification for covering all of Montana, Wyoming, and Colorado in the Flora includes: 1) it will be useful to many more people, 2) many of the species in the plains to the east also occur in foothills adjacent to the Rockies and in intervening basins, 3) the western plains were not covered well in the Flora of the Great Plains (Great Plains Flora Association 1986) as most of the specimen studied were from herbaria in the central tier of the region, 4) and finally, there are several examples of major floras which overlap in coverage such as the Manual of Vascular Plants of Northeastern United States and Adjacent Canada (Gleason and Cronquist 1991) with the Flora of the Great Plains (Great Plains Flora Association 1986) and the Vascular Flora of the Southeastern United States (Cronquist 1980). Such overlaps in coverage are valuable in providing different taxonomic and regional perspectives on a flora.

Following the publication of the Flora, an atlas of the region is planned using specimen databases established at herbaria throughout the region and the latest technologies in GIS and computer cartography. This will be of tremendous use in phytogeographic, ecological, and systematic studies.

The Flora of the Rocky Mountains project represents a collective effort by the systematics community in the region and throughout North America. If the proposed Flora is to represent accurately the geographical range, ecological amplitude, morphological variability, and other aspects of the vascular plants of the region, additional floristic projects will be necessary. A coordinated effort by a number of herbaria and museums is needed. Although much of the fieldwork may not be completed in time to be reflected in all of the volumes, it is hoped that the knowledge gained from these efforts will be included in a supplement to or a revision of the Flora. Such information certainly can be incorporated into specimen and taxon databases.

In conclusion, the goal is to have the Flora published within the next 12 to 15 years with help from systematists throughout North America. To this end, a Rocky Mountain Flora Association is being formed to coordinate efforts for completing of fieldwork, for the data basing of specimens, and for the preparation of the Flora.

ACKNOWLEDGMENTS

I would like to thank past and present employees, graduate students, and associates for their dedication to the program at the RM and to benefactors who have helped to make it possible.

Likewise, I wish to acknowledge the support of the Department of Botany, the College of Arts and Sciences, the University of Wyoming, and state and federal agencies.

I thank B. Ernie Nelson, Walter Fertig, Greg Brown, Timothy K. Lowrey, and Michael D. Windham for their helpful comments on the manuscript.

LITERATURE CITED

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Coulter, J. M. 1885. Manual of the Botany (Phaenogamia and Pteridophyta) of the Rocky Mountain Region, from New Mexico to the British Boundary. American Book Co., New York, xvi + 452 pp.

Coulter, J. M. and A. Nelson. 1909. New Manual of Botany of the Central Rocky Mountains (Vascular Plants). American Book Co., New York, 646 pp.

Cronquist, A. 1980. Vascular Flora of the Southeastern United States. Volume 1, Asteraceae, University of North Carolina Press, Chapel Hill, xiii + 261 pp.

Cronquist, A., A. H. Holmgren, N. H. Holmgren, and J. L. Reveal. 1972. Intermountain Flora: Vascular plants of the Intermountain West, U.S.A. Hafner Publishing Co., Inc., New York, Vol. 1, iii + 270 pp.

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