Great Basin Naturalist Memoirs

Volume 9 A Utah Flora Article 1

1-1-1987 Introduction Stanley L. Welsh Monte L. Bean Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602

N. Duane Atwood USDA Forest Service, Intermountain Region, Ogden, Utah 84401

Sherel Goodrich Intermountain Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Ogden, Utah 84401

Larry C. Higgins Herbarium, Department of Biology, West Texas State University, Canyon, Texas 79016

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Recommended Citation Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; and Higgins, Larry C. (1987) "Introduction," Great Basin Naturalist Memoirs: Vol. 9 , Article 1. Available at: https://scholarsarchive.byu.edu/gbnm/vol9/iss1/1

This Chapter is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist Memoirs by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Great Basin Naturalist Memoirs A Utah Flora

Stanley L. Welsh', N. Duane Atwood", Sherel Goodrich^, and Larry C. Higgins^, editors

No. 9 Brigham Young University, Provo, Utah 1987

Abstract — A comprehensive treatment of the vascular flora of Utah is presented. Keys are provided to famihes, genera, species, and infraspecific taxa

(when present). Taxa are described, ecological data is given, and geographical information is provided. County distribution in Utah is given for each species and infraspecific taxon General geographical information is given for taxa that extend beyond the boundaries of Utah. Chromosome numbers are provided for each taxon, where that information was available in literature Indigenous taxa include some 2,572 species and 355 infraspecific taxa, or a total of 2,927. Introduced species number some 580. and the total taxa treated in the flora is 3,507. New nomenclatural proposals include lanceolata var. nivalis (Wats.)

Higginsstat. nov. ; C/irysot/iamnosnauseosus (Pallas) H, &C. var. uintahensis (L. C. Anderson) Welsh stat. nov.; Mac/ia?ran(/ieracanexccn.s(Pursh) Cray var. latifolia (A. Nels.) Welsh stat. nov., and var. aristatus (Eastw.) Turner comb, nov.; Vif;,uiera longffolia (Robins. & Greenm.) Blake var annua (Jones) Welsh comb. nov. ; Dudleya pulverulenta (Nutt. ) Britt. & Rose var. arizonica (Rose) Welsh stat. nov. ; Cordytanthun kintlii Wats. var. densiflorus (Chuang & Heckard)

Atwood stat. nov ; Cymopterus acaulis (Pursh) Raf. var. fendteri (Gray) Goodrich comb. nov.

INTRODUCTION

Stanley L. Welsh

Historical Basis Thus, the present work is not only a fruition of the years

of labor by its editors and contributors, but it represents the culmination of than Study of Utah flora began in September 1843, when John more a century and four decades of exploration collection Charles Fremont took the first collections from the and of the of Utah. Begin- region later included within the state (Fremont 1845). In the ning with Fremont, the exploration has gone foreward to late 1860s Sereno Watson collected in Utah and Nevada and the present, not in a steady rate but at a varied one as the authored the most important contribution to the understand- personnel involved have waxed and waned. Hundreds of ing of Utah flora in the 19th century (Watson 1871). His work collectors have contributed specimens that form the basis was used by such important Utah botanists as Marcus E. of this flora. The few specimens taken by Fremont repre- Jones, who published an important summary treatment of his sent only a very small but significant part of the tremen- own 1894 fieldwork in 1895 (Welsh 1982a). A. O. Garrett dous number of collections made in the intervening vears (Welsh (1909 and subsequent) published the Spring Flora of the 1986). Wasatch Region early in the 20th century, which was used by Perhaps 400,000 specimens from Utah reside within generations of students to as late as midcentury. His work the herbaria of the state, and possibly almost that many stimulated the subsequent publication of other local floras, more examples from Utah are present in herbaria else- including the Handbook of the Vascular Plants of the North- where. More than a thousand type specimens have been ern Wasatch by Holmgren (1948, 1957), and the Flora of the taken from within Utah (Welsh 1982a). There has been a Central Wasatch Front, Utah bv Arnow, Albee, and Wvcoff renaissance within classical plant in Utah dur- (1977, 1980). ing the past three decades. That rebirth of taxonomic Tidestrom (1925) wrote a summary treatment of the flora of emphasis was involved with the improvement in trans- Utah and Nevada, which has long been out of print and badly portation and accessibility of Utah, in the increased num- dated by changes in nomenclature and new information. ber of personnel, in the devotion of those persons, in the Welsh and Moore (1973) attempted a more comprehensive environmental movement aided by federal laws, and in an treatment of the Utah flora. Their work involved some 2,500 increased emphasis for production of state and regional taxa and included all of Utah. floras.

Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. "USDA Forest Service, Intermountain Region, Ogden, Utah 84401. Intermountain Forest and Range Experiment Station, Forest Service, U.S. Department of .\griculture. Ogden, Utah 84401. Present addr Ranger District, Ashley National Forest, Vernal, Utah 84078. Herbarium. Department of Biology, West Texas State University, Canyon. Texas. 79016 .

Great Basin Naturalist Memoirs No. 9

A flora of the Uinta Basin has been prepared by Sherel phytochemistry, breeding mechanisms, and other items Goodrich and EUzabeth Neese, and its pubhcation is important to its interpretation. anticipated in 1986. An atlas of distibution maps of Utah Simple recombinant types should not be taken for taxa plant species has been prepared by Beverly Albee and necessarily. Such unrelated examples might well appear Leila Shultz, and it is expected to be published shortly. again and again within a group, and keys written to such plants will provide means of identification of these recom- Philosophical Basis and Species Concept l)inant types, but do they represent taxa? Therein lies perhaps the greatest dilemma of plant classification, i.e., The underlying principle in this work is reality and its the taxonomic character should reflect relationship, not representation as nearly as possible. Reality is inter- haphazard trivial preted differently by each worker, of course, and one's merely a recombination of features, but the pursuit of the truly reflective characteristics is often concept of reality changes as new information is forthcom- merely an ideal. ing. Taxonomic concepts in this work are derived from the study of the plants themselves, both in the herbarium and Ideally, taxonomic treatments represent a translation in the field. All contributors to the taxonomic text are of the reality of nature into graphic representation. Diffi- familiar with the plants as they occur in nature, and that culty arises, and taxonomists are held in disrepute, when reverse familiarity is transferred to the plants as they exist on they try to the situation by imposing a system of herbarium sheets. In the natural context plant taxa occur their own creation onto the plants as they are in the real in correlation with geography, geology, and climatic or world. positional factors. The correlations are important in de- And, the flora is only imperfectly understood. Many of termination of the nature of the taxon, and it is to these the plant taxa are known from a few collections only, and it correlations that we have looked for understanding. The is impossible to represent them beyond the limits of the juxtaposition of taxa in the field (or the lack thereof) forms information available. Many taxa have been discovered another parameter that must be determined, because recently and we know nothing of their chromosome com- many of the taxa form morphological intermediates when pliments, breeding mechanisms, and other features. in contact. Hybridization and its potential must be re- They occur in the treatments presented below as best as garded in taxonomic decisions. Presence of cleistogamy possible, but modification and reinterpretation will follow and apomixis within a group or complex must be consid- the accumulation of other data. Other workers will evalu- ered. Other representations of reality, the floras, revi- ate the same data differently, resulting in the presenta- sions, and monographs must be studied to supplement tion of restructured taxonomic opinions. Some will wish the information gained from field and herbarium studies. to dissect the genera into ever smaller units, based on Thus, taxonomic concepts in this work are derived both various tangible and intangible features, as if such re- from the study of the plants themselves and from the structuring was of tremendous importance. All presenta- works of other students of the flora. tions will be based in fact, as the worker will interpret that All indigenous plant species known to occur in Utah are fact, but the weighting of the data and the biases of the included in the flora. Introduced plants are also covered, worker will determine, in some part, the presentation but not so intensively as are the native ones. Plant tax- followed. onomists have been conditioned to work with the indige- Prior to the present manuscript, which is acknowl- nous plants alone, often to the exclusion of introduced edged beforehand as imperfect and stained with the bi- ones. The introduced taxa are, however, a portion of the ases of its authors, routine identification of a collection of flora of the state and cannot be ignored. Many are estab- plants from anywhere in Utah has involved the use of as lished within native or human-induced habitats. Others many as a dozen or more floras and monographs. The are merely waifs that grow occasionally in the state. Culti- summary works of Tidestrom (1925) and Welsh and vated plants offer still another group that has been largely Moore (1974) functioned merely to tantalize the serious neglected by botanists. student of Utah plants. The local floras by Garrett (1909 This flora attempts to present coverage of all estab- and subsequent), Holmgren (1948, 1957) and Arnow et al. lished species, all common adventive taxa, and many of (1977, 1980), served the local areas more or less ade- the commonly grown cultivated species. No attempt has quately, but they left students of surrounding areas grasp- been made to provide coverage for all the cultivated ing for tools for identification. plants. There are a great many of them, and they are Literature used for determination of plant taxa depended typically poorly represented in herbaria. Future workers on the locality where the plant was collected. Those taken should focus more attention on the cultivated flora. A from southern Utah were identified using the Arizona Flora more reasonable taxonomy of those plants will then be (Kearney and Peebles 1951), those from eastern Utah were possible. determined by vise of the Manual of the Plants of Colorado

A taxon is based on the totality of its characters, but the (Harrington 1951 ) and others (Munz 1969; Correll & Johnston character does not make the taxon. When workers have 1970; Cron(iuist et al. 1972, 1977, 1982; Dorn 1977; Hitch- become intrigued with thi- value of a character they have cock et al. 1955- 1969). Specific groups were checked against been led to absurdity. Within this century fruit differ- the most modern revisionary works and monographs. Serious ences were used by one worker to separate the species of workers soon ac()nired a library oi books and separates if their Astragalus into numerous genera, often with closely re- detenninations were to hv at least partially adecjuate. Now, lated and otherwise practically identical species placed in for the first time, there will he one work that can he used for all

separate genera (e. g. , A . heckwithii in Phacomcne and A but an unknown niiniher ol plants taken iroin within tlie state oophorus in Phaca). The character must be considered in of Utah. A general i)ii)!i()grapli\' oi works used in identification context with other features of the plant, with its geogra- of Utah plants previously and on which concepts ol this flora

phy, geological substrates, elevational range, features of are based in part is presented below. Specific citations are slope, sympatry or allopatry, chromosome condition. included with the taxonomic treatments. 1987 Welsh etal: A Utah Flora, Introduction

Methodology Spring begins in Utah at the lowest and southernmost elevations in Washington, Kane, and San counties as Each taxonoinic treatment was written in its entirety by Juan early as January in some years, but more commonly in its author or authors, beginning with a checkHst of taxa March and April. As the relationship of the earth to known or thouglit to occur within the state. Keys were the sun moves from vernal constructed based on specimens taken from the herbar- the toward the autumnal equinox, flowering and fruiting of plants move both northward ium and from concepts in hterature. Many have been and upward in elevation. Spring arrives at some of the high improved by comparison with hving plants in the field. mountain summits only to meet autumn moving down. However, most were written in their entirety from Summer at high elevations is a fleeting condition herbarium specimens, and their similarities to published that hardly exists in some years. The autumn-flowering plants treatments are coincidental. Where keys were derived in reverse the trend of flowering time, finally closing out the part from publications, the source is included in literature season in the lowermost elevations of the citations, typically listed following the generic descrip- southernmost portions of the state typically in October tion. and November (December). Descriptions are likewise based on specimens, except There are plants that begin to flower in southern Utah in where those were inadequate for the purpose (some taxa November and December and continue flowering to spring of the following are known from few and sometimes imperfect speci- year (e.g., Tham- nosrna and Garnja). For most of Utah spring arrives in mens), and the circumscriptions then came, at least par- April and May, but there are areas where frost can (and tially, from literature, especially from original descrip- often does) occur at all months of the year. tions. Format for the descriptions is essentially that Such ecological data as slopes and valleys (or standard for much of taxonomic literature. Duration, rocky or dry) have been omitted because practically all plants root, stem (including habit and aspect), , inflores- grow in such sites. The term saline has used in most cence, flowers, and fruits are described in essentially that been cases

in place of the familiar alkaline , since one can observe order. Chromosome number is included at the end of salinity in the field but must test for alkalinity. descriptions, where that information has been available in literature. We have not attempted to do routine counting Measurements of chromosomes as a part of the basic research. Type specimens of most taxa described from Utah have Plant parts are measured in metric units. Parts measur- been examined by one or more of the authors. Decisions ing less than 1 cm long are always cited in millimeters on critical groups have been based on the type specimens. (mm) and tenths of millimeters. Millimeter accuracy is Authors of taxa are cited routinely for each taxon of the routine for parts up to about 25 mm in length or breadth, rank of family or below. The author names are abbrevi- and then the unit of measurement is changed to centime- ated following the list by Chatterley, Welsh, and Welsh ters (cm). Decimeters (dm) are used for plants or plant (1982) and supplemented by the work of Halliday, parts that generally exceed 50 centimeters. Meters (m)

Meikle, Story, and Wilkinson (1980). We have chosen not are used for plants that exceed about 1 . 5 meters and for all to use all the abbreviations in the latter work because of elevational measurements. Kilometers are used for long brevity; e.g., M. E. Jones is cited merely as Jones in the lineal measurements (except those cited in type locality Utah flora and A. Gray is Gray. A complete list of author information where miles are used). abbreviations and their given names and surnames, along Plant height is taken as the length of the plant above with year of birth, year of death, or year of publication, is ground level and includes the inflorescence and flowers. appended to this work. measurements are from the petiole base to the blade Discussions following the description provide clarifica- apex, unless otherwise indicated. Leaf breadth is taken at tion in many instances, but they routinely include plant the widest position on the blade. Petioles are measured community data (typically from low to high elevations), from point of insertion on the stem to the blade base. elevational range (in meters), counties from which the Stipules are measured from the point of insertion on the taxon is known in Utah (arranged in alphabetical order), stem or petiole base to the apex. Peduncles are measured distribution beyond Utah (or whether they are endemic from the uppermost bract to the first flower or branch of a to the state), the number of specimens seen from Utah compound inflorescence. Total flower length is taken during the study (Arabic numerals) and the number col- from the point of insertion of the pedicel (peduncle) to the lected in Utah by the author of the particular group apex of the longest petal. Calyx measurements are taken

(Roman numerals). The number examined is an index to from the point of insertion of the pedicel to the tip of the both the familiarity of the author with the taxon and its longest calyx teeth, except in legumes, and then the abundance; the number collected by the author provides measurement is to the length of the longest lateral tooth. an index to that worker's understanding of the taxon in the Calyx teeth and corolla lobe measurements are from the field. base of the sinus to an adjacent lobe or tooth apex. Petal Taxa known to be adventive or introduced purposefully length is determined from the point of insertion of the are so noted in the discussion, and the region from which petal to its apex. Fruit length measurements include from the plants originated is indicated. the base of the ovary (point of insertion on the receptacle) Flowering and fruiting dates have been routinely omit- to the tip, excluding style and stigma measurements, ted, because practically everything in Utah flowers in unless otherwise noted. Gynophores are excluded from either spring (extending to early summer) or (late summer ovary measurements but stipes are not. to) autumn; and many plants that flower in springtime Nomenclature again flower in autumn. Only when the situation is un- usual has the flowering date been included. Names given for taxa in this work are consistent with Midsummer is a poor time to collect plants in Utah, stipulations of the International Code of Botanical except in high mountains, where it is still springtime. Nomenclature (Voss 1983). The principle of priority is the Great Basin Naturalist Memoirs No. 9

overriding consideration, except where expressly limited taxonomic categories arranged in a phylogenetic se- by mandate ofthe Code. Species and generic concepts are quence follows (the arrangement of the mainly conservatively interpreted, with the definition of families is modified from Cronquist 1981); species considered multifaceted because of differences in breeding mechanisms and point in evolutionary time. GROUP TRACHEOPHYTA The concept of subspecies is herein restricted to those species with usually large geographical and morphologi- Approximation of a phylogenetic arrangement of families. cal representation. They typically consist of two or more varieties. Varietal designation is more often used alone as LYCOPODIOPHYTA the principal category below the species level, especially Family name Author Genera Species Infra in less widely ranging and less complex species wherein designation of subspecies is unwarranted. However, no 1. Isoetaceae slw effort has been made to routinely propose new nomen- 2. Selaginellaceae hig clatural transfers of all subspecies to varietial level. Such a course is not indicated in this provincial treatment that EQUISETOPHYTA cannot encompass the variation in all taxa beyond the boundaries of Utah. Family name Author Genera Species Infra

The names utilized for genera and species are based on I. Equisetaceae hig the best information available to us and on our under- standing of the groups as they occur throughout their FOLYPODIOPHYTA natural and adoptive ranges. Major checklists, which at- tempt to categorize and standardize generic and specific Family name Author Genera Species Infn are followed only in part Soil Conserva- names, (USDA 1. Ophioglossaceae hig 1 4 tion Service 1982). The choice of generic and specific 2. Polypodiaceae hig 15 35 concepts are best left to competent taxonomists, not legis- 3. Marsileaceae hig 1 2 lated by adoption of standardized lists. Workers on 4. Salviniaceae hig 1 1 specific areas of a country as large and diverse as the United States are certain to know more about the plants of PINOPHYTA their area than are workers who labor more casually on a broader problem. However, that tendency of more inti- mate knowledge does not relieve the worker in a limited area from reviewing all the information available from whatever source. We hope we have reviewed the infor- mation from all sources carefully, and our taxonomic and nomenclatural decisions result from that review. The final word is not to be found, however, in the following pages. As additional collections are made, and as summary monographs and revisions become available, the views presented here will be subject to change. The flora of the region and its nomenclature still require much under- standing and great labor.

Common Names

Common names have been taken, with modification, from the USDA Forest Service General Technical Report INT-.38 and from general nonpublished sources. No at- tempt has been made to provide common names through- out, especially when they are not known or where they are not in general usage.

Classification

This hook is dedicated to the many students of l)otany

who will use it as a reference. Much of its use will be to find names of genera and species of examples in hand. Because of that use and the need for ease and economy of

time, a phylogenetic arrangement is carried only to the class level. The book begins with a key to the divisions,

and is followed by the lern allies (Lyeopodiophyta and K(juisetophyta), ferns (Folypodiophyta), conifers (Fino- phyta or gymnosperms), and finally the flowering plants

(Magnoliophyta), with Magnoliopsids (dicots) first, fol- lowed by the Liliopsids (nionocots). Within each major group the families are arranged in alphabetical order, as

are the genera and species within each family. A list of the 1987 Welsh etal; A Utah Flora, Introduction

28. Giittiferae big 1 2 101.

29. Tiliaceae slw 1 7 (intro 7)

30. Malvaceae slw 10 25 (intro 7) 1

3L Violaceae .sKv I 11 (intro 4)

32. Tamaricaceae slw 1 3 (intro 3)

33. Frankeniaceae .slw 1 1 (intro 1)

34. Passifloraceae slw 1 1 (intro 1) 35. Cucurbitaceae hig 7 12 (intro 10) 36. Loasaceae kht 3 20 37. Salicaceae sg 2 35 (intro 10) 38. Capparaceae slw 3 5 39. Cruciferae slw 46 165 (intro 42) 39

40. Rcsedaceae slw 1 1 (intro 1) 4L Ericaceae slw 5 12 42. Pyrolaceae slw 4 9

43. Ehenaceae slw 1 1

44. Primulaceae slw 6 17 (intro 1) 45. Saxifragaceae sg 13 41 (intro 4) 46. Crassulaceae slw 3 6 47. Rosaceae slw 33 111 (intro 45) 9 48. Leguminosae slw 46 244 (intro 51) 65

49. Elaeagnaceae slw 2 5 (intro 1)

50. Haloragaceae slw 1 2 5L Lythraceae higslw 3 4 (intro

52. Punicaceae hig 1 I (intro 53. Onagraceae slw 10 59

54. Cornaceae slw 1 1

55. Garryaceae hig 1 1

56. Santalaceae slw 1 1 57. Viscaceae nda 2 58. Celastraceae slw 4 9 (intro 6) 59. Aquifoliaceae slw 1 2 (intro 2) 60. Euphorbiaceae higslw 6 28 (intro 5) 6L Rhamnaceae slw 3 8 (intro 3) 62. Vitaceae slw 2 5 (intro 3) 63. Linaceae slw 1 6

64. Polygalaceae slw 1 3

65. Krameriaceae slw 1 2

66. Sapindaceae slw 1 1

67. Hippocastanaceae kht 1 4 (intro 4) 68. Aceraceae slw 1 12 (intro 9) 69. Anacardiaceae slw 5 8 (intro 5)

70. Simaroubaceae hig 1 1

7L Meliaceae hig 1 1 (intro 1) 72. Rutaceae higslw 2

73. Zygophyllaceae slw 4 4 (intro 1)

74. Oxalidaceae slw 1 2 (intro 2) 75. Geraniaceae slw 3 9 (intro 1)

76. Limnanthaceae slw 1 1

77. Tropaeolaceae slw 1 1 (intro 1) 78. Araliaceae slw 3 4 (intro 3) 79. Umbelliferae sg 25 70 (intro 5) 10 80. Gentianaceae hig 5 18

8L Apocynaceae hig 5 10 (intro 3) 1

82. Asclepiadaceae hig 3 20 (intro 1) 83. Solanaceae hig 11 33 (intro 14) 84. Convolvulaceae hig 5 8 (intro 4)

85. Cuscutaceae gib 1 12 (intro 1)

86. Menyanthaceae slw 1 1 87. Polemoniaceae slw 13 67 (intro 3) 10 88. Hydrophyllaceae nda 9 49 6 89. hig 17 97 90. Verbenaceae hig 3 8 91. Labiatae hig 24 43 (intro 14)

92. Hippuridaceae slw 1 1

93. Callitrichaceae slw 1 3

94. Plantaginaceae hig 1 7 (intro 2)

95. Buddlejaceae hig 1 2 (intro 1) 96. Oleaceae slw 6 19 (intro 13) 27 97. Scrophulariaceae nda/ecn 18 137 (intro 12) 98. Orobanchaceae hig 1 5

99. Pedaliaceae hig 1 1 100. Bignoniaceae hig 3 5 (intro