Distribution and Relationships of Utah Brome Grasses in Western North America

Home , Arundinoideae, Bromus, Bromus arvensis, Bromus ciliatus, Bromus commutatus, Bromus diandrus

Brigham Young University BYU ScholarsArchive

Theses and Dissertations

1973-04-01

Distribution and relationships of Utah brome grasses in western North America

Jerry B. Anderson Brigham Young University - Provo

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BYU ScholarsArchive Citation Anderson, Jerry B., "Distribution and relationships of Utah brome grasses in western North America" (1973). Theses and Dissertations. 8012. https://scholarsarchive.byu.edu/etd/8012

This Thesis is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. DISTRIBUTIONAND RELATIONSHIPS OF UTAHBROME GRASSES IN WESTERNNORTH AMERICA

A Thesis

Presented to the

Department of Botany and Range Science

Brigham Young University

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

by Jerry B. Anderson

April 19'i3 ii

This thesis by Jerry B. Anderson is accepted in its present form by the Department of Botany and Range Science ·of Brigham Young

University as satisfying the thesis requirement for the degree of Master of Science. iii

ACKNOWLEDGMENTS

Grateful acknowledgment is made for the valuable suggestions and assistance given by the chairman of my advisory committee, Dr.

Stanley Welsh, and to Dr. Wilmer W. Tanner, the other member of my graduate committee.

Appreciation is extended to Dr. Walter P. Cottam, without whose encouragement I would never have entered the field of Botany.

Special thanks are due the curators of the many herbaria of the

Intermountain Region for making specimens available for study. Thanks also to Larry C. Higgins who drew and furnished copies of the map of

Utah. To my wife, LaPrele, deepest appreciation is expressed for her untiring devotion, patient help for these several years, and cooperation.

She also assisted in the preparation of the drawings. TABLEOF CONTENTS

Page ACKNOWLEDGEMENTS . . . iii f· LIST OF TABLES V LIST OF FIGURES ...... vi INTRODUCTION 1 MATERIALSAND METHODS . . 2 Historical account 3

Distribution and ecology 5

General morphology 6

Cytology and generic relations 8

TAXONOMICTREATMENT_. . 15 LIST OF REFERENCES ...... 72 APPENDIX-Index of Synonyms 81 V

LIST OF TABLES

Table Page 1. Nomenclature and historical development of the genus and sub generic taxa in Bromus ...... 7

2. Sub-families and Tribes of the Family Gramineae 11

3. Chromosome numbers reported for Utah Bromus 13 vi

LIST OF FIGURES

F_igures Page

1. Illustration of Bromus marginatus .. 26

2. Illustration of Brornus willdenowii 30

3. to 6. Maps of distribution for Bromus marginatus, B. marginatus var. polyanthus, ~- willdenowii, and B. 1.nermis ...... • . • . . . . 32

7. Illustrations of Bromus carinatus, B. porteri, and B. trinii ...... - . . . . 38 8. Illustration of Bromus canadensis 41

9. Illustration of Bromus vulgaris 44

10. to 13. Maps of distribution for Brcmus porteri, ~- canadensis, ~-· j aponicus, and B. commutat:us 48 14. Illustrations of Bromus commutatus and B. japonicus so

15. Illustration of Bromus hrizaeforr.tis ... 56

16. to 19. Maps of distribution for Bromus racemos 1.1s, ~- mollis, B. brizaeformis, and~- trinii .. 58

20. Illustrations of Bromus tectorum and B. rubens 62

21. Illustration of Bromus sterilis 66

22. to 25. Maps of distribution for Bromus tectorum, B. sterilis, .Q_.rub~~~-., and B. dianarus .... 68 1

INTRODUCTION

Since the revision of Bromus by Shear (1900) and publication of the Manual of Grasses of the United States (Hitchcock, 1935) a considerable amount of new study material has become available. Wagnon (1952) did a revision of one of the five sections, and there have been some cytological studies of parts of the genus.

There has been a variety of treatments by a number of authors, and some disagreement in interpretation of entities. Specific and generic delimitations have varied from one author to another with a resulting proliferation of name proposals. Bromus has continued to be a problem genus in Utah, and there has been a need for a comprehensive study.

This study is an investigation of the taxonomy and biogeography of Bromus in Utah. In gen~-ral the treatment is concerned with plants at the species level, but some infraspecific taxa have been included.

I have attempted to present a useful definition of the species within the genus for those concerned with problems of classification in Utah.

In some aspects it is preliminary since detailed and lengthy investigations of individual species and s-pecies complexes will be required for definitive solution to a number of problems.

The problems concerning nomenclature of the entities involved in this study have been partially solved by previous authors (Stapf, 1928; Hitchcock, 1935; Wagnon, 1952) and I have adocted their work as a basis for my nomcnclatural treatment. 2

MATERIALSAND METHODS

The materials used in this study are largely the dried and

pressed specimens in herbarium collections. Over 1000 specimens were carefully examined to provide accurate and consistent descriptions of ,t the taxa, to obtain information on habitat and other useful data, and

to determine distribution and variability in the constituent groups.

Because of the large number of specimens examined, citation of all of

them has not been attempted.

The original descriptions of our species were examined with illustrations and commentaries concerning the type specimens wherever

possible. Lack of time and funds precluded any first-hand study of the

type specimens, most of which are available only in Europe.

Taxonomic criteria relating to size, length, and shape were

evaluated through measurements of plant structures with the aid of low-

power binocular microscopes and a 15 cm, ruler graduate~ in millimeters.

Herbaria from which spedmens have been seen, with the standard abbreviations (Lanjow & Stafleu, 1964) by which they are referred to in the text are as follows:

ASC Arizona State College, Fl2.gsta.ff, Arizona.

BRY Brighar.1 Young University, Provo. Utah.

CCH Colorado College, Pueblo, Colorado.

DAV University o_f California, Davis California.

DIX Dixie Junior College, St. George, Utah

DS Dudley Herbarium, Univ. of California, Berkeley, Calif. ISC Iowa State College, Alnes, Iowa. 3

JEPS .Jepson Herbarium, Univ. of Calif., Berkeley, Calif.

MIN . University of Minnesota, Minneapolis, Minnesota.

NCU University of North Carolina, Chapel Hill, N. Carolina.

NY New York Botanical Garden, New York; New York.

OKLA Oklahoma State University, Stillwater, Oklahoma.

RIK Rick's College, Rexburg, Idaho.

RM Rocky Mountain Herbarium, Univ. of Wyoming, Laramie, Wyo.

SLP Slippery Rock State College, Slippery Rock, Penn.

SUSC Southern Utah State College, Cedar City, Utah.

UC University of California, Berkeley, Califor·nia.

UC! University of California, Irvine, California.

UT University of Utah, Salt Lake City, Utah.

UTC Intermountain Herbarium, Utah State University, Logan, Ut.

USFS Forest Service Herbarium, U.S. Dept. of Agriculture.

WES Weber State College, Ogden, Utah.

WTS West Texas State University, Canyon, Texas.

Historical Account

The modern origin of the name Brom~ is revealed by Linneaus in his Philosophia Br::itani~a (1751:188). Bromus was derived from the ancient Greek word ~pwµ.a:meaning food, and from that w:ird also comes the name l3rot"-oi;meaning ,: oat". Linneaus indicates that Theophrastus and Dioscorides used the latter term. It appears that Linneaus was the first modern author to apply the name as currently used, but he does cite the earlier works of •Monti and Scheuchzer as showing representa-- tives of Bromus on various plates of their separate works. 4 Professor S.M. Tracy at the AMS meeting ln 1893 reported on the distribution of the 20 species known in the United States at that time, and noted that half of the number were innnigrants (Beal, 1896:668).

Shear (1900) describes 40 species and 20 yarieties, while 42 species and 12 varieties are listed for the United States in Hitchcock's manual

(1951). Shear listed 18 species as indigenous, and Hitchcock's manual recognizes 22. A compilation of the species in the Genus, as at present recognized in the wider sense throughout the world, would yield a total probably just short of 100.

The first revisionary treatment of Bromus was that of Shear

(1900). He treated the North American species occurring North of Mexico.

A very significant stride forward in agrostology crune with the publication of Hitchcock's (1935) Manual of Grasses of the United States. The publication of this manual has given a degree of confidence to students of the grass family, and herbarium collections and the number of research papers on grasses h;ive increased enormously since the advent of that treatment.

Both Shear (!900) and Hitchcock (1935) accepted Bromus secalinus as the type of the genus on the l:,asis of its being the first species listed in the first edition of Species Plantarum of Linneaus (1753).

Wagnon (1952) in his revision of the section Bromopsis argues that

B. arvensis is the type of the genus. The controversy has been a factor leadirtg to the acceptance of the concept of a "standard" species for Bromus. Stapf '(1928), Hitchcock (1935), and i'!agnon (1952) all accept

B. sterilis as the standard species, that is, the species that best represents the taxonomist's concept of a brome grass. Stapf (1928) reviewed the nomenclature of B-ro:r.usand presented a table in his work that shows the historical development of the concept 5 of the genus from. 1737 to 1913. It points out the trend that has de-

veloped since the work of Parlatore in 1844, to replace Linneaus'

B. secalinus and its allies with the B. sterilis group as the true Bromi.

Some authors entirely eliminated the B. secalinus group from the genus,

but since about 1935 there has come to be an increasing amount of

evidence that the genus as it is now recognized in its wider sense is probably a valid natural grouping. The differences noticed are those of

' the treatment of the higher taxa during the past several years.

Distribution and Ecology

Grasses of the genus Bromus and others of the sub-family

Festucoideae are found consistently in the cool or cold regions of the

earth, and are found growing and reproducing in the subtropics only

during the cool season of the year. The genus Bromus is most widely

distributed in the temperate and mountain regions of the world, and in

the United States reaches its greatest development in the Rocky Mountains.

In the temperate zone, various representatives of the brome grasses can

be found from sea level to over 12,000 feet elevation. In the tropics

the species are mostly restricted to the higher mountains (Shear, 1900).

The native bromes are referred to the sections Pnigma and

Ceratochloa, and the two sections show rather differ-ent distribution

patterns. Within the section Ceratochloa there are abundant representatives from the southern tip of South America to Western Alaska, but

they are not found in the Eastern Hemisphere except where sparingly

introduced. Several of the species in the section Pnigma are important forage grasses of our mountainous and northern regions. This section is represented in circumboreal distribution. 6 The several species of weedy annual bromes in the United States

are mainly adventive European and Mediterranean species referrable to the sections Brornium and Stenobromus. These grasses are distributed in

almost any disturbed locality throughout the country, but mainly in the lower elevations of the Northern and Western States. Some of these brome

grasses qualify as really troublesome weeds. Those of the section Stenobromus are, at maturity, a serious pest, and the narrow, sharp-

pointed minutely barbed florets with their long awns, work into the

eyes, nostrils and mouths of livestock. Serious injury and inflammation

can result, and sometimes the intestines are pierced causing death

(Klemmedson and Smith, 1964). The ripgut grasses, Bromus diandrus and

B. rigidus, are the chief offenders in this ~egard.

The relatively barren plains and overgrazed foothills and ranges of the mountains in the western states are often covered with an extens-

ive green blanket of B. tectorum in the early spring. At this stage it

is widely grazed, but very soon becomes dry and straw colored when the plant matures. The great stands of this grass constitute a major fire hazard in the late summer. In June and July along the Wasatch Front this grass is the host fo-r smut fungus, and the a·ir becomes filled with the spores thc1.t are carried by the wind from the infested cheat grass on the low hills. Peck (1959) reported a count of over 2000 smut spores on one exposed slide for t~e day of June 12, 1958 in downtown Salt Lake

City. The weedy grasses also harbor i:isect pests, and may be hosts for plant diseases that could be transfe:rred to crop plant hosts.

General !-forphology

One of the diagnostic chaTacteristics of the genus Bromus that 7

Table 1. Nomenclature and historical development of the genus and sub generic taxa of Bromus.

1 Authors Year Bromus sens. lat. and possible segregates~ 1 2 3 4 s

Linn. Gen. Pl. 1737 BROMUS Sp. Pl. 1753 BROMUS BROMUS Beauvais 1812 BROMUS BROMUS BROMUSand CERATOO:ILOA BRACHYPODIUM Panzer 1813-14 BROMUS ZERNA ZERNA TRAGUS Dumortier 1823 Bromium Bromium Bromopsis and Pnigma Lej. 1825 ' Dedebour 1829 Zema Bertolini 1833 Bromi Bromi Bromi secalini genuini festucacei !Coch 1835 " II II Drejer 1838 FESTIJCA Bromochloa Hooker 1840 BRO~rus CERATOCHLOA Parlatore 1840-42 SERRAFALCUSBROMUS BROMUS. Koch 1844 Serrafalcus Bromi Bromi genuini festucarei Grisebach 1843-45 Zeobromus Stenobromus Fries 1845 SCHEDONORUS Kittel 1847 Festucula Koch 1848 ANISA.NTHA Ledebour 1853 Schedonorus Schedonorus Steud 1854 TRINIUSIA Grenier & Godr. 1855-56 SERRAFALCUSEubromus Festucaria (B. danth.) Casson &Duri 1856 Festucoi1es Willkomm 1861 SERRAFALCUSEubromus Festucaria Fourneau 1869 BROMOPSiS Bentham 1883 Zeobromus Stenobromus Festucoides Ceratochloa Boissier 1884 Serrafalcus Eubromus Festucaria Hackel 1887 Zeobromus Stenobromus Festucoides Ceratochloa Shear 1900 Bromus prpr. Stenobromus Zerna Ceratochloa Sub-genus Buhani 1901 FORASACCUS Neobromus Asch. & Graeb. 1901 Serra.falcus Eubromus . Festucaria Ceratochloa (B. trinii) Groves 1904 SERRAFA!..CUSBRO'!US BRO~rus Rouy 1913 SERRAFA!..CUSEubromus Festucoides Brit. & Brown 1913 BROMUS BRO~lUS BR~rus BRQMllS Lunell 1915 FORASACCUS FORASACCUS Stapf 1928 Serrafalcus Eubromus Zema Ceratcchloa Nevski 1934 BRO~rus A:HSA.Ylr!A ZEm'iA TRISETCBROMUS Hitchcock 1935 Eromium Eubrolll!1S Bromopsis Ceratochloa Neobromus Gould 1968 Bromium Eubromus Pnigma Ceratochloa Ovadiahu-Yavin 1969 Zeobromus Stenobroruus Zema Present paper 1973 Bromium Stencbromus Pnigma Ceratoc~loa Neobromus Dumort. • Grisebach, Dt.mortier, (Beauv.) Benth. (Shear) Hitch.• 1823 1343-45 1823 1883 1935

1Groups treated as genera by the autho,s cited are printed in capital letters, and those treated as sub-divisions of the genus are in regular type. 8 is mentioned in various treatments of the family Gr~~ineae in regard to

separation of genera, is the 2-toothed apex of the lemma. Parodi (1947)

has indicated that there are exceptions to the 2-toothed condition. In

my observations I find that only in the sections Stenobromus and Neo- bromus is the 2-toothed condition generally found. As Wagnon (1950)

points out, the perpetuation of the belief that the 2-toothed condition

exists throughout the genus can perhaps be attributed to the delicate

•· nature of the apex of the lemma, and to the fact that the student of

grasses often will split the tip in an attempt to determine if it is

2-toothed or not, and will create the condition that he is looking for.

Cytology and Generic Relat'ions

Gould (1968:89) gives the typical basic (x) chromosome numbers

for the six sub-families of grasses presently recognized among grass

cytotaxonomists as follows:

Festucoideae X = 7 Panicoideae X = 5, 9, 10 Eragrostoideae X = 9, 10

Bambusoideae X = 12

Oryzoideae X = 12 Arundinoideae X = 6, 12

The taxonomic significance of the grass chromosomes was first

realized by Avdulov (1931), who ~ade chromosome counts for 232 species. He recognized three types of chromosomes in grasses: the first having a

basic number of x = 10 or 9, and small chromosomes; the second with x =

12, and small chromosomes; but the third with x = 7 and large chromosomes. These chromosome types were found to be closely associated with

some other characteristics such as organization of the resting nucleus, 9 the kind of starch grains in the endospe1.·m, geographical distribution,

and especially the leaf structure.

Bromus falls under the sub-family Festucoideae; has the basic

chromosome number of x = 7_; and simple starch grains of elliptic,

elliptic-round, or rarely reniform shape. Wagnon (1952:421-425) dis-

cusses the various proposed affinities of Bromus, and cites some early

work by Nageli (1858) and Harz (1880-82) on starch grains and hybrid-

, ization. Harz pointed out that species with compound starch grains will

not hybridize with those having simple ones. The majority of the grasses

in the tribe Hordeae have simple starch grains while most of the Festu-

ceae tribe have compound grains. For this reason Harz classified the

bromes close to Hordeum, Elymus, and the other genera of the tribe

Hordeae. Tateoka (1962) examined starch grains of 766 grasses and con-

cluded that the type of starch grains varied within some species and

could not be used for differentiation of the major taxonomic categories.

At best, we can say that there has been a lack of agreement in

regard to the affinities of Bromus to othe~ grasses. Linneaus had no

doubts about any of the grasses that he lumped together with his B. secalinus group of 1737 being closely related. The original clear con-

cept of the genus became somewhat muddied in later years with the addi-

tion of species that we recognize now as Festuca, Vuluia, and Brachy-

podiu~. Many authors pla-.::e ~:lror~us_next to Festuca in their scheme of classification, as they readily recognize the superficial resemblance.

Krause (1903), Holmberg (1926), and Hayek (1932-33) followed Harz in placing Bromus next to members of the Hordes.e.. Holmberg further

felt that Bromus and Brachypodium should be placed together because of the similarity of the hairy terminal appendage of the ovary, and he 10 created the subtribe Brachypodiinae within the tribe Hordeae for the two

genera. Hayek gave the Brachypodiinae tribal rank, and placed it between

the tribes Bambuseae and Triticeae (Hordeae).

The classification system of Bentham (1881) had 13 grass tribes

grouped in two sub-families, the Festucoideae and·Panicoideae. Bews

(1929) used the Bentham system in his treatment of the grasses of the

world, as did Hitchcock (1920, 1935, 1951) with minor modifications for

, the classification of the grasses of the United States. Until recently

it has been easier to simply adopt some former popular treatment of the

higher taxa, rather than add to the confusion with further speculation.

These former systems are all based primarily on morphological characters,

as are the works of those who would dismember the genus. Principal

among the latter is Nevski (1934) who recognizes Bromus only in the

narrow sense of including just the~- secalinus group.

About the time that Hitchcock was completing his manual, there

were two significant systems of grass classification published: in

Russian by Avdulov (1931), and in French by Prat (1936). These two were

based to a large extent on microscopic characters, and arranged according

to the genetic and evolutionary relationships. The various divisions

into sub-families and tribes as treated furing the past century is

presented very briefly in Table 2. Furth~r information concerning the

treatment of the higher taxa is beyond the scop0 of this thesis and can be found in Gould's Grass Systematics (1968). Evidence to show unity of the genus c2.me from the work of mycologists Ward (19O2a &b), Freeman (1902), and Salmon (1903 & 1904). They indicate that the uredo stage of the parasites Puccinia dispersa

Erikss. & Herin. and Erysiphe gray.:.inis DC., are closely restricted to 11

Table 2. Sub-families and Tribes of the Family Grarnineae.

Avdulov Hitchcock Stebbins Gould 1931 1935 1956. 1968 Subfamily Subfamily Subfamily Subfamily POATAE FESTUCOIDEAE FESTUCOIDEAE FESTUCOIDEAE Festuciformes Bambuseae Hordeae Festuceae (Agrosteae, Festuceae Festuceae Aveneae Aveneae, Hordeae Phalarideae Triticeae Phalarideae, Aveneae Aveneae Meliceae Festuceae, Agrostideae Agrostideae Stipeae and Hordeae Zoysieae Brachyelytreae of Hackel'· s Chlorideae PANICOIDEAE Diarrheneae Nardeae System of Phalarideae Paniceae 1887) Monermeae Oryzeae Andropogoneae Zizanieae Maydeae PANICOIDEAE Phragmitiformes Arundinelleae (Arundineae, PANICOIDEAE Danthonieae Paniceae Bambuseae, & Melinideae Andropogoneae etc.) Paniceae ERAGROSTOIDEAE ERAGROSTOIDEAE SACCHAFIFERAE Andropogoneae Zoysieae Tripsaceae Chlorideae Eragrosteae Sporoboleae Eragrosteae Chlorideae Chlorideae Pappaphoreae Zoysieae Andropogoneae Aeluropodeae Zoysieae BAMBUSOIDEAE Unioleae Tristegineae Pappaphoreae Maydeae Bambuseae Orcuttieae Paniceae Aristideae ORYZOIDEAE Oryqeae BA..~BUSOIDEAE Bambuseae ARUNDINOIDEAE Phareae Arundineae Stipeae ORYZOIDEAE Oryzeae

ARUNDINOIDEAE Arundineae Danthonieae Centotheceae 12 certain allied species. They found intermediaries or "bridging species"

which served in the passage of the fungi fro~ one section of the genus

to another. The study needs to be carried out using modern techniques

and a better understanding of the parasites and host plants.

Another study bearing upon the relationship of the various

entities within the genus also provided indirect evidence that Bromus

may be more closely related to Agronyron than to the remainder of the

Triticeae (Hordeae) tribe. Cugnac and Beval (1941) discovered that f· Bromus contains a levoratatory glucoside which differs chemically and

physically from that found in many other grasses, including Festuca,

but was related to the triticin found in species of Agropyron.

Cytogenetic studies by Stebbins, Tobgy, and Harlan (1941),

Knowles (1944), and by Stebbins (1947, 1949) advance further the evidence

of relationships among the sections of the genus Bromus. 13

Table 3. Chromosome numbers reported for Utah Bromus.

Section Name of Species 2n Studied by*

Bromium B. japoniaus Thunb. 14 Taeoka, 1934; Marie et Kondo, 1949a; Ovadiahu-Yavin, 1969. 28 Avdulov, 1928; Schulz-Schaeffer, 1956, 1960. B. aommutatus Sehr. 14 Felfoldy, 1947a; Schulz-Schaeffer, 1962; Sakamoto &Nuram, 1962. 28 Litardiere, 1950. 56 Nielsen, 1939. B. Paaemosus L. 28 Knowles, 1944; Hill, 1965; Schulz-Schaeffer, 1956. B. seaaUnus L. 14 Nielsen, 1939. 28 Knowles, 1944; Schulz-Schaeffer, 1960; Cugnac & Simonet, 1941. B. squal'Y'OSUSL. 14 Schulz-Schaeffer &Markarian, 1957; Schulz-Schaeffer, 1960. B. moUis L. 28 Knowles, 1944; Bowden & Sen, 1962; Stebbins & Love, 1941. B. bPizaeformis 14 Schulz-Schaeffer, 1960; F. &M. Avdulov, 1928.

Stenobromus B. rubens L. 28 Knowles, 1944; Reese, 1957; Ovadiahu-Yavin, 1969. B. teato!'Um L. 14 Cugnac & Simonet, 1941; Knowles, 1944; Barnett, 1955; Taeoka, 1956; . Bowden, 1960; Ovadiahu-Yavin,1969 . B. steriUs L. 28 Stahlin, 1929; Taeoka, 1955; Schulz-Schaeffer, 1956; Ovadiahu- Yavin, 1969. B. Pigidus Roth. as 42 Cugnac & Simonet, 1941; B. vn,.Zosus 56 Stebbins and Love, 1941. Forssk. 56, 70 Beck & Horton, 1932. as B. rigens L. 56 Schulz-Schaeffer, 1957. B. rigidus Roth. 42, 56 Schulz-Schaeffer, 1956. 56 Taeoka, 1955; Taeoka &Mark, 1957; Bowden & Sen, 1962. B. r1:gidus Roth. ssp. rigidus 42 Ovadiahu-Yavin, 1969. ssp. gussonei 56 Ovadiahu-Yavin, 1969. B. diandrus Roth. 28 Hubbard, 1954. as B.rigidus Roth. 42 Schulz-Schaeffer, 1956. 28 Stebbins & Love, 1941.

*Cited .in Darlington and Wylie, 1955; Cave, 1958-60 & 1961; Munz, 1965; and Ovadiahu-Yavin, 1969. 14

Table 3. (continued)

Section Name of Species 2n Studied by

Pnigma B. anomaZus Rupr. 14 Wagnon, 1952; Reeder, 1971 (Mexican material). B. inermis Leyss. 56 Hill &Meyers, 1948; Schulz- Schaeffer, 1960; Bowden, 1960 (for all ssp. &var.). B. porteri (Coult) Nash as 28 Schulz-Schaeffer, 1960. B. anomaZus Rupr. 14 Wagnon, 1952; Bowden, 1960. B. canadensus Michx. 14 Stebbins &Tobgy, 1944; Schulz- as B. aiZiatus L. Schaeffer, 1960; Wagnon, 1952; Bowden, 1960. B. vuZgaris (Hook.) 14 Stebbins &Tobgy, 1944; Wagnon, Shear. 1952.

Ceratochloa B. marginatus Nees 42 ~ielsen & Humphrey, 1937. 56 Stebbins &Tobfy, 1944; Stebbins, 1947. 70 Nielsen, 1939 (Wyo., prob. rare). B. marginatus var. 56 Stebbins &Tobgy, 1944; Schulz- poZyanthus as Schaeffer, 1960; Schulz- B. potyanthus Ser. Schaeffer and Markarian, 1957. B. carinatus Hook. & 56 Stebbins &Tobgy, 1944; Stebbins Arn. Tobgy &Harlan, 1944b; Stebbins, 1947; Stebbins & Love, 1941. B. wiZZdenowii .Kunth 42 Stebbins & Tobgy, 1944; Stebbins, as B. catharticus 1947; Schulz-Schaeffer, 1960. Vahl. B. arizonicus (Shear) 84 Stebbins, Tobgy, &Harlan, Stebbins 1944b. B. brevia:r>istatus 56 Stahlin, 1929. Buckl. 14 Avdulov 1928; Schulz-Schaeffer 1960. B. unioZoides HBK 42 Schulz-Schaeffer & Markarian, 1957; Bowden &Sen, 1962.

Neobromus B. trinii Desv. 42 Knowles, 1944; Bowden & Senn, 1962. 15

TAXONOMICTREATMENT

Bromus L. Genera Plantarum 15. 1737.

Calliagrostis Ehrhart, Beitrage 4:146. 1789.

Lasiopoa Ehrhart, Beitrage 4:147. 1789.

Avena Salisb., Prodr. Stirp. 22. 1796. pro parte. Ceratochloa Beauvais, Agrost. 75, t. 15 f. 7. 1812.

Schedonorus Beauvais, Agrost. 99. 1812.

Brachypodium Beauvais, Agrost. 101, 1S5-. 1812. pro parte.

Zerna Panzer, Denkschr. Acad. Munch. 296. 1813.

Tragus Panzer ex Jacks., Ind. Kew. 2:1099. 1895. pro parte.

Michelaria Dumortier, Obs. Gram. Belg. 77; t. 16. 1823.

Libertia Lejeune, Nov. Act. Nat. Cur. 12:755; t. 65. 1825.

Trisetum Trinius, Linnaea 10:300. 1836. pro parte.

Festuc~ Drejer, Fl. Exe. Hafn. 43-44. 1838. pro parte.

Aechmophora Spreng. ex Steudel, Norn. ed. 2, 1:29. 1840.

Serrafalcus Parlatore, Pl. Nov. 75. 1842.

Anisantha Koch, Linneae 21: 394. 1848.

Triniu.sia Steudel, Syn. Plantarum Graminearum 328. 1854.

Danthonia C.- \'iuell., Bot. Ztg. 14:348. 1856. pro parte.

Bromopsis Fourneau, Ann. Soc. Linn. Lyon n. ser. 17:187. 1869.

Forasaccus Bubani, Fl. ?yr. 4:385. 1901.

Tricetobromus Nevski, Sc.hed. Herb. Flor. Asiae Mediae VIIIb, Bot., Fa·sc. 17: 14-20. 1934. 16 Plants annual, biennial or perennial, caespitose or less

commonly rhizomatous; culms slender, erect; leaves with broad, flat

blades, sheaths closed below the collar forming a cylinder; inflor -

escence a panicle, or sometimes a raceme, bearing large spikelets with

several (usually S to lO))florets; disarticulation above the glumes and

between the florets; glumes unequal, and usually awnless, acute, first

glume 1 or 3 nerved, the second 3 or 5 nerved; lemmas 5 to 9 nerved, the

,. awn a continuation of the center nerve arising from below the rounded

apex or from the entire or 2-toothed apex; palea about 2/3 the length of

the lem.~a, ciliate on the keels, keels in marginal position, and palea

adnate to the caryopsis; stamens 3; stigmas plumose and sessile, arising

laterally below the hairy cushionlike apex of the ovary; basic chromo-

some number x = 7. Type species Bromus secalinus.

Key to the Species of Bromus

1. Plants perennial or biennial; mostly native species (B. carinatus and B. willdenowii of the section Ceratochloa often appear as annuals. Check those· species if the spikelet is flattened and 4 mm. or more in width, and the lermnas keeled and broad rather than accuminate). (2). 1. Plants annual; mostly adventive weeds. (13). 2. Spikelets strongly flattened and at least 4 mm. wide; lemmas keeled and acute at apex. (3). Section Ceratochloa. 2. Spike lets 1;1ore er less terete, usually 3 to 4 mm. wide (spread to 5 or 6 mm. ir. some pr0ssed specimens); lemmas rounded on the back, not keeled, awned from the back just below the rounded apex. (6). Section Pnigma. 3. Lemmas awned, awn over 2 mm. long. (4). 3. Lemmas awnless or near·ly so . , 3. B. willdenowii 4. Plant perennial; bunchgrass of our mountainous areas; mostly over S dm. tall; awns of lemma up to 5 or 6 mm. long; panicle 17 erect to spreading. (5). 4. Plant annual or biennial; up to 4 or 5 elm. tall; awns 6 mm. or longer; panicle open with spreading or drooping branches ...... 2. B. carinatus ' 5. Sheaths mostly conspicuously pilose, sometimes only on collar or lower sheath; lemmas pubescent on back; mature spikelets about 6 mm. wide ...... 1. !· marginatus 5. Sheaths glabrous; lemmas shiny glabrous to scabrous; mature spikelets 4 to 5 mm. wide . . la. !· marginatus var. polyanthus 6. (2) A\'ms very short or lacking; plants with creeping rhizomes; spikelets held erect. (7). 6. Awned lemmas; plants without rhizomes. (8). 7 • Lemmas g 1abrous ...... • ...... • . 4. B. inermis 7. Lemmas pubescent near the margin; blades pilose on upper surface ...... 4a. B. ·inerrnis var. purpurascens 8. Second glume 5 to 7 nerved with acute apex. (3). 8. Second glume 3 nerved and somewhat rounded at apex. (9). 9. Lemmas pubescent rather evenly over the back; first glume 3 nerved. (10). 9. Lemmas with conspicuous pubescence on margin and glabrous on the back toward the apex; first glume 1 nerved. (12). 10. Panicle drooping, usually not more than 10 cm. long. (11). 10. Panicle open and larger, branches drooping; sheaths glabrous • • ...... • • 6. B. ftondosus 11. Sheaths sparsely pilose to sub-glabrous 5. !· porteri 11. Sheaths woolly ...... Sa. !· porteri var. lanatives 12. Ligule inconspicuous, about 1 mm. long; awns 3 to 5 mm. long ...... • 7. B. canadensis 12. Ligule prominent, 3 to 5 mm. long; awn:; 5 mm. or longer ...... 8. B. vulgaris 13. (1) Awn straight or divaricate, or nearly lacking (somewhat flattened and with a single twist sometiines in!· japonicus). (14). 13. Awn geniculate and tidsted at the base; teeth 0f lemma aristate. (26). Section Neobromu.s. 14. Spike lets and lem;nas broad; lemmas rounded at the apex. (15). Section Bromium, 18 14. Spikelets and lemmas narrow, accuminate; awn as long or longer than the lemma. (22). Section Stenobromus. 15. Lemmas awned from the back 1 or 2 mm. below the rounded apex. (16). 15. Lemmas awnless or nearly so; spikelets large and lemmas inflated, rattling when moved 14. · B. brizaeformis 16. Lemmas glabrous to scabrous. (17). t6. Lemmas pubescent; panicle compact ..•. 13. B. mollis 17. Panicle branches erect to stiffly spreading. (18). 17. Panicle branches lax or flexuous. (21). ,. 18. Foliage pubescent. (19). 18. Foliage glabrous ...... 12. B. secalinus 19. Panicle open; branches spreading or drooping. (20). 19. Panicle contracted and dense; branches erect to ascending .... 11. B. racemosus 20, Awn of first lemma 4 to 5 mm. long and·others about twice as long . . • . . • . . . 9. ~- j aponicus 20. Awns about equal on upper and lower florets, mostly 5 to 6 mm. long ...... 10. B. commutatus 21. (17) Panicles mostly 10 to 25 cm. long; awn often twisted and divergent at maturity; awn of lowest floret only half the length of the awns of the upper florets 9. ~- j aponicus 21. Panicles mostly- less than 10 cm. long; awns straight and all about the same length ...... 10. B. commutatus 22. Panicle erect and contracted into a dense head; in dry and often saline soils of the south and west in Utah ... 15. B. rubens 22. Panicle open; branches erect to nodding. (23). 23. Pedicels flexuous and very slender; panicle nodding; spikelets, including awns, about 3 cm. long. (24). 23. Pedicels stiff; spikelets over 4 cm. long. (25). 24. Spikelets pubescent 16. B. tectorum 24. Spike] ets glabre:Js . . .16a. B. tectorum var. glabratus 25. Mature spike lets 4 to 6 cm. long, including awns; first glume about 8 mm. long . . . 17. B. sterilis 25. Mature spikelets over 6 cm. long; first glume about 15 mm. long 18. B. diandrus 26. (13) Only one species in th:i_s section 19. B. trinii 19 Section 1. Ceratochloa (Beauv.) Benth., Genera Plantarum. 1883.

Plants perennial, biennial or annual; Native grasses of the

Americas with large, distinctly flattened spikelets; glumes and lemmas

keeled; the South American representatives of this section are hexa-

ploid species, and the North American ones are octoploids. The octo-

ploid species are thought to have arisen through allopolyploidy invol-

ving hexaploid species o~ this section and dipicid species of the sec-

, tion Pnigma (Bromopsis). Stebbins (1947) provided the evidence to

indicate such crossing in the past through his studies of chromosome

pairing.

1. Bromus marginatus Nees

Bromus marginatus Nees ex Steudel, Synopsis Plantarum Graminearum 1:322. Stuttgartiae. 1855. Columbia River, Douglas.

Bromus hookeri var. marginatus (Nees) Fourn., Mex. 2:127. 1886.

Certachloa marginata (Nees) Jacks., Ind. Kew. 1:487. 1893.

Bromus marginatus var. seminudus Shear, U.S. Dept. of Agric., Div. of Ag~ost. Bull. 23:55. 1900.

Bromus marginatus var. latior Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:55. 1900.

Bromus flodmanii Rydb., Torrey Bot. Club Bull. 36:538. 1909.

Forasaccus marginatus (Nees) Lunell, Amer. Midland Nat. 4:225. 1915.

Bromus latior Rydb., Flora of tr..e Rocky Mountains 89. 1917.

Perennial; culrns mostly S to 10 dm. tall; leaf sheaths variously

pubescent to retrorsely pilose, pubescent only at collar of sheath or on

lower sheaths in some, ligule about 2 mm. and somewhat ragged, leaf

bladc-s spar.sely pubescent on lower surface, mostly 4 to 8 mm. wide (up

to 12 mm.), 1.5 to 3 dm. long; p_anicle open at maturity but not widely 20 spreading; spike lets mostly 25 to .35 rnrn. long including awns, short pubescencP. extending to the panicle branches in many specimens, often with purplish cast; glumes unequ.:1-l, acute, the first 3 nerved and 7 to

9 mm. long, the second 7 to rarely 5 nerved and 8 to 10 mm. long; lemmas pubescent at least near the base, mostly 13 to 16 mm. long, awned from the tip of lemma, the awn up to 5 or 6 mm.; Chromosomes 2n = 56.

Mountain brome is a native to the Rocky Mountains a.nd Pacific coast regions. It is closely related to California brome, B. carinatus, and differs mainly in being a perennial and larger in most respects.

Mountain brome is a valuable range forage crop, and is probably our most common native brome grass. It is found throughout the State in the mountainous regions at elevations from about 4000 to over 9000 feet. !· marginatus is found in all of the Western States, and as far east in the central pains states as Illinois.

The Indians of California found that the grain of this grass was large enough to be used as Pinole (Chesnut, 1902).

Figure 1 on page 26, indicates.the range of types that are found in Utah. The lemma labelled Dis pubescent only at the base, shows the typical hyaline maTgin, a.nd the long awn that approaches the ~- carinatus complex. LemmaEis pubescent all over, and has the more typical short awn. Every intermediate can be found. Note also the diminutive nature of the nerve next to the keel in drawing C of the glumos. The distribution map is Fi~re 3 on page 32.

Representative ccllections. UTAH: Beaver Co.; Buss 156, Puffer

Lake, 24 July 1964 (BRY). Box Elder Co.; Harrison 12.659, Raft River

Mtns., 21 June 19$6 (BRY). Cache Co.; Cronquist 519-37, Logan, 4 June

1937 (UTC). Davis Co.; Hull 33, Centerville Cnyn., 26 June 1936 (UTC). 21 Duchesne Co.; Hobson 3, Iron Mine Creek, 25 June 1939 (NY, BRY, UTC).

Garfield Co.; Harrison 9147, Calf Creek Falls, 11 May 1939 (BRY). Grand Co.; Welsh &Moore 2818, Courthouse Wash, 30 May 1964 (BRY). Iron Co.; Parley Dalley 2, Cedar City,' 17 July 1939 (SUSC). Juab Co.; Stanton 691,

Deep Creek Mtns., 8 July 1932 (BRY,UT). Millard Co.; Cottam 13372,

Pioneer Ranger Station, 3 Aug. 1952 (BRY, UT). Kane Co.; Stanton 749, Long Canyon, 25 June 1932 (UT). Morgan Co.; Anderson 933, East Canyon, 22 June 1968 (BRY). Rich Co.; Harrison &Larsen 7972, Laketown, 22 June 1934 (BRY). Salt Lake Co.; Anderson 393, Emigration Canyon, 22 June 1968 (BRY). San Juan Co.; Maguire &Redd 1669, Abaja Mtns., 28 June 1932 (UTC). Summit Co.; Hobson 145, Hewinta Ranger Station, 8 July 1939 (BRY).

Tooele Co.; Frischknecht 74, Bennion Creek, 8 June 1961 (USPS, BRY). Uintah Co.; Vickery &Wiens 1626, East Tavaputs Plateau, 13-15 July 1956 (UT). Utah Co.; Larson 7060, N. Provo Park, 9 June 1933 (BRY). Wasatch

Co.; Matthews 109, Strawberry Valley, 23 July 1967 (USPS, BRY). Washing- ton Co.; Cottam 8779, Santa Clara Creek, 17 June 1941 (BRY). Weber Co.;

·cardun s.n., Ogden Canyon, 8 June 1937 (UTC). ARIZONA:Coconino Co.; Mirand & Deaver 5715, W. of Ashfork, 26 May 1960 (ASC, BRY). Maricopa

Co.; Higgins 3988, 20 ~i. NE of Tempe, 23 Apr 19.71 (WYS, BRY). Mohave

Co.; Cottam 13679, Mt. Emma, 25 May 1953 (UT, BRY). Gila Co.; Gould 4699 Reynold's Creek, 31 ~ay 1948 (UTC). Pima Co.; Crossin 34, Picture Rock

Pass, 27 Mar 1963 (ASC, BRY). CALIFORNIA:Marin Co; Long 211a, Mill

Valley, 4 May 1933 (UC, BRY). Tulare Co.; Long 247a, Sequoia Nat'l Park,

8 Aug 1934 (UC, BRY). Ventura Col; Beetle 4310, Mt. Pine, S June 1946

(BRY). COLORADO:Montezuma Co.; Welsh 1810, N. of Mancos, 5 Aug 1962

(BRY). IDAHO: Camas Co.; Patton 153, 25 Mi. N. of Fairfield, 25 June 1970 (BRY). NEVADA:Humboldt CQ.; Holmgren & Reveal 1362, Sonoma Range, 22 · 10 July 1964 (NY,. BRY). Lincoln Co.; Hall s.n., Deer Lo.dgy, 3 July 1935

(BRY). Nye Co.; Beatley 5657, Forty-mile Canyon, 21 May 1968 (BRY). White

Pine Co.; Cotta.'Tl 8101, Mt. Wheeler, 12 June 1940 (UT, BRY). OREGON:

Crook Col; Erdman 2344, Orhoco Mtns., 24 July 1965 (SLP, BRY). WYOMING: Carbon Co.; Beetle 4816, Brush Creek Ranger Sta., ·16 June 1947 (BRY).

Fremont Co.; Anderson 1101, Atlantic City, 5 Aug 1968 (BRY). Lincoln Co. Maguire & Piranian 22780, Smith's Fork Ranger Sta., 6 Aug 1935 (UTC).

Teton Co.; Hitchcock et al. 3859, Moran, 30 June 1938 (UTC).

la. Bromus marginatus Nees var. polyanthus (Scribn.) Anderson, stat. nov. Bromus polyanthus (Scribtler) Shear, U.S. Dept of Agric., Div. of Agrost. Bull. 23:56. 1900. Based on B. multiflorus Scribn. Bromus multiflorus Scribn., U.S. Dept. of Agric., Div. of Agrost. Bull. 13:46. 1898. non.£.· multiflorus Weigel, 1772. Bromus polyanthus var. paniculatus Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:57. 1900. Bromus paniculatus Rydb., Fl. Rocky Mountains 90. 1917.

This variety differs from the species only in pubescence. All measurable morphological features were entirely useless as constants for

separation of these two intergrading varieties. This variety, with its

"shiny glabrous" lemmas is slightly more restric.ted in its range than

var. marginatus. Its distribution extends barely beyond the eastern

slope of the Rocky Mountains, and in Utah it is found primarily along

the mountainous backbone through the center of the State. See Fig. 4

on page 32 for a map cf the distribution.

Separation of the two varieties becomes completely arbitrary and

subject to the interpretation of the observer when these are studied

closely, as pubescence can be found on practically every one of these

grasses. In his revision of th~_genus, Shear (1900) states, "The above 23 species is very ~losely related to!~ marginatus, into which it passes and of which it should perhaps be regarded as a variety. It differs

chiefly from the typical form of~; marginatus in being smoother throughout." I agree. Type locality: Wyoming..

Representative collections. UTAH:Cache Co.; Passey 72, Logan

Cnyn., 18 July 1935 (UTC). Davis Co.; Anderson 95, Bountiful Peak, 29 Sep 1958 (UT). Duchesne Co.; Harrison 10452, Mirror Lake, 13 July 1941

, (BRY). Grand Co.; Harrison 12484, LaSal Mtns., 18 Aug 1954 (BRY). Iron

Co.; Bruhn G57A, Cedar Breaks Nat'l Mon., ca. Aug 1954 (BRY). Juab Co.; Rydberg & Carlton 7595, Mount Nebo, 15 Aug 1905 (UT). Morgan Co.; Anderson 932, East Canyon, 22 June 1968 (BRY). Salt Lake Co.; Anderson 1000, Butterfield Canyon, 29 June 1968 (BRY). Garfield Co.; Weights KEWB-31/69, Bryce Canyon, 16 June 1931 (UT). Piute Co.; Rydberg & Carlton 7176, near Marysvale, 23 July 1905 (R14). San Juan Co.; Cottam

12732, AbajoMtns., 18 Aug 1951 (UT, BRY). Sanpete Co.; Collotzi & Clark 429, Manti Canyon, 10 Aug 1965 (USFS, UTC). Sevier Co.; Coles 62, Solitude Guard Sta., 15 July 1964 (BRY). Tooele Co.; Vickery 1054,

Stansbury Mtns., 20 July 1957 (UT). Summit Co.; Hutchings 100, Uintah Forest, 23 July 1927 (USFS, BRY). Utah Co.; Thomas 187; Dia.mend Fork,

21 June 1969 (BRY). Wasatch Co.; Despain 29, Cascade Springs Road,

5 Aug 1969 (BRY). COLORADO:Garfield Co.; Stewart 16, East Tavaputts Plateau, 21 June 1966 (UT). IDAHO:Franklin; Smith 2175, Preston,

13 June 1910 (RM). MONTANA:Beaverhead Co.; Calvert 195, Odell Creek, , 21 July 1959 (BRY).

2. Bromus carinatus Hook. and Arn.

Bromus car1natus Hooker and Arnott, Botany of Captain Beechey's Voyage -403-:--1s40. California. 24 Ceratcchloa F,"Tandiflora Hook., __Fl. Bor. A1Uer. 2:25.3. 1840. · Bromus oregonus Nutt. ex Hook., Jour. Bot. Kew Misc. 8:18. 1856. · Bromus virens Buckl., Acad. Nat. Science Phila. Proc. 1862:98. 1862. · Bromus californic:us Nutt. ex Buck1., Acad. Nat. Sci. Phil a. Proc. 1862:336. 1862. Bromus nitens Nutt. ex A. Fray, Acad. Nat. Sci. Phila. Proc. 1862:336. 1862. · Bromus hookerianus Thurb. ex Wildes, U.S. Expl. Bot. 17:493. 1874. Bromus hookerianus var. minor Scribn. ex Vasey, Descr. Cat. Grasses U.S. 92. 1885. Bromus virens var. minor Scribn. ex Beal, Grasses N. Amer. 2:614. 1896. Bromus carinatus var. californicus Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:61. 1900. Bromus carinatus var. hookerianus Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:60. 1900.· Brorr,us carinatus var. densus Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:61. 1900.

Erect annual or mostly biennial; culms mostly 5 to 7 elm. tall; sheaths sparsely retrorse-pilose or glabrous; blades sparsely pilose, 5 to 10 mm. wide; panicle erect, 10 to 20 cm. long, the branches rather stiffly ascending_ to slightly reflexed and bearing 1 or 2 (rarely 3) spikelets, the lower branches as much as 10 cm. long; spikelets 2.5 to

3.5 cm. long, 3 to 6 florets; glu.mes sub-equal, :the first 3 nerved, the second 5 or indistinctly 7 nerved; lemmas broadly lanceolate, 7 nerved, scarious margined, smooth to scabrous pubescent, about 15 mm. long; awn

7 to 10 mm. long; Chromosomes 2n = S6.

California brome has been cited in numerous collections in the

Intermountain States, and has been treated in various ways in combin- ation with Bromus marginatus and Bromus polyanthus. The second edition of Hitchcock's Manual treats B. carinatus as more of a coastal species, and the original description of the plant would certainly indicate that 25 our Mountain brome was not the grass that the authors had in mind when they named Bromus carinatus. This problem was carefully studied, and all of our specimens have been ass_igned to the Bronius marginatus group, except for an occasion al rare specimen from the St. Geo_rge area of Utah.

Figure 7A on page 38 shows the typical open nature of the panicle and spikelets. Bromus carirt.atus is facultatively cleistogamous, producing both chasmogamous and cleistogamous florets on the sa.."'lleplant (Harlan,

1945). This grass does not produce fertile hybrids with Bromus arizonicus·, another closely allied member of this complex, even though the two look very much alike· and have very close common ancestory

(Stebbins, Tobgy and Harlan, 1944b). I expect that somewhat the same situation exists between B. carinatus and our mountain species of the section Ceratochloa. There may not be as much genetic exchange between these s_pecies as has been supposed in the past.

Representative collection. UTAH: Washington Co.; Galway s.n.,

St. George, 11 May 1935 (BRY). CALIFORNIA: Alameda Co.; Long 180a,

Berkeley, 2 June 1932 (UC, BRY); Beetle 1717, Berkeley, 29 Mar 1941 (UC, BRY).

3. Bromus willdenowii Kunth

Bromus willdE:nowii Carl Sigismund Kunth, Revision des Graminees 1:134. 1829. Based on Festuca unioloides Willd., Hort. Berol. 3: pl. 3. 1803. Type grown at Berlin; seed from "Carolina".

Bromus cathaTticus Vahl., Symb. Bot. 2: 22. 1791 Lima, Peru. Based, at least in pert, on the pre-Linnaean "Graman bromoides catharticum valgo guilno" of Feuillee, Jour. Obs. Phys. 705, pl. 2. 1714. The plate shows a mixture of two different plants and no type specimen is available. Rejected as nomen confusem (Raven, 1960).

Ceratochloa unioloides (Willd.) Beauv., Ess. Agrost. 75, pl. 15, f. 7. 1812. Fig. 1. Illustration of Bromus marginatus

A. Habit sketch of panicle

B. Sheath, ligule, and culm joint

C. Empty glumes

D. Side view of lemma

E. Back view of a pubescent lemma /smm 27 · Brornus unioloides (Willd.) Raspail, Ann. Sci. Nat., Bot. 5:439. 1825. Non H.B.K. ·1815.

· Ceratochloa pendula Schrad., Linnaea 6: Litt. 72. 1831. Grown at Gottingen from seed £ram Carolina.

Bromus schraderi Kunth, Enum. Pl.. 1:416. 1833. Based on Ceratochloa pendula Schrad. Ceratochloa breviaristata Hook., Fl. Bar. Amer. 2:253, pl. 234. 1840. Lewis and Clark River and near the sources of the Columbia. Douglas in 1826.

Bromus breviaristatus (Hook.) Thurb. in Wilkes, U.S. Expl. Exued. Bot. 17(2):493. 1874. Non Buckl., 1862.

· Tragus unioloides (Willd.) Panz. ex Jacks., Ind. Kew. 2:1099. 1895.

Forasaccus brebia.ristatus (Hook.) Lunell (e-rror for breviaristatus), Amer. Midl. Nat. 4:225. 1915.

· Ceratochloa cathartica (Vahl.) Herter, Rev. Sudamer. Bot. 6:144. 1940.

Annual or biennial; culms erect or spreading, mostly 4.5 to 6.5 dm. tall, with some specimens ranging from 2 to 10 dm.; sheaths usually pubescent--at least lower ones--with upper sheaths having at least a fringe of pubescence along.the margin where sheath is open, sheath encloses culm to within 2 or 3 cm. of collar; culm leaves generally 10 to 25 cm. long, 4 to 7 mm. wide--some to 10 mm., other leaves smaller; leaf blades blabrous or sparsely pilose; ligule conspicuous, rounded and membranous, 3 mm. long; panicle open at maturity, mostly 15 to 25 cm. but as much as 35 cm. long in some, with branches to 15 cm., few spikelets at base of branches; spikelets 4 to 7 mm. wide and 2 to 3 cm. long, mostly 8 to 10 flowered; glumes accuminate, first 9 to 12 mm., second barely longer; lemmas scabrous or pubescent at least on nerves and keel towards the tip, often glabrous below, 9 or 11 distinct nerves, 2 mm. wide from the keel to the scarious margin. accuminate, 13 to 15 mm. long including short terminal awn of 1 to 2 nun.; lemmas closely over- 28 lappi_ng and conceq.li_ng the short rachilla joints; rachilla joints 2 mm. long, rounded~-or sometimes angular--and pubescent on the side away from palea; palea two-thirds as long as lemma; chromosomes 2n = 42. This plant has been widely known as Bromus unioloides HBK, and also as Bromus catharticus Vahl, since Hitchcock (1934) took up the latter name. A discussion of the problem and a citation of the correct name can be found in the Raven (1960) article concerning Rescue grass.

This grass is known as rescue grass and Schrader's bromegrass.

Because of the high quality of rescue grass as a winter forage, it has been cultivated in the Southern States for over 100 years, and is found in those states and rerely northward in humid regions with mild winters.

Bromus willdenwii was collected in Price and St. George in the early 1930s, and since that time has been found in many of the low cultivated valleys of the State as far north as Provo. Utah distribution is shown on map Figure 5 on page 32. The illustrations on page 30 show: A. the habit, B. the sheath and ligule, and C. the lemma.

Rescue grass is a. native of South America, and grows commonly in the pampas of southern Brazil, Uraguay, and north eastern Argentina.

Representative collections. UTAH: Carbon Co.; Stanton 567,

Price, 12 Sep 1931 (UT). Garfield Co.; Anderson 923, Henrieville, 24 Aug

195 7 (UTC). Iron Co.; Dalley 1, Cedar City, 10 Aug 1938 (SUSC); Anderson

96, Cedar City, 14 Aug 1937 (UTC). Kane Co.; Anderson 1183, Kanab, 22

June 1958 (UTC). Piute Co.; Harrison 11037, Circleville, 21 Aug 1946 (BRY). Utah Co.; Nixon 27, BYUCampus, 27 May 1959 (BRY). Washington Co;

Woodbury 8, Pine Valley Mtns., 5 June 1913 (SUSC); Higgins 511, Beaver- dam Mtns., 25 Apr 1966 (BRY); Maguire & Blood 4356, Hurricane, 18 May 29 1933 (UTC). Wayne Co.; Cottam 9294, Fruita, 27 May 1944 (UT). ARIZONA:

Maricopa Co.; Marcus s.n., East Phoenix, 7 Apr 1963 (ASC, BRY).

CALIFORNIA: Inyo Co.; Alexander & Kell_og 2879, Saline Valley Trail,

27 May 1942 (UC, BRY). OKLAHOMA:Payne Co.; Erwin 5802/46, Campus of Oklahoma A &M, 21 Mar 1938 (OKLA, BRY). NORTHCAROLINA: New Hanover Co.; Leonard &Radford 3089, Smith Creek W. of US 117, 14 Apr 1970 (NCU, BRY). T~XAS: San Patricio Co.; Sanderson 92, Encino Lake, 1 June

1965 (BRY). Randall Co.; Higgins 3194, Palo Dura Canyon State Park, 8

May 1970 (WTS, BRY). Harris Co.; Higgins 3916, Houston, 31 Mar 1971

(WTS, BRY). Ochiltree Co.; Headlee 498, ca. 22 miles South of Perryton,

13 May 1971 (WTS, BRY).

Section 2. Pnigma Dumort., Obs. Graminees Belgique. 1823.

Plants perennial; native grasses of the northern hemisphere with more or less terete spikelets; glumes and l~m,~as rounded.

4. Brornus inermis Leyss

Bromus inermis Leysser, ·Flora Halensis~ 16. 1761. Europe.

Festuca inermis (Leyss) DC. Lam., Fl. France. 3:49. 1805.

Schedinorus ineTJ11is (Leyss) Beauv., Ess. Agrost. 99:177. 1812.

Festuca inermis var. vilios::i. Mert. and Koch, Deutschl. Fl. 1:675. 1823.

Bromus inermis var. aristatus Schu!', Enum. Pl. Transsilv. 805. 1866.

Bromus inopinatus Brues, Trans. Wis. Acad. Sci., Arts, and Letters 17:73. 1911.

Forasaccus inermis (Leyss) Lunell, Amer. Midl. Nat. 4:225. 1915.

Zerna inermis (Leyss) Lindm., Svensk Fanerogamfl. 101. 1918.

Bromus inermi s :forma villoc:;nc:; FP.rn.::t1d, Rhodora 3S: ~16. 19~3. Fig. 2. Illustration of Bromus willdenowii

A. Habit sketch

B. Sheath and Ligule

C. Individual lemma

31 Bromus inermis forma aristatus Fernald, Rhodora 35:316. 1933.

Bromus inermis forma bulbiferus Moore, Rhodora 43:76. 1941.

Erect perennial from creeping rhizomes, 5 to 10 dm. tall; leaf sheaths and blades smooth or nearly so, blades 10 to 20 cm. long, 6 to 8 nun. wide; the panicle erect and somewhat open, especially at anthesis; spikelets 5 to 9 flowered, 2 to 3 cm. long, narrowly oblong to lanceolate in outline, terete, frequently purplish; glumes acute to obtuse, first one 4 to 5 mm., second 5 to 6 mm.; lemmas obtuse, and often with a short awn, apex usually smooth and entire but sometimes torn in pressi_ng and drying of specimens; chromosomes 2n = 56.

Smooth or Hungarian brome is native to Europe, Siberia and

China. It was introduced into the United States in 1884 (Wagnon, 1952;

USDA, 1948) and has been grown widely throughout this country. Our native grass of the same type has been recognized in the past as Bromus pumpellianus .. The two were probably segregates from the same origin in

Asia, and were well on theiT way toward speciation, one in the Old World and one in the New World, when they were reintroduced into close prox- imity. B. inermis seems to be the more vigorous of the two varieties, and is displacing the sub-species pumpellianus and interbreeding freely.

Where there is any contact of the two entities they are indistinguish- able, but still are found as distinct varieties in other places

(Elliott, 1949b).

Smooth brome is cultivated as a pasture grass, used for re- seeding and now naturalized and running wild in the Intermountain States from Minnesota and Kansas to Washington and California. It is adapted especially to regions of moderate rainfall and low to moderate summer temperatures. This long-lived perennial has strong creeping rhizomes Figs. 3 to 6. Maps of distribution

3. Brornus rnarginatus

4. B. rnarginatus var. :eolrartthus 5. B. willdenowii

6. B. inerrnis UTAH UTAH County Lines and Cour,ty Lines and • river drainages river drainages Fig.3 Fig.4 A ..I _J.. ..I ..I

I' ------.,,I - ,-' __,,""'-1 __ .1-, __ .l __

UTAH UTAH Cownty lines and County lines and rh,cr d,ainaaes river drainages Fig. 5 .. 33 and fonns a good sod to protect the watersheds. As a pasture or hay

grass there are few that will equal Smooth Brome when in the area of its

best adaptation. There have been hundreds of pages written extolli.ng

the virtues of this grass'for many uses (Scribner, 1894; Yearbook of

Agric., 1948; Komarcv, edit, 1934; and others). In Utah it is widely

distributed in rich noist soil from 4500 to 11000 feet elevation.

For distribution see map Figure 6 on page 32.

Representative collections. UTAH: Beaver Co.; Maguire 19667,

East of Beaver, 19 July 1940 (RM, UTC). Cache Co.; Wasatch Range Project,

Cottam et al. 15387, Beaver Creek, 7 Aug 1958 (UT); Anderson 787, Logan,

12 June 1957 UTC). Daggett Co.; Richins s.n., N. slope of Uintah Mtns.,

.July 1959 (UTC). Davis Co.; Garrett 7629, Bountiful Canyon summit, 4 &6 Sep 1937 (UT). Garfield Co.; Anderson 175, Hatch, 1 Sep 1958 (UT, BRY); Buchanan 54, Bryce Canyon Nat'l Park, 20 July 1956 (UT, BRY). Iron

Co.; Bruhn G60, Cedar Breaks Nat'l Mon., 24 Aug 1954 (BRY, DIX). Juab

Co.;Diehl s.n., Robinson, 10 Aug 1913 (BRY). Millard Co.; Cottam 13381,

Pioneer Ranger St.? 3 Aug 1952 (UT). Piute Co.; Higgins 1172, Monroe

Mtn., 9 Sep 1967 (BRY). Morgan Co.; Anderson 950, W. of Morgan, 22 June

1968 (BRY). Salt Lake Co.; Anderson 892, Emigratiqn Canyon, 22 June 1968 (BRY). San Juan Co.; Cottam &Hutchings 2209, LaSal, 13 June 1927 (BRY).

Sanpete Co.; Stutz 24, Hayfield, 2 July 1966 (BRY); I

13202, Alpine Sta. in Ephraim Cnyn., 15 Aug 1934 (fu~). Sevier Co.;

Anderson 1070, Salina Canyon, 18 July 1968 (BRY); Cottam 9205, Hancock

Flat, 30 June 1943 (UT). Summit Co.; Anderson 941, W. of Henefer, 22

June 1968 (BRY). Uintah Co.; Welsh 505, Dinosaur Nat'l Mon., 2 June 1956

(BRY). Utah Co.; Harrison 306H, Hobblecreek, 23 June 1938 (BRY); Rerun

8667, Aspen Grav~, 10 Sep 1938 (BRY). Wasatch Co.; Foster 187, Provo 34 River, 8 Oct 1964 (BRY). Washi_ngton Co.; Gould 1938, Pine Valley Mtns ..,

26 .July 1942 (DIX). Wayne Co.; sans collector 19283, E. of Fruita (UTC).

Weber Co.; Hermann 12110, SW. of Huntsville, 12 Aug 1955 (UTC).

4a. Bromus inermis var. purpurescens (Hook.) Wagnon

Bromus inermis var. putpurescens (Hook.) Wagnon, Rhodora 52:211. 1950.

Bromus purgans var. purpurescens Hook., Fl. Bor. Amer. 2:252. 1840.

Bromus purgans var. longispicata Hook., Fl. Bor. Amer. 2:252. 1840.

Bromus pumpellianus Scribn., Bull. Torrey Bot. Club 15:9, as to the type. 1888.

Bromus ciliatus var. coloradensis Vasey· ex Beal, Grasses N. Amer 2:619. 1896.

Bromus pumpellianus var. tweedyi Scribn. ex Beal, Grasses N. Amer. 2:622. 1896.

Btomus purnpellianus var. melicoides Shea~, U.S. Dept of Agric., Div. of Agrost. Bull. 23:50. 1900.

Forasaccus pumpeJ lianus (Scribn.) Lune 11, Arner. Midl. Nat. 4:225. _ 1915.

This variety is best identified by its blades which are pilose

at least on the upper surface, by the pubescent culm nodes, the glabrous glumes, and lemmas which are pubescent along the margins and on the nerves. This grass is not at all common in Utah, if present at all, and

can be expected only in the north. It is found from well within the

arctic circle south in the Rocky Mountains to Colorado. I have seen

specimens from Canada and Alaska, but none from Utah.

5. Brornus porteri (Coult.) Nash.

Bromus portcri (Coult.) Nash, Torrey Botanical Club Bull. 22:512. 1895. 35 Bromus kalmii var. porteri Coult., Man. Rocky Mount. 425. 1885.

Btomus ciliatus var. porteri (Coult.) Rydb., U.S. Nat'l Herb. Contrib. 3:192. 1895.

· Btomus ciliatus var; montarius Vasey ex Beal, Grasses N. Amer. 2:619. 1896.

Btomus kalmii var. occidentalis Vasey ex Beal, Grasses N. Amer. 2:624. 1896.

Bromus ciliatus var. scariosus Scribn., U.S. Dept of Agric., Div. of Agrost. Bull. 13:46. 1898.

Btomus scabratus Scribn., U.S. Dept. Agric., Div. of Agrost. Bull. 13:46. 1898.

Bromus kalmii var. major Vasey ex Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:35. 1900.

Stout perennial, mostly 5 to 8 dm. tall; leaf sheaths glabrous below, sometimes hairy at the summit, ligule small, blades flat, smooth to sparsely pubescent, 1 to 2 dm. long, 3 to 4 mm. wide; panicle spreading, 9 to 12 cm. long, and spikelets drooping on flexuous branches up to 5 ·or 6 dm. long; panicle branches pubescent; spikelets plump, not compressed, 5 to 11 flowered, 2 to 3 cm. long; glumes 3 nerved, lower one 5 to 6.5 mm. long, upper one .5 mm. longer, pubescent; lemmas 9 to

11 mm. long with rounded tip, 7 to 9 nerved, evenly pubescent over the back; awn less than 3 mm., an extension of the midnerve; chromosomes

2n = 28 (as~- anomalus Rupr.).

Nodding brome of the Intermountain West has been referred to as

Bromus anomalus Rupr., which is a species with type locality in Hidalgo,

Mexico, and an extended range into western Texas and northward into New

Mexico and Arizona. B. porteri extends south to New Mexico, with its type locality being at Twin Lakes, Lake County, Colorado. B. porteri has first glumes that are consistently 3-nerved, auricles are never present, and the midrib of the foaf is not tapered just below the collar 36 as it is in B. anomalus. .. anomalus has 1 or 3 nerves on the first

glume and frequently has auricles. See Illustration on page 38, Fig. 7B.

Nodding brome is found in the mountains thr~ughout the State at . elevations from 6000 to 9500 feet, and in similar habitat in all of the Intermountain States. See map Figure 10 on page 48. Representative collections. UTAH: Carbon Co.; Welsh &Christen- sen 6533, West Tavaputs Plateau, 10 Aug 1967 (BRY). Cache Co.; Passey 93,

Logan Canyon, 18 July 1935 (UTC). Daggett Co.; Cottam 6140, Manila area,

30 June 1935 (BRY). Daggett Co.; Cottam 6140, Manila area, 30 June 1935 (BRY). Duchesne Co.; Harrison &Nissan 8800, Blind Stream, 3 July 1938 (BRY). Garfield Co.; Boyle 1139, Bryce Canyon, 19 July 1942 (BP.Y). ·Stanton 133, Henry Mountains, 25 June 1930 (BRY). Grand Co.; Holmgren

et al. 2310, Hill Creek, 30 July 1965 (UTC); Payson 4024, LaSal Mtns.,

24 July 1924 (RM). Iron Co.; Maguire 19440, E. of Cedar City, 12 July

1940 (B~Y, UTC); Seaman 125, Lowder Ranger Stat., 18 Aug 1930 (SUSC).

Kane Co.; Gierisch 32, Harris Flat Ranger Stat., 4 July 1937 (RM, UTC).

Millard Co.; Cottam 13375, Pavant Mtns., 3 Aug 1952 (UT, BRY). Piute Co.;

Higgins 1154, Monroe Mtn., 9 Sep 1967 (BRY). Salt Lake Co.; Garrett 2013,

Big Cottonwood, 25 Aug 1906 (UT). San .Juan Co.; Rydberg & Garrett 9205,

E. of Monticello, 25 July 1911 (RM). Sanpete Co.; Hull 9578, Ephraim Canyon, 16 July 1939 (BRY); Collotzi &Clark 430, Manti Canyon, 10 Aug 1965 (UTC). Sevier Co.; Markhu.m 8923, Fish Lake area, Aug 1938 (BRY);

Rickenback 159, S. of Glenwood, 25 Aug 1965 (UTC). Summit Co.; Peterson

94, Eridger Lake, 27 Aug 1965 (UTC). Uintah Co.; Garrett 7830, Barker

Spring, 25 July 1938 (BRY, UT). Utah Co.; Thomas 191, Diamond Fork, 21

June 1969 (BRY); Garrett 5649, Mnt. Timpanogus, 1 Aug 1930 (UT). Wasatch

Co.; Eastmond s.n., Strawberry Re.s., Aug 1966 (BRY). Wayne Co.; Harrison 37 .12374, N. of Cycl_one Lake, 6 Aug 1954 (BRY). Washington Co.; McAllister

41, Arizona Strip, summer19.38 (UTC). COLORADO:Gunnison Co.; Hall 459,

12 July 1961 (UT, BRY). Montezuma Co.; Welsh 1807, 5 Aug 1962 (BRY).

Saguache Co.; Higgins 2270, 19 July 1969 (BRY). NEVAD: Clark Co.; Clokey 8277, 28 July 1939 (BRY). Elko Co.; Holmgren 1552, 27 July 1941 (BRY). Humboldt Co.; Holmgren &Reveal 1362, 10 July 1964 (BRY, UTC). WYOMING:Fremont Co.; Anderson 1100, 5 Aug 1968 (B~Y).

Sa. Bromus porteri var. lanatipes Shear

Bromus porteri var. lanatipes Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:37. 1900.

Bromus lanatipes (Shear) Rydb., Colo. Agr. Expt. Sta. Bull. 100:52. 1906.

Bromus anomalus var. lanatipes (Shear) Hitchc., Wash. Acad. Sci. Jour. 23:449. 1933.

TI1is variety of nodding·brome may not be present in Utah, but is reported for Colorado and Arizona to Western Texas. It is said to be more robust, with woolly sheaths and usually broader blades (Hitchc.,

1952:46).

6. Bromus frondosus (Shear) Woot. & Standl.

Bromus frondosus (Shear) Woot. & Stand!., New Mex. Exp. Stat. Bull. 81: 144. 1912.

Bromus Porteri var. frondosus Shear, U.S. Dept. of Agric., Div. of Agrost. Buli. 23: 37. - 1900.

, This grass has beel' reported in Utah collections, but the specimens cited have been identified with other species in this treatment.

In the original description of this plant, it is reported to differ from

Bromus porteri in having "very weak, leafy culms, rather narrower and Fig. 7. Illustrations of Bromus carinatus, !- porteri, and B. trinii A. Portion of Bromus carinatus panicle

B. Spikelet of Bromus porteri C. Spikelet of Bromus trinii cm 39 less densely flowered spike lets with the empty glumes smooth'.' (Shear,

1900:38). Wagnon (1952:471) gives a distribution limited to New Mexico,

Arizona, .and northern Mexico. A description of this grass as it occurs in Utah must of necessity be deferred until such time as a veri- fied collection is made.

7. · Brom.us canadensis Michx.

Bromus canadensis Michaux, Flora Boreali-Americana 1:65. 1803.

Bromus ciliatus auct. Fl. N. A.~er. non L.

Bromus richardsoni Link, Hort. Berol. 2:281. 1833.

Perennial, slender and tall, 6 to 12 dm.. ; Leaf sheaths on culm glabrous to pubesc~nt, often with fringe of long pubescence at collar, auricle somewhat developed, leaves broad, 4 to 12 mm., and 1 to 3 or sometimes 4 dm. long; panicle open and spreading, some cf the branches drooping, usually glabrous but some specimens with scabrous to pubescent branches; spikelets 5 to 10 flowered, 2 to 3 cm. long; lower glume 1 nerved, upper with 3 nerves, glabrous or sometimes scabrous on the nerves only; upper glume has small mucro in some specimens; lemma 5 or 7 nerved, 7 to 12 mm. long, long soft pubescence on margins and base, glabrous on back toward the tip; awn 3 to 7 mm. long; chromosomes 2n = 14.

Fringed brome grass is frequent in the mountains from 6000 to

11000 feet in meadows and on rocky slopes and ridges. It is widely distributed in Utah (Figure 11 on page 48), and in most of the northern and western states. This grass is known also as Bromus richardsoni Link, or

Richardson's brome grass. In his revision of the section Bromopsis

(=Pnigma), Wagnon (1952) recognizes both as distinct species. In exam- ining the specimens I was unable.to find any characteristics constant 40 eno_ugh to separate them into groups, even though I followed the descrip-

tions with care. I rather hold with the opinion of Hitchcock (1935:42,

44), that "it (Bromus richardsoni) grades freely into B. ciliatus and

can scarcely be ranked even as a variety." See Figure 8 on page 41.

The recognition of the existence of a holotype of B. ciliatus

and its previous identification by Hitchcock as the grass we know in

America as!· latiglumis (Shear) Hitchc. has made some nomenclatural

changes necessary. Bromus ciliatus L. becomes the correct name for B.

·1atiglumis (Shear) Hitchc. and fringed brome, which we have known as

B. ciliatus must receive another epithet. B. ciliatus of the North

American botanists should now be called B. canadensis Michx. (Baum, 1967}

Representative collections. UTAH: ~eaver Co.; Maguire 19843,

Big John Flats, 24 July 1940 (RM). Cache Co.; Passey 81, Logan Canyon,

2 Aug 1934 (UTC). Carbon Co.; Van Cott 131, Helper-Duchesne Summit, 9

Aug 1940 (UTC). Ducnesne Co.; Harrison et al. 10017, Third Chain Lake,

27 July 1940 (BRY). Garfield Co.; Stanton 148, Henry ~1tns., 25 June 1930

(BRY); Holmgren et al. 2459, Aquarious Plateau, 13 Aug 1965 (BRY, UTC);

Hall 13593, SW. of Escalante, 17 Aug 1951 (UT). Grand Co; Maguire et al.

16405, LaSal Mtns., 22 July 1933 (UTC). Iron C0.; Cottam 14144, Cedar

Breaks, 6 Aug 1955 (UT). Juab Co.; McMillan 1361, Deep Creek Mtns., 6

Sep 1947 (UTC). Millard Co.; Cottam(?) 13375, Pavant Mtns., (UT).

Piute Co.; Coles 100, Grindstone Plat, 14 Aug 1964 (BRY). Salt Lake Co.;

Maguire 17352, Brighton, 7 Aug 1939 (UTC, BRY). San Juan Co.; Cottam

12727, Abajo Pass, 18 Aug 1951 (UT). Sanpete Co.; Wasatch Range Project,

Cottam et al.16061, Hop Creek, 3 Sep 1959 (UT). Sevier Co.; Maguire

19914, U.O. Pass Springs, 5 Aug 1940 (UTC, BRY). Summit Co.; Maguire et

al. 14627, Henry's Fork, 16 Aug 1936 (~~. UTC). Utah Co.; Ream 8679, Fig. 8. Illustration of Bromus canadensis

A. Habit sketch

B. Glumes and lenL~a

c. Back view of lemma \

I I 42 Aspen Grove, 30 July 19.38 {BRY),; Flowers 606, Timp Creek, 6 Aug 1925

(UT)~ Uintah Co.; Walkup 118, Mosby Ctn., 19· Aug 1930 (UTC). Wasatch

Co.; Adair 43, Wolf Creek, 7 Sep 1929 (BRY). Washi_ngton Co.; Cottam 5692, Pine Valley Mtns., 23 Aug 193.3 (BRY). ARIZONA:Coconino Co.;

Romans 9b, 13 Aug 1968 (UTC). COLORADO:Routt Co.; Sans collector 162,

22 July 1966 (UTC). Mesa Co.; Hennann 12194, 17 Aug 1955 (UTC). Boulder

Co.; Murdock 412, 27 June 1962 (BRY). IDAHO: Benewah Co.; Cronquist &

Jones 5964, 17 July 1949 (UTC). Blaine Co.; Cronquist 3695, 8 Aug 1941

(UTC). Clark Co.; Cronquist 867-37, 29 Aug 1937 (UTC). Clearwater Co.; Cronquist &Jones 5989, 24 July 1949 (UTC). NEVADA: Clark Co.; Clokey

8135, 6 Aug 1938 (BRY). Elko Co.; Holmgren 1791, 16 Aug 1941 (UTC, BRY).

Lincoln Co.; Galway 8423, 16 July 1935 (BRY).• White Pine Co.; Holmgren & Reveal 1625, 12 Aug 1964 (BRY). WYOMING:Teton Co.; Anderson 835, 10 Aug 1957 (UTC).

8. Bromus vulgaris (Hook.) Shear

Bromus vulgaris (Hook.) Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:43. 1900.

Bromus purgans var. vulgaris Hook, Fl. Bor. Am. 2:252. 1840.

Bromus ciliatus var. ligulat1.,s Vasey ex Macoun, Cat. Can. Pl-:-T(4):238. 1888.

Bromus ciliatus var. pauciflorus Vasey ex Macoun, Cat. Can. Pl. 2(4):238. 1888.

Bromus debilis ~{utt. ex She:i.r, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:43. 1900.

Bromus vulgaris var. eximus She~r, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:44. 1900.

Brornus vulgaris var. robustt,s Shear, U.S. Dept. of Agric., Div. of _Agrost. Bull. 23:44. 1900. Bromus ciliatus var. glaberrimus Suksdorf, Deuts. Bot. Monats. 19: 93. 1901. 43 Bromus eximius Piper, Contrib. U.S. Nat'l Herb. 11:144. 1906.

Btomus eximius var. umbraticus Piper, Contr. U.S. Nat'l Herb. 11:144. 1906.

Zerna vulgaris (ljook.) Henrard, Bltm1ea 4(3) :498. 1941.

Perennial; culms erect or spreading, 6 to 12 dm. tall, the nodes pubescent; leaf sheaths pilose, blades to 25 cm. or occasionally longer and 6 to 12 mm. wide; ligule conspicuous, 3 to 5 mm. long; panicle 10 to 15 cm. long, the branches drooping; spikelets narrow, 1.5 to 3 cm. long with 4 to 9 florets; lower glume acute, 1 or occasionally 3 nerved, 6 to 8 mm. long, upper glume 3 nerved, 8 to 11 mm. long; lemmas 9 to 12 mm. long, pubescent on the margins and glabrous to pubscent over the back, 7 nerved; awn 5 to 11 mm. long; chromosome number 2n = 14.

This grass is found growing in the dense shade, and is seen only rarely in Utah. It is found generally north to Alberta and British

Columbia. Figure 9 on page 44 illustrates the distinctive ligule (B), the loose nature of the spikelets (A), and the habit (C & D). Representative collections. UTAH: Salt Lake Co.; Arnow 2007, Red Butte Canyon (UT). IDAHO: Elmore Col; Hitchcock &Muhlick 10215 (UTC). WYOMING:Teton Co.; Anderson 467 (UTC). Park Co.; Anderson 1232, Yellow- stone Park (UTC).

Section 3. Bromium Dumort., Obs. Graminees Belgique. 1823.

Plants annual; int::...·oduced weedy spe.:ies with broad spike lets; lemmas broad and rounded at the apex.

9. Bromus japonicus Thunb.

Bromus japonicus Thunberg, Flora Japonica, Lipsiae. 1784.

Bromus patulus Mert. and Koch, Deut. Fl. 1:685. 1823. Fig. 9. Illustration of Bromus vulgaris

45 Bromus arvensis var. patulus (Mert. & Koch) Mutel., Fl. Franc. 4:134. 1837.

Se:rrafalcus patulus (Mert. & Koch) Parl., Fl. Ital. 1:394. 1848.

· · Bromus squarrosus var. patulus (Mert. & Koch) Regel, Act. Hort. Petrop. 7:602. 1881. Forasaccus patulus (Mert. & Koch) Bubani, Fl. Pyr. 4:387. 1901. Bromus japorticus var. porrectus Hack., Magyar Bot. Lapok (Ungar. Bot. Bl.) 2:58. 1903. Bromus janonicus var~ subsquarrosus (Borb.) Suval. & Rays., (Rumania) Min. Agr. Bull. 4(2):39. 1924.

Plant annual; culms erect or geniculate, 1 to 8 dm. tall; leaf blades 1 to 4 mm. wide and 7 to 20 cm. long; sheaths and blades pilose except upper sheaths glabrous, ligule short to 1.5 mm. long; panicle diffuse and somewhat drooping, mostly 5 to 16 cm. long, some depauperate plants with only single or a very few spikelets, and some plants with panicle up to 25 cm. long, branches mostly simple, lower ones 3 to 9 cm., slender and flexuous; spikelets fusiform-eliptic in shape, awns of lower florets about half the length of the upper ones, spikelets 3 to 5 mm. wide and 15 to 28 mm. long; lower glume acute, 3 to 7 nerved, 4 to

5.5 mm. long, upper glume S to 7 nerved, 5.5 to 7 mm. long; lemmas broad obtuse, 7 nerved with pair next to center nerve rather weak, 6 to 9 mm. long; awn of lowest floret mostly 4 to 5.5 mm. long, awn of other florets mostly 9 to 11 mm., straight to twisted a~d deflexed, scabrous, semi-circular or flattened at base in cross-section, arising from just below tip of floret; palea shorter than lemma by 1.5 to 2 rmn.; chromosomes 2n = 14 & 28. Japanese hrome is a cool season, shallow-rooted annual that was introduced from Asia. It is a common invader on abused ranges where the climax grasses are thinned out. It is found on the foothills and bench- 46 lands in Utah from 2500 to 7000 feet elevation, and thro_ughout the

United States except in extreme south eastern states. Utah distribution is shown in Figure 12 on page 48.

Careful observation of the comparative length of the awns should solve any difficulties of identification between this species and Bromus commutatus. They are quite ~imilar in appearance. Hofler and Knoll

(1956:60) noted, "Wird auch afters mit B. commutatus verwechselt." The spikelet is illustratec! on page SO.

· Representative collections. UTAH: Box Elder Co.; Wasatch Range Project, Allen et al. 16209, Perry Canyon, 23 June 1960 (UT). Cache Co.;

Maguire 20386, Logan Canyon, 28 June 1939 (BRY, RM, UTC). Davis Co.;

Anderson 310, Mueller Park, 14 June 1959 (UT). Grand Co.; Harrison 10394, near Moab, 15 June 1941 (BRY). Iron Co.; Dalley 5, Cedar City, 5 Aug

1938 (SUSC). Kane Co.; Stanton 748, Long Canyon, 25 June 1932 (UT).

Morgan Co.; Anderson 946, W. of Morgan, 22 June 1968 (BRY). Salt Lake

Co.; Garrett 7667, Salt Lake City, 15 June 1938 (BRY, US, UT); Arnow

1433, Red Butte, 9 .June 1968 (BRY, UT) . San Juan Co.; Harrison, Marshal and Nielson 10346, W. of Bluff, 13 June 1941 (BRY). Sevier Co.; Anderson

1069, Salina Canyon, 18 July 1968 (BRY). Summit Co.; Anderson 957, SW. of Henefer, 22 June 1968 (BRY). Utah Co.; Harrison 10415, Spanish Fork

Canyon, 18 June 1941 (BRY). Washington Co.; Galway s.n., St. George, 1

June 1934 (BRY). Weber Co.; Hull 9554, Ogden, 16 June .t9:S6 (BRY).

CALIFOIUHA: Lake Co.; Bacigalupi & Robbins 5679, 1 June 1956 (BRY,

JEPS). Modoc Co.; Whitney 3351, 30 June 1935 (BRY, UC). COLORADO:

Boulder Co.; Murdock 432, 27 June 1962 (BRY). Jefferson Co.; Fambrough 473, 28 July 1961 (BRY). LaPlata Co.; Flowers &Hall 155, 1 July 1960 (BRY, UT). Pueblo Co.; William & Penland 4470, 22 June 1955 (BRY, CCH). 47 IDAHO: Blaine Co.; Pyrah 122, 29 June 1963 (BRY). ILLINOIS: Champaign

CO.; Ahles 7395, 12 June 1953 (BRY, UC). MI1'."NESOTA: Pipestone Co.;

Moore & Sanger 26933, 21 June 1966 (BRY,. MIN). NEWMEXICO: Lincoln

Co.; Steyennark 38185, 7 July 19.38 (BRY). Rio Arriba Co.; Higgins .3584,

3 July 1970 (BRY, WTS). NEVADA:Lincoln Co.; Hall s.n., 2 July 1935

(BRY). TENNESSEE: Knoxville; Snow 1105S, 6 June 1938 (BRY) .. TEXAS:

Randall Co. ; Higgins 4.323, 3 June 19 71 (BRY,. \\11'S). WYOMING: Sheridan

Co.; Bird 69B, 11 July 1940 (BRY). Weston Co.; Porter 3982, 4 July 1946

(BRY,. RM).

10. Bromus commutatus Schrad.

Bromus commutatus Schrader, Flora Germanica, tom. 1, Gottingae. 1806.

Bromus pratensis Ehrh., Beitrage 6:84. 1791.

·s:tachynodium corrunutatum (Schrad.) Beauv., Ess. Agrost. 101, 155. 1812.

Serrafalcus commutatus (Schrad.) Bab., Man. Brit. Bot. ed. 1:374. 1843.

Bromus mutabilis var. commutatus (Schrad.) Schultz, Flora .32: 234. 1849 ..

Bromus racemosus var. commutatus (Sch~ad.) Coss. &Dur., Expl. Sci. Alger. 2:165. 1855.

Bromus mollis var. conmmtatus (Schrad.) Sanio, Verh. Bot. Ver. Brand. ~3: Abh. 31. 1882.

Serrafalcus racemosvs var. corm1ut:itus (Sch rad.) Husnot, Gram. Fr. Belg. 72. 180D.

Forassaccus coromutatu.s (Schrad.) Bubani, Fl. Pyr. 4:387. 1901.

Bromus secalinus var. gladewitzii Farwell, Amer. Midl. Nat. 10: 24. 1926.

Plant annual; culms 2. 5 to 8 and occa:;ionally 10 dm. tall; leaf

sheaths softly pubescent to retrorsely pilose, blades mostly 3 to 6 mm. F_igs. 10 to 13. Maps of distribution

10. Bromus porteri

11. Bromus canadensis

12. ·sromus ja:eonicus

13. Bromus ·commutatus \ I UTAH \ UTAH I County Lines and I County Unes 4nd • river drainages river drainage3 Fig.11 Fig.JO A

-.• •

O\ UTAH UTAH I County lines and County Lines and I river drainages river drainages Fig.13 I I Fig. 12 I A .. A .. ..

J I

--~ -----1 49

wide and 6 to 15 cm. long,. pubescent; ligule. small and inconspicuous;

panicle open, mostly 10 to 15 cm. long but ranging from 5 to 25 cm••

pedicels mostly with only 1 spikelet, some pedicels on lower whorl up to

6 cm. long but mostly shorter; spikelets 3 to 4 mm. wide and 12 to 18 mm.

long; first glume 3 or weakly 5 nerved, 5 to 7 mm·. long, second glume 5

to 7 nerved and 6 to 8 mm. long with alternate nerves weak; lemma 7 to

10 mm. long with hyaline margin, rounded apex without teeth, 7 nerved;

awn of first lemma 4.5 to 6 mm. long, upper awns 2 or 3 nun. longer,

sometimes slightly divergent but mostly straight; chromosomes 2n = 14

and 28.

Hairy chess is distributed in the lowland foothills of Utah from

2000 to 8000 feet (see Figure 13 on page 48). It is an introduced weed

from Europe found from Washington to California and eastward through the northern states. In Europe, where these plants are native, the closely allied members of the section Bromium, including Bromus commutatus, !· japonicus !_. secalinus, !· mollis and !· racemosus, 2.re treated as ~eparate spe- ies. They can be separated by some rather arbitrary characteristics

after a little practice. The glu,~es and lemma are illustrated in

Figure 14, A and B. Representative collections. UTAH: Box Elder Co.; Greenhalgh 11,

Bear River Refuge, 17 June 1949 (UT). Cache Co.; Cronquist 946, Logan,

12 June 1938 (BRY, RM, UTC). Davis Co.; Flowers s.n., Centerville, 15

June 1929 (UT). Garfield Co.; Stanton 140, Henry Mtns., 25 June 1930

(BRY); Buchanan 451 (mixed with!· tectorum), Bryce Canyon, 23 June 1959

(UT). Grand Co.; Harrison et al. 10395, upstream from Moab, 16 June 1941

(BRY). Iron Co.; Plummer 14S, . Burro Flats, 13 July 1941 (USFS, UT) . Fig. 14. Illustrations of Bromus commutatus & B. japonicus

A. Empty glumes of Bromus commutatus

B. Lemma of Br6mus commutatus

C. Spikelet of Bromus japonicus I 51 Salt Lake Co.; Garrett 5885, Williams Lake, 18 June 19.31 (PJ.1, UT).

Sanpete Co.; Plummer 254, Ephraim Canyon, 21 Aug 1942 (UT). Sevier Co.;

Lindsay s.n., Fish Lake, 17Aug 1962 (RIK). Summit Co.; Anderson 940, Little East Canyon, 22 June 1968 (BRY). Utah Co.; Harrison & Cottam 163, Provo River, 6 July 1925 (BRY). Wasatch Co.; Harrison 10870, N. of

Charleston, 2 Aug 1945 (BRY). Washington Co.; Higgins 717, Beaverdam

Wash, 18 June 1966 (BRY). Weber Co.; Collotzi 71, South Weber, 14 Oct

1963 (UTC). IDAHO: Elmore Co.; Hockaday 2, 20 June 1930 (BRY). Owyhee

Co.; Davis 2151, 21 June 1940 (BRY). NEVADA: Elko Co,; Holmgren 1001,

9 June 1941 (BRY). Washoe Co.; McMillan & McKnight 154, 21 June 1942

(BRY).

11. Bromus racemosus L.

Bromus·rac8mosus L., Sp. Pl. ed. 2, .1:114. 1762.

Btomus mollis v2.r. leiostachys Hartm., Skand. Fl. Handb. ed. 2·:33. 1832.

Serrafalcus racemosus (L.) Parl., Rar. Pl. Sic. 2:14. 1840.

Bromus arvensis var. racemosus (L.) Neilreich, Fl. Nieder- Oesterr:-s1. 1859.

Bromus ·squarrosus var. racemosus (L.) Regel, Act. Hort. Petrop. 7: 602. 1881.

Forasaccus racemosus (L.) Bubani, Fl. Pyr. 4:387. 1901.

Bromus mollis forma leiostachys (Hartm.) Fernald, Rhodora 35:316. 1933.

Bromus hordeaceus var. glabrescens (Coss.) Shear, U.S. Dept. of Agric., Div. of Agrost. Bull. 23:20. 1900. For the specimens referrecl by Sh0ar.

Annual weed with.slender erect stems, 3 to 6 dm. tall; culm nodes densely long pubescent; leaf sheaths and blades pubescent, at least the lower ones, blades narrow, mostly 7 to 14 cm. long; inflorescence a 52 raceme or simple panicle, contracted, the branches erect to ascending;· spikelets few, rather distant, 5 to 9 £lowered, 15 to 20 lillil. long; glumes rather broad and broadly acute at tip, first glume 3 to 5 nerved and 5 to 6 mm. long, the second 5 to 7 nerved and 6 to 8 mm. long; lemmas blunt to obtuse ac the apex, 7 nerved, 6 to 9 mm. long, smooth or minutely scabrous; awn 5 to 8 mm. long, arising 1 or 2 mm. below the rounded tip of the lemma; chromosomes 2n = 28.

This weedy brome grass is another that was introduced from

Europe and is found in a few localities along the foothills from 4000 to 8000 feet elevation in Utah. Figure 16 on page 58 shows distribution in the State. Its distribution is primarily on the Pacific coast and eastward to Montana, Colorado, and Arizona, as well as some few locations in the north eastern states.

ReEresentative collections. UTAH: Box Elder Co.; Flowers 844,

Willard, 19 June 1929 (UT). Davis Co.; Anderson 27, Centerville Canyon,

24 May 1958 (UT); Hull 9?53, Farmingtor., 17 June 193i (BRY). Garfield

Co;; Stanton 771, Escalante Mtns., 24 June 1924 (UT). Juab Co.; ½cMillan

9_27, Deep Creek Range, 1 July 1947 (UT, UTC). ARIZONA:Coconino Co.; Silveus 1937 (BRY).

12. Bromus secalinus L.

Bromus·secalinus L., Sp. Pl. 76. 1753.

Bromus molljs var. secalinus (L.) Huds., Fl. Angl. ed. 2. 49. 1778.

Avena secalinus (L.) Salisb., Pro

Serrafalcus secalinus (L.) Bab., Man. Brit. Eot. ed. 1:374. 1843.

Forasaccus secalinus (L.) Bubani, Fl. Pyr. 4:388. 1901. 53 Very similar in description -co Bromus commutatus except that

the sheaths and blades of the leaf are glabrous; the lemmas a little

more plump and somewhat more distant so that light passes through the

small·openings at the base of each floret when the spikelet is viewed

from the side; and the awn tends to be somewhat shorter; chromosomes

2n = 14 &28. This grass, often called Chess or Cheat grass, is reported to be more or less throughout the United States. All of the specimens reported for Utah were collected during the early years when!· secalinus was the only recognized species of this complex thought to be in the western United States.

Cited specimens examined. UTAH: Cache Co~; Smith 1697, Logan,

23 June 1909 (UTC). Salt Lake Co.; Jones 1009, Salt Lake City, 8 July 1879 (UT, UTC); University of Deseret collections in 1880 & 1887, 4 sheets, Salt Lake City area, (UT); Clemens s.n., Fort Douglas, 28

Sep 1911 (UT).

All of these have been carefully examined and should be referred

to other species of this complex, as they do not fit the description of

B. secalinus.

13. Bromus mollis L.

Bromus mollis L., Sp. Pl. ed. 2, 1:112. 1762.

Serrafalcus mollis (L.) Parl., Pl. Rar. Sic. 2:11. 1840.

Forasaccus moll is (L.) Bubani, Fl. Pyr. 4: 386. 1901.

Bromus hordeaceus L. subsp. mollis (L.) Hylander, Uppsala Univ. Arskr: 7: 84 .. 1945.

Annual; simple to slightly tufted, culms slender and erect, 2 54 to 8 dm. tall; leaf blades spar~ngly pilose to nearly smooth~ sheaths softly pubescent, blades 3 to S mm. wide and 5 to 15 cm. long; panicle erect and contracted, 5 to 10 cm. lo_ng, pedicels short; spikelets 10 to 16 mm. lo_ng, 5 to 12 flowered, coarseiy pilose to scabrous pubescent; glumes broadly acute, the first 3 to 5 nerved and 4 to 6 1IDil. long, the second 5 to 7 nerved and 7 to 8 mm. long; lemmas broadly obtuse, 7 nerved, 8 to 9 mm. long; rather stout awn, 6 to 9 1IDil. long; chromosomes 2n = 28.

Soft Chess is a weed in fields and waste places in most of the north western part of the United States, and from the northern plains states throughout New England. It is found in Utah along the roadsides and fields of the Wasatch front and on the saline flats bordering the

Great Salt Lake from 4000 to 5000 feet elevation. See Figure 17 on page 58 for details of distribution in Utah.

Representative collections. UTAH: Cache Co.; Maguire 2897, Pine Canyon, 18 June 1932 _(UTC). Davis Co.; Anderson 804, S. of Farming- ton, 11 May 1968 (BRY); Maguire 13201, Farmington, 13 May 1934 (RM,

UTC); Arnell s.n., Farmington Bay, 8 July 1966 (UT). Salt Lake Co.;

Flowers 1347, Salt Lake City, 18 June 1933 (UT). Utah Co.; Larsen 117,

Provo, 30 June 1933 (BRY). CALIFORNIA: Alameda Co.; Beetle 2610,

Berkeley, 25 Apr 1941 (BRY, UC). Santa Clara Co.; Keck 1439, Stanford Univ., 23 Apr 1932 (DS, BR~). IDAHO: Elmore Co.; Keller I-223,

Hammett, 15 June 1940 (USFS, BRY). NEVADA:Elko Co.; Holmgren 590,

Midas, 29 May 1940 (USFS, BRY).. OREGON: Jackson Co.; Gierisch 3015,

Ashland, 7 June 1965 (USFS, BRY):

14. Bromus b:tiiaeformis Fisch. &Mey.

Bromus brizaeformis Fischer &Meyer, Index seminum, quae Hortus botan- 55 botanicus imperialis Petropoli tanus; . . • . Akademia Nauk. Botanicheskii Institute part 3:30-31. 1837.

Annual; simply to densely tufted, culms mostly 2 to 3 run.tall but some plants to 6 dm. tall; leaf sheaths with soft pilose pubescence, blades 2 to 5 mm. wide, 3 to 12 cm. long, sparingly to densely pubescent; panicle erect when young, gracefully droopi_ng after flowering; spikelets few, 8 to 18 florets, 1 to 2 cm. long, broad and inflated with papery texture which produces a characteristic rustling sound when agitated; glumes obtuse, the first 4.5 to 6 mm., the second 6 to 7 mm., 3 to 5 and sometimes 7 nerved; lemmas 7 nerved, mostly 8 to 9 mm. long, broadly ovate, shiny and straw colored when mature, minutely hairy; awn less than 1 mm. long; chromosomes 2n = 14.

Rattlesnake grass or Quake brome is a commonweed of the foot- hill areas throughout the state and is also found in canyons and around farms. This gracefully attractive grass is illustrated on page 56. It was introduced from Europe, and spread from Alaska and Canada south through most of the western states, but is rare in the eastern states.

It is found at elevations from 4000 to 6500 feet in our area. The distribution map is on page 58.

Representative collections. UTAH: Box Elder Co.; Williams and

Pierson 625, Cold Spring Canyon, 15 June 1932 (R.M,UTC); Erdman 1448,

Promontory Mtns., 7 June 1965 (BP.Y). Cache Co.; Maguire 2899, Wellsville

Range, 18 June 1932 (RM, UTC). Daggett Co.; Collotzi 64, Sheep Creek,

11 June 1963 (UTC). Davis Co.; Anderson 309, Mueller Park, 14 June

1959 (UT). Salt Lake Co.; Arnow 2813, Red Butte, 18 June 1969 (BRY, UT,

UTC); Rydberg 6163, N. of Salt Lake City, 12 June 1905 (NY, RM). Sevier

Co.; Stanton 742, Salina, 17 Jun~ 1932 (UT). Summit Co.; Rydberg 2353, Fig. 15. Illustration of Bromus brizaeformis

A. Habit sketch B. Empty glumes C. Lemma A 57 Echo, 13 ~ug 1895 (RM). Utah Co.; Larson 118, Provo, 15 June 19.'33 (BRY);

Larson·L105, Payson Canyon, 14 July 1932 (UTC); Saling 57838, Hobble

Creek, 18 July 1927 (UT). ARIZONA: Mohave Co.; Atwood 1775, Olsen

Canyon, 27 May 1969 (BRY).. IDAHO: Bearlake Co.; Calvert 197, SE. of Preston, 19 June 1959 (BRY). NEVADA: Elko Co.; Holmgren, NE of Elko,

29 June 1938 (BRY). Washoe Co.; McKnight and McMillan 106, W. of Gerlach, 14 June 1942 (BRY).

Section 4. Stenobromus Grisebaugh, Spicel. Fl. rumel. 2:448. 1844.

Tufted annual plants; spikelets compressed; glumes and lemmas narrow, accu.~inate; lower glume 1 nerved and the second 3 nerved; lemma

5 to 7 nerved, with 2 hyaline teeth at the apex; the long awn arises in the cleft between the teeth; callus of the floret is hard and sharp at maturity; introduced from Europe.

15. Bromus rubens L.

Bromus rubens L., Cent. Pl. 1:5. 1755.

Festuca rubens (L.) Pers., Syn. Pl. 1:94. 1805.

Bromus scoparius var. rubens (L.) St. Amans. Fl. Agen. 45. 1821.

Bromus madritensis subsp. rubens (L.) Husnot, Gram. Fr. Belg. 71. 1899.

Anisantha rubens (L.) Nevski, Act. Univ. Asiae Med. VIIIb. Bot. 17:19. 1934.

Zerna rubens (L.) Grossh., Akad. tfauk S.S.S.R. Bot. Inst. Trudy Axerbaidzh. Fil. 8:306. 1939.

Plant a simple to tufted annual; culms 1.5 to 4 dm. tall, puberulent below the panicle; leaf sheaths and blades pubescent, blades mostly 2 to 4 mm. wide and 5 to 7 cm. long; the panicle erect and compact with purplish color, 4 to 8 cm. long; spikelets 4 to 11 flowered, I I UTAH 'I UTAH ' I CJunty lines and I County lines and river drainages river drainages Fi'g.16 Fig.17 I • ..I ..I .I ti. ..

,, ,,( ------.../~y \ I ~-~~~~:~:;~/--1~,_-----1 ------,LI IJ 1-' ,- 0 _ __, I,- _ _...."'... -. __ ,- I, ______...___ _,,.- ·------

'I UTAH UTAH I I County Lines and County Lines ar..d ri.,,er draicages river drainages fig.18 Fig.19 ti. ti. 59 2.5 cm. long; first gltL~e 7 to 9 mm. long, the second 10· to 12 mm. long; lemmas 12 to 16 mm. long with teeth 4 to 5 mm., scabrous to pubescent; awn 18 to 22 mm. long and usually spreading somewhat at maturity; chromosomes 2n = 28.

This grass is abundant over extensive areas in the southern and western parts of Utah. It is found in dry and often saline soils on the hills or in waste or cultivated ground. In the western states it is found from Idaho to western Texas and westward to the coast. It is also reported in Massachusetts. Map Figure 24 on page 68 shows the distribution.

The illustration on page 62, Fig. 20, E, shows the habit of the plant. Sketches A, B, & Dare specifically_of Bromus tectorum, however, the glumes and lemmas are very similar in shape and size for the two species, and the illustrations can serve as a close approximation for the details of B. rubens. Representative collections. UTAH: Garfield Co.; Welsh &Moore

7123, Hite, 28 May 1968 (BRY). Juab Co.; Christensen 115, Fish Springs,

24 Apr 1965 (BRY). Kane Co.; Harrison 12124, Hole in the Rock, 14 May

1953 (BRY). Millard Co.; Fautin 32F, \\bite Valley, 10 May 1940 (BRY). Tooele Co.; Saccomanno 7666, Stansbury Island, ca. 1940 (UT); Maguire & Holmgren 21842, Simpson Springs, 10 July 1943 (UTC). Washington Co.;

Maguire & Blood 1301, E. of Ht1rricane, 1 May 1932 (RM, UTC); Christian

784, Beaverdam Mtns., 3 Apr 1963 (UT, UTC); Olson 4, Toquerville, 4

Apr 1970 (BRY); Higgins s.n., St. George, 14 Apr 1961 (DIX); Dalley 6,

Purgatory Valley, 22 Apr 1939 (SUSC). ARIZONA: Maricopa Co.; Higgins

3979A, 20 miles NE of Tempe, 23 Apr 1971 (WTS, BRY). Mohave Co.; Cottam

1123, Virgin Narrows, 14 Apr 1927 (BRY). Yavapai Co.; Graham s.n., 60 Sunset Point, 18 Apr 1965 (ASC, BRY),- CALIFORNIA: El Dorado Co.; Pohn

7230, E. of Placerville, 16' May 1953 (ISC, BRY). Modoc Co.; Whitney

3113, Lake Joaquina, 21 May 19..35 (UC, BRY. Riverside Co.; Crosswhite ' 1707, Riverside, 21 Jan 1961 (ASC, BRY). San Bernardino Co.; Lepper 37,

Strawberry Canyon Rd., 21 Mar 1964 (DAV, BRY). San Luis Obispo Co.;

Hendrix 24, Pozo, 17 Apr 1937,(UC, BRY). IDAHO: Owyhee Co.; Nadeau

I-247, Givens Hot Springs, 11 May 1940 (USFS, BRY). NEVADA: Clark Co.;

Cottam 8399, Key West Mine, 7 Apr 1941 (BRY). Elko Co.; Holmgren 508,

Izzenhood Ranch, 22 May 1940 (USFS, BRY). Lander Co.; Stahmann 19, S. of Dunphy, 23 May 1934 (USFS, BRY).

16. Bromus tectorum L;

Bromus tectcrum L., Sp. Pl. 77. 1753.

Schedonorus tectorum (L.) Fries, Bot. Not. 131. 1843.

Bromus setaceus Buckl., Acad. Nat. Sci. Phila. Proc. 1862:98. 1862.

Zerna tectorum (L.) Panz. ex Jacks., Ind. Kew. 2:1249. 1895.

Slender erect annual; culms single to tufted, 3 to 6 dm. tall; leaf sheaths and blades pubescent, blades mostly 3 -::o 5 mm. wide and 5 to 10 err.. long; panic le some,,hat di-ooping, often with some purple color,

5 to 15 cm. long, pedicels very thin; spikelets nodding, 12 to 20 mm. long; first glumP- ,1 to 6 1rnn. long, 1 nerved, second glnme 8 to 10 mm. with 3 nei-ves, a-::cuminate, pubescent to villous; lemmas lanceolate, villous or pilose, 10 to 12 1mn. long with teeth 2 to 3 mm. long; awn 12 to 14 mm. long; chromosomes 2n = 14.

Cheatgrass brome or Downy Chess i£- a very well known and common weed in Utah. 'rhe distribution map is Figure 22 on page 68. It is .61 probably found on more acres o~ ground in the State than any other j

species of plant. Herbaritnn collections have been seen for all counties

in the state except Carbon, Wayne, and Rich. The grass is very likely

to be found in those counties as well as in the others, but botanists

are loath to pick up such a common weed when collecting plants for

study. Cheatgrass is found throughout the United States except in the

extreme south east. A very extensive coverage of Cheatgrass was done by Klemmedson

and Smith (1964). Anyone desiring further information relative to the usefulness, control, growth habit, distribution, or any other topic relative to this grass is recommended to that source.

Bromus tectorum normally sprouts in the autumn and is well est-

ablished by early spring when it forms a lush green cover over the hills and over-used lands of the State. During the springtime Cheatgrass is a nourishing diet for our grazing animals, but by early summer it has finished flowering and begins to turn brown and straw colored. It is then that Cheatgrass becomes a very real fire hazard in many localities.

On several occasions during the sl.lI!Ur.ersof 1958 and 1959 I had the oppor- tunity to measure the effect of fire on Cheatgrass. In most cases the

Cheatgrass gained in percent frequency and percent cover when regrowth occurred after er..ch successive fire. Br0•vse plants are generally severely injured or killed by the fires, but the grasses are able tc recover or quickly become re-established. The grass is illustrated in

Figure 20 on page 62. Fig. 20 A and Bare the glumes; Dis the lemma; and C is the habit.

Cheatgrass brome is a good indicator of land misuse, and in Utah we have had a history full of abuse of the land. Beetle (1950) reports Fig. 20. Illustration of Bromus tectorum & B; rubens A. First. glume of B. tectorum B. Second glume C. · B. tectorum habit sketch D. Floret of B. tectorum

E. B. rubens habit sketch A

E ~- '1 I l

\ \ \\\\ 63 on the very low occurrence of this_ grass in Wyoming, and notes that it is found only on roadcuts and other disturbed sites. He then concludes that Wyominghas had a relatively good record for grazing man_agement that has kept the rangelands from being overrun by weedy species. He says, "any general or widespread increase in cheatgrass brome should be viewed with alarm." Representative collections. UTAH: Beaver Co.; Stahman & Hutchings 37, Wah-WahRange, 25 Apr 1934 (BRY, UT). Box Elder Co.;

Slade 4, 17 May 1966 (UTC). Cache Co.; Maguire 3233, Blacksmith Fork

Bench, 5 June 1932 (RM, UTC). Daggett Co.; Williams 449, Flaming Gorge,

30 May 1932 (RM). Davis Co.; Anderson 805, S. of Farmington, 11 May

1968 (BRY). Duchesne Co. ; Brotherson 851 & . 970, Rock Creek, 27 :May 1966 & 23 July 1965 (BRY). Emery Co.; Welsh 3916, S. of Highway junction of Utah 24 with 50 &6, 3 May 1965 (BRY). Garfield Co.; Stanton 629, Henry Mtns., 28 May 1932 (UT); Hall 13598, Upper Valley, 15 June 1950 (BRY,

UT). Grand Co.; Anderson 761, E. of Green River, 30 Apr 1968 (BRY).

Iron Co.; Ogden s.n., CSU State fam, 24 June 1958 (SUSC). Juab Co.;

Anderson 826, W. of Jericho, 8 June 1968 (BRY); Goodding 1106, Nephi,

13 June 1902 (RM); Maguire & Becraft 2527, Troutcreek Ranch, 19 June 1933 (UTC). Kane Co.; Castle 126, W. edge of Dunes, 19 June 1953 (BRY).

Millard Co.; Patten 119, Desert Range Expe:r. Stat., 2 May 1970 (BRY);

Worthen 100, S. of Deseret, 1 May 1965 (UT). Morgc1.nCo.; Anderson 931,

East Canyon, 22 June 1968 (BRY). Salt Lake Co.; Anderson 101, Univ. of Utah campus, 9 June 1958 (BRY, UT); Anderson 973, Bluffdale, 27 June

1968 (BRY). San Juan Co.; Wilson 21, Fry Canyon mesa·, 13 June 1965 (UTC). Sanpete Co.; · Peterson s. n ,., Snovr College campus, 9 May 1961

(DIX). Sevier Co.; Jeffrey s .n., Willow Creek, 29 Aug 1962 (UTC). 64 Tooele Co.; Foster 21, S. of Vernon, 20 June 1964 (BRY). Smmnit Co.;

Anderson 937, W. of Henefer~ 22 June 1968 (BRY). Uintah Co.; Brotherson

813, Dinosaur Nat' 1 Mon., 15 July 1965 (BRY). Utah Co.; Anderson 239,

N. of Provo, 8 June 1959 (BRY,.UT);. Liddle 6819, Payson Canyon, 30 July 1933 (BRY). Wasatch Co.; Brotherson 690, Strawberry Valley, 16 July

1965 (BRY). Washington Co.; Gould 1741, NW. of Pine Valley, 10 May 1942

(RM); Higgins 652, Beaverdam Mtns., 26 May 1966 (BRY). Weber Co.;

Salimeno 6, Sullivan's Pond, 10 May 1965 (WES, BRY).

16a. · Bromus tectorum var. glabratus Spenner.

Bromus tectorum var. glabratus Spenner, Fl. Friburg. l:152. 1825. Bromus tectorum var. nudus Klett. &Richt., Fl. Leipzig 109. 1830.

Artisantha tectorum (L.) Nevski, Act. Univ. Asiae Med. VIII. Bot. 17:20, 22. 1934. ·Bromus tectorum forma nudus (Klett. & Richt.) St. John, Fl. Southeast. Wash. &Adj. Idaho 36. 1937.

This variety differs from the species only in having glabrous spikelets. The variety'is far less common than the species, but has much the s~~e range.

Representat5ve collections. UTAH: Cache Co.; Maguire 13200,

Logan Canyon, 29 Apr 1934 (RM, UTC). Kane Co.; Harrison 12098, 50 mi.

E. of Kanab, 12 May 1953 (BR\'). Millard Co.; Harrison 10125, Beaver

Mtns., 4 May 1941 (BRY). Piute Co.; Buss 196, W. of Junction, 25 July

1964 (BRY). Salt Lake Co.; Arr.ow 1209, Red Butte, 12 May 1968 (BRY, UT).

San Juan Co.; Moore 362, Canyonlands Nat'l Park, 15 May 1965 (BRY); Spear &Markham Unit 18-37, Goodman Wash, 3 Aug 1939 (BRY). Sevier Co., Stevens 35, Salina Canyon, 10 July 1964 (BRY). Washington Co.; 65 Christian 881, near 3 mile wash &Santa Clara confluence, 3 May 1963 (UT); Higgins 497, Beaverdam Mtns., 24 Apr 1966 (BRY); Hall s .n.,

Enterprise, 27 May 1935 (BRY)_.

17. · Bromus sterilis L.

Bromus sterilis L., Sp. Pl. 77. 1753.

Schedonorus sterilis (L.) Fries, Bot. Not. 131. 1843.

Zerna sterilis (L.) Panz. ex. Jacks., Ind. Kew. 2:1249. 1843.

Anisantha sterilis (L.) Nevski, Act. Univ. Asiae Med. VIIIb. Bot. 17:20. 1934.

Plant a tali annual; culms 5 to 10 dm. tall; leaf sheaths pubescent, blades mostly 10 to 20 cm. long, narrow; panicle 10 to 20 cm. lo_ng, with drooping branches; spikelets 2.5 to 3.5 cm. long, 6 to 8 and sometimes 10 florets; glumes lanceolate-suhulate, the first about 8 mm. long; the second one just over 1 cm.; lemmas 17 to 20 mm. long, scabrous to scabrous-pubescent, teeth on lemmas 2 mm. long; awn 2 to 3 cm.; chromosomes 2n = 28. Bromus sterilis has become established in scattered locations from California to British Columbia and eastward to Colorado and New

Mexico. It is found in the New England States and Illinois to Virginia and Arkansas. Utah distribution is shown in Figure 23 on page 68. It can be found in cultivated gardens, fields and waste places.

Representative_ collections. UTAH:Salt Lake Co.; Anderson 219,

Salt Lake City, 19 May 1959 (BRY, UT); Garrett 5896, City Creek, 23

June 1931 (UT). Utah Co.; Anderson 246, N. of Provo, 8 June 1959 (BRY,

UT). Washington Co.; Harrison 10200, Veyo, 7 May 1941 (BRY); Dalley

61, Veyo, 29 May 1939 (SUSC). Fig. 21. Illustration of Bromus sterilis

67 18. · Bromus diandrus Roth.

Bromus diartdrus Roth .• Bot. Abhl. 44. 1787.

Bromus rigidus auct. Fl. N. Amer. non Roth.

Robust annual; culms 4 to 7 dm. tall; leaf sheaths and blades pilose; panicle 7 to 15 cm. long, erect to nodding, branches simple with few spikelets, short; spikelets mostly 5 to 7 flowered, 3 to 4 cm. long, awns excluded; glumes smooth, lanceolate-subulate, the first 1.5 to 2 cm. long; lemmas scabrous to puberulent, 2.5 to 3 cm. long with teeth to

4 mm. long; awn stout, 3.5 to 5 cm. long; chromosomes 2n = 28. Ripgut grass or rigid brome is of Mediterranean origin and has been introduced into Europe, North and South America and South Africa.

Utah distribution is shown on map Figure 25 on page 68. This species resembles Bromus rigidus Roth. in its general appearance and has been confused with it. The distinctive character of B. diandrus as compared with~- rigidus is the shape of the callus at the base of the lower lemma in each floret (Bor, 1968); it is short, spheroid and rounded at the tip, while in B. rigidus the callus is elongate, compressed and acute at the tip. We probably have both species.in the Uriited States, however, all of the specimens that I have examined have examined have been B. diandrus. ~- rigidus is limited to sandy soils, mainly along the coasts, in other parts of the world (Ovadiahu-Yavin 1969:199).

This grass is known as a troublesome nest when the spikelets are mature. The narrow, sharp-pointed minute} y barbed florets with their long awns can work into the eyes, nostrils and mouths of livestock causing considerable injury.

Representative collections. UTAH: Iron Co.; Goodrich s.n., Figs. 22 to 25. Maps of distribution 22. · Bromus tectorum

23. Bromus sterilis

24. ·B:tomus·rubens

25. Bromus diandrus I \ UTAH I UTAH I Countv Lines and I County lines and Fig.22 river drainages Fig.23 rivetr drainages

•

/ ______..,,,..,) ~2h---:------, l ___------,- I,~ __,- '------

, \ UTAH I UTAH I Coun'ty lines ar.d I County lines and Fig.24 river draimiges river drainages I ) Fig. 25 ) .. .. ~:, .. _... .,,,---\----,.:-,, ~- r ' I 69 Cedar City, 15 May 1965 (SUSC). Salt Lake Co.; Anderson 218, City Creek,

19 May 1959 (BRY, UT); Flowers 797, Salt Lakebenchland, 17 June 1927 (UT)~ Millard Co.; Kimball s. n., Kanosh, 1 May 1965 (SUSC). Washi_ngton . Co.; ~iggins 4175, Beaverdam Mtns., 17 May 1971 (BRY); Weight KEW828Z,

Zion Nat'l Park, 5 June 1935 (UT); Anderson 753, Zion Nat'l Park, 8

June 1957 (UTC). Weber Co.; Hull 28, Ogden, 1 July 1936 (UTC). ARIZONA:

Coconino Co.; Drake-Welsh 47, Oak Creek Canyon, 6 May 1961 (ASC, BRY). Mojave Co.; Welsh &Atwood 9700, Pipe Springs, 4 May 1970 (BRY). CAL- IFOR.NIA: Alameda Co.; Beetle 2727, Berkeley, 20 May 1941 (BRY, UC).

Kern Co.; Smith 1044, on Greenhorn, 22 June 1961 (JEPS, BRY). Riverside Co.; Isely & Bragonier 7190, Valle Vista, 21 Mar 1960 (ISC, BRY). San · Luis Obispo Co.; Nordstrom 1092, SE. of Adelaida, 19 Apr 1937 (BRY, UC).

Santa Clara Co.; Keck 1445, Stanford Univ., 23 Apr 1932 (BRY). Santa

Cruz Co.; Beetle 2685, W. of Pacheco Pass, 13 Apr 1941 (BRY, UC). Shasta Co.; Long 60a, Anderson, 11 May 1931 (BRY, UC). Yolo Co.; Hackett 32,

Davis, 14 May 1960 (DAV, BRY). OREGON: Jackson Co.; Erdman 1972, White

City, 22 June 1965 (SLP, BRY). Multnomah Co.; Rummell F.S., Portland, 20 May 1945 (USFS, BRY).

Section 5. Neobromus (Shear) Hitch., Manual of Grasses U.S. 56. 1935. Annual; lemm~s lanceolate, deeply bifid, the teeth aristate; awn twisted, geniculate. Approaches Trisetum.

19. Bromus trinii Desv.

Bromus trinii Desv. in Gay, History of Chile 6:441-443. 1853.

Trisetum hirtum Trin., Linnaea 10:300. 1836. Bromus trinii var. pallidiflorus Desv. in Gay, Fl. Chili 6:441. 1853, 70 Trisetwn parbaturn Steud., Syn. Pl. Glum. 1:229. 1854. Dartthonia pseudo-spicata C. Muell., Bot. Ztg. 14:348. 1856.

Trisetum barbatum var; major Vasey, U.S. Dept. of Agric., Div. Bot. Bull. 13(2):60. 1893.

Bromus ba:tbatoides Be~l, Grasses N. Amer. 2:614. 1896.

Bromus ba:tbatoides var. sulcatus Beal, Grasses N. Amer. 2:615. 1896.

Trisetum trinii (Desv.) Louis-Marie, Rhodora 30:243. 1928. Trisetum trinii var. majus (Vasey) Louis-Marie, Rhodora 30:243. 1928.

Ttisetobromus hurtus (Trin.) Nevski, Acta Univ. Asiae Med. VIII b. Bot. 17:15. 1934.

Plants annual; culms 3 to 6 dm. mostly, sometimes to 10 dm., erect or tufted and spreading below, often pubescent at the nodes; ieaf sheaths and blades smooth to pilose; panicle 8 to 20 cm. long, erect and narrow, branches erect to spreading (lower ones); spikelets 1.5 to 2 cm. long, narrow, 5 to 7 florets; glurnes lanceolate, accuminate, the

lower one ulually 1 nerved; 8 to 10 mm. long; the second mostly 3 nerved and 12 to 16 mm. long; lemmas 5 nerved, pubescent, 12 to 14 mm. long, accuminate with narrow teeth 2 to 3 mm. long; awn 1.5 to 2 cm. long, twisted near the base and strongly :iivaricate when mature; chromosomes

2n = 42.

Chilean Chess is found on the dry pains and rocky or wooded slopes from Oregon south to Baja California and rarely eastward to

Colorado. As the name suggests, it was introduced from Chile. In Utah it is found occasionally in Washington County, as Figure 19 on page 58 indicates.

An illustration of the spikelet can be seen in Figure 7C on page 38. 71 · ·Representative collections. UTAH: Washington Co.; ,Harrison

10213, S. of Veyo, 8 May 1941 (BRY, UT); Galway s.n., W. of St. George,

38 Apr 1935 (BRY, UT); Harrison 10249, top of Sugarloaf Mtn., 9 May

1941 (BRY). CALIFORNIA: Riverside Co.; Hitchcock 5924, 2 Apr 1940 (UTC); Munz 15706, 24 Mar 1940 (UTC). Co. unknown, Orcutt s.n., 1886 (UTC). COLORADO:Boulder Co.; Murdock 420, Bluebell Canyon, 27 June

1962 (BRY). NEVADA:Clark Co.; Train 1358, 11 Apr 1938 (UTC). 72

LIST OF REFERENCES

Abrams, LeRoy. 1940. An illustrated flora of the Pacific States, Vol.

1. 2nd Ed. Stanford University Press, Stanford.

Arnold, Chester A. 1947. An introduction to paleobotany. McGraw-Hill,

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APPENDIX·

Index of Synonyms

The names presented in the following list consist of the synonyms of the species treated in the foregoing work and also names of species previously reported for the area. The names in the left column are the synonyms, and the nai11es in the column on the right are those which are used for the species in the present treatment.

ANISANTHA BROMUS P_age

A. !'Ubens (L.) Nevski. 1934. B. !'Ubens L. 1755. 57 A. steriZis (L.) Nevski. 1934. B. steriZis L. 1753. 65 A. tectorum (L.) Nevski. 1934. B. tectorum var. gZabratus Spenner. 1825. 64 AVENA A. secaZinus (L.) Salisb. 1796. B. secaZinus L. 1753. 52 BRACHYPODIUM B. comrrrutatum(Schrad.) Beauv. B. commutatus Schrad. 1806. 47 1812.

BROMUS B. anomaZus Rupr. 1840. ·Rejected. The range of this taxon lies to the south of Utah. The nam0 has been applied to B. porteri of the Rocky Mountain area.

B. arvensis var. patulus B. japonicus 1bunb. 1784. 43 (Mert. & Koch) Mute 1. 1837. B. arvensis var. racemosus (L.) B. racemosus L. 1762. 51 Neilreich. 1859. B. barbatoides Beal. 1896. B. trinii Desv. 1853. 69 B. barbato·ides var. suZcatus B. trinii Desv. 1853. 69 Beal. 1896. B. breviaristatu.s Buckl. 1862. B. ma.rginatu.s Nees. 1855. 19 82 BROMUS BROMUS Page B. breviaristatus (Hook.) B. u?iZZdenowii Kunth. 1829, 25 Thurb. ex Wilkes. 1862. B. brizaeformis Fisch. &Mey. 1837. B. aaliforniaus Nutt. ex B. aarinatus Hook. &Arn. 1840. 23 Buckl. 1862. B. aanadensis Michx.11803. B. aarinatus Hook. &Arn. 1840T B. aarinatus var. caZifornieus B. aarinatus Hook. & Arn. 1840. 23 Shear. 1900. B. aarinatus var. densus B. aarinatus Hook. & Arn. 1840. 23 Shear. 1900. B. aarinatus var. hookerianus B. aarinatus Hook. & Arn. 1840. 23 Shear. 1900. B. aatharticus Vahl. 1791. Rejected as nomen confusum. Previously applied to our species B. hlilldeno-wii. 'B. ailiatus L. 1753. Rejected. Previously applied to B. aanadensis, and now correctly applied to the eastern grass which has been known as B. "latiglv.mis. B. ciliatus auct. Fl. N. Amer. B. aanadensis Michx. 1803. 39

B. ailiatus var. glaberrimus B. vulgaris (Hook.) Shear. 1900. 42 Suksdorf .. 1901. B. ailiatus var. ligulatus E. vulgaris (Hook.) Shear. 1900. 42 Vasey ex ~lacoun. 1888. B. aiZiatus var. montanus B. porteri (Caul t.) Nash. 1895. 34 Vasey ex Beal. 1896. B. aiZiatus var. pauaiflm.•i,:.,s B. vulgaris (Hook.) Shear. 1900. 42 Vasey ex Macoun. 1888. B. ailiatus var. porteri B. poj."'ter•i (Caul t.) Nash. 1895. 34 (Coult.) Rydb. 1895. B. ciliatus var. scay,iosus B. porteri (Coult.) Nash. 1895. 34 Scribn. 1898. B. commutatus Schrad. 1806. B. debilis Nutt. ex Shear. 1900. B. vulgaris (Hook.) Shear. 1900. 42 B. diandrus Roth. 1787. B. eximius Piper. 1906. B. vulga1•is (Hook.) Shear. 1900. 42 B. eximius var. umbraticus B. vulgaris (Hook.) Shear. 1900. 42 Piper. 1906. 83 BROMUS BROMUS Page B. fl,odmanii Rydb. 1909 B. marginatus Nees. 1855 19 B. froruiosus (Shear) Woot. & Rejected. The range of this species Standl. 1912. is reported to be Arizona, Colorado, and New Mexico. B. hookeri var. marginatus B. marginatus Nees. 1855 19 (Nees.) Fourn. 1886. B. hookerianus Thurb. ex B. aarinatus Hook. &Arn. Wisdes. 1874. 1840. 23 B. hookerianus var. minor B. aarinatu.s Hook. & Arn. Scribn. ex Vasey. 1885. 1840. 23 B. hordeaeeus var. glabreseens B. raaemosus L. 1762. 51 (Coss.) Shear. 1900. B. inermis Leyss. 1761. B. inermis var. aristatus B. inermis Leyss. 1761. 29 Schur. 1866. B. inermis forma aristatus B. inermis L~yss. 1761. 29 (Schur.) Fernald. 1933. B. inermis forrna bulbiferus B. inermis Leyss. 1761. 29 Moore. 1941. B. inerrr.is forma villosus B. inermis Leyss. 1761. 29 Fernald. 1933. B. inopinat'u.s Brues. 1911. B. inermis Leyss. 1761. 29 B. japoniaus Thunb. 1784. B. japoniaus var. porreetus B. japonicus Thunb. 1784. 43 Hack. 1903. B. japonieus var. subsquarrosus B. japonieus Thunb. 1784. 43 (Borb.) Suv. & Rays. 1924. B. kalmii var. major Vasey ex B. porteri (Coult.) Nash. 1895. 34 Shear. 1900. B. kalm·ii var. oeeidentaUs B. porteri (Coult.) Nash. 1895. 34 Vasey ex Beal. 1896. B. kalmii var. porteri Cot:lt. B. porter•i (Coult.) Nash. 1895. 34 1885. B. latior Rydb. 1917. B. mai>gina:tus Nees. 1855. 19 B. mad.1:>ite.nsissubsp. rubens B. rubens L. 17SS. 57 (L}. Husnot. 1899. B. marginatus Nees. 1855. B. marginatus var. latior B. marginatus Nees. 1855. 19 Shear. 1900 .. B. marginatus var. seminudus B. marginatus Nees. 1855. 19 Shear. 1900. 84 BROMUS BROMUS P_a.ge

B. moZZis L. 1762. 53 B. moZZis var. commutatus B. commutatus Schrad. 1806. 47 (Schrad.) Sanio. 1882. B. moZZis var. leiostachys B. racemosus L. 1762. 51 Hartm. 1832. B. moZZis forma Zeiostachys B. racemosus L. 1762. 51 (Hartm.) Fernald. 1933. B. moZZis var. secalinus (L.) B. secalinus L. 1753. 52 Huds. 1778. B. muZtifZorus Scribn. 1898. B. marginatus var. polyanthus 22 (Scribn.) Anderson. 1973. B. mutabilis var. comrrrutatus B. commutatus Schrad. 1806. 47 (Schrad.) Schultz. 1849. B. nitens-Nutt. ex A. Fray. B. carinatus Hook. & Arn. 1840. 23 1862. B. oregonus Nutt. ex Hook. B. carinatus Hook. & Arn. 1840. 23 1856. B. panicuZatus Rydb. 1917. B. marginatus var. polyanthus 22 (Scribn.) Anderson. 1973. B. patuZus Mert. & Koch. 1823. B. japonicus Thunb. 1784. 43 B. poZyanthus (Scribn.) Shear. B. marg·inatus var. poZyanthus 22 1900. (Scribn.) Anderson. 1973. B. poZyanthus var. panicuZatus B. marginatus var. poZyanthus 22 Shear. 1900. (Scribn.) Anderson. 1973. B. porteri (Coult.) Nash. 1895. 34 B. porteri var. frondosus B. frondosus (Shear) Woot. & 37 Shear. 1900. Stand!. 1912. B. pratensi-s Ehrh. 1791. B. commutatus Sthrad. 1806. 47 B. pumpeZZianus Scribn. 1888. B. inermis ssp. pumpeUianus 34 (Scribn.) Wagnon. 1950. B. purgans var. vulgaris B. vulgaris (Hook.) Shear. 42 Hook. 1840. 1900. B. racemosus L. 1762. 51 B. racemosus var. commutatus B. comrrn1.ta'tU.sSchrad. 1806. 47 (Schrad.) Coss. &Dur. 1855. B. richardsoni Link. 1833. B. canadensis Michx. 1803. 39 B. rigidus auct. F. N. Amer. B. diandrus Roth. 1787. 67 B. rigidus Roth. Rejected. Probably to be found only in coastal sands of lower elevations. Not found in Utah. 85

BROMUS BROMUS P_age 8. r'Ubens L. 1755. 57 B. scabratus Scrubn. 1898. B. porteri (Coult.) Nash. 1895. 34 B. sahPaderi Kunth. 1833. B. wiZZdenowii Kunth. 1829. 25 B. scoparius var. rubens (L.) · B. rubens L. 1755. 57 St. Amans. 1821. B. secalinus L. 1753. 52 B. secaZinus -var. gZadewitzii B. aommutatus Schrad. 1806. 47 Farwell. 1926. B. setaceus Buckl. 1862. B. tectorum L. 1753. 60 B. squarrosus var. patulus B. japonicus Thunb. 1784. 43 (Mert. & Koch) Regel. 1881. B. squarrosus var. y,acemosus B. racemosus L. 1762. 51 (L.) Regel. 1881. B. sterilis L. 1753. 65 B. tectorum L. 1753. 60 B. tectorum var. gZabratus 64 Spenner. 1825. B. tectorum var. nudus Klett. B. tectorum var. glabratus 64 & Richt. 1830. Spenner. 1825. B. tectorum forma nudus (Klett.· B. tectorum var. glabratus 64 & Richt.) St. John. 1937. Spenner. 1825. B. trinii Desv. 1853. 69 B. trinii var. pallidiflorus B. trinii Desv. 1853. 69 Desv. 1853. B. unioloides (Willd.) Rasp. B. willdenowii Kunth. 1829. 25 1825. B. virens Buckl. 1862. B. carinatus Hook. &Arn. 1840. 23 B. virens var. minor Scribn. B. carinatus Hook. &Arn. 1840. 23 ex Beal. 1896. B. vulgaris (Hook.) Shear. 1900. 42 B. vulgaris var. ex~:»zius B. vulgaris (Hook.) Shear. 42 Shear. 1900. 1900. B. vulgaris var. robustus B. vulgaris (Hook.) Shear. 42 Shear. 1900. 1900. B. wiUdenow·ii Kunth. 1829. 25 CERATOCHLOA B. brevia,ristata Hook. 1840. B. willdenowii Kunth. 1829. 25 B. cathartica (Vahl.) Herter. B. willdenowi{ Kunth. 1829. 25 1840. 86 CERATOCHLOA BROMUS Page C. graruiiflora Hook. 1840. B. oarinatus Hook. erC Arn. 1840. 23 C. marginata (Nees.) Jacks. B. marginatus Nees. 1855. 19 1893. C. penduZa Schrad. 1831.· B. uJiZZdenowii Kunth. 1829. 25 C. unioloides (Willd.) Beauv.- B. wilidenowii Kunth. 1829. 25 1812. DANTHONIA D. pseudo-spiaata C. Muell. B. trinii Desv. 1853. 69 1856.

FESTUCA F. inermis (Leyss.) DC. & B. inermis Leyss. 1761. 29 Lam. 1805. F. inermis var. vilZosa Mert. B. inermis Leyss. 1761. 29 & Koch. 1823. F. rubens (L.) Pers. 1805. B. rubens L. 1755. 57 F. unioZoides Willd. 1803. B. wiZZdenowii Kunth. 1829. 25 FORASACCUS F. brebiaristatus (Hook.) B. uJi.Zldenowii Kunth. 1829 25 F. aommutatus (Schrad.) B. aommutatus Schrad. 1806. 47 Bubani. 1901. F. inermis (Leyss.) Lune 11. B. inermis Leyss. 1761. 29 1915. F. marginatus (Nees.) Lunell. B. marginatu.s Nees. 1855. 19 1915. F. moZZis (L.) Bubani. 1901. B. moUis L. 1762. 53 F. patuZus (Mert. & Koch) B. japoniaus Thunb. 1784. 43 Bubani. 1901. F. raa0~osus (L.) Bubani. B. r-aaemosus L. 1762. 51 1901. F. seaaZinus (L.) Bubani. B. seaalinus L. 1753. 52 1901. SCHEDONORUS S. inermis (Leyss.) Beauv. B. inermis Leyss. 1761. 29 1812. S. steriZis (L.) Fries. 1843. B. steriZis L. 1753. 65 S. teatorum (L.) Fries. 1843. B. tectOl"Um L. 1753. 60 SERRAFALCUS S. aorronutatus (Schrad.) Bab. · · B. aommutatus Sch rad. 1806. 47 1843. 87 SERRAFALCUS BROMUS Page S. moZZis (L.) Parl. 1840. B. moZZis L. 1762. 53 S. patu.Zus (Mert. & Koch) Parl. B. japonicus Thunb. 1784. 43 1848. S. raaemosus (L.) Parl. 1840. B. raaemosus L. 1762. 51 S. racemosus var. aommutatus B. comrnutatus Schrad. 1806. 47 (Schrad.) Husnot. 1899. S. secaZinus (L.) Bab. 1843. B. secalinus L. 1753. 52 TRAGUS T. unioZoides (Willd.) Panz. ex B. uJiZZdenowii Kunth. 1829. 25 Jacks. 1895. TRISETOBROMUS T. hirtus (Trin.) N~vski. 1934. B. ti>inii Desv. 1853. 69 TRISETUM T. ba.J>batwnSteud. 1854. B. trinii Desv. 1853. 69 T. barbatwn var. major Vasey. B. trinii Desv. 1853. 69 1893. T. hu:PtumTrin. 1836. B. trinii Desv. 1853. 69 T. trinii (Desv.) Louis-Marie. B. trinii Desv. 1853. 69 1928. T. trinii var, majus (Vasey) B. t-Pinii Desv. 1853. 69 Louis-Marie. 1928. T. trinii var. paZZidiflorus B. trinii Desv. 1853. 69 (Desv.) Loui~-Marie. 1928. ZERNA z. inermis (Leyss.) Lindm. 1918. B. ine.mzis Leyss. 1761. 29 z. P'ubens (L.) Grossh. 1939. B. .r>UbensL. 1-755. 57 z. sterilis (L.) Panz. ex B. sforiZis L. 1753. 65 Jacks. 1843. z. teatorum (L.) Panz. ex B .. tectorum L. 1753. 60 Jacks. 1895. z. vulgari.s (Hook.). Henrard. B. v-i,1,lga11is (Hook.) Shear. 1900. 42 1941. DISTRIBUTIONAND RELATIONSHIPS OF UTAHBROME GRASSES

IN WESTERNNORTH AMERICA

Jerry B. Anderson

Department of Botany and Range Science

Brigham Young University

M. S. Degree, April 1973

ABSTRACT

A considerable amount of study material has beccme available since the revision of the genus in 1900 by Shear, and the revision of the section Bromopsis in 1952 by Wagnon. A number of cytological studies have been done in recent years; and Wagnon's work pointed out the need for a comprehensive study of the entire genus.

This study is a taxonomic revision of the Genus Bromus for the State of Utah, and contains illustrations and descriptions of the species; synonomy; distribution maps; and discussions of many of the important aspects of the economics, ecology, and historical treatment. I have recognized 17 species and 2 varieties as being present in the State, and these, along with another 4 closely allied or formerly cited species or varieties reported for Utah, are separated by means of a comprehensive dichotomous key based on the natural grouping of these grasses into sections within the genus.

Much of the research has been based 0n herbarium study of the morphological variation for the separation of taxa, and was supplemented with extensive field observation and collection during the past decade.

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