A Revision of the Hedysarum Boreale Complex

Brigham Young University BYU ScholarsArchive

Theses and Dissertations

1969-05-01

A revision of the hedysarum boreale complex

Terry Edwin Northstrom Brigham Young University - Provo

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BYU ScholarsArchive Citation Northstrom, Terry Edwin, "A revision of the hedysarum boreale complex" (1969). Theses and Dissertations. 8102. https://scholarsarchive.byu.edu/etd/8102

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]. THE GENUS HEDYSARUM IN NORTH AMERICA

A Dissertation Presented to the Department of Botany Brigham Young University

In Partial Fulfillment of the Requirement for the Degree Doctor of Philosophy

by Terry Edwin Northstrom August 1974 This dissertation by Terry Edwin Northstrom, is accepted in its present form by the Department of Botany of Brigham Young University as satisfying the dissertation requirement for the degree of Doctor of Philosophy.

August 16, 1974 Date

ii ACKNOWLEDGEMENTS

My cordial appreciation is expressed to Dr. Stanley L. Welsh, Brigham Young University, under whose supervision this study was carried out. Special thanks are given to Mrs. Kay Thorne for her help in preparing the illustration, Jim Allen for his help with the scanning electron microscopy and to the curators of the following herbaria who generously loaned their specimens: University of Alaska Herbarium (ALA); University of Arizona Herbarium (ARIZ); Brigham Young Uni- versity Herbarium (BRY); British Museum (BM); California Academy of Science Herbarium (CAS); National Museum of Can- ada (CAN); University of Colorado Museum (COLO); Dudley Herbarium of Stanford University (DS); Gray Herbarium of Harvard University (GH); Komarov Botanical Institute Herbar- ium (L); University of Michigan Herbarium (MICH); University of Montana Herbarium (MONT); Montana State University Her- barium (MONTU); New York Botanical Garden (NY); North Dakota State University Herbarium (NDA); University of Notre Dame Herbarium (ND); Bebb Herbarium of the University of Oklahoma (OKL); Oregon State University Herbarium (OSC); Herbarium of the Philadelphia Academy of Sciences (PH) ; Rocky Mountain Herbarium (RM); South Dakota State University Herbarium (SDU); United States Forest Service Herbarium (USFS); United States National Museum (US); University of Utah Herbarium iii (UT); Utah State University Herbarium (UTC); University of Washington Herbarium (WTU); Washington State University Herbarium (WS).

iv TABLE OF CONTENTS

Page ACKNOWLEDGEMENTS...... iii LIST OF TABLES. • • . • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • vi LIST OF FIGURES ••.•••••••.•••••.•.••••...•...•.•.•.•... vii

INTRODUCTION. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 1 HISTORICAL ACCOUNT...... 9 DISTRIBUTION...... 11 TllE GROWTH CYCLE • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 12 GENERAL MORPHOLOGY. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 14 ANATOMY OF HEDY SAR UM. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 2 2

CHRO.MOSOMES. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 3 5

PH'YTOCHEMISTRY. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 52 TAXONOMY OF HEDYSARUM. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 7 4 INDEX TO EXSICCATAE...... -160 BIBLIOGRA.PHY • ••••••••••••••••••••••••••••••••••••••••• • 17 5

v LIST OF TABLES

Table Page 1. Taxa recognized for North America by various Authors since 1803 •••••••••••••••••••••• 4 2. Index of Hedysarum taxa described for North .Alnerica • ••••••••••••••••••••••••••••••• -. • • • • 6 3. Range and mean seed weights for Hedysarum taxa 21 4. Pollen grain measurements of Hedysarum taxa ••• 34 5. Chromosome reports for North American Hedy- sarUin taxa...... • ...... 3 7 6. Phytochemical summary of Hedysarum taxa ••••••• 72 7. Quantitative variation of Hedysarum alpinum ••• 111 8. Character scores of Hedysarum alpinum ••••••••• 117

vi LIST OF FIGURES

Figure Page

1. Flower of Hedysarum boreale ssp. mackenziei.. 38 2. Scatter diagram depicting the relationship of the width to length of the loment articles i~ H. alpinum, ~- sulphurescens and ~- ~~ cidentale...... 3 9

3. Stem cross section of H. sulphurescens...... 40 4. Vascular bundle of H. occidentale...... 40

5. Xylem elements of H. boreale ssp. boreale var. rivulare ...... :-...... 41 6. Endodermis of H. occidentale...... 41

7. Leaf cross section of H. occidentale...... 42 8. Leaf cross section of H. boreale ssp. boreale 42 9. Cross section of petiole of H. boreale ssp. boreale var. rivulare ••••• 7...... 43 10. Cross section through receptacle of H. sulphurescens • ••••••••••••••.•••••••• 7 .. :-:-:-. . . 44 11. Androecium cross section of H. boreale ssp. bore ale ...... 7...... • ...... 44 12. Ovule of H. boreale ssp. boreale...... 44 13. Cross section of young anther of H. boreale ssp. boreale •.••••••.•....•••.• 7...... 45 14. Mature anther wall of H. boreale ssp. boreale 45

15. Cross section of filament of ~· alpinum ••••••, 46 16. Cross section of corolla of H. alpinum...... 46

17. Cross section through calyx tube of li· ~- ~ ssp. boreale...... 47 vii Figure Page 18. Cross section through calyx tooth of H. boreale ssp. boreale ••••••••••••••• 7...... 47 19. Cross section of mature root of H. occidentale ...... :- ...... 4 8 20. Root periderm of H. occidentale...... 48

21. Pericarp cross section of H. alpinum...... 49 22. Seed cross section of H. alpinum...... 49

23. Pollen morphology of selected Hedysarum taxa. SO

24. Chromosomes of selected Hedysarum taxa...... Sl

25. Densitometer scan of H. boreale...... 54 26. Densitometer scan of H. occidentale...... SS

27. Habit of ~· boreale ssp. boreale var. rivulare. . . . • ...... • ...... 7 8 28. Distribution of H. boreale ssp. boreale var. rivulare ...... :-...... 80

29. Lament of ~· boreale ssp. boreale var. gremiale...... • ...... 87

30. Habit of H. boreale ssp. boreale var. boreale 89 31. Habit of H. boreale ssp. mackenziei...... 94

32. Polygonal graph comparing 8 characteristics of H. dasycarpum (U.S.S.R.) and H. boreale ssp. mackenziei (U.S.)...... 97 33. Polygonal graph comparing 8 characteristics of Old and New World H. alpinum ...... 100

34. Ecological variation in H. bore·a1e ••••••••••• 102

35. Ecological variation in H. sulEhurescens ••••• 103

36. Ecological variation in H. occidentale ••••••• 104

37. Ecological variation in H. alEinum ••••••••••• lOS

38. Polygonal graph comparing 8 characteristics of H. hed~saroides (Europe) and H. al}2inum var. g:randiflo·rum (U.S.) ••••••••••••••••••• 108 viii INTRODUCTION

Considering the fact that within the genus Hedysarum in North America some 46 epithets have been bestowed upon its various forms in American floras since 1803, {Table 1) Hedy- sarum lends itself well to a detailed morphological and biosystematic study. In fact, not only are the systematics of Hedysarum taxa widely disputed, but their geographical distribution, their ecological and physiological responses and their anatomy and morphology are poorly known. It has thus seemed worthwhile to undertake such a study.

The genus Hedysarum in th~ family Leguminosae is a somewhat large genus of Old World origin which is largely north temperate and circumpolar in distribution. Estimates of the number of species within the genus vary widely.· B. A. Fedtschenko in his monograph of 1902 recognized 78 species for tbe world, but in a later work on the flora of the u. S. s. R. (1948), he recognized 88 species for the Soviet Union alone. At the other extreme, Willis (1973) estimates the total number of species at 150. After having critically examined the North American.members of the genus, and assuming that the Old World entities have been treated equally, it appears that the total number of species does not exceed 100. In North America, Hedysarum taxa span an altitudinal 2 range of from sea level to 13,000 feet, a latitudinal range which extends from southern Arizona to Banks Island, and a longitudinal range extending from northwestern Alaska to Newfoundland. It occupies a vast variety of habitats ranging from gravel bars, sandy areas, roadsides and woodlands at low and moderate elevations, to alpine tundra and stony outcrops at high elevations or extreme northern latitudes. Thus, as one might expect, Hedysarum taxa exhibit a considerable degree of polymorphism which is usually expressed in the form of a gradient in morphological and physiological characteristics, the extremes of which often appear as distinct entities occupying relatively large geographical areas. This appears to account for the numerous species, subspecies, and varieties as well as recombinations which have been described over the past 150 years (Table 2). Previous workers have dealt mainly with taxonomic considerations and little work has been done on the morphology, anatomy, and general biology of Hedysarum. This study is thus oriented from a somewhat biosystematic point of view. In order to correctly interpret taxonomically such a highly divergent group of taxa, a thorough knowledge of the anatomy, chemistry, cytology, morphology, and reproductive biology of at least one representative member of the genus is necessary. The study includes the following topics: 1) an anatomical study of each member of the genus which clearly merits recognition as a distinct taxon; 2) a detailed mor- 3 phological study of all major variants throughout their range using herbarium specimens from major American and European herbaria; 3) a study of their geographical distribution; 4) an analysis of phytohemaglutinins,; S) a chromosome analysis; 6) a phytochemical survey; 7) a quantitative and qualitative analysis of crude protein content. 4

TABLE 1 TAXA RECOGNIZED FOR NORTH AMERICA BY VARIOUS AUTHORS SINCE 1803

Micheaux 1803 ~~ alpinum L. var. americanum Michx. Hooker 1830 H. boreale Nutt., H. mackenziei Richards. Torrey & Gray 1840 H. boreale Nutt., H. mackenziei Richards., H. canescens Nutt. Gray 1857, Watson 1878, H. boreale Nutt., H. mackenziei Macoun 1883, Coulter 1885 Richards. Heller 1900 H. americanum (Michx.) Britt., H. carnosulum Greene, H. cinerascens Rydb., H. lancirolium Rydb., H. leucanthum Greene, H. mackenzI'ei Richards., H. mar= ginatum Greene, ~· occidentale Greene, ~· sulphurescens Rydb. Robinson & Fernald 1908 H. boreale Nutt. Britton 1913 H. boreale Nutt., H. mackenziei Richards. Piper & Beattie 1915 H. occidentale Greene Rydberg 1922 H. sulphurescens Rydb., H. americanum (Michx.) Britt., H. boreale Nutt., H. lancifoliUm R'Y

Graham 1937 H. cinerascens Rydb., ~· ~ hense Rydb. 5 TABLE 1 (continued) Dole 1937 H. boreale Nutt. Polunin 1940 H. mackenziei Richards. Tidestrom & Kittell 1941 H. pabulare A. Nels. Peck 1941 H. mackenziei Richards. Hulten 1946 H. alpinum L., H. mackenziei Raup 1947 Richards. Stevens 1950 H. boreale Nutt. Davis 1952 H. occidentale Greene, H. sulphurescens Rydb., H. boreale Nutt. Gleason 1952 H. alpinum L. Weber 1953 H. boreale Nutt. Harrington 1954 H. occidentale Greene, H. boreale Nutt. - --- Porsild 1955 !· alpinum L., H. mackenziei Richards. Anderson 1959 H. alpinum L., H. mackenziei Polunin 1959 Richards. Kearney & Peebles H. boreale Nutt. Hitchcock 1961 H. alpin um L. , ! . boreale Nutt., !· sulehurescens Rydb., !· occidentale Greene Gleason & Cronquist 1963 H. alpinum L. Boivin 1967 H. sulphurescens Rydb., H. alpinum L. ,· !· boreale Nutt. Weber 1967 H. boreale Nutt. Hulten 1968 H. mackenziei Richards., H. alpinum L. , H. hedrsarioides (L.) Schinz.-& The 1. Seymour 1969 H. alpinum L. 6 TABLE 2 INDEX OF HEDYSARUM TAXA DESCRIBED FOR NORTH AMERICA

H. albiflorum (Macoun) Fedtsch., Acta Hort. Petrop. 19:252. 1902. H. alpinum L., Sp. Pl. 750. 1753. H. alpinum L. subsp. americanum Fedtsch., Acta Hort. Petrop. 19:255. 1902. H. alpinum L. var. americanum Michx., Fl. Bor. Arn. 2:74. 1803. H. alpinum L. var. americanum Michx. f. albiflorum Fernald, Rhodora 35:275. 1933. H. alpinum L. var. grandiflorum Rollins, Rhodora 42:233. 1940. H. alpinum L. var. philoscia (A. Nels.) Rollins, Rhodora 42: 233. 1940. H. americanum Britt., Mem. Torr. Bot. Club. 5:201. 1894. H. americanum Britt. var. mackenziei Britt., Mem. Torr. Bot. ClUb. 5:202. 1894. -H. auriculatum East., Bot. Gazette 33:205. 1902. H. boreale Nutt., Gen. N. Arn. Pl. 2:110. 1818. H. boreale Nutt. subsp. boreale var. gremiale (Northstrom & Welsh) Rollins, Gr. Basin Nat. 30:125. 1970. H. boreale Nutt. subsp. mackenziei (Richards) Welsh, Gr. Basin Nat. 28:152. 1968. H. boreale Nutt. var. albiflorum Macoun, Cat. Canad. Pl. 1:510. 1884. H. boreale Nutt. var. cinerascens (Rydb.) Rollins, Rhodora 42:235. 1940. H. boreale Nutt. var. cinerascens (Rydberg) Rollins f. album B. Boi., Nat. Canad. 87:34. 1960. H. boreale Nutt. var. flavescens Coult. & Fisch., Bull. Herb. Boiss. 7:256. 1899. H. boreale Nutt. var. leucanthum Jones, Proc. Calif. Acad. Sci. 5:677. · 1S95. 7

TABLE 2 (continued) H. boreale Nutt. var. mackenziei (Richards) c. L. Hitchc., Vase. Pl. Pac. N. w. 3:275. 1961. H. boreale Nutt. var. obovatum Rollins, Rhodora 42:235. 1940. !• boreale Nutt. var. typicum Rollins, Rhodora 42:232. 1940.

H. boreale Nutt. var. utahense (Rydb.) Rollins, Rhodora 42: 232. 1940. H. canescens Nutt., T. & G. Fl. N. Am. 1:357. 1838.

H. carnosulum Greene, Pitt. 3:212. 1897.

H. cinerascens Rydb., Mem. N.Y. Bot. Gard. 1:257. 1900. H. flavescens Coult. & Fisch., Bot. Gaz. 18:300. 1893.

H. gremiale Rollins, Rhodora 42:230. 1940.

H. hedysaroides (L.) Schinz & Thellung, Viert. Nat. Ges. Zurich 58:70. 1913. !· lancifolium Rydb. Mem. N.Y. Bot. Gard. 1:256. 1900. H. mackenziei Richards. var. canescens (Nutt.) Fedtsch. Acta. Hort. Petrop. 19:362. 1902. (sic) !· mackenziei Richards. Frankl. 1st. Journ. Bot. App. 745. 1823.

H. mackenziei Richards. var. fraseri B. Boi., Canad. Field. Nat. 65:20. 1951.

H. mackenziei Richards. var. mackenziei f. niveum B. Boi., Canad. Field Nat. 65:20. 1951.

H. mackenziei Richards. var. pabulare (A. Nels.) Kearney & Peebles, J. Acad. Sci. 29:485. 1939.

H. marginatum Greene, Pitt. 4:138. 1900.

H. occidentale Greene Pitt. 3:19. 1896. H. eabulare A. Nels., Proc. Biol. Soc. Wash. 15:185. 1902.

H. eabulare A. Nels. var. rivulare Williams, Ann. Mo. Bot. Gard. 21:344. 1934. 8

TABLE 2 {continued) H. philoscia N. Nels., Proc. Biol. Soc. Wash. 15:185. 1902. H. roezlianum Prantl., Ind. Sem. Hort. Wirceb. 8. 1873. H. sulphurescens Rydb., Bull. Torr. Bot. Club 24:251. 1897. H. truncatum East., Botan. Gaz. 33:205. 1902. H. uintahense A. Nels., Proc. Biol. Soc. Wash. 15:186. 1902. H. utahense Rydb., Bull. Torr. Bot. Club 34:424. 1907. 9

HISTORICAL ACCOUNT

The genus Hedysarum was established by Linnaeus in 1753 on the basis of g. alpinum L. The generic name origi-

. nates from the Greek word 11 o v u a p o P (Hedysaron) found in the fifteenth century herbal of Dioscorides. The first syllable 11 o v u (hedys) = sweet, and the second syllable a po P

(~) = smell, derived obviously from the sweet smell which many species of Hedysarum emit from their flowers. Four species of Hedysarum are recognized in this treatise for North America. The first species was described by Micheaux (1803) as ~· alpinum-americanum. Micheaux considered the New World plants to be a variety of the Old World H. alpinum L. Fedtschenko (1902) thought the American material to be less closely related than did Micheaux and divided the Asian and American material subspecifically, while Britton (1894) considered the differences to be suf- ficiently distinct to warrant specific segregation (as H. americanum Britt.). The second North American species of Hedysarum was described by Thomas Nuttall as H. boreale Nutt. in his . - Genera of North American Plants (1818). A definite misunder- standing of g. boreale by Nuttall together with a poor description led later workers into considerable confusion as to the true identity of this species. 10 Hedysarum mackenziei, a close relative of H. boreale, was described by Richardson (1823) as a distinct species. In recent years, however, it has been recognized not only as a species but also as a subspecies (ssp. mackenziei (Richards) Welsh) and as a variety (var. mackenziei (Richards) c. L. Hitchcock) • The third species recognized in this treatise, [·

J sulphurescens, was described as H. flavescens by Coulter and Fischer (1893) who were apparently unaware of the previous description of H. flavescens Regel & Schmalh. (1882). Ryd- berg corrected this misnomer (1897) with the description of H. sulphurescens Rydb. The fourth species, H. occidentale, was described by Greene (1896) from the Olympic mountains of Washington. No comprehensive study of Hedysarum for the entire North American continent has been made. Fedtschenko (1902) published a world revision of the genus Hedysarum, but his treatment of American material was highly unsatisfactory, undoubtedly the result of an inadequacy of North American specimens. The only other significant study was that of Rollins (1940), although, Alaska, the Northwest Territories, and the Yukon were not included in his monograph. Moreover, no distribution maps, illustrations or experimental data were in- eluded. 11

DISTRIBUTION

In North America, Hedysarum taxa occur from North- western Alaska, Banks and Victoria Islands, across much of the Mackenzie and Keewatin districts to Labrador and New- foundland, south to New Brunswick, Northern Maine and Ver- mont, then west through Quebec, Ontario and the prairie provinces to British Colwnbia, south to Washington and eastern Oregon, then eastward through Idaho, Montana, and Wyom- ing to North and South Dakota. They then extend southward along the Rocky Mountains through Utah, Colorado and northwestern Oklahoma to northern New Mexico, Arizona and Nevada. 12

THE GROWTH CYCLE

The perennating buds of Hedysarurn are found at or near the level of the soil and are usually covered with litter during the dormancy period. The required length of the dormancy period is apparently quite short as plants transplanted to pots in the fall and kept at s0 c. for as little as three weeks will break dormancy with an increase in temperature. The relationship of the photoperiod to the breaking of dormancy in Hedysarurn is not known, although transplanted and refrigerated specimens of ~· boreale subsp. boreale will break dormancy throughout the fall and winter regardless of photoperiod. On the foothills of the Wasatch Mountains in northern Utah, vegetative growth assumes immediately after the breaking of dormancy and expansion of the buds in the spring, with flowering stalks appearing from 7-15 days thereafter, and anthesis after an additional five to ten days. The plants remain in flower for three to five weeks with loments maturing after an additional two to three weeks. In the late fall, a small amount of vegetative growth occurs from the caudex just before the onset of dormancy. The seeds apparently do not require a dormancy period and will germinate on moist filter paper immediately after dispersal, although in the field, they seldom, if 13 ever, germinate until late in the following spring. This is undoubtedly due to the thickness of the seed coat and the resistance to germination which it renders on the seed. This resistance appears to be only a mechanical one as cracking the seed coat reduces germination time by fifty to seventy percent. The seeds of all four species of Hedysarum germinate readily in two to four days after mechanically or chemically disrupting the testa. Whether or not the seeds of all Hedysarum taxa germinate in the spring season is not known. Surely latitude and elevation play some part, and plants from extreme southern latitudes or low elevations could and may germinate in the fall and over winter in a vegetative state. 14

GENERAL MORPHOLOGY

Habit

The North American species of Hedysarum are all long- lived perennials. Their relative life span depends upon various environmental conditions, however, tap roots as old as six years are conunonly observed in the field. The solitary to numerous aerial shoots arise from a woody caudex at about the level of the soil with the degree of branching varying to a large extent with environmental conditions. In habit they range from nearly prostrate to decumbent, ascending or erect.

Roots The root system of Hedysarum is a branched ligneous taproot. The young plant forms a taproot which develops rapidly reaching more than a meter in length during the first season of growth. The main branch roots are formed in the first few centimeters of soil and spread laterally for only a short distance before assuming a course similar or parallel to the main taproot. After maturation of the primary tissue, a cambium develops which forms large sectors of secondary tissue easily discernable as rings of seasonal growth. Thus, in the main roots, a massive woody axis is formed particularly 15 in H. occidentale whose root system is more extensive than that of any other of the included species. The roots of H. alpinum are reported to be utilized as a source of food by Indians and mammals in subarctic and boreal North America. They also served as a food source for early expeditions into Alaska and Northern Canada (Richardson 1823).

Stems The solitary to numerous aerial shoots arise from a stout woody caudex consisting of the stem bases of previous years growth. The caudex nodes are surrounded by thin, papery, sheathing stipules. The caudex, or crown, is initiat- ed from buds in the axils of the cotyledons and the first foilage leaves. These buds develop coincident with the primary axis which forms from the epicotyl. The second and following years growth results from perennating buds found in the axils of the primary shoots. Shoots may also arise adventitiously from buds developing at or near the base of the lateral branches. Adventitious roots sometimes develop also from the base of the caudex, particularly in transplanted plants. The young stem is terete and solid, but disintigra- tion of much of the pith parenchyma late in the growing season results in the formation of a hollow cylinder. The mature stem is somewhat angular in cross section and often becomes longitudinally grooved late in the growing season due to the differentiation of strips of thick walled collen- 16 chyma at the stem angles. The main vascular bundles are situated at the stem angles and more collenchyma develops between the main bundles and the epdiermis than at any other position in the stem.

The Leaf The pinnately compound petiolate leaves are alter- nately arranged and the elliptical to lanceolate, oblong, or ovate leaflets are in two to twelve pairs with a single terminal leaflet. In texture, the petiolulate leaflets are herbaceous with entire margins and reticulate venation. The North American species of Hedysarum can be divided into two well defined natural groups on the basis of leaflet venation. In H. sulphurescens, ~· occidentale, and H. alpinum, the veins and veinlets are positioned close to the abaxial epidermis forming prominent ridges and thus giving a marked reticulate appearance. In H. boreale, the veins and veinlets are embedded deep in the mesophyll giving a hidden appearance when viewed from the surface. Leaflet characters other than venation are less stable but nevertheless are useful as diagnostic features. The leaflets of H. occidentale and H. sulphurescens are quite similar in shape ranging from ovate to elliptic or more rarely oblong but differ in size, number, and pubescence. H. occidentale has 11-19 leaf lets which are usually pubescent on both surfaces while the 9-17 leaflets of H. sulphurescens are glabrous above and pubescent below. H. alpinum is characterized by broadly lanceolate or rarely elliptic or oblong 17 leaflets which are glabrous above and very sparsely pubescent below. The 5-15 leaflets of H. boreale range from ovate to elliptic, linear or rarely obovate and are glabrous above and pubescent below except in H. boreale ssp. boreale var. boreale which has both leaflet surfaces pubescent. The stipules are triangular to subulate and pubescent. In texture, they vary from membranous and white-translucent to dark brown and chartaceous. Hedysarum taxa exhibit considerable infraspecific variation in the degree of connation of the stipules. The lower stipules are usually united, although sometimes partially free, while the upper ones are usually partially free and occasionally entirely free.

The Flower The purple, yellow, or rarely white flowers are born in compact to elongate axillary racemes with elongate pedun-. cles. Each flower is subtended by a single subulate bract. Albino forms are occasionally found in each species. (Fig. 1) The bracteolate, pubescent calyx consists of five undiverged sepals terminated by five teeth. The bracteoles occur in symmetrical pairs at the base of the campanulate tube. In !!· alpinum, !!· sulphurescens, and H. occidentale, the calyx teeth are markedly unequal and shorter than the tube while in !!· boreale, the teeth are equal or nearly so and longer than the tube. In shape, the teeth range from nearly deltoid in H. alpinum to linear subulate in H. boreale. The papilionaceous corolla consists of five glabrous 18 petals: a broad spathulate-cuneate standard which encloses the others in the bud: a keel composed of two small petals ·which are united along their ventral margins and surround the androecium and gynoecium: and two lateral wings which extend obliquely outward. The wings possess a small auricle which

in H. alpinum, H. occidentale and ~· sulphurescens are united under the standard and are equal to or exceed the wing claw in length. In H. boreale, the wing auricles are free, short and do not exceed one third the length of the wing claw. The diadelphous androecium consists of nine terete, threadlike filaments which are undiverged at their bases forming a subcylindric tube which encloses the gynoecium, and a tenth stamen which is free. The anthers are two lobed with four sporangia. The gynoecium consists of a single carpel with an elliptical and laterally flattened one-celled superior ovary. One to six ovules arise in a single row on one parietal a- daxial placenta. The ovules are amphitropous when young, but campylotropous when mature. The glabrous, slender style is curved, erect and topped by a minute stigma.

The Fruit The fruit of Hedysarum is a flat loment which is constricted into several indehiscent, one-seeded articles. The surface of each article bears elevated reticulations which form prominent aerolae at maturity. In H. boreale ssp. boreale var. gremiale, distinct spines arise from 19 these reticulations which may reach five nun. in length. Small tubercles occasionally occur on the loments of H. boreale ssp. boreale var. boreale although they are never as prominent as in var. gremiale. Morphology of the loment article has proven to be of considerable value in delimiting taxa. Two groups of species can be separated on this basis. The first group, containing ~· sulphurescens, ~· occidentale and H. alpinum, has winged margins and polygonal aerolae. A scatter diagram (Fig. 2) depicts the relationship of the width to length of the loment articles in H. alpinum, H. sulphurescens and H. occidentale. The loments of ~· occidentale are pubescent and larger than any other North American species, ranging from 6.1-10.5 nun. in width, and 8.0-17.6 mm. in length. H. sulphurescens has glabrous loments ranging from

5.5-8.7 mm. in width and 7.2-13.5 mm. in length, and~· alpinum has loments which range from 3.1-7.9 nun. in width and 5.0-10.5 nun. in length. H. alpinum var. philoscia has pubescent loments while var. grandiflorum and var. alpinum usually have glabrous loments. The second group, containing both subspecies of H. boreale, has usually glabrous loments which range from 4.2- 6.4 mm. in width, and 5.0-8.3 mm. in length. In general, the width of the wing margin varies in proportion to the size of the loment.

The Seed The kidney shaped, golden brown to dark brown seeds 20 range from 2.1 to 5.2 mm. in length. The range and mean seed weights for all taxa are summarized in Table 3. 21

TABLE 3 *SEED WEIGHTS OF HEDYSARUM TAXA

Taxon Weight Range Weight Mean

H. alpinum var. .0022-.0044 .0036 al:e.:l.num H. al:einum var. .0031-.0056 .0040 grandiflorum H. al:einum var. .0024-.0049 .0037 Ehiloscia H. boreale ssp. .0061-.0108 .0091 boreale var. bore ale var. rivulare .0059-.0112 .0095 var. g:remiale .0069-.0110 .0095 H. boreale ssp. .0053-.0116 .0096 mackenziei -H. occidentale .0052-.0132 .0098 H. sul12hurescens .0045-.0140 .0081

*All we.ights in grams. 22

ANATOMY OF HEDYSARUM

Fresh plant material was killed and fixed in ten times its volume of FAA (50% ETOH, glacial acetic acid, for- malin, 90:5:5) for 24 hours, washed with two changes of 50% ETOH, then dehydrated using a graded series of ethyl and tertiary butyl alcohols following the methods of Johansen (1940). Following dehydration, the tissue was infiltrated and imbedded with paraplast, sectioned, then stained with safranin and fast green. Herbarium material was soaked in absolute ETOH for 24 hours, hydrated in a graded series of ETOH, then soaked in a 4% solution of potassium hydroxide for several days to soften. The softened tissue was then dehydrated and embedded using the same procedure as with fresh material. Voucher specimens for all anatomical preparations are deposited in the herbarium of Brigham Young University (BRY) •

The Stem The stem of Hedysarum is angular in cross section, eustelic, and has a pith of large, compactly arranged, thin walled parenchyma cells which occupy a large portion of the young stem and are several times larger than the cortical parenchyma (Fig. 3). The eleven to fourteen collateral 23 vascular bundles lie in a ring toward the perifery of the stern. The major bundles are located at the axis angles with smaller ones at the intervals between. External to the phloem in each vascular bundle is a strand of thick walled protophloern fibers (Fig. 4). In the mature stern, these fibers may form a nearly continuous zone around the stele. The phloem consists of long narrow sieve tubes with simple sieve plates, companion cells and parenchyma. The primary xylem elements have spiral thickenings while the secondary vessel elements have oval bordered pits {Fig. S). Fibers and parenchyma are also present. An endodermis, or starch sheath, consisting of a single continuous row of rectangular cells delimits the cortex centripetally {Fig. 6). Adjacent to the endoderrnis centrifugally is a zone of parenchyma, followed by a zone of collenchyma of variable thickness. The collenchyma zone is usually only one or two layers thick except at the angles of the stem where it may be six or more layers thick. These angular patches of collenchyma are responsible for the longitudinal ridges often projecting from the mature stem. The epidermal cells are axially elongated and thick walled ·when mature.

As the stem matures and· secondary thickening pro- ceeds·, an interfascicular cambium develops which produces a continuous uninterrupted zone of xylem. Although the interfascicular cambium produces mainly lignified parenchyma, the fascicular cambium continues to ~roduce vessels, fibers and 24 parenchyma. As maturation continues, much of the pith parenchyma disintegrates so that the mature stem becomes hollow. Stem anatomy is essentially the same for all the species of Hedysarum examined. Minor infraspecific variation was observed in the degree of interfascicular cambium development, the abundance and distribution of phloem fibers, and the amount of secondary phloem produced.

The Leaflet The midvein extends the entire length of the leaf let while the pinnately arranged lateral veinlets form a net- veined system ending in the mesophyll. In H. alpinum, H. occidentale, and H. sulphurescens, the midvein and the lateral vein branchlets lie very close to the epidermis forming prominent ridges on the abaxial leaflet surface (Fig. 7). Each vein is surrounded by a bundle sheath of thick walled cells which become lignified on the abaxial surface. The degree of abaxial lignification decreases toward the distal part of the leaflet. In H. boreale the midvein is situated close to the abaxial epidermis but does not possess the degree of lignification found in the preceding species (Fig. 8). The lateral veinlets are separated from the epidermis by one or two layers of parenchyma and are positioned near the center of the leaflet. The mesophyll is dorsiventral with an adaxial palisade which makes up slightly more than half of the thickness of the leaf. The palisade parenchyma are several times 25 taller than broad and somewhat loosely arranged except at areas between the veins and the epidermis where they are quite compact. The spongy mesophyll consists of two or three layers of variously shaped parenchyma cells. The cells are loosely arranged and connected by lobes, thus forming a network surrounding the intercellular spaces. The vascular bundles are collateral with a small amount of secondary thickening occurring in the larger veins. Stomata occur on both epidermal surfaces but are about twice as numerous on the abaxial surface. Unicellu- lar hairs are abundant on the abaxial surface while the adaxial surface is frequently glabrous.

The Petiole The petiole is triangular or subtriangular in cross section with a longitudinal groove extending radially up to one-half of its diameter (Fig. 9). There are five main bundles with the largest one located on the abaxial side of the petiole. Numerous small vascular bundles occur between the larger ones giving a total of from eleven to fourteen. The epidermis is similar to that of the blade with a thin cuticle and numerous stomata. Adjacent to the epidermis is a single layer of collenchyma except at points between the epidermis and the vascular bundles. Here the collenchyma may reach eight or more layers of cells in thickness. The re- mainder of the subepidermal region consists of several layers of loosely arranged chlorenchyma. The central part of 26 the petiole is made up of large thin walled parenchyma cells, many of which may disintegrate at maturity. The vascular bundles resemble those of the stem. A thick band of proto- floem fibers caps each bundle.

The Receptacle A transverse section of the pedicel taken just below the receptable reveals an incompletely dissected siphono- stele in which numerous vascular bundles can be seen differ- entiating (Fig. 10). A section taken slightly above this point but below the point of departure of the calyx primordia, reveals three series of traces diverging from the vascular ring. An outer series of ten traces supply the perianth, followed centripetally by ten stamina! traces and three carpel traces.

Floral Development The flower primordium first appears as a dome shaped primordium in the axil of a small bract. The sepal primordia appear first followed by the petal, stamen and carpel primordia. Growth from tissue beneath the sepal primordia results in the formation of a protective tube which surrounds the other floral parts. The first petal primordia to appear are those of the keel, followed by those of the wings and standard. After the appearance of the petal primordia the remaining growth of the petals is very slow, with the petals remaining very small until after the differentiation of the stamens. 27 The stamens appear in two separate whorls, an inner and an outer each consisting of five primordia (Fig. 11). Soon after their appearance, growth of tissue beneath the stamina! primordia produces a collar of nine undiverged stamens, with a tenth stamen free. The free stamen origi- nates from the inner whorl of primordia and elongates at the same rate as the collar so as to maintain its anther at the same level as the other members of the whorl. In the mature flower the androecium is diadelphous with the stamens appearing to be in a single whorl with nine of them united by their filaments and a tenth one free. At maturity, the · double whorled nature of the androecium can still be detected by the earlier maturation and pollen production of the outer whorl of stamens. Before the appearance of the ovule primordia, the point of fusion of the carpel borders is still evident. During ontogeny, however, the carpellary margins become intimately fused and no evidence of fusion persists. The vascular system of the carpel consists of a single abaxial bundle and two adaxial bundles. The ovules arise from only one side of the adaxial carpel+ary margins on a single parietal placenta. The one to six crassinucellate ovules are amphitropous when young but campylotropous when mature, with two integuments (Fig. 12). The inner integument arises first but the outer integument grows more rapidly and soon encloses it. 28 The Anther A hypodermal cluster of archesporial cells divide periclinally and the outer cells form the tapetum while the inner cells form the primary sporogenous cells (Fig. 13). The epidermis and wall layers arise from tissues outside the archesporium. The anther wall consists of an epidermis, a fibrous layer, and a parietal layer adjacent to the tapetal cells. Both the tapetum and the parietal layer are absorbed by the developing microspores so that at maturity, the anther wall consists only of an epidermal layer and the fibrous layer (Fig. 14). The filament is terete and threadlike with a single trace vascular supply which continues to the anther (Fig. 15). The epidermis is cutinized, glabrous, and lacks stomata. The ground tissue consists of large somewhat loosely arranged and vacuolated parenchyma cells many of which are pigmented.

Corolla The standard, keel and wings are anatomically similar and differ only in the thickness of the mesophyll. The cutinized epidermis is lightly striated and glabrous. The epidermal cells are transversely elongated with slightly undu- lated anticlinal walls. The mesophyll consists of loosely arranged parenchyma from two to six layers in thickness. The vascular system consists of a system of small dichoto- mously branched veinlets. The mesophyll of the wings varies from two to four cells in thickness while the banner and keel varies from three to six layers thick (Fig. 16). 29 Calyx The calyx tube is strongly leaflike in anatomy with a thickly cutinized epidermis and numerous stomata. The mesophyll is undifferentiated and consists of isodiametric loosely arranged parenchyma with large intercellular spaces (Fig. 17). The anatomy of the calyx teeth is identical with that of the tube (Fig. 18). The branched system of collateral vascular bundles is similar to that of the leaf, but much less elaborate. Each collateral vascular bundle is surrounded by a slightly lignified sheath. The primary xylem elements have spiral thickenings. The bracts and bractioles are triangular in cross section with a single trace vascular supply. The epidermis is cutinized and the mesophyll consists of loosely arranged parenchyma with large intercellular spaces.

The Root The vascular cylinder of the primary root is a triarch protostele with exarch primary xylem. Each arch consists of three or four protoxylem vessels with spiral or annular thickenings. One or two reticulate metaxylem vessels occupy the center of the stele. The primary phloem cells differentiate between the radial arms of xylem. The xylem arms radiate outward and abut the uniseriate peri- cycle. The cortex consists of relatively large, loosely arranged parenchyma cells. It is limited centripetaly by an endodermis with well developed casparian strips and centrifugally by the epidermis. 30

An active cambium produces considerable secondary thickening in Hedysarum and a large woody axis often forms, particularly in H. occidentale. Three or four broad wedge shaped zones of secondary xylem are produced which are separated by parenchymatous rays of pericyclic origin which radiate centrifugally from the protoxylem points. Annual rings are well demarcated by the relatively small vessels and parenchyma together with little or no development of fibers in the summer wood, while spring growth contains numerous fibers and large vessels (Fig. 19}. Secondary phloem elements are also well differentiated with numerous fibers in the spring growth, and mostly sieve tubes and parenchyma in the summer and fall growth. Most of the cortical tissue disintegrates, and the outer portion of the mature root consists of a small amount of pericyclic tissue with a well developed periderm (Fig. 20}.

The Loment The pericarp of Hedysarum has three well developed layers (Fig. 21). The exocarp consists of a single layer of elongated thick walled epidermal cells often possessing tri- chomes. Stomata are frequently present adjacent to large air spaces. The mesocarp consists of three to five layers of large thin walled, compactly arranged parenchyma. Vascu- lar bundles with lignified sheaths are situated in the mesocarp. The endocarp contains three layers of cells. Adja- cent to the mesocarp parenchyma are two layers of fibers 31 which differ in their longitudinal orientation. The first layer is oriented parallel to the short axis of the loment, while the second is parallel to the long axis. The two sclerenchymatous layers of the endocarp are lined on the inside by a single layer of thin walled epidermal cells lacking stomata.

The Seed The outermost layer of the testa consists of a row of radially elongated macrosclereids or Malpighian cells with unequally thickened walls and tapering ends (Fi~ 22). The outer ends of these cells are covered by a thin cuticle. The hilum is surrounded by two layers of macrosclereids and a layer of thick walled schlerenchyna which extends from the hilum deep into the subepidermal layer of parenchyma. Beneath the macrosclereid layer is a layer of thick walled osteosclereids with rather large intercellular spaces. Below the subepidermal layer is a zone of thin walled parenchyma cells which are periclinally elongated and lack intercellular spaces. The endosperm consists of thin walled cells with large vacuoles which are nearly wholly absorbed by the developing embryo. The epidermal cells of the cotyledons are small, ·rectangular, and arranged end to end. The mesophyll consists of two layers of palisade parenchyma and four or five layers of spongy parenchyma. The cotyledons are high in 32 protein content and starch grains are also present. 33

POLLEN MORPHOLOGY

Pollen was obtained from dried herbarium specimens and examined using both light and scanning electron microscopy. For light microscopy, the microspores were first boiled for one minute in a 9:1 acetic anhydride-sulphuric acid solution to remove the fibrillar matrix and any extran- eous material, and then measured with an ocular micrometer. For scanning electron microscopy, the specimens were mounted on the specimen stub as chemically untreated raw pollen and then shadowed with a thin film of gold under a vacuum to de- crease distortion. Selected electron micrographs are presented in figure 23. The microspores of Hedysarum are tritreme, zonocol- porate and apocolpate. In shape, they are generally perpro- late, prolate or rarely subprolate. The sexine is subtec- tate with a reticulate tectum and baculate muri. Small bacules are also present in many of the lumina. The colpi are long and narrow with tapering acuminate ends. Ranges in pollen grain size overlap considerably as illustrated in Table 4 but some size differences are evident between certain taxa at the specific and subspecific level. 34

TABLE 4 *POLLEN GRAIN MEASUREMENTS OF HEDYSARUM TAXA

Width Width Length Length Taxon Range Mean Range Mean

H. boreale ssp. bor- 12.2-14.3 13.5 28.0-30.3 29.2 - eale var. rivUI'ai:-e -H. boreale ssp. bor- 12.8-14.6 13.2 28.6-30.0 29.3 -eale var. boreale !!.· boreale ssp. bor- 13.1-13.9 13.4 27.9-29.7 28.9 -eale var. gremiale H. boreale ssp. mack en- 14.8-17.1 16.2 26.4-32.1 29.3 ziei H. alpinum var. alpin um 14.8-16.4 15.2 27.0-29.4 28.0 H. al;einum var. ;ehil- 14.2-16.1 15.0 28.1-29.3 28.5 oscia H. al;einum var. g:rand- 13.9-15.8 14.8 27.7-30.2 28.7 iflorum -H. occidentale 13.9-16.2 15.1 21.6-28.4 25.6 H. sulEhurescens 14.4-15.8 15.3 27.8-31.4 29.S

*All measurements in microns. 35

Chromosomes Either meiotic or mitotic chromosome counts for all North American Taxa were determined from buds or seeds collected in the field. Seeds were germinated on moist filter paper. The roots were then washed and pretreated with either a 0.1% colchicine or 0.002 M. 8-hydroxy quinolin solution in an ice bath for 5-7 hours. After fixation in a 1:3 acetic-alcohol solution for 12 hours, the root tips were stained and squashed using either the acetolacmoid or acetocarmine technique. Buds were killed and fixed in a 3:1 ethanol-acetic acid solution for 12 hours, then transferred to 70% ETOH. They were then stained and squashed using the acetocarmine technique. Counts determined during the course of this study as well as counts determined by previous workers which are relative to this study are listed in Table 5. Photographs of selected mitotic and meiotic config- urations are illustrated in figure 24. No meiotic irregu- larities were noted. Voucher specimens for all counts are deposited in the herbarium of Brigham Young University. Chromosome evidence supports the taxonomic position of the species as they are delimited on the basis of morphological and anatomical features. Hedysarwn alpinum, H. occidentale, and H. sulphurescens in addition to sharing similar morphological features all have chromosome numbers of 36 n=7, while the two subspecies of H. boreale both have numbers of n=8. Johnson and Packer (1968, p. 446) reported a tetra- ploid race of H. alpinum from Ogotoruk Creek at Cape Thompson, Alaska which is characterized by having ernarginate leaflets. 37

TABLE 5 CHROMOSOME REPORTS FOR NORTH AMERICAN HEDYSARUM TAXA

Taxon n 2n Reference

H. bore ale ssp. 16 Northstrom {herein} bore ale var. 8 Northstrom {herein) boreale H. bore ale ssp. boreale var. 16 Northstrom (herein) 2rem1ale H. boreale ssp. boreale var. 16 Northstrom (herein) rivulare H. boreale ssp. 16 Johnson and Packer (1968) mackenziei 16 Knaken (1968) 16 Mulligan (1971) 16 Zhukova (1969) 16 Ledingham (1960) 16 Northstrom (herein) H. alpinum var. g:rand:lflorum 14 Northstrom (herein) H. alpinum var. Ehiloscia 14 Northstrom (herein) H. al;einum var. a112l.num 14 Northstrom (herein) 7 Holman (1962) 7 Ledingham (1957) 7 Taylor (1967) 7 Mulligan (1967) 14 Packer (1968) 14 Taylor and Brockman (1966) 14 Ledingham (1960) 14 Johnson and Packer (1968) 28 Johnson and Packer ( 1968) 16 Hedberg (1967) H. sulEhurescens 14 Northstrom (herein) H. occidentale 7 Northstrom (herein) 14 Northstrom (herein) 38

Figure 1. Flower of Hedysarum boreale ssp. mackenziei. A. Flower, B. Banner, c. Keel, D. Wing. 39 18 T 17 16 ' 15 T 14 •' ' T' • •T • T 13 TT•T• T • ' T TTT Tf 12 ·• TtT T .TT • • T• • T • : t .,-r,T, T TT •..-.. .. • • T f•6' • • ' • T • • ~ • 'l.• •:.... :.~ ' • •• , .·,~• ' 8 ...·::,:-~·· . . ••• .. '· • •I •J. • e - glabrous 7 .,,,._,.,. .. ,.. • , . • - marginally pubescent 6 • : T•:T,'•• f - pubescent • • 5 ' .' 4 ~ - H. sulphurescens 3 • - H. occidentale e - H. alpinum 2

3 4 5 6 7 8 9 10 11 12 Loment Width Figure 2. Scatter diagram comparing the loment width, length and pubescence of !!· alpinum, H. occidentale and H. sulphures·cens. 40

Figure 3. Stern cross section of H. sulphurescens.

Figure 4. Vascular bundle of H. occidentale. 41

~igure 5. Xylem elements of H. boreale ssp. boreale var. rivulare.

F: gure 6. Endodermis of H. occidentale. 42

Figure 7. Leaf cross section of H. occidentale.

Figure 8. Leaf cross section of H. boreale ssp. boreale. 43

Figure 9. Cross section of petiole of H. boreale ssp. boreale var. rivulare.

Figure 10. Cross section through receptacle of H. sulphurescens. 44

Figure 11. Androecium cross section of H. boreale ssp. boreale.

Figure 12. Ovule of H. boreale ssp. boreale. 45

Figure 13. Cross section of young anther of H. boreale ssp. boreale.

Figure 14. Mature anther wall of H. boreale ssp. boreale. 46

: ~igure 15. Cross section of filament of H. alpinum.

F:.gure 16. Cross section of corolla of H. alpinum. 47

Figure 17. Cross section through calyx tube of H. boreale ssp. boreale.

Figure 18. Cross section through calyx tooth of H. boreale ssp. boreale. 48

Figure 19. Cross section of mature root of H. occidentale.

Figure 20. Root periderm of H. occidentale. 49

Figure 21. Pericarp cross section of H. alpinum.

Figure 22. Seed cross section of H. alpinum. 50

H. sulphurescens H. alpinu~ var. alpinum

H. occidentale H. boreale ssp. boreale var. rivulare Figure 23. Pollen morphology of selected Hedysarum taxa. 51

H. sulphurescens H. boreale ssp. boreale var. rivulare

H. occidentale H. alpinum var. alpinum

Figure 24. Chromosomes of selected Hedysarum taxa. 52

PHYTOCHEMISTRY

Introduction

The contribution of phytochemistry to plant systematics has been tremendous in recent years. The Legumino-

~' in particular, has been found to be a chemotaxonomists delight with an abundance of uncommon amino acids, xantho- nes, flavanoids, hemaglutinins and various other compounds of taxonomic importance. A comprehensive review of Legume chemotaxonomy was recently written by Harborne et al (1971).

All taxa included in this treatise were examined for the presence of phytohemaglutinins, canavanine, mangiferin and various ureides. In addition, leaf element accumulation characters and seed protein electrophoretic patterns were determined for each species.

Total Protein and Protein Electrophoresis

One gram of powdered seed was extracted with pe- troleum ether at room temperature and the solvent removed by aeration. The air dried meal was then extracted at lo0 c. with three successive portions of barbiturate buffer. The buffer (Beckman B2) consists of 2.76 grams of diethyl barbituric acid and 15.40 grams sodium diethyl barbiturate diluted to one liter with distilled water giving a pH of

8.6 and an ionic strength of 0.075. Irving et al (1945) 53 showed that peanut meals extracted with an alkaline buffer contained an average of 98% of the total nitrogen in the meal, by comparison with Kjeldahl nitrogen analysis. The combined extracts were then concentrated lOOX using an Amicon macromolecular concentrator. Electrophoresis was performed on cellulose acetate strips for 30 minutes at a constant voltage of 250 volts. After electrophoresis, the strips were stained with panceau S, washed with a 5% acetic acid solution and air dried. The- strips were then scanned with a Beckman microzone densitometer with digital integrator. Two well defined electrophoretic patterns were obtained which are reproducible using this procedure. The two patterns correspond to the two sections of the genus now delimited on the basis of morphological, anatomical and cytological characteristics. Both subspecies of H. boreale produced a pattern of three well defined peaks - ' (Fig. 25) consisting of approximately 38, 54 and 8% of the total protein of the seed. ~· sulphurescens, H. alpinum and !· occidentale produced a pattern of three peaks (Fig. 26) consisting of approximately 3%, 4% and 93% of the total protein of the seed. Total seed protein was estimated by determining N by the Kjeldahl method and then multiplying by 6.25. H. alpinum contained the highest percentage of protein at 50.7% followed by H. boreale 44.2%, H. sulphurescens 42.9% and H. occidentale 39.3%. 54

'f~~~: VTON- .-~~=ii----- ~~: ~-:~~ltif5i:~~~ii: f: ~:""" •.' .~ ~ ~ . ~)~''!'.!Tu :·••ffe•~1:•.;.E £••.i;•:c: . j •C·."

.~ . '

--=...... -"" :~ ·-.· -· i=::= =--

Figure 25. Densitometer scan of H. boreale ssp.· boreale var. rivulare. 55

,_Oii 1811111 8 ·.-TQ 'N0.111111W 't1K[ 'tJU81Un.f09S 61 -F:_ l==j_ :3:: --+-+--E-~---( --!=-±c _~ .- -~ ::.-:: c_ i::~:;. F=i:_c~

Figure 26. Densitometer scan of H. occidentale. 56

Canavanine Most amino acids are rather widely distributed in nature and thus only quantitative data concerning them can be utilized as taxonomic criteria. Also, as many previous workers have shown, amino acid concentrations within a plant may vary considerably with enviromental conditions, stage of development, availability of nutrients and other factors, so that quantitative information of this type is of limited value in taxonomic studies. However, many plants have been shown to contain uncommon amino acids some of which are found scattered in diverse groups throughout the plant kingdom, while many are restricted to a particular taxon. The presence in the Leguminosae of many of these amino acids has stimulated considerable research into their distribution patterns within the family as well as simple methods for their detection. This has led to a considerable body of information which was recently reviewed by Bell (1971). One of these amino acids, canavanine H2N.C(:NH).NH.O.CH2.CH2.CH (NH2).C02H, has shown extensive value as a phylogenetic marker. Tschiersch (1961) and Nakatu et al. (1962) surveyed 97 plant families for the presence of canavanine and found it to occur only in the Leguminosae. Other workers, Turner and Harborne (1967), Birdsong et al. (1960), Bell (1958), have shown that within the family it is found only in the subfamily Papilionoideae. To date, a total of 540 species have been examined for canavanine with 60% of them positive (Bell 1971) •. ,;rnsome, genera, such as Astragalus and Vicia, 57 both positive and negative species can be found. In those plants which contain it, canavanine is usually found in fairly high concentrations. Bell (1958) reported concentrations as high as 3.8% of the total seed weight. Recent workers have suggested that canavanine serves as a nitrogen storage compound and may be active in the transport of nitrogen. Whatever its function, it is clear that canavanine is an important metabolite and its distribution within the family, when thoroughly known, will be of considerable taxaonomic and phylogenetic value. Kitagawa et al. (1929a, 1929b, 1937) demonstrated the presence of canavanine in the seeds of Canavalia ensiformis, Canavalia lineata and Canavalia obtusifolia. They also observed that canavanine gave a magenta color when exposed to a solution of sodiwn nitroprusside which had been previously exposed to sunlight (Kitagawa et al. 1932, 1936). It was later shown by Fearon (1946) that compounds which contain the guanidoxy grouping, H2N.C(:NH). NH.O.R, react with trisodiwn pentacyanoammonioferrate in a neutral solution to give colors ranging from orange-red to magenta. Canavanine is the only known naturally occurring compound found in plants that gives a magenta color with PCAF. A few compounds have been found which inhibit color formation such as creatinine and ascorbic acid, but this problem can be easily overcome by using paper chroma- tography to separate PCAF reacting compounds and then spray- ing the chromatograph with the reagent (Bell 1958). 58 From these findings have been developed simple quantitative methods for determining canavanine concentrations (Archibald 1943, Archibald 1946, Bell 1958). The PCAF reagent can be prepared by treating a solution of sodium nitroprusside with an oxidizing agent such as superoxal (30% H202), or can be purchased from a commercial distributor (Fisher Scientific Co. #S-659.). Seeds were finely ground and dried overnight. Fifty mg. of powdered seed was extracted with 2 ml. of 0.1 N HCl for 24 hours. The mixture was neutralized with 0.1 N NaOH and filtered. To 1 ml. of the filtrate was added 0.5 ml. of 1.0 M phosphate buffer and 0.5 ml. of 1% PCAF. Standard solutions of canavanine and a reagent blank were treated similarly with buffer and PCAF. After 2 hours of color development, the solutions were read in a coleman spectrophotometer in small cuvettes at 520 mu. The amount of canavanine in each extract was read from a standard curve prepared from the standard solutions and their respective optical densities. Reagents were prepared as follows. PCAF, a 1% solution, Fisher certified reagent. 1.0 M phosphate buffer, pH 7.2, 0.25 mole NaH2Po4 • H 2 ~ + 0.75 M K2HP04 • Adjust volume to 1 L. A stock solution of canavanine was prepared from urease powder using the methods of Cadden (1940), and Damodaran and Narayanan (1939). The results are presented in Table 6. All species show fairly high concentrations of canavanine. Only a few of the estimated 100 or more species 59 within the genus Hedysarum have been examined for canavanine, and a complete survey would undoubtedly provide considerable insight into the phylogeny of the genus. Tschiersch and Hanelt (1967) surveyed the genus Vicia for uncommon amino acids, and were able to subdivide it into three major groups of species which corresponded to the accepted subdivisions of the genus based upon the classical methods of morphology and cytology. It appears that the presence of canavanine in species at such high concentrations represents more than just minor genetic differences between taxa which lack it. It has been found that after accumulating at high concentrations, it then disappears during germination suggesting a role in the storage and transport of nitrogen. Furthermore, the syn- thesis of canavanine must require an appropriate set of enzymes even though it has been shown that the enzyme arginase, besides hydrolyzing arginine, hydrolyzes canavanine to canalin and urea (Damodaran and Narayanan 1940). In order to account for the evolutionary advantage a plant storing canavanine has over one that does not, Bell (1971) suggested that canavanine may confer some additional advantage on a plant, besides nitrogen storage, such as toxicity to animals or insects which would confer some selective advant~ge over a plant storing arginine.

Phytohemaglutinins The phenomenon of phytohemaglutination was first 60 observed by Stillmark (1888) using an extract of Ricinus comunis which agglutinated certain animal erythrocytes. Boyd (1947) observed that saline extracts of certain species of lima beans would specifically agglutinate erythrocytes of certain blood groups. The substances responsible for this agglutination were termed lectins by Boyd and Sharpleigh (1954) due to their selective activity. To date, lectins have been discovered which specifically agglutinate erythrocytes of groups A, B, H, M, N, as well as numerous animal, nonspecific, and multispecific lectins. On the basis of their specificity, lectins have found use in blood grouping and subgrouping, forensic medicine, immunological research and as taxonomic markers. Other biological actions include toxicity, therapeutic effects on certain anemias, and stimulation of cell division in lymphocyte cultures (Toms and Western 1971). Lectins have now been found in 79 plant families, although they are largely confined to the Leguminosae as 95% of the species reported to be positive are legumes (Toms and Western, 1971). Until recently, relatively little was known of the chemical and physical characteristics of lectins, but they are now known to be heat labile proteins of a pseudo- globulin nature and are readily soluble in water, saline, and dilute mineral acids. Recent studies have revealed a muco or glycoprotein structure for certain lectins, as well as amino acid compositions for a few (Toms and Western, 1971). 61 The highest concentration of lectins are found in viable seeds, although Eisler and Portheim (1926) reported a lectin in both the seeds and leaves of Vicia cracca. Krupe and Einsgraber (1958) found a lectin present in root, leaf and stem tissue. Toms and Western (1971) reviewed the distribution of lectins within the Leguminosae and concluded that lectin studies are of considerable taxonomic value. With regard to lectin activity, legume species can be classified as nonhemaglutinating, specific hemaglutinating, nonspecific hemaglutinating, and animal hemaglutinating species. Toms and Western (1971), noted that different investigators occasionally reported different results with the same species, which they attributed either to incorrect identification of the species, inadequate or aged seeds, or inefficient methods of extraction. It was noted that many workers have limited their studies to human erythrocytes of groups o, A, and B while only a few investigations have been made using the A and B subgroups or the many other known human blood group antigens. The above discrepancies were particularly noted with regard to the lectin studies previously made on Hedysarum species. Each extract in this study was tested against a battery of at least five A cells, five A cells, five AB 1 2 cells, five B cells and two commercial panels composed of a total of eighteen group o bloods. In addition, a panel composed of 5% cell suspensions from sheep, dog, cat, 62 guinea pig, rat and monkey bloods were tested also. Each extract was tested by saline, enzyme (Bromeline-Dade) and albumin techniques. Both seed and leaf tissue were powdered in a mortar, then extracted overnight with ten times their weight of physiological saline at 4 to 37 degrees c. the residue removed by centrifugation and the supernatant tested for lectin activity. Two drops of extract and one drop of 5% cell suspension were added to a sero-tube, mixed and incubated for 15 minutes, spun for 15 seconds and read both macroscopically and microscopically for agglutination and/or hemolysis. For the bromelase test, one drop of _enzyme is added to the cell-extract mixture, incubated and read as was the saline. To the albumin tube, one drop of 3% bovine albumin was added, incubated, centrifuged and read. Each test was carried out at 4 degrees c., 25 degrees c., and 37 degrees c. to detect cold agglutinins, or other temperature sensitive lectins. Gray (1961) has shown that bromelin increases the avidity and titer of anti A and B so as to intensify lectin activity. With the above pro- cedures, all of the North American species of Hedysarum were found to be non hemaglutinating.

Mangiferin Mangiferin is a phenolic pigment which occurs in a number of widely different plant groups including the Legu- minosae. Its distribution pattern within the family, however, has not been determined. 63 To detect mangiferin, two dimensional paper chro- matographs were developed using extracts prepared from H. alpinum var. alpinum, H. occidentale, ~· sulphurescens and H. boreale ssp. boreale var. rivulare. Twenty grams of finely ground dried leaf and stem tissue was extracted with 100 mLof 25% aqueous cold methanol for 72 hours at room temperature. The extract was filtered and then evaporated in a vacuum. 0.1 grams of the residue was dissolved in 1 ml. of methanol. This solution was then spotted on whatman 3 MM chromatographic paper. The chromatogram was developed descendingly using TBA (3:1:1 reagent grade tertiary butanol: reagent grade glacial acetic acid: water) as the solvent. After drying, the chromatogram was developed descendingly in the second direction using HOAC (15 ml. reagent grade glacial acetic acid with 85 ml. distilled water) as the solvent. Mangiferin was tentatively identified on each chromatograph from its Rt value, its appearance under u v light, and its appearance over ammonia vapor.

Ureides and Related Compounds

Introduction In a consideration of the nitrogen metabolism of plants, ureides are proving to be of increasing importance, due mainly to their only recently discovered widespread occurence throughout the plant kingdom. Of these compounds which are common to most plant species, only quantitative 64 differences or differences with respect to the mode of their metabolism may serve as criteria. However, compounds such as canavanine or citrulline which are restricted in their distribution may be of supreme value as taxonomic markers since they are often genus or even species specific. From a quantitative point of view, they may be extremely important in plant nitrogen metabolism. Bell (1958) reports canavanine concentrations of 3-% of dry weight in the seeds of Canavalia ensiforrnis and Canavalia arborescens. In Acer sp., the sap exuding from severed roots may contain up to 99% of its nitrogen in the form of allantoin and allantoic acid (Mothes and Engelbrecht 1952). The importance of these compounds has probably been greatly underestimated, and future research will undoubtedly find them playing a vital role in nitrogen metabolism, particularly in the Leg- urninosae. Ureide plants tend to be concentrated in groups. In particular, the papilionoideae of the Leguminosae. It has been thought that ureides were essentially products of purine catabolism, which they may b~but recently found. high concentrations of these compounds have led many investigators to suggest that these amounts are too large to arise in this way. It has now been shown that in some plants they play a major role in the storage and transportation of nitrogen (Mathes and Engelbrecht 1952b).

Uric Acid Uric acid has been isolated from many plant sources 65 in recent years (Tracey 1955). Fosse et al (1932) isolated uric acid from Melilotus officinalis, and later from Trifolium sativum and Faba vulgaris .(1932). They reported uric acid in the range of 30-250 mg. per kg. of dry seed. Uric acid was detected in seed extracts by its ability to reduce alkaline phosphotungstate to a tungsten blue using sodium carbonate as an alkali. All four of the species recognized in this treatise were positive.

Arginine Arginine has been shown to be a regular component of proteins, and some seed proteins, such as those of Pisum sativum contain large amounts (Holmes 1953). As a free amino acid, it is widely distributed in the plant kingdom particularly in storage organs (Zacharius et al 1957, Reuter 1957). Arginine was detected in seed extracts using the Sakaguchi (1925) reaction as modified by Dubnoff and Bor- sook (1941). The only other compound known to react with the Sakaguchi reagent is glycocyamine but it has not been foun~ in plants. All four species were positive for arginine.

Allantoin and Allantoic Acid Allantoin was first isolated from plants by Schu1ze and Barbieri (1881) from the shoots of Platanus orienta1is. , Fosse (1926) found both allantoin and allantoic acid in the sap of Phaseolus vulgaris. Allantoin has also been 66 found in Phaseolus mungo (Fosse et al 1933) and in Glycine hispida (Brunel and Echevin 1938). Allantoic acid has been found in Melilotus officinalis {Fosse et al 1933) and in Glycine hispida (Brunel and Echevin 1938). Seed extracts were examined for the presence of allantoin and allantoic acid using the methods of Young and Conway (1942). All taxa were negative.

Thiourea Klein and Farkass (1930) isolated thiourea from the seeds of Laburnum, and Wehmer and Hadders (1933) reported it from various organs of Laburnum. Seed extracts were examined for thiourea using the blue color it produces with PCAF reagent. All species were negative.

Hydantoin and Creatinine Both hydantoin and creatinine give an orange color when treated with alkaline picrate. Seed, leaf, and root extracts were tested for creatinine and hydantoin using this reaction on a technicon autoanalyser. All species were negative.

Urea The presence of urea in plant tissues has been reported by many workers (Tracey 1955) • It now appears that urea probably rarely occurs in a free state in vascular plants, and previous reports of its presence can probably 67 be attributed to the common occurence and relative lability of the many urea precursors found in plants, particularly allantoin and allantoic acid. Allantoic acid is readily hydrolysed to urea and glyoxylic acid in an acid medium. Further, much of the urease which in previous years was sold as pure has now been shown to contain large quantities of canavanine and arginase, giving the preparation a system capable of forming urea. Many previous workers were undoubtedly unaware of the common occurence of allantoin in plants. Seed extracts were examined for urea using recry- stallized urease. The urease splits urea to co2 and ammonia, and the ammonia determined photometrically by the phenol- hypochlorite reaction. Sodium nitroferricyanide is used as a catalyst. EDTA was used to complex any interfering metals and to buffer the urease. All species were negative. 68

ELEMENTAL ANALYSIS

Plant analysis surveys have demonstrated representative nutrient concentration levels for many plants and are used in agriculture for evaluating mineral deficiency and toxicity in crop plants. From such studies it has become evident that nutrient concentration is under genetic control. Gorsline et al (1968) observed that the leaf concentration of boron is under genetic control and Clark (1970) found that a balance exists between micronutrients and that the nutrient concentrations of corn plants varied as one nutrient was varied from deficiency to near excess. The element accumulation character of a plant can have considerable importance from an evolutionary stand- point, as it could provide a characteristic for selection, since in general, high element accumulators are less sensitive to stress than low accumulators. The elemental concentrations presented herein were determined for two reasons. First, it appears that no elemental study of these species has been made and second, it has been demonstrated that the element accumulation characters of some plants are under genetic control and thus they may be of systematic value. As micronutrient concentration levels can vary considerably with such factors as stage of development, the 69 particular plant part being analysed, and environmental factors, the method of sampling becomes extremely important. Gorsline et al {1965) found that the Mn concentration of corn plants decreases with development, varies from leaf to leaf, and increases in concentration from lower to upper leaves. Also, it accumulates in the margins to a level of about twice that of the blade. Jones {1970) found that in corn leaves, the Mo concentration of the blade was higher than the margins, and the B concentration of the leaf petiole is much less than the blade. Jones and Mederski {1964) showed that B concentration in leaves increases gradually until maturity and then drops. When a plant begins to develop fruit, the micronutrient concentration of its vegetative portions changes considerably so that sampling after pollination should not be done. Each sample had leaflets from at least fifty plants. Mature leaf let blades were taken about one third the way down from the top of the plants which were at about the one tenth bloom stage. No leaves were used which were dust covered, treated, insect damaged nor mechanically damaged. Quantitative determinations were made for Na, P, K, Ca, Mg, Fe, B, Cu, Zn, Mo, s, and N using either atomic absorption spectrophotometry, flame photometry, or color- imetry. Several methods of extraction were utilized de- pending on the particular elements being analysed for. 70 For Na, P, Ca, K, Mg, Fe, Cu, Zn, and Mo, a wet digestion was carried out. Two grams of finely ground tissue was placed in a 500 ml. erlenmeyer flask and ten ml. of con-

centrated HN0 3 was added. The mixture was heated for 15 minutes at loo0 c. Then 10 ml. of a ternary acid mixture

(10:1:4 HN03, H2so4 , HCl04) was added and the heating continued until the solution turned colorless. The clear solution was then diluted with distilled water. Sodium and potassium determinations were made on a technicon flame photometer and recorder. Calcium, mag- nesium and iron were determined on a Perkin Elmer 290B atomic absorption spectrophotometer. Zinc and copper were determined using the methods of Page (1965). Sulphur was determined turbidimetrically by pre- cipitation of S04 with Barium chloride on a technicon autoanalyser using the methods of Basson and Bohmer (1972) • This procedure requires a special wet oxidation procedure for preparing the samples. 0.25 g. of sample was placed in a flask. 15 ml. of concentrated HN0 3 was added and heated for 20 minutes. Then 2 ml. of 70% perchloric acid was added and the heating continued until white fumes appeared. The solution was evaporated to a small volume and diluted with distilled water. It was then filtered and the residue washed with hot 0.1 M HCl. Molybdenum was determined using the colorimetric method of Ssekaalo (1971). For boron, two grams of ground plant material and 71

five ml. saturated Ca(OH) 2 solution was dried in a plati- num dish then dry ashed at soo 0 • After cooling, 10 ml. of 0.36 N H2so 4 was added, the ash was broken up, stirred and filtered. Two ml. of the filtrate and 15 ml. of quinalizarin reagent were added to a cuvette, mixed, allowed to stand for 24 hours at room temperature, then read at 620 mu. against a reagent blank. Nitrogen was determined by the Kjeldahl method. All results are presented in Table 6. 72

TABLE 6

PHYTOCHEMICAL SUMMARY OF HEDYSARUM

Compound or Species* Element A B c D

Urea

Thiourea

Allantoic Acid

Allantoin Arginine + + + + Hydantoin

Canavanine ** 0.84 1.08 0.75 1.12 Uric Acid + + + + Creatinine

Mangiferin + + + + Phytohemagglu- tin ins

S** .27 .32 .21 .37 N** 3.89 3.55 3.92 4.91

Na*** 460 366 304 287

P** .54 .37 .46 .42 Fe*** 304 209 260 197

K** 1.84 1.68 1.29 2.01

Mo*** 1.29 .43 .81 2.01 Zn*** 40 21 35 36

Cu*** 16 7 14 11 B*** 28 45 39 81

Mg** .30 .29 .33 .36 73 TABLE 6 {continued)

Compound or Species* Element A s· c D

Ca** 1.01 .81 .93 1.24 Total Seed Protein** 50.7 42.9 39.3 44.2

*Species A - H. aleinum var. alpinum B - H. sulphurescens C - H. occidentale D - H. boreale ssp. boreale var. rivulare **Results in percentage. ***Results in ppm. 74

TAXONOMY OF HEDYSARUM

Hedysarum L. Sp. Pl. Ed. 1, 745. 1753. Perennial, caulescent, terrestrial herbs arising from ligneous taproots; stems few to many, decumbent to erect, usually branched, solid, terete, usually pubescent; leaves odd-pinnate, petiolate, with petiolulate leaflets; stipules membranous, usually united, sometimes free; inflorescence racemose, axillate, pedunculate, bracteate; flowers few to many, yellow, pink or purplish; calyx campanulate, connate, five toothed, pubescent with two subulate basal bracteoles; corolla papilionaceous, glabrous, the keel exceeding the banner and auriculate-based wings; stamens 10, diadelphous; pistil one, 1-carpellary with parietal placentation; style slender, curved, erect; fruit a flat loment constricted into 2 to 6 I-seeded articles; loment articles glabrous or pubescent, areolate, stipitate, indehiscent, usually breaking transversely.

Key to the Species of Hedysarum A. Leaflet veins obscure or hidden; loment articles with wingless margins; calyx teeth sub- equal to markedly unequal in size; wing auricles free from each other, blunt, shorter than the claw ...... 1. H. boreale 75

A.A. Leaflet veins conspicuous; loment articles with winged margins; calyx teeth equal to nearly equal in size; wing auricles united, linear, nearly ·equal or equal to the claw •••••••••••• B. B. Loment articles 3.5-5.8 mm. broad, narrowly wing margined; flowers 11-18 mm. in length; leaflets mostly 4-10 mm. wide, oblong to oblong-lanceolate •••••••••••••••••• 2. H. alpinum B.B. Loment articles 5.6-10.2 mm. broad, conspicuously wing margined; flowers 15-22 mm. in length or if shorter then sulphur-yellow; leaflets mostly 6-17 mm. wide, ovate to ovate-

lanceolate ...... ·...... C. C. Flowers sulphur-yellow, 15-18 mm. long; loment articles mostly 5.5-7 mm. broad, glabrous ••••••

•.•..••.••....•.•..•..••... 3. H. sulphurescens c.c. Flowers reddish-purple, 16-24 mm. long; loment articles mostly 7-10 mm. broad, usually pube-

scent .•...••...... •....•....•... 4. H. occidentale

1. Hedysarum boreale Nuttall, Gen. N. Am. Pl. 2:110. 1818. Common name: Sweetvetch. Terrestrial, herbaceous perennials; roots stout, ligneous, 4-12 mm. in diameter; stems 2 to many, decumbent to erect, longitudinally grooved, not rooting, 2.5-7 dm. long, 1.5-5 mm. in diameter, usually branched above, solid, terete, sparsely pubescent to canescent; stipules tan to 76 brown, or white-translucent with brown streaks, with a subulate tip, usually united, upper sometimes free, 2-10 mm. long; leaves odd-pinnate, 3-12 cm. long; leaflets 5- 15 ovate, elliptic, linear, or obovate, canescent to glabrous above, canescent to pubescent below, entire,veins usually hidden, 10-35 mm. long, 3-19 mm. wide, apices round to obtuse; inflorescence racemose, 2-15 cm. long, interrupted to compact; bracts subulate, brown, pubescent, 2-5 mm. long; bractioles pubescent, linear, 1-3 mm. long; flowers 5-45, erect to spreading, pink to purple, 10-26 mm. long; calyx tube campanulate, 4-8 mm. long, bractiolate, pubescent, calyx teeth 0.5-1.5 mm. wide, 2-5 mm. long, subulate, greenish, erect; standard obovate to cuneate or·spatulate, apex emarginate, base cuneate, 8-16 mm. long, 4-10 mm. wide; wings 6-16 mm. long, 2-5 mm. wide; wing auricles blunt, round, free, 1-2 mm. long; stamens 10, diadelphous; style slender, curved; loments pendulous to spreading; articles 2-8, pubescent or setaceous, stipitate, not wing-margined, prominently transversely reticulated.

Key to the Subspecies 1. Flowers mostly 10-19 mm. in length, pink to purple; racemes 8-23 cm. long; articles mostly

2-5 ..••••.•...... •...•.•..••...• ssp. boreale 1. Flowers mostly 15-26 mm. in length, usually purple; racemes compact, usually 2-10 cm. long; articles mostly 3-8 ••••••••••• ssp. mackenziei 77 Hedysarum boreale Nutt. ssp. boreale Stems 2.5-7 dm. long, usually branched above; stipules usually pale brown, occasionally brown streaked; flowers 13-45, 10-19 mm. long, pink to magenta or purple; racemes elongated, 8-23 cm. long; leaflets usually pubescent on both surfaces, occasionally glabrous above; articles 2-5, seldom black pigmented.

Key to varieties of ssp. boreale 1. Articles of the loment covered with spines; leaves pubescent on both surfaces or glabrous above ••••••••••••••••••••••••• var. gremiale 1. Articles of the loment lacking spines; leaves pubescent to canescent on both surfaces or occasionally glabrous above. 2. Leaves glabrous to sparsely pubescent on upper surface ••••••••••••••• var. rivulare 2. Leaves markedly pubescent on both surfaces, whole plant grayish hairy or canescent •••• •••••••••••••••••••••••••••• var. boreale

Hedysarum boreale Nutt. ssp. boreale var. rivulare (Williams) Northstrom, comb. nov. Fig. 27 (Along the Snake River, Teton County, Wyoming, 31 July 1932, Williams 975). H. carnosulum Greene, Pittonia 3:212. 1897. H. pabulare A. Nels. Proc. Biol. Soc. Wash. 15: 185. 1902. H. utahense Rydb. Bull. Torrey Bot. Club 34:424. 1907. 78

Figure 27. Hedysarum bore·a1e ssp. boreale var. rivulare. 79

H. pabulare var. rivulare Williams, Ann. Missouri Bot. Gard. 21:344. 1934.

H. mackenziei var. pabulare (A. Nels.) Kearney and

Peebles, Wash. Acad. Sci. 29:485. 1939. H. boreale var. typicum Rollins, Rhodora 42:232.

1940.

H. boreale var. utahense (Rydb.) Rollins, Rhodora

42:232. 1940.

H. mackenziei var. fraseri B. Boi., Canad. Field. Nat. 65:20. 1951.

Distribution: In grasslands or on sagebrush slopes in the lowlands, to open fields or woodlands in the mountains. British Columbia to Alberta, south through Idaho,

Montana, and North Dakota to Arizona and New Mexico. Also in eastern and central Nevada. It merges in the north with the ssp. mackenziei (Fig. 28).

Representative specimens: ARIZONA. Apache County:

Salahkai Mesa, 7 August 1952, C. F. Deaver, s.n., (ARIZ),

Luka-Chukai Mountains, 29 June 1936, George J. Goodman

(MONT); Navajo County: 15 miles southeast of Snowflake,

26 June 1962, Richard H. Hevly s.n. (UT, ARIZ), Black Mesa,

21 June 1939, R. H. Peebles 14359 (ARIZ), North Base of

Black Mesa, 21 June 1939, Forest Shreve 8968 (ARIZ, UT,

COLO). BRITISH COLUMBIA. Yoho Park on Mt. Stephen, 2

August 1927, Titus Ulke 175 (MONTU), Marble Mountains, 20

June 1938, William Thompson 195 (PH). COLORADO. Boulder

County: north of Boulder, 22 July 1921, E. Bethel 4170, 80

ICM.& I,,_T •=... ___ : ·• l. _,,_,_ •=-:.:,:

... 110 ... .. Figure 28. Distribution of H. boreale. 81

(MONTU, OTC, COLO, PH), 6 miles north of Boulder, 20 July

1948, William A. Weber 4226 (COLO); Dolores County: T. 39

N., R. 14 W., 26 August 1937, Marion OWnbey (PH, UTC, MONT), near Mancos, 27 May 1901, F. K. Vreeland 849 (MONTU); Eagle

County: Holy Cross Forest Station, 18 July 1941, Ben F.

Rice 351 (USFS); Fremont County: south of Canon City, 12 June 1942, Joseph Andorfer Ewan 14225 (COLO), Canon City,

June 1895, G. E. Osterhout s.n. (COLO); Jefferson County:

Hog Back near Mt. Morison, 16 June 1940, J. H. Ehlers 8052

(ARIZ); Laplata County: 1 mile northeast of Durango, 12

June 1954, Margaret Douglass 54-219 (COLO); Las Animas

County: mouth of Berwind Canyon, 14 June 1941, Joseph A.

Ewan 12966 (COLO), Region of the Mesa De Maya, 29 June

1948, C. M. Rogers 5831 (COLO), 20 miles northwest of Branson, 14 July 1937, R. c. Rollins 1864 (UTC, USFS, MONT); Mes County: 1 mile east of state line on Highway 6,

26 June 1968, James A. Erdman 691 (COLO), Colorado National Monument, 24 June 1938, F. w. Pennell 22144 (PH), Coke Ovens, 20 May 1962, w. A. Weber 11306 (COLO); Moffat County: Dinasaur National Monument, 7 June 1965, M. Mcleod 377

(COLO); Montezuma County: Mesa Verde National Park, 7 June 1963, s. L. Welsh 2082 (BRY, SDU), Switchback Road below Point Lookout, 13 June 1964, S. L. Welsh 3029 (BRY); Mont- rose County: Naturita, no date, G. Breivster s.n.(COLO);

Rio Blanco County: south side of White River, 31 May 1935,

E. H. Graham 9044 (NDA), 15 miles east of Rangley, Highway

54, 13 June 1967, S. L. Welsh 6221 (BRY). IDAHO. Butte 82

County: National Reactor Test Station, 8 June 1967, N.

D. Atwood 914 (BRY); Custer County: 4 miles northeast of Dickey, 15 July 1941, Arthur Cronquist 3125 (UTC),

Lost River Mountains, 14 August 1944, C. L. Hitchcock 11109 (UTC); Franklin County: Cache Forest, Trail Can- yon, 25 June 1923, James o. Stewart C-71 (USFS); Lemhi County: Warm Springs Ranger Station, 4 June 1928, N.

C. Schulze 71 (USFS). MONTANA. Gallatin County: Gal- latin Forest, Bear Creek, 31 July 1935, James c. Whitham 1811 (USFS), Glacier National Park, east entrance, 14

June 1952, W. E. Booth (MONT); Golden Valley County: northwest of Levina, 26 June 1955, W. E. Booth 55222

(MONT); Jefferson County: 23 June 1965, E. C. Moran s.n. (BRY); Judith Basin County: 28 June 1964, E. c. Moran s.n. (BRY); Lewis and Clark County: 25 miles north of Augusta, 6 July 1948, C. L. Hitchcock 18074 (UTC, COLO),

Gibson Dam on the Sun River, 2 July 1952, H. N. Metcalf s.n. (MONT); Madison County: near Ennis, 20 July 1966, E. c. Moran s.n. (BRY}; Pondera County: Conrad, 15 June 1952, w. E. Booth s.n.(MONT). NEW.MEXICO. Colfax County: east of Colfax, 13 June 1940, Leslie N. Goodding Gl43-40 (OSC), Raton, July 1895, Mrs. o. st. John 147 (UTC); Santa Fe County: location unknown, 18 June 1897, A. A. Heller

3732 (UT) • NEVADA. Elko County: Humboldt National For- est, near Clover Creek, 19 July 1941, J. H. Robertson 300

(USFS); Nye County: Toiyabe Forest, Cloverdale Canyon,

25 June 1938, J. L. Levy JLS-11 {USFS). NORTH DAKOTA. 83 Billings County: Medora, 29 June 1938, o. A. Stevens 11 (MONT); Dunn County: R. 92, TWP. 149, 11 June 1936, V.

J. Heidenwish s.n. (NDA); Grant County: location unknown,

10 July 1964, O. A. Stevens s.n. (NDA); Mackenzie County: between Missouri River and Alexander, 25 June 1967, Claude

A. Barr 973 (NDA); Mclean County: location unknown, 27 June 1949, o. A. Stevens s.n. (NDA); Ward County: location unknown, 7 July 1935, o. A. Stevens 132 (NDA). OKLA- HOMA. Cimarron County: 2 miles north of Kenton, 26 May 1966, w. Hess 723 (OKL), 4 miles north of Kenton, 13 June 1918, c. M. Rogers 5991 (OKL); Ellis County: southwest of Harmon, 20 July 1957, Jack Engleman s.n. (OKL); Roger

Mills County: Redlands, 24 May 1937, Jack Engleman 1596

(OKL), location unknown, 25 May 1964, Dwight Isely 8900

(BRY). UTAH. Box Elder County: Wellsville Mountains,

23 May 1932, Melvin Burke 3012 (UTC); Cache County: bank above canyon road, Logan, 4 June 1937, Arthur Cronquist

533-37 (UTC), south of Dry Canyon, 16 May 1962, N. H.

Holmgren 108 (UTC); Carbon County: Soldier Canyon, 2

June 1962, E. M. Christensen s.n. (BRY), Wellington, 5 June 1927, w. Cottam 2046 (BRY), West Tavaputs Plateau, 9 June 1940, Bassett Maguire 18515 (UTC, BRY); Davis

County: Mueller Park, 28 April 1940, K. Brizzee s.n.

(UT), Bountiful City Reservoir, 28 June 1964, L. Hogan

42 (BRY); Duchesne County: Red Creek, 20 July 1965, Jack Brotherson 489 (BRY), Tabby Mountain, 2 July 1938, B. F.

Harrison 8790 (BRY); Emery County: east of Ferron, 7 June 84 1932, W. P. Cottam 5275 (UT), Huntington Canyon, 21 July 1935, A. O. Garrett 7020 (UT); Garfield County: near Sun- rise Point, 4 July 1957, H. Buchanan 167 (UT), Bryce Can- yon, 11 July 1927, w. Cottam 2758 (BRY); Grand County: 20 miles north of Dead Horse Point, 27 May 1950, w. P. Cottam 12143 (UT), 15 miles south of Cisco, 6 June 1961, Arthur Cronquist 9222 (UTC), Arches National Monument, 2 May 1963, S. L. Welsh 1903 (BRY); Juab County: east of Diamond, 10 June 1900, I. E. Diehl s.n. (BRY), Junction west of Eureka, 20 May 1962, s. L. Welsh 1728 (BRY); Kane County: 43 miles south of Escalente, 24 June 1965, Albert W. Collotzi 503 (UTC}, Vicinity of Bryce, 24 May 1940, G. H. Vansell s.n. (UTC); Millard County: Pavant Mountain, 19 June 1966, R.H. Foster 408 (BRY); Rich County: south Sink Garden City, 10 July 1927, G. w. Craddock GWC-20, (USFS); Salt Lake County: Bell Canyon, 6 July 1939, A. Bleak 56 (UT), Red Butte Canyon, 13 July 1965, George Swan-

~ 174 (UT); San Juan County: 6 miles up Colorado River from Moab Bridge, 6 May 1938, Bryan and Deming 6-10 (BRY), Lasal Mountains, 13 June 1927, w. Cottam 2212 (BRY}, Nat- ural Bridges National Monument, 28 May 1968, s. L. Welsh 7135 (BRY); Sanpete County: Manti Canyon, 12 June 1964, A. H. Barnum 1294 (BRY), Manti Forest, 6 July 1934, A. Perry Plummer 304 (USFS); Sevier County: Fishlake For- est, Convulsion Canyon, 11 July 1930, Albert Albertson 41 {USFS), Moroni Peak, 28 June 1947, w. P. Cottam 9667 (UT); Tooele County: Sheeprock Mountains, 28 July 1963, 85 E. M. Christensen s.n. (BRY), Stansbury Range, 25 June 1943, Bassett Maguire 21804 (UTC); Uintah County: 5 miles northwest of Whiterock, 21 July 1965, Jack Brotherson 565 (BRY), 1 mile west of Rainbow, 4 June 1965, N. H. Holmgren 1819 (BRY, UTC); Utah County: Rock Canyon, 18 June 1964, M. L. Fairbourn 3 (BRY), 6 miles east of Soldier Sununit, 2 June 1961, S. L. Welsh 1564 (COLO); Wasatch County: East Rim of Strawberry Valley, 16 July 1965, Jack Brother-

~ 663 (BRY), Little Deer Creek, 15 July 1964, Jack Brotherson 17 (BRY); Wayne County: Fruita, 14 June 1938, D. E. Becks. n. (BRY); Weber County: mouth of Ogden Canyon, 18 May 1965, R. J. Bragonje 27 (BRY), Saddle of Ben Lomond, 20 June 1940, w. Call 33 (UT). WYOMING. Carbon County: 8 miles west of Medicine Bow, 15 June 1948, C. L. Porter 4584 (COLO); Fremont County: 10 miles west of Dubois, 4 June 1948, C. L. Porter 5780 (COLO); Natrona County: Pathfinder Reservoir, 4 June 1948, c. L. Porter 4506 (COLO); Park County: 20 miles north of Cody, 11 July 1950, C. L. Porter 5442 (COLO, ARIZ), Valley of the Clarks Fork of the Yellowstone River, 20 July 1948, Jean G. Witt 1284 (COLO); Sublette County: Pine Grove Ridge, 23 July 1931, s. E. Cazier C-124 (USFS); Teton County: 1 mile below mouth of Coburn Creek, 11 July 1928, Charles H. McDonald 837 (USFS), 10 miles south of Jackson, 2 July 1938, R. v. Rethke 3894 (UTC); Uintah County: 14 miles east of Evanston, 22 July 1939, Reed c. Rollins 2875 (USFS) • 86

The type specimen of Hedysarum boreale could not be located by previous workers, nor by the present author in an earlier study (Northstrom and Welsh, 1970). However, this specimen has now been found in the British Museum along with the type specimen of Hedysarum mackenziei (H. boreale ssp. mackenziei). These were obtained on loan and photographed (Figs. 52 and 53). It was immediately recognized that the specimen which Nuttall described as Hedysarum boreale is indeed homologous with specimens now passing as H. boreale, but corresponds to material now described as variety cinerascens, instead of variety boreale. This of course, ne- cessitates some recombining of varietal names. ~pecimens previously assignable to variety cinerascens must now be placed under variety boreale while specimens formerly described as variety boreale require a new combination. Variety rivulare Williams has priorty within rank so the new combination H. boreale Nutt. ssp. boreale var. rivulare (Williams) Northstrom must be made. This variety is usually assumed to inhabit only the plains regions of Canada, and the lower mountain slopes of the Rockies. But it is also commonly found at elevations of 8000 feet or more in many areas throughout its range. Flowering occurs from early May to early August.

Hedysarum boreale Nutt. ssp. boreale var. gremiale (Rollins) Northstrom and Welsh, Fig. 29, Great Basin Natural- ist 30:125. 1970. (14 miles west of Vernal, Uintah County, Utah, 16 June 1937, Reed C. Rollins 1733 (G)) 87

Figure 29. Hedysarum boreale ssp. boreale. a. loment of var. gremiale, b. flower of var. boreale, c. leaflets of var. boreale. 88 Distribution: Dry slopes, ravines, and pinyon- juniper communities from 5000-6000 ft. elevation in the

Uinta Basin of Utah (Fig. 28).

Representative specimens: UTAH. Uintah County:

Moenkopi Shale, Split Mountain Gorge, Dinasaur National

Monument, 15 July 1956, Jack Brotherson 787 (BRY), west of Vernal, 16 June 1937, R. C. Rollins 1733 (G, type,

PH, OKL isotype); Duchesne County: North of Myton, 1

July 1938, Pennell 22497 (PH).

Hedysarum boreale Nutt. ssp. boreale var. boreale, Fig. 30 (Fort Mandan, on the Banks of the Missouri, Nuttall sn (BM))

Hedysarum canescens Nutt. ex. T. and G. Fl. N. Am.

1:357. 1838.

Hedysarum cinerascens Rydb. Mem. New York Bot.

Gard. 1:257. 1900.

Hedysarum macguenziei (sic) var. canescens (Nutt.)

Fedtschenko, Acta Hort. Petrop. 19:364. 1902.

Hedysarum boreale var. cinerascens (Rydb.) Rollins,

Rhodora 42:234. 1940.

Hedysarum boreale var. obovatum Rollins, Rhodora

42:235. 1940. Hedysarum boreale var. cinerascens f. album B. Boi.,

Nat. Canad. 87:34. 1960.

Distribution: Alberta and Saskatchewan, south through Idaho and Montana to Nevada, southern Wyoming and eastern Utah. Dry banks and ravines from 5000 to 8000 feet elevation (Fig. 28). 89

Figure 30. Hedysarum boreale ssp. boreale var. boreale. 90 Representative specimens: ALBERTA. Milk Mountains,

26 June 1952, B. Boivin 9502 (OKL), East of Waterton River,

23 July 1953, A. Breitung 16486 (COLO), Rosedale Trail, 22

June 1915, Marion Moodie 1020 (OSC). IDAHO. Custer Coun- ty: West of the Salmon River, 15 June 1944, C. L. Hitch- cock 8989 (MONT, UTC), West of Clayton, 5 August 1944, c. L. Hitchcock 10794 (UTC); Jefferson County: Locality unknown,

15 June 1967, E. C. Moran s.n. (BRY). MANITOBA. Near Churchill, 6 July 1963, Alice Brues s.n. (OKL). MONTANA.

Beaverhead County: Lima, 9 June 1958, D. Ryerson 599 (MONT);

Bighorn County: 4 miles east of Busby, 12 May 1952, D. K.

Scharff s.n. (MONT), Flint, 3 August 1918, R. B. Streets

441 (MONT); Broadwater County: 7 miles east of Townsend,

1 July 1945, C. L. Hitchcock 11824 (PH, UTC); Cascade Coun- ty: Volta Dam, 22 June 1935, v. L. Marsh 739 (MONT), Belt Canyon, 11 July 1887, R. s. Williams 91 (MONT); Dawson County: Bloomfield, 24 July 1958, Bessie Ming s.n. (MONT); Fergus County; Big Snowy Range, 10 July 1958, s. Bamberg s.n. (COLO), 1 mile north of Heath, C. L. Hitchcock 11929

(UTC, PH, COLO); Gallatin County: Bear Creek Road, 8 June

1952, J. £·Wright s.n. {MONT); Golden Valley County: Rye- gate, 17 June 1957, w. E. Booth s.n. (MONT); Glacier Coun- ty: Midvale, 14 July 1903, L. M. Umbach 378 (MONT)' Gla- cier National Park, 3 miles west of eastern entrance, 14 June 1952, w. E. Booth s.n., {ARIZ); Lewis and Clark Coun- ty; Ear Mountain Ranger Station, 1 July 1921, c. A. Butler D3-12 (USFS), Sun River Acquisition, 12 August 1952, R. L. 91

Hodder s.n. (MONT); Livingston Park County: 25 miles north of Field, 17 June 1936, D. B. Single s.n. (MONTU); Madison

County: Beaverhead, 30 June 1937, L. ~- Short S-521 (USFS), Hammond Creek, 23 July 1947, c. L. Hitchcock 16721 (COLO, UTC); Meagher County: West of Harlowton, 26 June 1955, w. E. Booth 55360 (MONT), 6 miles northwest of White Sulphur Springs, 11 July 1945, c. L. Hitchcock 12182 (PH, UTC, OSC, MONT); Musselshell County: 10 miles northwest of Roundup, 18 June 1957, w. E. Booth s.n. (MONT); Park County: Li- vingston, 6 July 1901, E. Schuber s.n. (MONT); Phillips

County: Malta, 9 June 1900, J. W. Blankinship s.n. (MONT);

Stillwater County: Absarokee, 21 June 1914, P. H. Hawkins s.n. (MONT), 2 miles west of Columbus, 26 May 1952, J.C.

Wright s.n. (MONT); Teton County: 1 mile west of Choteau,

6 July 1948, c. ~·Hitchcock 18109 (COLO); Wheatland Coun- ty: Shawmut, 26 June 1955, w. !·Booth 55314 (MONTU}, Haymaker Game Range, 24 June 1959, John Kirch 107 (MONT); Yellowstone County: Pompey's Pillar, 23 May 1952, w. E. Booth 52205 (MONT), Yellowstone National Park, Lamar Val- ley, 7 August 1958, G. Denton s.n. (MONT), Swan Lake, 4

July 1928, P.H. Hawkins 384 (USFS). NORTH DAKOTA. Bill- ings County: north side of Butte, 3 August 1923, o. A. Stevens s.n. (NDA); Medora, 29 June 1938, Q· A. Stevens 38-011 (NDA); Dunn County: 10 miles north of Butte, 19 August 1954, o. A. Stevens s.n. (NDA); Golden Valley County: Sentinel Butte, 11 July 1962, O. A. Stevens s.n.

(NDA); Mercer County: North Missouri River Valley, 29 92 June 1941, O. A. Stevens 906 (NDA); Mountrail County:

Spanish, 28 July 1923, o. A. Stevens s.n. (NDA); Kerr- ville County: location unknown, 12 June 1939, Mrs.

Malt Meissuer s.n. (NDA); Kidder County: 8 miles north

of Lake Williams, 27 July 1956, D. R. Moir s.n. (NDA); Slope County: Saddle Butte, 14 June 1963, o. A. Stevens s.n. (NDA); Ward County: 2 miles west of Minot, 25 June

1968, J. E. Bare 1053 (NOA); Wells County: Hurdsfield, 27 June 1943, o. A. Stevens 696 (NDA); Williams County: Buford, 21 June 1945, o. A. Stevens s.n. (NDA). QUEBEC. Riviere Vaureal, 28 July 1925, P. Louis-Marie 20871 (PH).

SASKATCHEWAN. Battle Creek Ranger Station, Cypress Hills,

25 July 1947, A. J. Breitung 5054 (UTC). UTAH. Summit County: Elk Horn Ranger Station, 6 June 1921, J. o. Stewart 20 (USPS), Shake John Springs, 28 May 1926, George w. Walkup GW-14 (USPS); Uintah County: 2 miles north of Vernal, 26 June 1951, C. L. Porter and R. c. Rollins 5673 (PH). WYOMING. Big Horn County: Big Horn Forest, Day- ton-Kane Road, 17 June 1932, B. Dickson 250 (USPS), Red

Bank, 20 July 1901, Leslie N. Goodding 332 (MONT, UTC,

ARIZ); Carbon County: Medicine Bow Mountain, 2 August

1929, F. B. Warn 44 (UTC); Fremont County: Washakie

Forest, Horse Creek, 21 June 1914, R. J. Guthrie 7 (USPS),

10 miles west of Dubois, 27 July 1951, C. L. Porter and R. c. Rollins 5780 (PH); Natrona County: near Pathfinder Reservoir, 4 June 1948, C. L. Porter 4506 (MONT); Park

County: 20 miles north of Cody, 11 July 1950, C. L. Porter 93 5442 (PH, OKL); Sublette County: Bridger Forest, 16 June 1926, c. H. McDonald 625 (USFS), 20 miles west of Big Piney, 9 July 1922, E. B. Payson 2617 (PH, OSC); Teton County: 20 miles south of Jackson, 2 August 1935, Bassett Maguire 12845 (UTC), Teton Forest, lower Blackrock, l July 1916, Rudolph Rosencrom 39 (USFS).

Hedysarum boreale Nutt. ssp. mackenziei (Richards.) Welsh, Fig. 31, Great Basin Naturalist 28:152. 1968. (Barren grounds, Point Lake to the Arctic Sea, Richardson s.n. (BM)) Hedysarum mackenziei Richards. in Frankl. Journal Bot. App. 745. 1823. Hedysarum americanum var. mackenziei Britt. Mem. Torrey Bot. Club. 5:202. 1894. Hedysarum mackenziei var. mackenziei f. niveum a. Boi. Canad. Field Nat. 65:20. 1951. Hedysarum boreale var. mackenziei (Richards.) c. L. Hitchcock. University Wash. Publ. Biol. 17(e): 275. 1961. Hedysarum dasycarpum Turcz. Bull. Soc. Nat. Mose. 1840.

Stems 1.5-5 dm. long, usually unbranched above; stipules pale tan, or white-translucent with brown streaks1 flowers 5-20, 15-26 mm. long, usually purple occasionally pink or pale pink; racemes compact, 2-10 cm. long; leaflets usually green and glabrous above, grayish-green and pubescent below1 articles 3-8, margins of articles and aerolae often black pigmented. 94

Figure 31. Hedysarum boreale ssp. mackenzii 95

Distribution: Alaska, Yukon, Banks-Victoria Is- lands to Newfoundland, south to northeastern Oregon, and east through southern British Columbia to Quebec. Also in eastern Siberia (Fig.28).

Representative specimens: ALASKA. Milepost 265,

Alaska Highway, 16 June 1953, D. Bolinger s.n. (OSC),

Gakona, 28 June 1966, S. L. Welsh (BRY), King River, 19

June 1965, S. L. Welsh 4244 (BRY), Mount McKinley National Park, 21 July 1965, s. L. Welsh 4834 {BRY), 5 miles north of Nome, 14 July 1966, s. L. Welsh (BRY), Fairbanks Quad- rangle, Tanana River opposite Sam Charley Island, 8 July

1966, L.A. Viereck 8029 (USFS), Bering Strait District,

28 July 1960, L.A. Viereck 4365 (COLO). ALBERTA. Vi- cinity of Banff, 8 June 1906, s. Brown 75 (PH), Ptarmigan Valley, 4 July 1906, s. Brown 401 (PH), Jasper National Park, 22 July 1962, R. J. Davis 5082 (BRY), south of Peyto

Lake, 12 July 1941, W. A. Weber 2429 (COLO). BRITISH

COLUMBIA. Vicinity of Field, 16 July 1906, S. Brown 538

(PH) , North end of Azouzetta Lake, 4 August 1954, .J. A. Calder 14022 (OSC), Summit Lake, 22 June 1966, s. L. Welsh 5447 (BRY), mouth of Racing River Bridge, 26 June

1968, S. L. Welsh 7340 (BRY). MANITOBA. Churchill, 10

July 1934, T. M. Cope s.n. (PH), Fort Churchill, 30 June

1948, J.M. Gillett 1913 (OKL), Limestone River Banks near Gillam, 12 July 1950, w. B. Schofield 1099 (COLO). NORTH- WEST TERRITORIES. opposite Fort Simpson, 30 June 1955, W.

J. Cody 8466 (ARIZ), Moraine Point, Great Slave Lake, 30 96

June 1951, W. H. Lewis 479 (COLO, OKL). OREGON. Union County: half mile north of East Eagle Creek Falls, 30 June 1957, s. c. Head 1045 (OSC); Wallowa County: Wallowa Mountains, 9 August 1911, W. c. Cusick 3694 (OSC), Hurri- cane Creek, 29 July 1951, Conrade Head 127 (BRY), Border of Ice Lake, 8 August 1963, Georgia Mason 6461 (OSC). QUEBEC. Ungava Bay, 27 July 1948, E. K. Bonde 283 (COLO). SASKATCHEWAN. near Bear Creek, 14 June 1908 s. Brown 909 (PH). YUKON TERRITORY. Haines Junction, 1 June 1967, K. Crook 318 (OKL), Mount Mcintyre, 21 June 1949, J.M. Gillett 3453 (COLO), Mile 1019, Alaska Highway, 9 June 1965, s. L. Welsh 4056 (BRY), Kluane Lake, Milepost 1054, 4 July 1968, s. L. Welsh 7839 (BRY), Mile 117.7 Dawson Highway, 25 June 1966, s. L. Welsh 5560 (BRY). Jurtsev (in Hulten 1973, p. 498) claims that H. boreale ssp. mackenzie is specifically distinct from its

Siberian counterpart (~. dasycarpum Turcz.). According to Jurtsev, H. boreale ssp. mackenziei differs from H. dasycarpum in having a keel which exceeds the banner in length and in the darkness of the flowers. A polygonal graph comparing specimens of H. dasycarpum from the Soviet Union and H. boreale ssp. mackenziei from northern Alaska was prepared from material on loan from the Komarov Botanical Institute at Leningrad and from specimens deposited in the Brigham Young University herbarium (Fig. 32). The graph illustrates that Siberian and American material cannot be separated on the basis of rela- 97 H. mackenziei H. dasycarpum

Keel Width

Bract Length

Wing Banner Width Length

Wing Length Figure 32. Polygonal graph comparing 8 characteristics of H. dasycarpum (U.S.S.R.) and H. boreale ssp. mackenziei. 98 tive lengths or widths of floral parts. It appears that the differences between the Old World and New World plants are only trivial and thus the two populations should be regarded as conspecific. Essentially restricted to rocky slopes, roadsides, gravel bars, .sandy places and other areas which are in early stages of succession. Occasionally occurs in tundra and open woodlands. Flowering from early June to August. {Fig. 31).

2. Hedysarum alpinum L., Sp. Pl. 750. 1753. {"Habitat in Sibiria") Terrestrial herbaceous perennials; roots stout, ligneous, elongate; stems few to numerous, erect, longitudinally grooved, not rooting, 1-7 dm. long, 1-3 mm. in diameter, usually branched above, solid, terete, glabrous or appressed pubescent; stipules brown, connote, chartaceous, 5-15 mm. long, lanceolate, obtuse to acute; leaves odd-pinnate, 5-16 cm. long; leaflets 7-27, oblong to lanceolate, sparsely pubescent below, glabrous above, conspicuously veined, 10-30 mm. long, 4-10 mm. wide, obtuse to acute; inflorescence racemose, 2-30 cm. long, elongate to compact; bracts linear, 1-4 mm. long; brae- tioles pubescent, linear, 1-3 mm. long; flowers 5-50, deflexed to erect, lilac-purple or rarely pale purple or white, 10-18 (22) mm. long; calyx campanulate, tube 2-5 mm. long, bractiolate, pubescent, teeth 1-2.5 mm. long, unequal, triangular, greenish, erect; standard obovate to 99 spathulate, emarginate, 11-14 mm. long; 5-8 mm. wide; wings 11-15 mm. long, 2-3 mm. wide; wing auricles linear, united,

2~3 mm. long; stamens 10, diadelphous; style slender, curved; loments pendulous, stipitate; articles 2-5, glabrous or pubescent, wing margined, 3.5-6 mm. wide, 5.2-9 mm. long, reticulations polygonal. Rollins (1940) indicated that North American specimens of ~· alpinum differ from those of Asia and Europe in "certain minor ways" and thus "should be considered as separate varieties of a wide ranging species." The nature of these minor differences, however, was not indicated.

An examination of representative Old World specimens from the British Museum, the u. s. National Herbarium, and the Komarov Botanical Institute at Leningrad revealed no differences which could justify any specific nor infraspecific segregation of Old and New World specimens. A polygonal graph was prepared with the results obtained from the above mentioned specimens (Fig. 33). No differences could be detected to support any infraspecific segregation. Furthermore, a photograph obtained from the

British Museum of the type specimen of ~- alpinum L. appears homologous with North American material.

In North America, as in Europe and Asia, ~· alpinum L. exhibits considerable variation which is generally most apparent as contrasting ends of a variation continuum. Throughout southern Canada, it is found from the plains and lowlands of the prairie provinces to the mountain slopes of 100

H. alpinum, U.S.

------H. alpinum, U.S.S.R.

Keel Width

6 Bract Banner Length l------~.....11t--<4:~..-il--i-+---~----~------#!---ffiff+... --.L------f Width

Wing Width

Wing Length Figure 33. Polygonal graph comparing 8 characteristics of Old and New World H. alpinum. 101 Alberta and British Columbia, often at elevations exceeding 10,000 feet. It is found in such diverse habitats as exposed mountain slopes, calcareous cliffs, gravel bars, open fields, dry rocky foothills and alpine tundra. The variants produced in such diverse habitats often appear as distinct entities, qut on close examination are found to be interconnected by complete series of intermediates which very often cover vast geographical areas. This pattern of variation is evident in all four of the North American species of Hedysarum, but is most evident in H. alpinum. Figures 34-37 illustrate this pattern. Figure 34 illustrates a series of specimens of H. boreale ssp. mackenziei. Figure 34a illustrates a specimen collected from a favorable site at low elevation in British Colum- bia, while figure 34d was collected in tundra on the northern arctic coast of Alaska. Figures 34b and 34c are plants intermediate between these two extremes. Figure 35 illustrates the same pattern in H. sulphurescens, figure 36 in H. occidentale and figure 37 shows the same pattern in H. alpinum. In the latter species, however, the variation is the most pronounced. This can be accounted for by the much more extensive range of distribution exhibited by ~· alpinum than by any of the other American species. It is true that H. boreale occupies as vast a range as does g. alpinum, but the pattern of variation in H. boreale is exemplified by both subspecies independently and not by the species as a unit. In other words, this 102

A. B.

c. D.

Figure 34. Ecological variation in H. boreale. 103

~--~~~ !!:!~~ ~!.t~::.:.•.!!.~ ih•b. .... hl'a•u M~~ "~' ;, 1'1<1tl<-• •••! --- ww..-u. ' f" l 0 l \ .ll~(;.,n ; _A_,_ ..,_ •• ,_~_, 0 ~~~!::E:~J4~~i:ft 0.J<.(7'.h>.•• li.b. rl~~o l••• u,.;i i h. v ~:. J-1,. ... °'' ...... _1· ,... _. ~ .. u ~i-.- l~J A. B.

c. D. Figure 35. Ecological variation in H. sulphurescens. 104

A. B.

I I: i.jI j i: j

->1---,-...... 4- --'''1. _

.~>·•J;.J.~; "'~ · .,~, ·•r~ ·~·.'· f•'·•"'~ ,.,,,., ... _ ,_.u...,..;,;•,.,i._,.,, -:~!\""

c. D. Figure 36. Ecological variation in H. occidentale. 105 ...... -

A. B.

c. D. Figure 37. Ecological variation in H. alpinum. 106 continuum type of variation does not represent the basis for subspecific segregation in H. boreale. The nature of this variation is standard in all four species, but again is most pronounced in H. alpinum. Plants collected from high altitudes or extreme northern latitudes have short, compact and sometimes decumbent inflorescences bearing few flowers. These flowers are large, occasionally exceeding 19 mm. in length and usually much darker in color than their southern counterparts. The plants are much shorter in height, usually less than 25 or 30 cm. and have somewhat smaller leaves with fewer leaflets. Specimens exhibiting an extreme degree of this variation can be quite striking and readily explain the numerous synonyms bestowed upon them in recent years. Alice Eastwood described H. truncatum Eastwood from material collected from Nome Alaska in 1902. An examination of isotypes in the herbaria of the California Academy of Sciences, Stanford University and the Gray Herbarium which were identified by Eastwood as H. truncatum revealed typical boreal extremes of ~· alpinum L. Hulten (1968) considered these extreme high altitude populations to be specimens of H. hedysaroides (L.} Schinz. & Thell. which were hybridizing with the lower elevation dwelling populations of H. alpinum. The hybrids thus formed from such hypothetical crosses then introgress with both parents forming the variation continuum which is now exhibited. H. hedysaroides is an Old World species 107 which has been surrounded by considerable confusion for many years. It was first described by Linnaeus (1753) as Astragalus hedysaroides L. Later, Linnaeus recognized his error and changed its name to Hedysarum obscurum L. Then, Schinz and Thellung recombined the name, utilizing the specific epithet having priority, to Hedysarum hedysaroides

(L.) Schinz & Thell (1913). This entity exhibits nearly the same type of variation as does H. alpinum with a few exceptions. H. hedysaroides inhabits mostly higher altitudes, although it does have considerable elevational range. Specimens from lower elevations are quite large with elongated inflorescences while high mountain specimens have short, compact and few flowered inflorescences, but the differences in flower size found in H. alpinum is not exhibited to such a degree by H. hedysaroides and the flowers of the latter are much larger than even the most extreme alpine form of H. alpinum. A polygonal graph comparing some floral characteristics of Old World H. hedysaroides and New World alpine H. alpinum is presented in figure 38. The much larger flowers of the Old World Taxon are evident. Also, the loments of H. hedysaroides are often pubescent while the loments of a. alpinum are only rarely pubescent, except in var. ehiloscia. Rollins (1940) discussed the variation exhibited by H. alpinum and described the dwarfed arctic phase as var. grandiflorum, although he acknowledged the gradual transition from one phase to another. 108 H. hedysaroides ------H. alpinum var. grandiflorum

Keel Width

Bract Banner Length l--~~-l-~.1..'-"__..~~~;...~~~~~4t~~~~~-Jt-f.'-~-+Tf-+-~-t-~'--....,.~-t Width

Wing Width

Wing Length

Figure 38. Polygonal graph comparing 8 characteristics of H. hedysaroides (Europe) and H. alpinum var. srandif lorum. 109 All of the variation which has been and is now used to distinguish between the alpine and lowland phases of·Hedysarum alpinum is quantitative and continous. No qualitative nor discontinuous variation was observed during the course of this study, except for the prescence of considerable pubescence on the loments of plants inhabit- ing portions of Wyoming, Saskatchewan, South Dakota, and Alberta. Although here, too, their is a gradual transition of pubescent fruited plants into glabrous ones, with intermediates having pubescent margins or some other form of reduced vesture. This pubescent taxon was described by Rollins (1940) as variety philoscia. This appears to be a valid entity and is recognized herein. It occupies a distinct and unified range although it intergrades with var. alpinum. This is in harmony with the current use of the varietal rank. As for the subarctic phase, the situation is somewhat similar. An examination of all available specimens representing the extreme northern phase of ~· alpinum, referred to by Rollins (1940) as ~· alpinum L. var. grandiflorum Rollins, and by Hulten (1968) as "pure Hedysarum hedysaroides (L.) Schinz & Thell. (1913), "and specimens of H. alpinum from the most southerly portions of its range (Manitoba, Montana, Saskatchewan, Vermont, Maine, Alberta, British Columbia and Wyoming) for quantitative variation in seven morphological characteristics revealed the data tabulated below (Table 7) and illustrated in a 110 polygonal graph (Fig. 39). 111 TABLE 7 QUANTITATIVE VARIATION IN HEDYSARUM ALPINUM

NORTHERN PHASE SOUTHERN PHASE

CHARACTER RANGE MEAN RANGE MEAN

Inflorescence length* 2.2-6.1 3.4 8.6-18.5 13.9

Flower length** 16.0-19.0 17.7 11.0-15.0 13.4

Leaf let number 9.0-15.0 13.5 13.0-29.0 21.5 Leaf length* 6.8-8.9 8.4 7.5-16.2 11.8

Stature* 16.1-27.5 21.5 36.0-73.0 52.5 Leaflet length** 14.0-20.1 17.1 11.0-35.0 24.4 Flower number 10.0-18.0 13.4 29.0-56.0 41.0

*Measurements in centimeters. **Measurements in millimeters. 112 Northern Phase

------Southern Phase

, .. --- , I , I I I I , I E L..&~1&~SL...... --llL11~9!,._ __...______.,!.!~~~~--~14------=::::::~:=iiill"t---t~ ' ' ' ' ' ' ' ' ' '

A Leaf Length B Flower Length C Flower Number D Leaf let Number E Inflorescence Length F Plant Height

Figure 39. Polygonal graph depicting averages for each of 8 characteristics in the northern and southern phases of H. alpinum. 113 Leaflet length overlaps so greatly as to nullify the character. The inflorescence and flower length provad to be good characters. The northern phase only rarely (1 specimen) exhibited an inflorescence length of more than 4 cm., and no specimens had flowers shorter than 16 nun. The number of flowers and the height and growth habit also show no overlap between the groups. The northern phase usually has 11-13 leaflets, only rarely 15, while the southern form usually has 21 or more. Some overlap was exhibited in leaf length with the most variation occuring in the southern plants. However, they were only rarely less than 9 cm. From these figures, scores were assigned to each of six characters. An O for a grandiflorum characteristic, a 2 for an alpinum characteristic, and a 1 if the characteristic was intermediate. An additional 180 plants were then measured and tabulated for each character. The individual scores for each specimen are presented in the form of pictorialized scatter diagram (Fig. 40 ) and distribution map (Fig. 41), while the total point scores for each specimen are presented in the form of a histogram (Fig. 42). The categories used for assigning scores are illustrated in Table 8. 114 22 • 0 a a - Flower Number b - Leaf Length .. 1 21 d+b c - Plant Height c d - Leaf let Number --- 2

19 ... ,

18 .....

2 4 6 8 10 12 14 16 18 20 22 Inflorescence Length in cm. Figure 40. Scatter diagram depicting six morphological characteristics in H. alpinum. 115

30 ·---;I , ------I

a c d 1 2

-~ ..... ----- ''° HO •oo •o Figure 41. Distribution of variability within H. alpinum. 116

128 25 127

126

OJ J.f 0 0 Ul .µ20 .....s:: 154 MS 122 0 a. 1S3 1•• 121 .c: 152 1'3 120 0 RS OJ ISi 1'2 138 119

O'I IS9 ISO 1•1 137 118 -~ 15 118 1'9 1'0 J3ft 117 >RS .c: U1 MS Ma 139 135 11• .....rJl a:s ::s .....'tS ·.-tlO> 'tS 39 1a2 •7 79 92 131 112 .....s:: •• 'M 0 J.f OJ

§ 1a 22 29 IS ,. S2 83 7S 88 IOI 108 z 5 • 10 IS 21 28 14 4• SI •2 74 87 IOO 107 4 9 14 20 27 31 41 so II 13 8• 99 10•

3 8 13 19 2• 12 •2 49 ao 72 85 98 IOI 2 7 12 18 21 31 " 48 59 71 84 97 10• I • II 17 24 10 40 ,, •• 70 83 96 103 0 1 2 3 4 5 6 7 8 9 10 11 12 Total Points Figure 42. Histogram based on the total points for all characteristics. 117 TABLE 8 CHARACTER SCORES FOR HEDYSARUM ALPINUM

Character 0 1 2

Inflorescence less than 4.1- 8.5 greater than 8.6 Length* 4 Flower Length** 16-19 15.1-15.9 10-15 Flower Number less than 19-18 29-56 18 Leaf Length* less than 9.0-10.5 greater than 10.6 8.9 Leaflet Number 7-13 15-17 19-29 Height* decumbent 26-42 greater than 42.0 less than 25

*Measurements in centimeters. **Measurements in millimeters. 118 The distribution map is not intended to represent the complete distribution of the species, as many of the plants did not exhibit the proper stage of development or were in too poor a condition, but merely represents an attempt at outlining the distribution of variability within the species. A complete distribution map is illustrated in figure 44. It is noteworthy to mention that all of these characters are of the nature that they would normally be expected to vary together, particularly in a species with as broad a range of distribution as H. alpinum. The polygonal graph (Fig. 39) illustrates the extreme degree of quantitative variation separating the northern and southern phases of H. alpinum and readily explains their recognition as distinct taxa by many authors. The histogram and the pictorialized scatter diagram illustrate the gradual transition of one phase into another, and the continous nature of the variation. The distribution map illustrates that most of the specimens which were assigned low scores are restricted to the Seward Peninsula, parts of the northern Arctic Coast, portions of the Canadian Rockies, the southern coast of Labrador, and parts of the western and northwestern coasts of Newfoundland. These dwarfed, large flowered plants form a continuous series which grades into typical var. alpinum in more favorable southern and lowland habitats. Collections from the northern Arctic Coast and 119 especially the Seward Peninsula, indicate that considerable intrapopulational variation occurs in the northern phase of !· alpinum. The calyx teeth vary from deltoid to nearly linear, the spreading flowers range from 16 to 19 or more mm. in length and the leaflets vary from linear to nearly elliptic. It is the authors opinion that these highly variable and clearly arctic populations of H. alpinum, previously described as ~· truncatum, and as H. boreale var. grandiflorurn, represents the remains of a population which was isolated in parts of unglaciated North America for a great period of time under climatic conditions much more severe than found there at the present time. The present distribution of H. alpinum and of many species with an essentially similar geographical range can be explained by supposing a more or less continuous circurn- polar arctic-alpine distribution during the Tertiary. The absence in Tertiary deposits of arctic-alpine plant material does not preclude their former existence in such areas, as their herbaceous nature and upland existence was highly unfavorable to fossilization. Fossil and phytogeographical evidence suggests that the extant arctic-..alpine flora evolved during the late Terit~ry either from taxa which were preadopted to an arctic ~abitat by way of an·alpine existence in the high mountains .of An).erica and Eurasia, or from taxa which occupied the nor~hern Tertiary forests and evolved therein as a direct result of deteriorating climatic conditions. 120

With the onset of the Pleistocene, much of North America was covered by ice, having extended as far south as Kansas and southern Ohio. Geological, palynological and phytogeographical evidence, however, indicates that vast portions of land in North America were never glaciated and served as refugia for many plants throughout the Plei- stocene. These areas include the coast of the Bering Sea, the arctic coastal plain, parts of the Yukon Valley, portions of the Gaspe Peninsula, Anticosti Island, and parts of Newfoundland and Labrador. Ives (1963) concluded that large areas in Northern Labrador, Baffin Island, and New- foundland remained ice free throughout the Pleistocene period, while Hulten (1937, 1968) emphasized the importance of Alaska as a refugium for plant species during the Plei- stocene glaciations. A study of fossil pollen from the Seward Peninsula by Colinvaux (1963) indicates that during glacial times, dwarf Birch, Spruce and Alder were found much further south than at present and at each site he examined the vegetation was comparable to that now found several hundred kilometers to the north. At the close of the Wisconsin, there was an advance of trees and shrubs on the Seward Peninsula which continued until about 10,000 years ago when the tree line reached its present limits. This withdrawal of trees and shrubs during glacial times, with a corresponding advance during the interglacial periods, strongly suggests that climatic conditions were much colder than at present both 121 on the Seward Peninsula and on the arctic coast during the glacial periods. Johnson and Packer (1963) studied the flora of Ogotoruk Creek using the diploid-polyploid ratio. They were able to group the 300 species of angiosperms growing there into four geographical elements. A widespread oceanic element; a montane element which probably evolved in southern mountainous regions; an arctic element that probably evolved in the arctic and remained there; and a boreal element which is primarily associated with the boreal forest. They found a high degree of polyploidy in the arctic and boreal elements, while the oceanic and montane elements had a high degree of diploidy indica·ting that the boreal and arctic elements probably evolved under conditions of considerable stress. In recent years, considerable geological, palynological, oceanographic and paleontological evidence has accumulated supporting the existence of a land bridge (Beringia) between Alaska and Asia during glacial periods (Hopkins, 1967). During the middle Tertiary, Beringia was evidently part of an extensive mesophytic forest which occupied vast areas along the North Pacific. Beringia appears to have submerged for the first time in the late miocene, followed· by several restorals in the Pliocene and Pleistocene with the final closure about 12 or 13,000 years ago. During the Pleistocene, the bridge itself was probably a flat tundra plain which supported 122 lowland herbaceous species with a few low growing shrubs essentially very comparable to that now found at Cape Thompson. This land bridge may well have been the route of migration for many members of the present North Amer- ica flora and fauna, including g. alpinum, although long distance dispersal cannot be ruled out. To summarize the apparent geographic history of Hedysarum alpinum it appears that this species occupied an essentially circumpolar arctic-alpine distribution during the late Teritary. At the onset of the Pleistocene, when climatic conditions began to deteriorate and the ice sheets advanced over much of North America, it became restricted either to areas far south of the ice sheets or to isolated refugia in Alaska and along the Northeastern seaboard in parts of what is now Newfoundland, Labrador and Quebec. With each retreat of ice there was probably a corresponding advance of Hedysarum taxa into the areas laid bare by the retreating glaciers. Plants which were restricted to Beringia were subjected to climatic conditions considerably more severe than those which prevail in these regions at the present time. This resulted in an intense selection for morphological and physiological characteristics which were adaptive in such a climate. These include a dwarfed, prostrate habit, a short, compact inflorescence, larger and darker flowers, and deeply pigmented vegetative tissue, along with many physiological modifications the nature of which remain largely unclear at the present 123

time. H. alpinum, like all arctic-alpine plants, must adapt to conditions of low summer and winter temperatures, strong winds, short growing seasons, long photoperiods with low light intensities, little available moisture and nitrogen poor soil. These climatic and edaphic conditions tend to form a continuous gradation resulting from various degrees of latitude, altitude, topographic relief and various other features. In the arctic on sunny days, the temperature close to the ground is considerably higher than the air temperature and the low growing prostrate habit insures that meta- bolic activities procede at the most favorable temperature. It also insures that a small quantity of seed will mature each year even when the degree of warming is only slight by preventing carbohydrate wastage in the production of a tall flowering stem. On sheltered sites in moist well drained soils, ~· alpinum can reach 70 cm. or more in height with an elongated inflorescence bearing 50 or more rather small flowers, while on an exposed site the entire plant may be less than 15 cm. tall with a short compact inflorescence bearing 5 or 6 large dark blue flowers. Such dwarfing undoubtedly reduces seed production and probably dispersal but the advantage of insured seed production has a much greater selective advantage in the arctic. Under less extreme climatic conditions as are found in the more southerly portions of its range, such dwarfing would be a disadvan- 124 tage as air temperatures close to the ground might rise above that necessary for most efficient metabolism.' When intense pigmentation is combined with low stature and dense growth, the degree of warming increases considerably. Tikhomirov, et al, (1960) measured the air temperatures inside white and blue flowers. They found that inside white flowers the temperature was 0.7-2.o0 c. higher than the surrounding air temperature, while in blue flowers it was 3.4-4.2°c. higher. Deep pigmentation probably also aids in early spring growth under the snow by increasing the amount of light that is absorbed. H. alpinurn is also a cross polinated species, a condition which is relatively uncommon in arctic-alpine plants. This increases the selection advantage of darker and larger flowers to an even greater extent by raising the chances for insect pollination. The large storage roots of H. alpinum contain large amounts of food reserves and undoubtedly aid in the immed- iate resumption of growth in the spring. This is particularly effective in combination with deeply pigmented vegetative tissue. While isolated for long periods of time in Beringia under climatic conditions considerably more severe than at present, selection pressures for the above mentioned modifications would be extremely strong and considering the nature of the breeding system in H. alpinum such modifications would be easily obtained. Even after the retreat of 125

the ice, with relatively arctic-subarctic conditions main- tained in this region, one would expect a plant population exhibiting the most extreme degree of arctic-alpine adaptations to be present in this region. With glacial retreat from more southerly areas and the subsequent availability of large areas of land for recolonization, H. alpinum would be expected to radiate adaptively into such regions. With the boreal and temperate conditions found in the more southerly portions, selection would procede in a directional manner toward the habit which is currently found in the southerly portions of its range.

Key to the Varieties of H. alpinum 1. Loments markedly pubescent on both surfaces to only marginally pubescent .•.• var. philoscia. 1. Loments glabrous (2). 2. Flowers 11-15 mm. long; racemes elongate, flowers pendant •••••••••.••• var. alpinum. 2. Flowers 16-19 mm. long; racemes compact, flowers spreading •••••••• var. grandiflorum.

Hedysarum alpinum L. var. alpinum Fig. 43.

H. alpinum var. americanum Michx. Fl. Bor. Am. 2:74. 1803. H. alpinum subsp. americanum (Michx.) Fedtsch. Acta Hort. Petrop. 19:255. 1902 H. alpinum var. americanum f. albiflorum Fern. Rhodora 35:275. 1933. 126

H. americanum (Michx.) Britt. Mem. Torr. Bot. Club. 5:201. 1894. H. auriculatum East w. Bot. Gaz. 33:205. 1902. H. alpinum L. var. americanum Rollins, Rhodora 42:222. 1940. Stems few to many, erect 1.5-7 dm. long, 1-3 mm. in diameter, usually branched, glabrous or appressed pubescent; stipules brown, connate, 5-15 mm. long; leaves 5-16 cm. long; leaflets 13-27, oblong to lanceolate, pubescent below, glabrous above, 11-35 mm. long, 4-12 mm. wide; inflorescence 5-30 cm. long, elongate to compact, bracts linear, 1-4 mm. long; flowers 19 to 50 or more, usually pendant, lilac purple to pale ·purple or white, 10-15 mm. long; calyx tube 2-5 mm. long, pubescent, teeth 1-2.5 mm. long, triangular; loment articles 2-5, glabrous. Ecology and distribution. Alaska and the Yukon across the north to Labrador, Newfoundland, and Anticosti Island, south through British Columbia, Alberta, Ontario, Saskatchewan and Quebec to northern Montana. Also in the Allegenhy Mountains of northern Maine, Vermont and New Brunswick. Reported from New Hampshire and Massachusetts by Anderson (1959) and Seymour (1969) but specimens from these areas were not found during the course of this study Fig. 44). Var. alpinum is a boreal rather than an arctic- alpine variety. In Alaska, it inhabits mainly lowland areas in well drained alluvial soils, sand and gravel bars, 127

Figure 43. Hedysarum alpirtum var.· ·alpinum. 128

.. ·--- -

- IOO .... latO MOO •LOMDIM ---- ,...... •• Figure 44. Distribution of H. alpinum var. alpinum. 129 stream banks, open woodlands and occasionally wet tundra. Southward, it occupies a wide variety of habitats, in- clu~ing lake shores, river banks, open prairies and woodlands. Flowering from early June to early September. From sea level to 7000 feet elevation. Representative specimens: West of Killik River, 10 July 1946, R. M. Chapman 84 (US), Kenai Peninsula, mouth of six mile creek, 11 July 1953, John H. Beaman 601 (WS), Tok River Bridge, 10 miles east of Tok, 27

July 19~6, s. L. Welsh 6028 (BRY), 20 miles east of Slana- Tok Highway on road to Nabesna, 3 August 1965, s. L. Welsh 5088 (BRY), Fairbanks, 24 July 1968, P. Rutledge 18 (BRY), Mckinley National Park, 6 July 1936, Edith Scamman 304 (GH), Savage River, 22 June 1928, Ynes Mexia 2029 (US), Mckinley National Park, 30 June 1939, Aven Nelson 3594 (WS), Igloo Creek, 10 July 1939, Aven Nelson 3759 (WS), Glacier Bay, 14 July 1907, K. Stephens 51 (US), Rapids Lodge, 138 miles south of Fairbanks, 7 August 1936, Edith Scamman 302 (GH), northeast of Umiat, 27 July 1952, George Lindsay 2330 (GH), south bank of Colville River, below Umiat, 20 July 1951, Quentin Jones 637 (WS), mile 18 Slana-Tok Highway, 24 June 1944, J. P. Anderson 8749 (GH), arctic coastal plain, Ikpikpuk River, 30 June 1959, Otto w. Geist s. n. (BRY), Driftwood creek, 28 July 1952, George W. Wood 1266 (GH), Black Rapids Glacier Observation Point, 14 June 1965, s. L. Welsh 4325 (BRY), Kenai Lake 11 July 1951, J. A. Calder 5808 (WS), Palmer, 23 July 130 1965, s. L. Welsh 4899 (BRY), 44 miles south of Palmer on Glenn Highway, 15 July 1966, S. L. Welsh 5663 (BRY), Cir- cle Hot Springs, 17 July 1936, Edith Scamman 305 (GH), Circle, on the Yukon River, 20 July 1949, Edith Scamman 5333 (GH), Kobuck River, 20 July 1923, Palmer 657 (US), Chicken, 26 August 1949, Edith Scamman 5743 (GH), near Hot Sulphur Springs, 6 July 1926, A. E. Porsild 677 (GH), Franklin, 11 July 1941, J.P. Anderson 7166 (GH), north of Big Timber, 23 July 1939, Aven Nelson 3953 (GH), Manley Hot Springs, 19 July 1945, Edith Scamman 3725 (GH), Falls Creek, 4 August 1951, W. J. Cody 6318 (WS), Hurst Peak, 16 July 1917, G. C. Harrington, 54 (US), Umiat, 23 July 1960, Eric Hulten s.n. (US), Copper River Region, 23 June 1902, w. Poto 48 (US), Ophir, 26 June 1940, Edith Scamman 1878 (GH), mile 1022 Alaska Highway, 29 July 1957, w. B. Schofield 8046 (CAN), Kokrines Moun- tains, 6 July 1926, A. E. Porsild 677 (CAN), 60 miles east of McGrath, 6 July 1969, K. M. Reed 691 (BRY), Eagle Sum- mit, 12 July 1940, Edith Scamman 2117 (GH), Toklat River Bridge, 22 July 1965, S. L. Welsh 4842 (BRY). ALBERTA. Vicinity of Banff, 19 July 1916, F. W. Hunnewell 4449 (GH), vicinity of Rosedale, 5 June 1915, Marion E. Moodie 948 (US), Calgary, 18 June 1903, M. A. Barber 262 (GH), vicinity of Fort McMurray, 10 September 1935, Hugh M. Raup 7057 (GH), north of Jasper Park Lodge, August 1949, Louise R. Sieburth s.n. (WS), Craigmyle District, 11 July 1922, A. H. Brinkman 721 (GH), Jasper National Park, 31 131

July 1941, Edith Scamman 2361 (GH), 2 miles south of

Cardston, 21 June 1960, Keith Shaw 156 (BRY), Whitecourt,

16 July 1958, B. Boivin 12528 (NY), west of Pincher Creek,

30 June 1940, E. H. Moss 1027 (GH), Ptarmigan Lakes, 4 July

1906, Stewardson Brown 410 (GH), 15 miles west of Calgary,

·12 June 1965, C. L. Hitchcock 24032 (WS), Lee's Creek, 8

July 1960, R. K. Shaw 186 (BRY), Waterton Park, 4 August

1953, A. J. Breitung 17054 (NY), Devona, 13 August 1943,

Edith Scamman 3150 {GH), Peace River, 18 July 1941, E. H.

~ 6138 {GH), Fort Saskatchewan, 16 August 1940, G. H.

Turner, 2253 (GH), Rosedale, 5 June 1915, Marion Moodie

948 {GH}, 3 miles south of Islay, 27 June 1958, J. Lowther

66 (CAN), Primrose Lake, 17 July 1968, M. G. Dumois 3493

(CAN}, Cardston, 29 July 1911, M. Malte s.n. (CAN}, Red Deer River, 12 July 1917, c. H. Young s.n. (CAN}, Mcleod River, 19 June 1898, w. Spreadbough 19314 (CAN}, Salt River Region, 20 August 1916, Charles Camsell s.n. (CAN}, Bowness

Park, 16 July 1925, M. O. Malte 1296 {CAN}. BRITISH COLUM-

BIA. Pine Pass, 3 August 1954, J. A. Calder 13989 (WS),

along Swift River, 6 August 1960, J. A. Calder 27952 (WS),

Summit Pass, 20 July 1943, Hugh M. Raup 10687 (CAN}, vi-

cinity of Beatton River, 23 June 1943, H. M. Raup 10252

(CAN), Cassier District, 15 August 1925, G. W. Norman 53

(CAN), 2 miles south of junction of Goat Creek and Elk

River, 30 June 1958, R. Taylor 2268 (WS), Mt. Robson, 19

August 1943, Edith Scamman 3295 (GH), Coldfish Lake, 22

July 1969, K. Rigby 175 (BRY), vicinity of Hudson Hope, 132

20 June 1932, Hugh M. Raup 3626 (GH), Haines Road near mile 85, 17 July 1944, C. Clark 493 (CAN), 20 miles south of Land River, 26 June 1968, s. L. Welsh 7366 (BRY), Sikanni Chief River Valley, 21 June 1966, s. L. Welsh 5430 (BRY), Alaska Highway mile 369, 22 June 1966, s. L. Welsh 5438

(BRY), south end of Coldfish Lake, 26 July 1969, K. Rigby.

192 (BRY). LABRADOR. Dudley Lake near Labrador City, 12

July 1967, I. Hustich 328 (CAN). MAINE. Aroostook County:

Valley of Aroostook River at Fort Fairfield, 18 July 1902,

M. L. Fernald and E. F. Williams s.n. (GH, US), St. Francis, Banks of St. John River, 10 .July 1932, A. s. Pease s.n. (GH), Fort Kent, 15 July 1908, Kenneth K. Mackenzie 3536

(US), St. Francis, 17 August 1893, M. L. Fernald 26 (US),

Valley of the St. John River, 23 July 1917, Harold St.

John 2362 (US), Masardis, 8 September 1897, M. L. Fernald s.n. (GH). MANITOBA. Shoal Lake, 5 August 1951, H.J.

Scoggan 10170 (CAN), Brandon, 23 June 1953, H.J. Scoggan

10846 (CAN), Beulah, 10 July 1953, H.J. Scoggan 11239

(CAN), Foxwarren, 11 August 1951, H.J. Scoggan 10319

(CAN), Rossburn, 6 August 1951, H.J. Scoggan 10194 (CAN),

Porcupine Mountain, 15 July 1950, H. J. Scoggan 8071

(CAN), Clear Lake, 16 August 1948, G. de Ruyck s.n. (N¥),

Gillam, 29 July 1950, W. B. Schofield 1281 (WS), Lake Audy Road, 18 June 1932, w. Halliday 68-1932 (CAN), Hill Lake, 13 July 1948, H.J. Scoggan 3881 (CAN). MONTANA. Glacier County: Sec. 29, T. 37 N., R. 14 w., 16 July 1949, Dan Lynch 6095 (WS), 8 miles south of Babb, 2 August 133 1948, R. F. Daubenmire 48391 (WS), 6 miles west of Browning on highway 2, 17 July 1958, G. F. Payne 36 (US), vicinity of Kiowa, 13 July 1954, L. H. Harvey 5842 (MONTU), Rising Wolf

Ranch, 31 July 1952, L. H. Harvey 5224 (MONTU), St. Mary's

River, Glacier National Park, 26 July 1954, L. H. Harvey

6072 (MONTU), east side of Glacier National Park, 26 June

1930, Spear 5360 (BRY). NEW BRUNSWICK. Tobique River, 22

July 1884, N. G. Hay 1183a (CAN), Gorge of the Aroostook

River, 17 July 1902, M. L. Fernald s.n. (GH), Tidehead, 5 August 1926, M. o. Malte 771 (CAN), St. John River, 13 July 1903, A. s. Pease 2262 (GH). NEWFOUNDLAND. Grand Falls, Valley of the Exploits River, 3 July 1911, M. L. Fernald

5801 (CAN), Bishop Falls, 28 July 1911, M. L. Fernald 5800

(GH), Grand Falls, 22 July 1911, M. L. Fernald and K. M.

Wiegand 5799 (GH). ONTARIO. North shore of Lake Superior,

30 August 1937, R. c. Hosie 1586 (GH), Sibley Tp., Thunder Bay District, 29 June 1936, T. Taylor 682 (GH), Blackwater River, 1 mile from Lake Nipigon, 23 July 1960, c. E. Garton 7748 (WS), Smooth Rock Falls, 11 July 1957, A. s. Pease 38809 (GH), Fort Severn, 19 July 1956, I. Hustich 1459 (CAN), junction of Fawn and otter Rivers, 17 July 1952, D. Moir 633

(CAN), Caribou Island, 2 August 1952, o. Mojr 994 (CAN),

Cochrane, 12 August 1952, w. Baldwin 3956 (CAN), Hearst, 4 August 1952, W. Baldwin 3777 (CAN), Moose River, 10 August 1968, F. Miron s.n. (CAN), New Liskeard, 27 July 1953, w. Baldwin 5326 (CAN), Finlay Bay, 2 July 1936, T. Taylor 683 (CAN), northeast of Moosonee, 3 August 1968, W. s. Dickin- 134 son 557 (CAN), Mountain Lake, 4 August 1936, T. Taylor 680

(CAN). SASKATCHEWAN. 30 miles west of Touchwood, 17 July

1906, John Macoun 70784 (GH), Cypress Hills, 27 June 1895,

J. Macoun 10165 (CAN), Indian Head, June 1890, Spreadbo- rough 5378 (CAN), Prince Albert, 29 June 1896, J. Macoun

12506 (CAN). VERMONT. Franklin County: Mt. Mansfield, Pringles Ravine, L.A. Charette 2526 (BRY); Lamoille County:

Smugglers Notch, 29 June 1911, H. St. John 297 (WS), Stowe

Notch, 17 July 1891, J. Grant s.n. (NY). QUEBEC. Anticosti Island, Riviere Notiskotek, 8 August 1927, F. Marie Victorin and Rolland-Germain 27332 (GH), Anticosti Island, Riviere Du

Renard, 6 August 1927, F. Marie Victorin and Rolland-Germain

27354 (GH), Anticosti Island, Cap de L'ouest, 8 August 1927,

F. Marie Victorin and Rolland-Germain 27156 (GH), between

Balde and the Baie des Chaleurs, 5 August 1904, M. L. Fer- nald 6450 {GH) , east of Martin River on the Gulf of St. Law- rence, 26 July 1922, M. L. Fernald 25171 (GH), Tourelle, 20

July 1934, J. H. Pierce 7a {GH), Screes near Gros Marne, 12 July 1952, A. s. Pease 36623 (GH), west of Riviere a Pierre, 27 July 1927, s. Kelsey 97 (GH), gravel bar on the St. John River, 23 August 1904, M. L. Fernald 5076 (GH), Cap Gas, 17

July 1946, G. Proctor 2196 {GH), junction of Restigouche and Patapedia Rivers, 23 July 1929, J. Rousseau 32149 (CAN),

Ile Manitounouk, 12 July 1944, J. Rousseau 117 (CAN), sea cliffs near Bic, 1 July 1957, H.J. Scoggan 13778 {CAN),Bone- venture Formation, 4 July 1941, H. J. Scoggan 2066 (CAN),

Ile Du Massacre, 21 July 1939, H. J. Scoggan 201 {CAN), Fort 135

Chimo, 23 July 1963, Albert Legault 6789 (CAN), St. Donat, 7 June 1943, E. Lepage 4208 (GH), Lac Pleureuse, 8 July 1946, G. Proctor 2087 (GH), Lac St Jean, 23 July 1921, M• . - Victorin 16082 (WS), north of Point Bleue, 29 July 1941, A. S. Pease 28929 (GH), Ruisseau de Sorel, 4 August 1939, Un- gava, 11 August 1945, Rev. A. Dutilly 14627 (GH). NORTH- WEST TERRITORIES. Wood Buffalo Park, Slave River, 8 August 1928, Hugh Raup 2807 (GH), vicinity of Calumet Creek, 5 July 1926, H. M. Raup 998 (GH), Brintnell Lake, 5 July 1939, H. M. Raup 9423 (WS), Norman Wells, 4 August 1953, W. J. Cody 7797 (WS), Horn Lake, 14 July 1962, P. Youngman 115 (CAN), Fairchild Point, Great Slave Lake, 14 August 1927, Hugh M. Raup 997 (GH), north shore of Mcleod Bay, Great Slave Lake, 12 July 1927, H. M. Raup 1000 (GH), Peace Point, 2 August 1928, H. M. Raup 2803 {CAN), Heart Lake, 20 August 1928, H. M. Raup 2804 (CAN), east shore of Thelen River, 28 July 1965, G. B. Rossbach 6892 (CAN), Great Bear Lake, north shore, 6 July 1928, A. E. Porsild 4955 {CAN), Leith Peninsula, Great Bear Lake, 6 July 1948, H. T. Shacklette 2845 (CAN), Port Radium, 2 July 1948, H. T. Shacklette 2454 {CAN), Thelen Game Sanctuary, 27 July 1952, J. Tener 241 (CAN), Fort Providence, 6 July 1951, Alton Lindsay 163 {CAN), Campbell Lake, 16 June 1927, A. E. Porsild 2001 (CAN), Keewatin District, 62°30' N., 97° 30' w., A. E. Porsild 5834 {CAN), 10 miles upriver from Sans Sault Rapids, 3 August 1951, Alton Lindsay 602 (CAN), Baker Lake, 26 July 1957, Raymond D. Wood 90 (CAN), west 136

side of Bathurst Inlet, 30 July 1950, J. Kelsall and E.

McEwen 187 (CAN), origin of Great Bear River, 30 July

1951, Alton A. Lindsay 527 (CAN), Coppermine, 31 July

1951, W. Findlay 216 (NY), 23 miles west of Fort Smith,

9 July 1950 (WS). YUKON TERRITORY. 6 miles northwest

of Sam Lake, 6 July 1970, S. L. Welsh 10277 (BRY), Dawson, 19 June 1914, Alice Eastwood 306 (GH), Ladue Valley, 22

June 1939, Hugh Bastock 91 (CAN), mile 95 Canol Road, 11

July 1944, A. E. Porsild 10390 (CAN), vicinity of White- horse, 10 August 1943, H. M. Raup 11136, (CAN), Alaska

Highway at White River, 2 July 1944, J. P. Anderson 9282

(GH}, 33 miles southeast of Sam Lake, 1 July 1970, S. L.

Welsh 10116 (BRY), Old Crow Quadrangle, 26 July 1970, K.

Rigby 128 (BRY}, Ranch Creek, 10 June 1899, M. W. Gorman 1018 (US), mile 882 Alaska Highway, 24 June 1966, s. L. Welsh 5496 (BRY}, mile 660 Alaska Highway, 25 June 1962,

Ray J. Davis 5052a (BRY}, Dawson, 2 July 1914, Alice

Eastwood 502 (WS}.

Hedysarum alpinum L. var. grandiflorum Rollins, Rhodora 42:233. 1940. Fig. 45. (Fernald et al. 28625, Pistolet Bay Newfoundland, 11 August 1925 (G)}.

Hedysarum truncatum Eastw. Bot. Gaz. 33:205.

1902.

Stems one to few, erect to somewhat prostrate, 1-

2.5 nun. in diameter, often unbranched, glabrous or appressed pubescent; stipules brown, connate, 6-13 nun. long; leaves 7-10 cm. long; leaflets 7-17, oblong to lanceolate, 137

Figure 45. Hedysarum a1p·inum var. grandifl'o·rum. 138 pubescent below, glabrous above, 14-20 mm. long, 4-8 mm. wide; inflorescence 2-6 cm. long, compact; bracts linear, 2-5 mm. long; flowers 10-20, spreading, usually, 16-19 (22) mm. long; calyx tube 3-6 mm. long, pubescent, teeth 1-2 mm. long, deltoid or often nearly subulate; loment articles 2-5, glabrous. Ecology and distribution. Occuring in the Bering Strait District, St. Lawrence and Nunivak Island, east along the northern arctic coast of Alaska and Yukon, and south sporadically in the high mountains of Alaska, Yukon, British Columbia and Alberta. Also isolated in parts of Labrador, Newfoundland and Quebec (Fig. 46). Inhabits open stony tundra, rocky slopes and ridges in Alaska, mountain slopes at or above the timberline in the mountains of northern Alberta, British Columbia and the Yukon. In Newfoundland it occupies limestone barrens, calcareous sandstone cliffs and rocky ridges. Flowering from early June to late August. From at or near sea level to 7000 feet elevation. Representative specimens: ALASKA. Nome, 7 June 1970, M. Williams 2614 (BRY), Nome River Valley, 14 July 1966, s. L. Welsh 5863 (BRY), Teller, 6 August 1949, Edith Scamman 5566 (GH), Anvil Hill, 5 August 1945, Edith Scamman 3886 (GH), 1 mile north of Palmer, 13 June 1965, S. L. Welsh 4107 (BRY), Nugget, 13 July 1919, c. w. Thornton 414 (US), 6 miles northwest of Nome, 14 July 1966, Robert Pegau 239 (BRY). ALBERTA. Head of Smokey River, 5 August 1911, J. 139

•o

KAU j I T: 4T' I .. 1 I .,. 111t1.C• ·----·----~

<00

Figure 46. Distribution of a.· ·a1pinum var. grandiflorum. 140

H. Riley 36 (GH) , Headwaters of the Saskatchewan and Atha- basca Rivers, 11 July 1908, s. Brown 1218 (GH, NY}. BRITISH COLUMBIA. Slopes of Mt. Selwyn, 19 July 1932, Hugh Raup

3967 (GH}, Mt. Selwyn, 26 July 1932, Hugh Raup 4091 (GH).

LABRADOR. Near Forteau, 8 August 1893, A. C. Waghorne s.n.

(GH), Forteau, 1870, s. R. Butler s.n. (GH). NEWFOUNDLAND. East of Big Brook, 15 July 1925, K. M. Wiegand 28618 (GH),

Region of St. John Bay, 25 July 1929, M. L. Fernald 1833

(GH), Bay of Islands, 16 July 1929, M. L. Fernald 1832 (GH), north of St. Paul's Bay, Cow Head, 23 July 1910, M. L.

Fernald 3636 (GH), Pistolet Bay Anse Aux Sauvages, 11 Aug- ust 1925, M. L. Fernald, et al. 28625 (GH, type}, Burnt

Cape, 17 July 1925, M. L. Fernald 28621 (GH}, Table Moun- tain, Port a Port Bay, 16 July 1914, M. L. Fernald 10849

(GH). NORTHWEST TERRITORIES. Vicinity of Fort Simpson, 11

June 1939, Hugh M. Raup 9084 (GH}, Victoria Island, 8 Aug- ust 1949, A. E. Porsild 17316 (CAN}, Banks Island, 30 July

1949, A. E. Porsild 17588 (CAN), Banks Island, Sachs Har- hour, 16 July 1955, E. H. McEwen 257 (CAN). YUKON TERRITORY.

Kluane Lake, 2 July 1944, H. M. Raup 12128 (CAN}, Burwash

Landing, 3 July 1943, C. Clarke 259 (CANl, north slope of Buckland Mountains, 13 July 1970, s. L. Welsh 10613 (BRY).

Hedysarum alpinum L. var. philoscia (A. Nels.} Rollins, Rhodora 42:224. 1940 (Laramie Mountains, Albany County, Wyoming, Aven Nelson 2034, 1896)

Hedysarum philoscia A. Nels. Proc. Biol. Soc. Wash.

15:185. 1902. 141 Stems few to many, erect, 2-4 dm. long, 1-3 mm. in diameter, branched, pubescent; stipules brown connate, 5-13 mm._ long; leaves 6-13 cm. long; leaflets 13-25, oblong to lanceolate, pubescent below, glabrous above or occasionally pubescent on both surfaces, 11-27 mm. long, 7-11 mm. wide; inflorescence 9-23 cm. long elongate; bracts linear, 1.5-4 mm. long; flowers 20-40, pendant, pale purple to lilac purple, 11-14 mm. long; calyx tube 2-5 mm. long, pubescent, teeth 1-2.5 mm. long, triangular; loment articles 2-5, markedly pubescent on both surfaces to onl,y marginally pubescent. Ecology and distribution. Saskatchewan, Alberta, and Ontario then disjunct to South Dakota and Wyoming (Fig. 47). In the prairie provinces, variety philoscia inhabits dry grassy prairies, hills and occasionally stream banks, while southward in South Dakota and Wyoming, it occupies moist open woods, aspen forests and wet shaded stream banks. Flowering from late May to late August. From 2000 to 7000 feet elevation. Specimens examined: ALBERTA. Calgary, 30 May 1913, Marion E. Moodie s.n. (US), vicinity of Rosedale, 10 August 1915, Marion E. Moodie 948 (US), Banff, Bow River Valley, 20 July 1906, s. Brown 672 (GH), Banff, August 1900, Frances c. Prince s.n. (GH), Fort Saskatchewan, 3 August 1940, G. H. Turner 2129 (GH), near Pincher Creek, 31 July 1939, E. H. Moss 186 (GH). ONTARIO. near Beaver Falls, 15 August 142

Figure 47. Distribution of H. alpinum var. philoscia. 143

1959, C. V. Morton & E. Lepage 11626 (US), banks of Kopus- kosing River, 20 July 1952, W. Baldwin 3317 (CAN). MANITOBA.

Riding Mountain National Park, 27 July 1953, A. Love 6191

(GH), Clear Lake, 29 August 1941, Edith Scamman 2913 (GH),

Grand Rapids, 5 August 1948, H. J. Scoggan 4400 (GH), 10 miles south of Birtle, 10 July 1953, H. J. Scoggan 11239

(GH). SASKATCHEWAN. 32 miles west of Lloydminster, 17

July 1941, Susann Fry 564 (WS), 3 miles west of McKague, 3

July 1941, August J. Breitung s.n. (GH). SOUTH DAKOTA.

Lawrence County: Camp Judson, 17 June 1940, Philip Johnson

245 (GH), mountain slopes near Custer Peak, 26 June 1929,

E. Palmer 37554 (GH) ; Pennington County: Deerfield, 23

June 1929, E. Palmer 37509 (GH), Rochford, 12 July 1892,

P.A. Rydberg 640 (US), 5 miles south of Hill City, 3 July

1958, S. L. Welsh 755 (BRY). WYOMING. Albany County:

Willow Creek, 13 July 1897, Aven Nelson 3367 (US), Crow

Creek, 27 August 1903, Aven Nelson 8955 (US, GH), Pole Mountain Region, 8 August 1945, c. L. Porter 3742 (GH), Laramie Hills, 19 July 1901, Elias Nelson 622 (NY); Weston

County: Boyd, 19 July 1910, Aven Nelson 9436 (US, GH).

Hedysarum sulphurescens Ryd. Bull. Torrey Bot. Club. 24: 251. 1897. Fig 48. (Near Helena, Montana, F. D. Kelsey, May 1892)

Hedysarum boreale var. albiflorum Macoun, Cat.

Canad. Pl. 1:510. 1884.

Hedysarum flavescens Coult. & Fisch. Bot. Gaz. 18:

300. 1893, non H. flavescens of Regel & Schm.

1882. 144

Figure 48. Hedysarum: sulphurescens Rydberg. 145

Hedysarum boreale var. leucanthum, M. E. Jones. Proc. Calif. Acad. Sci. 5:677. 1895. Hedysarum boreale var. flavescens (Coult & Fisch.) Fedtsch. Bull. Herb. Boiss. 7:256. 1899. Hedysarum albiflorum (Macoun) Fedtsch. Acta Hort. Petrop. 19:252. 1902. Terrestrial herbaceous perennials; roots stout, ligneous, elongate; stems several to many, decumbent to erect, longitudinally grooved, not rooting, 1-6 dm. long, 1-5 mm. in diameter, branched above solid, terete, pubescent; stipules tan to brown, united, chartaceous, 8-15 mm. long, obtuse to acute or acuminate; leaves odd-pinnate, 7-12 cm. long; leaflets 9-17, elliptic to ovate-oblong, pubescent below, glabrous above, conspicuously veined, 9-33 (38) mm. long, 4.5- 15 (21) mm. wide, rounded to obtuse; inflorescence racemose, 2-12 cm. long, elongate to compact; bracts brown, pubescent, linear, 2-6 mm. long; bractioles pubescent, linear 1-3 mm. long; flowers 6-38, erect to spreading or pendent, yellow, 14-18 mm. long; calyx campanulate, tube 3-8 mm. long, bractiolate, pubescent, teeth 0.5-1.5 mm. wide, 1-3 mm. long, unequal, subulate, greenish, erect; standard obovate to cuneate, emarginate, 11-14 mm. long, 5-8 mm. wide; wings 11-14 mm. long, 2-3 mm. wide; wing auricles linear, united, 3-3.5 mm. long; stamens 10 diadelphous; style slender, curved; laments pendulous, stipitate; articles 1-4, gla~ brous, wing margined, 5.5-8 mm. wide, 7.5-12 (13.5) mm. long, reticulations polygonal. 146

Ecology and distribution. Found in a wide variety of habitats ranging from rocky ledges, talus slopes and limestone outcrops to dry banks, moist slopes and meadows from 3,000 to 10,000 feet elevations. Flowering from late May to late August. Alberta and British Columbia, south to Washington, Idaho and Wyoming (Fig. 49). Hedysarum sulphurescens exhibits the same variation continuum as does H. alpinum only to a much less degree due to its limited distribution. Alpine plants, however, exhibit the typical pattern of a short, compact few flowered inflorescnece, short stature, and larger flowers. Specimens examined. ALBERTA. Prairie west of Pincher Creek, 17 June 1940, E. H. Moss 763 (NY), mouth of Cameron Creek, 10 July 1953, A. J. Breitung 15664 (NY), Lake O'hara trail, 14 July 1949, L. R. Silburth s.n. (WT), Bow River Valley, 9 June 1906, s. Brown 127 (GH, CAN), Pipestone Valley, 7 July 1906, s. Brown 425 (GH), Sulphur Mountain near Banff, 28 June 1916, Marion E. Moodie 1261 (US), 113° 53' W, 49o 11' N, 3 August 1969, J. Nagy 2612 (CAN), Waterton Lakes National Park, 11 June 1969, W. Blais 979 (CAN), Sofa Creek, 17 June 1969, J. Nagy 1175 (CAN), Milk Mountains, 10 miles southeast of Milk River, 26 June 1952, B. Boivin 9504 (NY). BRITISH COLUMBIA. Paradise Mine, 15 miles west of Windermore, 1 August 1953, J. A. Calder 11290 (WT, NY), 8 miles south of Cran- brook, 2 July 1941, w. A. Weber 2240 (NY, WT), Burgess Trail, 1 September 1904, Charles H. Shaw 591 (NY), Sheep 147

120 110

Figure 49. Distribution of H. sulphurescens. 148

Creek, 19 July 1920, W. B. Anderson s.n. (WT), Banff, 10 August 1898, J. R. Anderson s.n. (WT), Burgess Trail, 1 September 1904, J. Macoun 591 (GH), Vermillion range, 10 August 1964, H.J. Scoggan 16220 (CAN), Sunburst Lake Camp, Mount Assiniboine, 7 August 1952, Edith Scamman 6669 (CAN). IDAHO. Kootenah County: south end of Lake Pend Oreille, 27 July 1892, J. H. Sandberg 748 (NY, GH, US). MONTANA. Beaverhead County: Anaconda Range, head of Pintlar Creek, 27 July 1945, c. L. Hitchcock 12740 (NY, GH, WT), east base of Black Lion Mountain, 30 July 1945, c. L. Hitchcock 12922 (NY, WT), head of Rock Creek east of Lake Waukena, 1 August 1945, c. L. Hitchcock 13142 (NY, WT), Lima, 14 July 1908, Marcus E. Jones 8221 (US); Carbon County: Red Lodge, Cooke City Road above Rock Creek, 12 August 1945, C. L. Hitchcock 13549 (NY, WT), 13 miles southwest of Red Lodge, 22 July 1955, Arthur Cron- quist 7967 (NY, WT), five miles east of Luther, 19 July 1947, C. L. Hitchcock 16627 (NY, WT), Bridger Mountains, 28 July 1896, J. H. Floodman 651 (NY), Beartooth Mountains, 17 miles southwest of Red Lodge, 17 July 1939, Reed Rollins 2828 (GH, NY); Cascade County: King's Hill, 45 miles southeast of Great Falls, 24 July 1948, R. F. Daubenmire 48216 (WT); Deerlodge County: Storm Lake, Anaconda Mountains, 21July1946, c. L. Hitchcock 14812 (NY, WT), Mount Haggin, 20 July 1905, Marcus E. Jones s.n. {GH}; Fergus County: Central Little Snowy Mountains, 4 July 1945, c. L. Hitchcock 11941 (WT), Half Moon Canyon, Big Snowy Mountains, 3 July 149 1945, c. L. Hitchcock 11920 (NY, WT), near Giltedge, 27 June 1946, R. C. Barneby 8222 (NY), 31 miles south of

Lewiston, 1 August 1957, Gerald B. Ownbey 2422 (NY);

Flathead County: Sperry Glacier, 21 August 1901, F. K. Vreeland 1150 (GH, NY), Lake McDonald, 22 August 1901,

L. M. Umbach 349 (NY), Columbia Falls, 27 August 1894, R. S. Williams 92 (US); Gallatin County: Boseman, 23 July 1895, P.A. Rydberg 2720 (NY), Fairy Lake area, 4

July 1963, T. Mosquin 11856 (NY), Bridger Pass, 26 Aug- ust 1916, W. M. Suksdorf 139 (GH); Glacier County: 14 miles south of St. Mary, 9 July 1941, Susann Fry 510

(WT), Glacier National Park, east facing slopes of Di- vide Mountain, 14 August 1957, A. R. Kruckeberg 4351 (NY, WT), Junction of Highways u. s. 2 and 89, Blackfoot Indian Reservation, 2 July 1952, H. H. Bartlett and J.

F. Grayson 464 (US), Glacier National Park, 1.5 miles north of Logan Pass, 10 August 1957, A. R. Kruckeberg 4338 (NY),

Cutbank Creek, 15 July 1934, George N. Jones 5513 (GH);

Lewis and Clark County: 22 miles west of Augusta, 25 July

1952, H. Bartlett 1177 (NY), Trident Peak, 22 July 1948, c. L. Hitchcock 18774 (NY, WT), Mount Ascension, 26 June 1909, B. T. Butler 4045 (NY); Madison County: Lazy Man's Hill, 26 July 1947, c. L. Hitchcock 16883 (NY, WT}, Taylor Mountains, 2 August 1946, c. L. Hitchcock 15165 (NY, WT}; Meagher County: Western edge of Little Belt Mountains, 11

July 1945, C. L. Hitchcock 12221 (NY, WT), 35 miles north of White Sulfur Springs, c. L. Hitchcock 16172 (NH); Park 150 County: vicinity of Cooke City, 11 August 1948, Jean G. Witt 1383 (NY, WT), 1 mile east of Silver Gate, 28 July 1951, C. L. Porter 5788 (GH, NY), north slope of Baldy Mountain, Absaroka Range, 25 June 1912, w. W. Eggleston 8079 (GH); Pondera County: Marias Pass, 7 July 1948, c. L. Hitchcock 18152 (WT); Powell County: 13 miles west of Lincoln, 28 June 1945, C. L. Hitchcock, 11643a (NY, GH, WT), 5 miles northeast of Big Prairie Ranger Station, 21 July 1948, c. L. Hitchcock 18603 (WT); Stillwater County: 11.4 miles northeast of Mystic Lake, 5 July 1963, T. Mosquin 11868 (NY); Teton County: 5 miles north of Chateau, 22 June 1968, Stanley L. Welsh 7227 (NY, BRY). WASHINGTON. Okanogan County: east of Winthrop, 2 July 1934, J. W. Thompson 10913 (NY, GH, WT), 10 miles northwest of Concon- cully, 2 July 1958, c. L. Hitchcock 21679 (WT, NY), south of Salmon Meadows, 28 June 1931, c. B. Piker 166 (WT), south slopes of Mount Chapaca, August 1897, A. E. Elmer 551 (NY, US, WT), along road to Salmon Meadows, 26 June 1931, J. w. Thompson 7024 (GH); Stevens County: northeast of Leadpoint, Frisco Mine Road, 7 July 1933, George G. Hedgecock s.n. (WT). WYOMING. Park County: Beartooth Butte, 6 August 1937, Louis o. Williams 3767 (GH, NY); Sheridan County: west of Dayton, 1 July 1953, c. L. Porter 6279 (NY), Big Horn Mountains, July 1899, Frank Tweedy 2371 (NY), Little Tongue River Canyon, 25 June 1936, L. o. Williams 3112 (NY), 20 miles west of Dayton, 6 July 1935, Louis o. Williams 2364 (GH, WT). Yellowstone National 151 Park: Blacktail Deer Creek, 1 August 1888, F. H. Knowlton s.n. (US), Electric Peak, 19 August 1897, P.A. Rydberg 4507 (NY).

Hedysarum occidentale Greene, Pittonia 3:19. 1896. Fig. 50 {Olympic Mountains, Washington, c. V. Piper, 1890, (ND)) H. lancifolium Rydb. Mern. New York Bot. Gard. 1: 256. 1900. H. rnarginaturn Greene, Pittonia 4:138. 1900. H. uintahense A. Nels. Proc. Biol. Soc. Wash. 15: 186. 1902. Terrestrial herbaceous perennials; roots stout, ligneous, elongate; sterns several to many, decurnbent to erect, longitudinally grooved, not rooting, 2-7 drn. long, 2-8 mm. in diameter, branched above, solid, terete, pubescent; stipules brown, united, chartaceous, 10-17 mm. long, obtuse to acute or acurninate; leaves odd-pinnate, 8-12 cm. long; leaflets 11-19, ovate, elliptical or oblong, pubescent on both surfaces or rarely glabrous above, conspicuously veined, 9-34 mm. (40) long, 5-16 mm. wide, rounded to obtuse; inflorescence racernose, 4-13 cm. long, elongated; bracts brown, pubescent, 2-8 mm. long; bractioles pubescent, linear, 1-3.5 mm. long; flowers 10-50, pendent or erect, reddish purple, 16-23 mm. long; calyx carnpanulate, tube 3-9 mm. long, bractiolate, pubescent, teeth 0.5-2 mm. wide, 1-3.5 mm. long, unequal, subulate to triangular, greenish, erect; standard obovate to spatulate, erna~ginate, 13-16 mm. long, 6-7.5 mm. wide; wings 13-16 mm. long, 2-3 152

Figure 50 • Hedysarum occidentale Greene. 153 mm. wide; wing auricles linear, united, 3-4 mm. long; stamens 10, diadelphous; style slender, curved; loments pendulous, stipitate; articles 1-5, pubescent or rarely glabrous, conspicuously wing margined, 5.5-10 mm. wide, 8-17 mm. long, reticulations polygonal.

Ecology and distribution. Flowering from early June to early September. Talus slopes, stony outcrops, rocky and clay banks to alpine open slopes and moist grassy areas from 3,000 to 11,000 feet elevation. Vancouver Island, Olympic Mountains of Washington, rare in the Cascades, then disjunct to the mountains of northern Idaho and Montana, south to Wyoming, Utah and Colorado (Fig. 51). Representative specimens: CANADA. British Colum- bia: Vancouver Island, Crest Mountain on Gold River Road, J. A. Calder and K. T. MacKay 31595, 19 July 1961 (DAO), Golden Hinde above Burman Lake, J. A. Calder and K. T. MacKay 32506a, 15 August 1961 (DAO). COLORADO. Hindsdale

County: 6 miles northwest of Rio Grande R~servoir, near Pale Creek, 8 August 1936, Reed c. Rollins 1503 (GH); Rio Grande County: 5 miles southwest of Summitville, 3 July 1939, R. K. Gierisch 1040 (NY); Saguache County: La Garita Hills, 14 July 1939, F. Ramaley and w. Gambill 16949 (WT); San Juan County: 4 miles southwest of Silverton, 26 August 1946, K. F. Parker 6411 (US, NY), Molas Divide, 29 July 1963. J.M. Gillett 12241 (NY). IDAHO. Bonneville County: Caribou Mountain, 18 July 1923, Edwin B. Payson 3538 (GH); 154

120 110

120 110 Figure 51. Distribution of H. occidentale. 155 Franklin County: 5 miles northeast of Preston, 29 June 1943, J. H. Christ 14166 (WT); Idaho County: Lolo Trail near Fox Creek, 17 July 1939, C. W. Sharsmith 4062 (WT), Sec. 17, T 37 N, Rl2 E., Indian Post Office Lakes, 27 July 1950, Quen- tin Jones 294 (WT), Parker Meadows, 12 July 1946, c. L. Hitchcock 14661 (WT), Lolo Trail, 14 July 1937, Lincoln Constance and F. W. Pennell 2005 (GH, NY, WT); Shoshone County: southwest of Quarles Peak, 12 July 1941, C. B. Wilson 181 (NY, WT); Teton County: Hills southeast of Vic- tor, 28 July 1920, Edwin B. Payson 2167 (NY). MONTANA. Granite County: 2 miles west of Skalkaho road summit, 7 July 1946, c. L. Hitchcock 14480 (WT); Lake County: southeast of McDonald Lake, 12 July 1948, c. L. Hitchcock 18292 (WT, US, NY); Mineral County: Region of Nine Mile Divide, 3 September 1951, Marion Bacon 239 (WT); Missoula County: 2 miles east of Holland Lake, 15 July 1948, C. L. Hitch- cock 18358 (WT); Powell County: Shale Mountain, 21 July 1948, c. L. Hitchcock 18604 (NY, WT), Gordon Mountain, 22 July 1948, c. L. Hitchcock 18847 (NY, WT); Ravalli County: 19 miles south of Hamilton, 10 July 1965, C. L. Hitchcock 23957 (NY, WT), 4 miles east of St. Mary's Peak, 10 August 1946, C. L. Hitchcock 15294 (WT), trail to St. Mary's Peak, 5 August 1947, c. L. Hitchcock 17084 (WT), 24 miles east of Hamilton, 18 July 1935, George G. Hedgecock s.n. (WT), ridge between Laquina Lake and Fish Lake, 2 August 1957, A. R. Kruckeberg 4302 (WT). UTAH. Carbon County: Soldier Canyon, 11 August 1967, s. L. Welsh 6614 (NY, BRY); Emery County: 156 13 miles west of Orangeville, 17 June 1948, H. D. Ripley

9213 (US, NY); Summit County: foothills of Uintah Moun- tains near Mill Creek, 4 July 1926, E. B. Payson 4881 (GH). WASHINGTON. Chehalis County: Baldy Peak, 25 July 1897, Frank H. Lamb 1318 (NY, WT); Clallam County: slopes of

Mount Angeles, 15 July 1933, J. W. Thompson 9471 (NY, GH), Hurricane Ridge, 7 September 1937, J. w. Thompson 14176 (GH, NY, WT), Seven Lakes Basin, 24 August 1935, George N. Jones 8273 (WT), ridge trail between Mount Angeles and

Hurricane Ridge, 11 August 1951, A. R. Kruckeberg 2793

(WT), Olympic Mountains, June 1900, A. E. Elmer 2529 (WT), alpine slopes of Mount Angeles, 15 July 1932, J. W. Thomp-

~ 8765 (GH), 4.5 miles east of Sal Due Hot Springs, 9

August 1937, F. G. Meyer 1024 (GH, WT), east of Obstruc- tion Point Ridge, 5 September 1937, F. G. Meyer 1273 (GH),

Bogachiel Peak, 4 August 1924, I. G. Otis 1344 (WT), Deer Park Recreation area, 8 August 1938, R. c. Rollins 2693 (NY, GH); Grays Harbor County: Mount Colonel Bob, 23 August 1933, J. w. Thompson 9968 (NY), Mount Colonel Bob Lookout, 9 July 1931, J. w. Thompson 7293 (GH), Mount Colonel Bob, 29 July 1965, c. L. Hitchcock 24103 (NY, WT); Jefferson County: above Heart Lake, 30 June 1938, G. B. and R. P. Rossbach 450 (GH); Mason County: ridges of high

Olympics, 16 July 1892 , L. F. Henderson 1850 (WT) I Mount

Gladys, 16 August 1939, F. G. Meyer 1697 (GH, WT), Mount

Steele, 14 August 1937, F. G. Meyer 1154 (GH, WT), Mount Ellinor, 15 August 1937, W. J. Eijerdam 1280 (WT). WYOMING. 157

Fremont County: 20 miles northwest of Dubois, 8 August 1958, u. T. Waterfall 15010 (US), Lava Mountain, 26 August 1965, Richard w. Scott 697 (GH}, 20 miles west of Dubois on u. s. 282, 21 July 1946, c. L. Porter 4052 (GH), 30 miles southwest of Dubois, 10 July 1958, C. L. Porter 7518 (NY), 2 miles northwest of Brooks Lake, 8 July 1966, Richard W. Scott 748 (NY); Johnson County: Big Horn Range west of Buffalo Circle Park, 23 July 1958, C. L. Porter 7559 (NY); Lincoln County: Moose Flats, 4 August 1949, c. L. Porter 5201 {GH, NY), Mount Wagner, 5 August 1923, Edwin B. Payson 3749 {GH), Sheep Mountain, 11July1923, Edwin B. Payson 3448 (GH); Sheridan County: west of Big Horn, 9 June 1934, Reed c. Rollins 503 {GH, NY); Sublette County: Green River Lakes, 27 July 1950, C. L. Porter 5549 (NY, GH, US),Sheep Mountain, 1 August 1925, Edwin B. Payson 4518 (GH, WT), near Middle Piney Lake, 17 July 1949, C. L. Porter 5015 {GH, NY), 20 miles west of Big Piney, 9 July 1922, Edwin B. Payson 2621 (GH); Teton County: Head of West Crystal Creek, 17 July 1955, H. K. Boechner 78 {WT), Teton Canyon, 30 July 1956, Loran c. Anderson 573 (NY); Uintah County: La Barge, 20 June 1894, E. Stevenson 29 {US), ranch south of Evanston, L. H. Panunel and R. E. Blackwood 4054 (GH), Evanston, 10 July 1897, T. A. Williams s.n. {NY), Evanston, 27 July 1897, Aven Nelson 3839 (NY), near Big Muddy Creek, 14 June 1938, Reed c. Rollins 2323 (NY, GH). 158

l I

Figure 52. Type specimen of H. boreale ssp. boreale 159

,) I / .r:,

/,; ,; .• ,f) :Io •

// ·'' ...... /... ~ - f ' ,,,,,f ...... ,.~ -' -

Figure 53. Type specimen of H. boreale ssp. mackenziei. 160

INDEX TO EXSICCATAE

1. Hedysarum alpinum a. var. al,einum b. var. Ehiloscia c. var. grandiflorum

2. Hed~sarum sul,ehurescens

3. Hed~sarum occidentale 4. Hedysarum bore ale a. ssp. bore ale var. bore ale b. ssp. bore ale var. rivulare c. ssp. boreale var. gremiale d. ssp. mackenziei The number following the collector's name refers to his field book number. The number in parenthesis refers to the herbarium in which the specimen is deposited. The last number refers to the taxon. 161

INDEX TO COLLECTORS

Albertson, A. 41 (USFS) 4b.

Anderson, J. P. 8749 (GH) la; 7166 (GH) la; 9282 (GH) la.

Anderson, J. R. sn (WT) 2. Anderson, L. c. 573 (NY) 3. Anderson, w. B. sn (WT) 2. Andorfer, J. E. 14225 (COLO) 4b.

Atwood, N. D. 914 (BRY) 4b.

Bacon, M. 239 (WT) 3. Baldwin, w. 3956 (CAN) la; 3777 (CAN) la; 3317 (CAN) la; 5326 (CAN) la. Bamberg, s. sn (COLO) 4a. Bare, J. E. 1053 (NOA) 4a.

Barber, M. A. 262 (GH) la. Barneby, R. c. 8222 (NY) 2; 8027 (NY) 2. Barnum, A. H. 1294 (BRY) 4b. Barr, c. A. 973 (NOA) 4b. Bartlett, H. 1177 (NY) 2.

Bartlett, H. and J. F. Grayson 464 (US) 2.

Bastock, H. 91 (CAN) la.

Beaman, J. H. 601 (US) la.

Beck, D. E. sn (BRY) 4b.

Bethel, E. 4170 (MONTU, UTC, COLO, PH) 4b. 162 Blais, W. 979 (CAN) 2. Blankinship, J. W. sn (MONT) 4a. Bleak, A. 56 (UT) 4b. Beechner, H. K. 78 (WT) 3. Bolinger, D. sn (OSC) 4d. Bonde, E. K. 283 (COLO) 4d. Booth, W. E. sn (MONT) 4a; sn (MONT) 4a; sn (ARIZ) 4a; 552205 (MONT) 4a; 55314 (MONTU) 4a; 55360 (MONT) 4a; sn (MONT) 4a; 55222 (MONT) 4b; sn (MONT) 4b; sn (MONT) 4b. Boivin, B. 12528 (NY) la; 9504 (NY) 2; 9502 (OKL) 4a; 8796 (NDA) 4a. Bragonje, R. J. 27 (BRY) 4b. Breitung, A. J. 17054 (NY) la; sn (GH) lb; 15664 (NY) 2; 16486 (COLO) 4a; 5054 (UTC) 4a. Breivster, G. sn (COLO) 4b. Brinkman, A. H. 721 (GH) la. Brizzee, K. sn (UT) 4b. Brotherson, J. 489 (BRY) 4b; 565 (BRY) 4b; 663 (BRY) 4b; 17 (BRY) 4b; 787 (BRY) 4c. Brown, S. 75 (PH) 4d; 401 (PH) 4d; 538 (PH) 4d; 909 (PH} 4d; 410 (GH) la; 1218 (GH, NY) le; 672 (GH} lb; 425 (GH) 2; 127 (GH) 2. Brues, A. sn (OKL} 4a. Bryan and Deming 6-10 (BRY) 4b. Buchanon, H. 167 (UT} ·4b. Burke, M. 3012 (UTC) 4b. 163 Butler, 4045 (NY) 2. Butler, c. A. D3-12 (USFS) 4a. Butler, s. R. sn (GH) le. Calder, J. A. 11290 (NY, WT) 2; 14022 (OSC) 4d; 5808 (WS) la; 13989 (WS) la; 27952 (WS) la. Calder, J. A. and K. T. MacKay, 31595 (DAO) 3; 32506A (DAO) 3; Call, W. 33 (UT) 4b. Cazier, s. E. L-124 (USFS) Chapman, R. M. 84 (US) la. Charette, L. A. 2526 (BRY) la. Chickering, J. W. sn (US) la. Christ, J. H. 14166 (WT) 3. Christensen, E. M. sn (BRY) 4b. Clark, c. 493 (CAN) la; 259 (CAN) le. Cody, w. J. 6318 (WS) la; 7797 (WS) la; 8466 (ARIZ) 4d. Collotzi, A. w. 503 (UTC) 4b. Constance, L. and F. W. Pennell 2005 (GH, NY, WT) 3. Cope, T. M. sn (PH) 4d. Cottam, w. P. 2046 (BRY) 4b; 5275 (UT) 4b; 2758 (BRY) 4b; 2212 (BRY) 4b; 12143 (UT) 4b; 9667 (UT) 4b. Craddock, G. GWC-20 (USFS) 4b. Cronquist, A. 7967 (NY, WT) 2; 3125 (UTC) 4b; 533-37 (UTC) 4b; 9222 {UTC) 4b. Crooke, K. 318 (OKL) 4d. Cusick, W. C. 3694 (OSC) 4d. Daubenmire, R. F. 48391 (WS) la; 48216 (WT) 2. 164

Davis, R. J. 5052a (BRY) la; 5082 (BRY) 4d. Deaver, c. F. sn (ARIZ) 4b. Denton, G. Sn (MONT) 4b.

DeRuyck, G. sn (~Y) la. Dickson, B. 250 (USFS) 4a. Dickinson, W. S. 557 (CAN) la. Diehl, I. E. sn (BRY) 4b. Douglass, M. 54-219 (COLO) 4b. Dutilly, Rev. A. 14627 (GH) la. Dumois, M. G. 3493 (CAN) la. Eastwood, A. 306 (GH) la; 502 (US) la. Eggleston, w. w. sn (US) la; 8079 (GH) 2. Ehlers, J. H. 8052 (ARIZ) 4b. Eijerdam, W. J. 1280 (WT) 3. Elmer, A. E. 551 (NY, US, WT) 2; 2529 (WT) 3. Engleman, J. sn (OKL) 4b; 1596 (OKL) 4b. Erdman, J. A. 691 (COLO) 4b. Ewan, J. A. 12966 (COLO) 4b. Fairbourn, M. L. 3 (BRY) 4b. Fernald, M. L. and K. M. Wiegand 5799 (GH) la. Fernald, M. L. 26 (US) la; sn (GH) la; sn (GH) la; 5800 (GH) la; 5801 (CAN) la; 6450 (GH) la; 25171 (GH) la; 5076 (GH) la; 1833 (GH) le; 1832 (GH) le; 3636 (GH) le; 10849 (GH) le; 28621 (GH) le. Fernald, M. L., Wiegand K. and B. Long 28625 (GH) le. Fernald, M. L. and E. F. Williams sn (GH, US) la. Fernald, M. L., J. Collins and A. s. Pease sn (GH) la. 165

Fiker, C. B. 166 (WT) 2. Findlay, W. 216 (NY) la. Floodman, J. H. 651 (NY) la. Foster, R. H. 408 (BRY) 4b. Fry, S. 564 (WS) lb; 510 (WT) 2.

Garrett, A. o. 7020 (U~) 4b. Garton, c. E. 7748 (WS) la. Geist, o. w. sn (BRY) la. Gierisch, R. K. 1040 (NY) 3. Gillett, J. M. 12241 (NY) 3; 3453 (COLO) 4d. Gleason, sn (WT) 3. Gooding, L. N. Gl43-40 (OSC) 4b; 332 (MONT, UTC, ARIZ) 4b. Goodman, G. J. 2807 (MONT) 4b. Gorman, M. W. 1018 (US) la. Graham, E. H. 9044 (NDA) 4b. Grant, J. sn (BRY) la. Guthrie, R. J. 7 (USFS) 4b. Halliday, w. 68-1932 (CAN) la. Harrington, G. L. 54 (US) la. Harrison, B. F. 8790 (BRY) 4b. Harvey, L. H. 5842 (MONTU) la; 5224 (MONTU) la; 6072 (MONTU) la. Hawkins, P. H. sn (MONT) 4a; 384 (MONT) 4a. Hay, N. G. ll83a (CAN) la. Head, C. 127 (BRY) 4d. Head, s. c. 1045 (OSC) 4d. Hedgecock, G. sn (WT) 2; sn (WT) 3. 166 Heidenwish, v. J. sn (NDA) 4d. Heller, A. A. 3732 (UT) 4b. Henderson, L. F. 1850 {WT) 3. Hess, w. 723 (OKL) 4b. Bevly, R. H. sn {UT, ARIZ) 4b. Hitchcock, c. L. 24032 (WS) la; 7778 {WT) 2; 12740 (GB, NY, WT) 2; 13549 (NY, WT) 2; 12922 (NY, WT) 2; 13142

(NY, WT) 2; 16627 (NY, WT) 2; 14812 (NY, WT) 2;

11941 (WT) 2; 11920 (NY, WT) 2; 18774 (NY, WT) 2; 16883 (NY,WT) 2; 15165 (NY, WT) 2; 12221 (NY, WT) 2; 16172 (NY) 2; 18152 (WT) 2; 21679 (NY, WT) 2; 11643 (NY, GB, WT) 2; 18603 (WT) 2; 18684 (NY, WT)

2; 14661 (WT) 3; 14480 (WT) 3; 18292 (WT, US, NY) 3; 18358 {WT) 3; 18604 (NY, WT) 3; 18847 (NY, WT) 3; 16721 (COLO, UTC) 4a; 18109 {COLO) 4a; 12182 (PH, MONT, UTC, OSC) 4a; 11929 (UTC, PH, COLO) 4a; 23957 (NY, WT) 3; 15294 (WT) 3; 17084 {WT) 3; 24103 {NY, WT) 3; 11109 (UTC) 4b; 18074 (OTC, COLO) 4b; 8989 (UTC, MONT) 4a; 10794 (UTC) 4a; 11824 (PH, OTC) 4a. Hodder, R. L. (MONT) 4a. Hogan, L. 42 (BRY) 4b. Holmgren, N. H. 108 (UTC) 4b; 1819 (OTC, BRY) 4b. Hosie, R. c. 1586 {GB) la. Hulten, E. sn (US) la. Hunnewell, F. w. 4449 (GB) la. Hustich, I. 1459 (CAN) la; 328 (CAN) la. 167 Isely, D. 8900 (BRY) 4b. Johnson, P. 245 (GH) lb. Jones, G. N. 5438 (NY) 2; 5513 (GH) 2; 8273 (WT) 3. Jones, M. E. 8221 (US) 2; sn (GH) 2; sn (GH) 2. Jones, Q. 637 (WS) la; 294 (WT) 3. Kelsall, J. and E. McEwen 187 (CAN) ia. Kelsey, s. L. 97 (GH) la. Kirch, J. 107 (MONT) 4a. Kirkwood, J. 2238 (NY) 2. Knowlton, c. H. sn (GH) la. Knowlton, F. H. sn (US) 2. Kruckeberg, A. R. 4351 (NY, WT) 2; 4351 (NY, WT) 2; 4338

(NY') 2; 4302 (WT) 3; 2793 (WT) 3. Lamb, F. H. 1318 (NY', WT) 3. Legault, A. 6789 (CAN) la. Lepage, E. 4208 (GH) la. Levy, J. L. S-11 (USFS) 4b. Lewis, w. H. 479 (COLO, OKL) 4d. Lindsay, A. 163 (CAN) la; 602 (CAN) la; 527 (CAN) la. Lindsay, G. 2330 (GH) la.

Louis-marie, P. 20871 (PH) 4a. Love, A. 6191 (GH) lb.

Lowther, J. 66 (CAN) la. Lynch, D. 6095 (WS) la; 6256 (WT) 2. Mackenzie, K. K. 3536 (US) la. Macoun, J. 70784 (GH) la; 10165 (CAN) la; 12506 (CAN) la; 591 (GH) 2. 168 Maguire, B. 18515 (UTC, BRY) 4b; sn (UTC) 4a; 21804 (UTC) 4b. Malte, M. sn (CAN) la; 1296 (CAN) la; 771 (CAN) la. Marsh, V. L. 739 (MONT) 4a. Mason, G. 6461 (OSC) 4d. McDonald, c. H. 837 (USFS) 4b; 625 (USFS) 4a. McEwen, E. H. 257 (CAN) le. Mcleod, M. 377 (COLO) 4b. Meissuer, Mrs. M. sn (NDA) 4a. Mexia, Y. 2029 (US) la. Metcalf, H. N. sn (MONT) 4b. Meyer, F. G. 1024 (GH) 3; 1273 (GH) 3; 1697 (GH, WT) 3; 1154 (GH, WT) 3. Ming, B. sn (MONT) 4a.

Miron, F. sn (CAN) la.

Moir, D. 633 (CAN) la; 994 (CAN) la; 222 (CAN) la.

Moir, D. R. sn (NOA) 4a. Moodie, M. E. 948 (US, GH) la; sn (US) lb; 948 (US) lb; 1261 (US) 2; 1020 (OSC) 4a. Moran, E. C. sn (BRY) 4b; sn (BRY) 4b; sn (BRY) 4b; sn (BRY) 4a. Morton, c. V. and E. Lepage 11626 (US) lb. Mosquin, T. 5186 (NY) 2; 11856 (NY) 2; 11868 (NY) 2. Moss, E. H. 1027 (GH) la; 6138 (GH) la; 186 (GH) lb; 763 (NY) 2 .. Nagy, J. 2612 (CAN) 2; 1175 (CAN) 2. Nelson, A. 3594 {WS) la; 3759 (WS) la; 3367 (US) lb; 8955 169 (GH, US) lb; 9436 (GH, US) lb; 3129 {GH) 2; 3839

{NY) 3. Nelson, E. 622 (NY) lb. Norman, G. W. 53 (CAN) la. Osterhout, G. E. sn (COLO) 4b.

Palmer, E. 37554 (GH) lb; 37509 (GH) lb. Pammel, L. H. and R. E. Blackwood 4054 (GH) 3. Parker, K. F. 6411 (US, NY) 3.

Payson, E. B. 3538 (GH) 3: 2167 (NY) 3; 4881 (GH) 3; 3749 (GH) 3: 3448 (GH) 3; 4518 (GH, WT) 3; 2621 {GH) 3; 2617 (PH) 4a. Payne, G. F. 36 (US) la. Pease, A. s. sn (GH) .la: 2262 (GH) la; 38809 (GH) la; 36623 (GH) la; 28929 (GH) la. Peebles, R. H. 14359 (ARIZ) 4b. Pegau, R. 239 (BRY) le. Pennell, 22497 (PH) 4c; 22144 (PH) 4b. Pierce, J. H. 7 (GH) la. Porsild, A. E. 677 (GH, CAN) la: 2001 {CAN) la; 5834 {CAN) la; 10390 (CAN) la; 4955 (CAN) la; 17316 (CAN) le; 17588 (CAN) le. Porter, c. L. and R. C. Rollins 5673 (PH) 4a; 5442 (PH, OKL) 4a. Porter, c. L. 3742 (GH) lb; 5788 (GH, NY) 2; 6279 (NY) 2; 4052 (GH) 3; 7518 (NY) 3; 7559 (NY) 3; 5015 (GH, NY) 3; 5201 (GH, NY) 3; 5549 (NY, GH, US) 3; 4584 (COLO) 4b; 5780 (COLO) 4b; 4506 (COLO) 4a. 170 Poto, ·w. 48 (US) la. Plummer, A. P. 304 (USFS) 4b. Price, H. 168 (USFS) 4a. Prince, F. c. sn (GH) lb. Pringle, c. sn (GH, US) la. Proctor, G. R. 2196 (GH) la; 2087 (GH) la. Ramaley, F. and W. Gambill 16949 (WT) 3. Raup, H. M. 7057 {GH} la; 10687 {CAN) la; 11136 (CAN) la; 2803 (CAN) la; 2804 {CAN) la; 997 {GH) la; 1000 {GH) la; 2807 {GH) la; 998 {GH) la; 9423 (WT) la; 10252 (CAN) la; 3626 (GH) la; 3967 (GH) le; 4091 (GH) le; 9084 (GH) le; 12128 (GH) le. Reed, K. M. 691 {BRY) la. Rethke, R. V. 3894 {UTC) 4b. Rice, B. F. 351 (USFS) 4b. Rigby, K. 175 {BRY) la; 192 (BRY) la; 128 (BRY) la. Riley, J. A. 36 {GH) le. Ripley, H. o. 9213 (US, NY) 3. Robertson, J. H. 300 {USFS) 4b.

Rogers, c. M. 5831 {COLO) 4b; 5991 {OKL) 4b. Rogers, T. H. 1098 (NY, WT) 2. Rollins, R. C. 2828 (GH, NY) 2; 1733 {PH, OKL) 4c; 1503 (GH) 3; 1864 {UTC, GH, MONT, USFS) 4b; 2875 {USFS) 4b; 2693 {NY, GH) 3; 503 {GH, NY) 3; 2323 {GH, NY)

3. Rosencrom, R. 39 {USFS) 4a. Rossbach, G. B. and P. R. Rossbach 450 {GH) 3. 171 Rossbach, G. B. 6892 (CAN) la. Rousseau, J. 32149 (CAN) la; 117 (CAN) la. Rydberg, P. A. 640 (US) lb; 2238 (GH) 2; 2720 (NY) 2; 4507 (NY) 2. Ryerson, D. 599 (MONT) 4a. Sandberg, J. H. 748 {NY, GH, US) 2. Scamman, E. 2117 (GH) la; 3295 (GH) la; 2361 (GH) la; 3150 (GH) la; 3295 (GH) la; 6798 (GH) la; 304 (GH) la; 302 (GH) la; 5743 {GH) la; 305 (GH) la; 5333 (GH) la; 1878 (GH) la; 3725 (GH) la; 5566 (GH) le; 3886 (GH) le; 2913 {GH) lb; 6669 (CAN) 2. Scharff, D. K. sn (MONT) 4a. Schofield, W. B. 8046 (CAN) la; 128 (WT) la; 1099 (COLO) la. Schuber, E. sn (MONT) 4a. Schulze, N. C. 71 (USFS) 4b. Scoggan, H. J. 13778 (CAN) la; 2066 (CAN) la; 2064 (CAN) la; 10170 (CAN) la; 10846 (CAN) la; 11239 (CAN) la; 10319 (CAN) la; 10194 (CAN) la; 8071 {CAN) la; 3881 (CAN) la; 4400 (GH) lb; 11239 (GH) lb; 16220 (CAN) 2. Scott, R. w. 697 {GH) 3; 740 (NY) 3. Shacklette, H. T. 2845 (CAN) la; 2454 (CAN) la. Sharsmith 4062 (WT) 3. Shaw, C. H. 591 (NY) 2. Shaw, K. 156 (BRY) la; 186 (BRY) la. Short, L. R. s-521 (USFS) 4a. Shreve, F. 8968 (ARIZ, UT, COLO) 4b. Sieburth, L. R. sn (WS) la. 172 Silburth, L. sn (WT) 2. Single, D. B. sn (MONTU) 4a.

Spear 5360 (BRY) la.

Spreadborough 5378 (CAN) la; 19314 (CAN) la. St. John, H. 2362 (US) la; 297 (US)· la. St. John, Mrs. O. 147 (UTC) 4b.

Stephens, K. 51 (US) la. Stevens, o. A. 11 (MONT) 4b; sn (NDA) 4b; sn (NDA) 4b; sn (NDA) 4b; sn (NDA) 4b; sn (NDA) 4a; 38-011 (NDA) 4a; sn (NDA) 4a; sn (NDA) 4a; 906 (NDA) 4a; 696 (NDA) 4a; sn (NDA) 4a; sn (NDA) 4a; sn (NDA) 4a. Stevenson, E. 29 (US) 3. Stewart, J. 20 (USFS) 4a. Stewart, J. O. C-71 (USFS) 4b. Streets, R. B. 441 (MONT) 4a. SUksdorf, w. M. 139 {GH) 2. Swanson, G. 174 (UT) 4b. Taylor, R. 2268 (WS) la. Taylor, T. 682 (GH) la; 680 (CAN) la; 683 (CAN) la.

Tener, J. 241 (CAN) la. Thompson, J. W. 10913 (NY, GH, WT) 2; 7024 (GH) 2; 195 (PH)

4b; 9471 (NY, GH) 3; 14176 (GH 1 WT, NY) 3; 8765 (GH) 3; 9969 (NY) 3; 7293 (GH) 3. Thornton, c. w. 414 (US) lo. Traphagen, F. w. sn (MONT) 4a. Turner, G. H. 2253 (GH) la; 2129 (GH) lb. Tweedy, F. 2371 (NY) 2. 173 Ulke, T. 175 (MONTU) 4b. Umbach, L. M. 349 (NY) 2; 378 (MONT) 4a. Vansell, G. H. sn (OTC) 4b. Victorin, F. M. and R. Germain 27332 (GH) la; 27354 (GH) la; 27156 (GH) la. Victorin, M. 16082 (US) la; sn (US) la. Viereck. 8029 (USFS) 4d; 4365 (COLO) 4d. Vrieland, F. K. 1150 (GH, NY) 2; 849 (MONTU) 4b. Waghorne,· A. c. sn (GH) le. Walkup, G. w. 14 (USFS) 4a. Warn, F. B. 44 (OTC) 4a. Waterfall, u. T. 15010 (US) 3. Weber, w. A. 2429 (COLO) 4d; 2240 (NY, WT) 2. Welsh, s. L. 4842 (BRY) la; 10277 (BRY) la; 10116 (BRY) la; 7366 (BRY) la; 5430 (BRY) la; 5438 (BRY) la; 5496

(BRY) la; 6028 (BRY) la; 5088 (BRY) la; 5663 (BR~) la; 4325 (BRY) la; 4899 (BRY) la; 4244 (BRY) 4d; 4834 (BRY) 4d; 5949 (BRY) 4d; 5447 (BRY) 4d; 7340 '(BRY) 4d; 5560 (BRY) 4d; 4056 (BRY) 4d; 7839 (BRY) 4d; 5863 (BRY) le; 4107 (BRY) le; 10520 (BRY) le; 10613 (BRY) le; 755 (BRY) lb; 7227 (BRY) 2; 4226 (COLO) 4b; 11306 (COLO) 4b; 2082 (BRY) 4b; 3029 (BRY) 4b; 7135 (BRY) 4b; 6221 (BRY) 4b; 1903 (BRY) 4b; 1728 (BRY) 4b; 1564 (BRY) 4b; 6614 (BRY, NY} 3. Whitham, J. C. 1811 (USFS) 4b. Wiegand, K. M. 28618 (GH) le. Williams, L. O. 3767 {GH, NY) 2; 3112 {NY) 2; 2364 (GH, WT) 2. 174 Williams, M. 2614 (BRY) le. Williams, R. s. 92 (US) 2; 91 (MONT) 4a. Williams, T. A. sn (NY) 3. Wilson, c. B. 181 (NY, WT) 3. Witt, J. G. 1284 (COLO) 4b; 1383 (NY, WT) 2; 1326 (NY, WT) 2. Wood, G. w. 1266 (GH) la. Wood, w. D. 90 (CAN) la. Wright, J. c. sn (MONT) 4a. Young, c. H. sn (CAN) la. Youngman 115 (CAN) la. 175

BIBLIOGRAPHY

Anderson, J. P. 1959. Flora of Alaska. Ames: Iowa State University Press.

Archibald, R~ M. 1946. Colorimetric determination of canavanine. J. Biol. Chem. 165:169-178.

Archibald, R. M. and P. B. Hamilton. 1943. Removal of canavanine from preparations of Jack Bean urease. J. Biol. Chem. 150:155. Baldwin, w. K. w. 1958. Plants of the clay belt of northern Ontario and Quebec. Nat. Mus. Can. Bull. 156.

Ballard, E. G. 1957. Translocation of organic nitrogen in the xylem. Aust. J. Biol. Sci. 10:292. Basson, w. D. and R. G. Bohmer. 1972. An automated procedure for the determination of sulphur. Analyst 97: 266-270. Bell, E. A. 1958. Canavanine and related compounds in Leguminosae. Biochem. J. 70:617.

Bell, E. A. 1971. "Comparative biochemistry of non-protein amino acids." In Chemotaxonomy of the Leguminosae, 179-206, Edited by J. B. Harborne, New York: Aca- demic Press.

Bell, E. A. and L. Fowden. 1964. Taxonomic Biochemistry and serology. New York: Ronald Press.

Benson, L. 1962. Plant Taxonomy. New York: Ronald Press. Bird, R. D. 1927. A preliminary ecological survey of the district surrounding the entomological station at Treesbank, Manitoba. Ecology 8:207-220.

Birdsong, B. A.; Alston, R.; and Turner, B. 1960. Distri- bution of canavanine in the family Leguminosae as related to phyletic groupings. Can. J. Bot. 38: 499-505. Boivin, B. 1967. Flora of the prairie provinces. Part 1. Phytologia 15:329-446. 176 Bowen, J. M. 1967. Comparative elemental analyses of a standard plant material. Analyst 92:124-131.

Boyd, W. C. 1947. Fundamentals of Immunology. 2nd ed. New York: Interscience. Boyd, W. C. and R. M. Reguera. 1949. Hemagglutinating substance for human cells in various plants. J. Immunology 62:333-339.

Boyd, W. C. and E. Sharpleigh. 1954. Specific Precipitat- ing activity of plant agglutinins (lectins). Sci- ence 119:419. Boyd, w. C.; Wasjczenko-Zacharczenko, E.; ands. M. Gold- wasser. 1961. List of plants tested for hemaglutinating activity. Transfusion 1:183. Britton, N. L. and A. Brown. 1913. An illustrated flora of the northern United States, Canada and the British possessions. Vol. 2. New York.

Brunel, A. and R. Echevin. 1938. Glyoxylic ureides in the flower and fruit of Acer ~seudo-platahus L. Compt. Rend. Acad. Sci. (Paris) 07:592-594.

Budd, A. C. 1957. The wild plants of the Canadian prairies. Can. Dept. Ag. Publ. 983. Burman, w. A. 1910. The flora of Manitoba. Handbook to Winnipeg and the province of Manitoba. Cadden, J. F. 1940. Spatial configuration and preparation of canavanine. Proc. Soc. Exp. Biol. Med. 45:224- 226. Candolle, A. P. de. 1825. Prodromus Systematis Naturalis Regni Vegetabilis. Vol. 2. Parisiis:treutel.

Clark, R. B. 1970. Effects of mineral nutrient levels on the inorganic composition and growth of corn. Ohio Agr. Res. Dev. Center Res. Circ. 181. Cody, w. J. 1960. Plants of the vicinity of Norman Wells Mackenzie District, Northwest Territories. Canad. Field-Nat. 74:71-100. 1965. Plants of the Mackenzie River Delta and ----reindeer grazing preserve. Ottawa: Canada Depart- ment of Agriculture.

Colinvaux, P. A. 1963. "Quaternary vegetational history of arctic Alaska." In the Bering Land Bridge, 206-261, 177

Edited by D. Hopkins, Stanford: Stanford Univer- sity Press. Cooper, W. s. 1959. Vegetation of the Prince William Sound Region, Alaska. Ecological Monographs 12:1-22.

Coulter, J. M. 1885. Manual of the Botany of the Rocky Mountain Region. American Book Co.

Criddle, N. 1927. A calendar of flowers. Can. Field-Nat. 41:48-55. Cromwell, B. T. 1955. "The Alkaloids," in Moderne Methoden der Pflanzenanalyse, band 4. Edited by Paech, K., and M. v. Tracey Ed., Berlin: Springer-Verlag. Davis, G. L. 1966. Systematic Embryology of the Angiosperms. New York: John Wiley.

Davis, R. J. 1952. Flora of Idaho. Provo: Brigham Young University Press.

Damodaran, M. and K. G. A. Narayanan. 1939. The Prepara- tion of Canavanine from Canavalia obtusifolia. Biochem. J. 33:1740.

1940. A Comparative Study of Arginase and Canava- nase. Biochem. J. 34:1449. Dole, E. J. 1937. The Flora of Vermont. Burlington: Free Press Printing Company.

Dubnoff, J. W. and H. Borsook. 1941. A Micromethod for the Determination of Glycocyamine in Biological Fluids and Tissue Extracts. J. Biol. Chem. 138:381.

Eisler, M. and L. Portheim. 1926. Agglutinins in Plants. Arz. Akad. Wiss. Wien. 63:35~36. Erdtman, G. 1966. Pollen Morphology and Plant Taxonomy. Vl. Angiosperms. New York: Hafner Publishing Co. Fearon, W. R. 1946. The Amidine-Pentacyanoferrate Reac- tion: Test for Guanidines, Urea and Thiourea. Analyst 71:562. Fedtschenko, B. 1902. A Revision of the Genus Hedysarum. Acta Hort. Petrop. 19:185-325.

Fosse, R. 1926. Formation by Heating Plant Juice of Urea and of a Substance Giving the Same Hydrazinic Color Reaction as Formaldehyde. Compt. Rend. Acad. Sci. (Paris) 182:175-177. 178 Fosse, R.; P. De Graeve; and P. E. Thomas. 1932. A New Plant Principle: Uric Acid. Compt. Rend. Acad. Sci. (Paris) 195:1198. 1933. The Role of Allantoic Acid in Higher Plants. ----Compt. Rend. Acad. Sci. (Paris) 196:883-886. Frink, c. R. and D. E. Peaslee. 1968. Spectrophotometric Determination of Micro Amounts of Aluminum in Plant Material with 8-Hydroxyquinoline. Analyst 93:469- 474. Fryzell, P. A. 1957. Mode of Reproduction in Higher Plants. Botan. Rev. 23:135-233. Gardner, G. 1937. Liste annotee des especes de pteriodotes, de phanerogames et d'algues recoltees sur la cote du Labrador, a la Baie a'Hudson et dans le Manitoba Nord, en 1930 et 1933. Bull. Soc. Fr. 84:1-33. Gillett, John M. 1967. Hedysarum occidentale Greene (Leg- uminosae), New to Canada. Can. Field-Nat. 81:2"!4:' Giterman, R. E. and L. V. Golubeva. 1963. "Vegetation of Eastern Siberia during the Arthropogene Period." In the Bering Land Bridge, pp. 232-244, Edited by D. Hopkins. Stanford: Stanford University Press. Gleason, H. A. 1952. The New Illustrated Flora of the Northeastern United States and Adjacent Canada. Vol. 2. Lancaster, Pa. Gleason, H. A. and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. New York: D. Van Nostrand and Co. Glyzin, V. I.; Bankovskii, A. I.; and N. s. Pavolova. 1970. Phenolic compounds of Hedysarum sachalinense and Hed,sarum brandtii. Khim. Prir. Soedin (Tashkent). 6:3 3-374. Gorsline, c. W.; Baker, D. E.; and w. I. Thomas. 1965. Accumulation of Eleven Elements by Field Corn. Penn. State Univ. Agr. Exp. Sta. Bull. 725. 1968. Major Gene Inheritance of Sr-Ca, Mg, K, P, ----Zn, Cu, B, Al-Fe, and Mn Concentration in Corn. Penn. State Univ. Agr. Exp. Sta. Bull. 746. Gray, A. 1857. Textbook of Western Botany. New York. Gray Herbarium of Harvard University. 1885-. Gray Her- barium Index. Vol. 5. Boston Mass. G. K. Hall Co. 179 Gray, M. P. 1961. Preparation of a stable bromelin solution. Transfusion 1:183. Graham, E. H. 1937. Botanical Studies in the Uinta Basin of Utah and Colorado. Ann. Carnegie Mus. 26. Graustein, J. E. 1967. Thomas Nuttall Naturalist. Cam- bridge: Harvard University Press. Hadoc, E. 1963. "On the History and Age of some Arctic Plant Species." in North Atlantic Biota and their History. pp. 207-219. Edited by A. and D. Love. New York: MacMillan Co.

Halliday, W. E. D. 1935. Vegetation and Site Studies, Clear Lake, Riding Mountain National Park, Mani- toba. Forest Service, Dept. of the Interior, Research Note 42.

Harborne, J.B.; Boulter, D.; and B. L. Turner, ed. 1971. New York: Academic Press.

Harrington, H. D. 1954. Manual of the Plants of Colorado. Denver: Sage Books. Hedberg, o. 1967. Chromosome Numbers of Vascular Plants from Arctic and Subarctic North America. Ark. F. Bot. Ser •. 2. 6:309-326. Hegi, G. 1964. Illustrated Flora von Mittel-Eur9pe. Band 4. part 3. pp. 1484-1487. Munchen: Carl Hansen. Heller, A. A. 1900. Catalogue of North American Plants North of Mexico. Henry, J. K. 1915. Flora of Southern British Columbia and Vancouver Island. Toronto. Henry, R. J. 1964. Clinical Chemistry. New York: Harper and Row. Hitchcock, c. L.; Thompson, J. W.; Ownbey, M.; and A. Cronquist. 1961. Vascular Plants of the Pacific Northwest, part 3, pp. 275-278. Seattle: Univer- sity of Washington Press. Holmen, K. 1962. Chromosome Studies in some Arctic Alaskan Leguminosae. Bot. Not. 115:87-92. Holmes, P. 1953. The Amino Acid Composition of Certain Seed Proteins. Aust. J. Exp. Biol. Med. Sci. 31: 595. 180 Hooker, J. D. and B. D. Jackson. 1885-. Index Kewensis: An Enumeration of the Genera and Species of Flower- ing Plants. London: Clarendon Press. Hooker, w. J. 1830. Flora Boreali-Americana. London. Hopkins, D. M. 1967. The Bering Land Bridge. Stanford: Stanford University Press. Hulten, E. 1937. Outline of the History of Arctic and Boreal Biota during the Quaternary Period. Stock- holm. 1946. Flora of Alaska and Yukon 7. pp. 1108-1113. ----Lund: Lund University Arsskrift. 1968. Flora of Alaska and Neighboring Territories. ----Stanford: Stanford University Press. 1970. The Circumpolar Plants 2. Dicotyledons. ----Almquist and Wiksell, Stockholm. 1973. Supplement to Flora of Alaska and Neighbor- ----ing Territories. A Study in the Flora of Alaska and the Transberingian Connection. Bot. Not. 126: 459-512. . Hutchinson, J. 1964. The Genera of Flowering Plants, An- giospermae. Oxford; Clarendon Press. -- Irving, G. W.; Fontaine, T. D.; and R. c. Warner. 1945. Electrophoretic Investigation of Peanut Proteins. Arch.' ~iochem. 7,475-489.

Isely, D. 1951. The Le~uminosae of the North-Central United States.. Hedysareae. Iowa State Jour. Sci. 30:33-118. Ives, J. D. 1963. "Field Problems in Determing the Maxi- mum Extent of Pleistocene Glaciation along the Eastern Canadian Seaboard - A Geographers Point of View." In North Atlantic Biota and their History, pp. 337-354. Edited by A. and D. Love. New York: Macmillan Co. Jackson, v. W.; Higham, J. F.; Groh, H.; and c. w. Lowe. 1922. Checklist of Manitoba Flora. Nat. Hist. Soc. of Manitoba. Johnson, A. W. and J. G. Packer. "Distribution, Ecology and Cytology of the Ogotoruk Creek Flora and the History of Beringia. 11 1963. in The Bering Land 181

Bridge, pp. 245-265. Edited by D. M. Hopkins. Stanford: Stanford University Press. Johnson, A. w. and J. G. Packer. 1968. Chromosome Numbers in the Flora of Ogotoruk Creek, N. w. Alaska. Bot. Not. 121:403-456. Jones, J. B. 1970. "Physiological Basis for Plant Analysis." in Proceedings from a Symposium on Plant Analysis, Edited by F. Greer. International Minerals and Chem. Corp., Skokie, Ill. Jones, J. B. and H. F. Mederski. 1964. Effect of Time and Soil Moisture Level on the Mineral Composition of Field Grown Soybean Plants. Agron. Abstr. Amer. Soc. of Agronomy, Kansas City, Missouri. November 15-19. Jurtsev, B.; Kozhevnikov, Yu. P.; and Nechaev, A. A. 1972. Interesting Floristic Finds in Easternmost Chukotka Peninsula. Bot. Zhurn. 57:765-778. Kearney, T. H. and R. H. Peebles. 1969. Arizona Flora. Berkeley: University of California Press. Kitagawa, M. 1937. A Diamino Acid, Canavanine, and a Monoamino Acid, Canaline. J. Biochem. Tokyo 25: 23-41. Kitagawa, M. and A. Takani. 1936. A Diamino Acid, Canava- nine. 4. The Constitution of Canavanine and Cana- line. J. Biochem. Japan. 23:181. Kitagawa, M. and Tomita, T. 1929a. A New Basic Compound in Jack Bean, from which Urea is Split off by a Liver Enzyme. Proc. Imp. Acad. Japan. 5:380-383. Kitagawa, M. and T. Tomiyama. 1929b. A New Amino-Compound in the Jack Bean and a Corresponding New Enzyme. J. Biochem. Japan. 11:265. Kitagawa, M. and H. Yamada. 1932. Canavanine - An Amino Acid. 2. J. Biochem. Japan. 16:339-349. Kirkbright, G. F. and J. H. Yoe. 1963. A New Spectro- photometric Method for the Determination of Micro- gram Amounts of Selenium. Analytical Chem. 35: 808-811. Klein, G. and E. Farkass. 1930. Der mikrochemische Nach- weis der Pflanze. 14. Der nachweis von cytisin. Oesterr. Bot. z. 79:107. Knaben, G. 1968. Chromosome Numbers of Flowering Plants 182 from Central Alaska. Nytt. Mag. Bot. 15:240-254.

Komarov, B. L. ed. 1972. Flora of the U.S.S.R. Vol. 13. Israel Program for Scientific Translations Ltd. Jerusalem: Keter Press. Kozlowski, T. T. 1972. Seed Biology. 3 Volumes. New York: Academic Press.

Kuvaev, V. B.; Glyzin, V. I.; Glyzina, G. S.; and A. I. Bankouski. 1972. The Prospects of the Search for Mangiferin in the Native Flora. Rastit Resur. 8: 367-371.

Lanjouw, J. and F. A. Stafleu. 1964. Index Herbariorum. Regnum Vegetabile 31:1-251. Ledingham, G. F. 1957. Chromosome Numbers of some Sas- katchewan Leguminosae with Particular Reference to Astragalus and Oxytropis. Can. J. Bot. 35:657-660.

1960. Chromosome Numbers in Astragalus and ~xy --~~~tropis. Canad. Journ. Genet. Cytol. 2:119-1 8.

Love, A. and D. Love. 1963. North Atlantic Biota and their History. New York: Macmillan Co.

Love, D. 1963. "Dispersal and Survival of Plants." in North Atlantic Biota and their History. 189-205. Edited by A. and D. Love. New York: Macmillan Co. Lovka, M.; Susnik, F.; Love, A.; and D. Love. 1971. IOPB Chromosome Number Reports 34. Taxon 20:785-787.

Luning, B. 1964. Studies on Orchidaceae Alkaloids 1: Screening of Species for Alkaloids. Acta Chem. Scand. 18:1507-1516.

Mackenzie, A. 1801. Voyages from Montreal on the River St. Lawrence, through the Continent of North America, to the Frozen and Pacific Ocean in the Years 1789 and 1793. 2 Volumes. London: T. Cadell. Macoun, J. 1883. Catalogue of Canadian Plants. Vol. 1 p. 117. Montreal.: Dawson Brothers. McDougall, W. B. 1973. Seed Plants of Northern Arizona. Flagstaff: The Museum of Northern Arizona. Mckee, H. s. 1962. Nitrogen Metabolism in Plants. oxford: Clarendon Press. 183 Mothes, K. and L. Engelbrecht. 1952. Uber geschlechts- verschildenen stoffwechsel zweihausiger pflanzen. 1. Untersuchungen uber den stickstoffumsatz beim Hanf (Cannabis sativa L.). Flora 139:1. 1952b. Uber allantoinsaure und allantoin. 1. Ihre ----rolle als wanderform des stickstof f s und ihre Beziehungen zum Erweistoffwechsels des ahorns. Flora 139:586. 1954. Uber allantoinsaure und allantoin 2. Ihr ----verhalten in den speicherwurzeln von Symphytum officinale. Flora 141:356. Mulligan, G. 1967. In IOPB Chromosome Number Reports 14. Taxon 16:552-571. Mulligan, G. A. and w. J. Cody. 1971. In IOPB Chromosome Number Reports 33. Taxon 20:609-614. Nakatu, S.; Matsugaki, z.; Simada, H.; Kashida, T.; Moto- mura, T.; Oshima, M.; and N. Sakei. 1962. Dis- tribution of Canavanine in Plant Seeds. Seikagaku 34:253. Nelson, Aven. 1902. The Genus Hedysarum in the Rocky Mountains. Pro. Biol. Soc. Wash. 15:183-186. Nuttall, T. 1818. The Genera of North American Plants. Vol. 2. Philadelphia; D. Heartt. Osburm, R. L.; Shendrikar, A. D.; and P. w. West. 1971. A New Spectrophotometric Method for Determination of Submicrogram Quantities of Selenium. Analyti- cal Chem. 43:594-597. Over, w. H. 1932. Flora of South Dakota. Vermillion: University of South Dakota Press. Packer, J. G. 1968. In IOPB Chromosome Number Reports. 17. Taxon 17:285-288. Paech, K. and M. v. Tracey. 1955. Moderne Methoden Der Pflanzenanalyse. Band 4. Berlin: Springer-Verlag. Page, E. R. 1965. Micro-Determination of Zinc and Copper in a Single Digest of Small Samples of Plant Mater- ials. Analyst 90:435-436. Peck, M. E. 1941. A Manual of the Higher Plants of Oregon. Portland: Binfords and Mort. Piper, c. v. and R. K. Beattie. 1915. Flora of the North- west Coast. Lancaster, Pa. 184 Polunin, N. 1940. Botany of the Canadian Eastern Arctic. Part. 1. Nat. Mus. Can. Bull. No. 92. 1959. Circumpolar Arctic Flora. 288-290. Claren- ----don Press. Porsild, A. E. 1951. Botany of Southeastern Yukon Adja- cent to the Cano! Road. Nat. Mus. of Can. Bull. 121.

1955. The Vascular Plants of the Western Canadian ----Arctic Archipelago. Nat. Mus. of Can. Bull. 135. 1957. Illustrated Flora of the Canadian Arctic ----Archipelago. Nat. Mus. of Can. Bull. 146. 1972. The Vascular Flora of Limestone Hills, North- ----ern Extension of the Ogilvie Mountains, Yukon Ter- ritory. Arctic 25:233-236. Porter, c. L. 1957. Leguminosae of Nevada. Part 3. Beltsville: u. s. D. A. Plant Industry Station. Radforth, M. w. 1950. Progress Report on Organic Terrain Studies. National Research Council of Canada. Tech. Mem. No. 16.

Raup, H. M. 1947. The Botany of Southwestern Mackenzie. Sargentia 6.

Reuter, G. 1957. Die Hauptformen des Loslichen Stickstoffs in Vegetativen pflanzlichen speicherorganen und ihre systematische Bewertbarkeit. Flora 145:3-26. Richardson, J. 1823. In Franklin, John. Narrative of a Journey to the Shores of the Polar Sea. Botanical Appendix. London: John Murray. Ritchie, J. c. 1962. A Geobotanical Survey of Northern Manitoba. Arctic Institute of North America Tech. Paper No. 9.

Robinson, B. L. and M. L. Fernald. 1908. Gray's New Man- ual of Botany. New York: American Book Co. Rollins, R. c. 1940. Studies in the Genus Hedysarum in North America. Rhodora 42:217-238. Rudd, v. E. 1951. Geographical Affinities of the Flora of North Dakota. Amer. Midl. Nat. 45:722-739. Rydberg, P. A. 1906. Flora of Colorado. Fort Collins: New Era Printing Co. 185 Rydberg, P. A. 1922. Flora of the Rocky Mountains and Adjacent Plains. Lancaster, Pennsylvania: In- telligence Printing Co. 1932. Flora of the Prairies and Plains of Central ~----~North America. New York. Sakaguchi, s. J. 1925. The Mode of Combination and De- termination of Arginine in the Protein Molecule. J. Biochern. Japan 5:133. Savile, D. B. O. 1972. Arctic Adaptations in Plants. Canada Dept. of Agriculture. Monograph No. 6. Schulze, E. and J. Barbieri. 1881. Ueber das Vorkornrnen von Allantoin irn Pflanzenorganisrnus. Ber. Dtsch. Chern. Ges. 14:1602. Scoggan, H. J. 1950. Botanical Investigations in Central Manitoba. National Mus. of Canada Bull. 118:84-92. 1950. The Flora of Bic and the Gaspe Peninsula, ~------Quebec. Nat. Mus. Can. Bull. 115. ------1957. Flora of Manitoba. Nat. Mus. of Can. Bull. 140. Seymour, F. C. 1969. The Flora of New England. Rutland, Vermont: Charles E. Tuttle Company. Sharples, A. w. 1938. Alaska Wild Flowers. Stanford, California: Stanford University Press. Shelter, s. G. 1964. Plants in the Arctic Alpine Environ- rnent. Srnithsonian Report for 1963. pp. 473-497. Simmons, H. G. 1913. A Survey of the Phytogeography of the Arctic American Archipelago. Kung!. Fysiogr. Sallsk. Handle. N. F. 24, No. 19. Soper, J. H. 1949. Vascular Plants of Southern Ontario. u. Toronto and Fed. Ont. Natur., Toronto. Soueges, R. 1956. Essai d'ernbryogenie cornparee dans Hedysarees. Ann. Sci. Bot. Biol. Veg. 11, 17:325- 352. Ssekaalo, H. 1971. A Rapid and Accurate Method for the Determination of Molybdenum in Plant Materials with Toluene - 3, 4 - Dithiol. Analyst 96:346-348. Stevens, o. A. 1950. Handbook of North Dakota Plants. Fargo, North Dakota: Knight Printing Co. 186

Stillmark, H. 1888. Uber Ricin, ein gifliges ferment. Inaugural Dissertation, Dorpat, Schnakenburg.

Taylor, R. C. and R. A. Ludwig. 1966. The Evolution of Canada's Flora. Toronto: University of Toronto Press.

Taylor, R. L. 1967. In IOPB Chromosome Number Reports 13. Taxon 16:445-461.

Taylor, R. L. and R. P. Brockman. 1966. Chromosome Numbers of some Western Canadian Plants. Canad. J. Bot. 44:1093-1103. Tidestrom, Ivar, and Sister Teresita Kittell. 1941. A Flora of Arizona and New Mexico. Wash. o. C.: The Catholic University of America Press.

Tikhomirov, B. A., et al. 1960. The temperature of arctic plants (in Russian). Acad. Sci. U.S.S.R. Biol. Ser. 3:429-442. Toms, G. c. and A. Western. 1971. "Phytohaemagglutinins." In Chemotaxonomy of the Leguminosae, pp. 367-462. Edited by J. B. Harborne, New York: Academic Press. Torrey, J. and A. Gray. 1838. A Flora of North America. Vol. 1. New York: Wiley and Putnam. Tracey, M. v. 1955. "Urea and Ureides." In Modern Methods of Plant Analysis, Vol. 4. pp. 119-141. Edited by K. Paech and M. Tracey. Berlin: Springer-Verlag. Tschiersch, B. 1961. Uber das Vorkommen von Canavanin. Flora 150:87. Tschiersch, B. and Hanelt, P. 1967. Die Freien Amino- sauren der Samen von Vicia L. und die systematische Gliederung der Gattung. Flora 157:389-406. Turner, B. c. and J. B. Harborne. 1967. Distribution of Canavanine in the Plant Kingdom. Phytochemistry 6: 863-866. Tutin, T. G. 1968. Flora Europaea, Vol. 2. 185-187. Cambridge, England: University Press. Watson, s. 1878. Bibliographical Index to North American Botany. Smithsonian Misc. Coll. #258. Wash., D. C. Weber, w. A. 1953. Handbook of Plants of the Colorado Front Range. Colorado: University of Colorado Press. 187 1967. Rocky Mountain Flora. Boulder, Colorado: ------University of Colorado Press. Wehmer, C. and M. Hadders. 1933. In Kleins Handbuch der Pflanzenanalyse, Bd. 4. Berlin: Julius Springer. Welsh, S. L. 1968. Nomenclature Changes in the Alaskan Flora. Great Basin Naturalist 28:147-156. Wiggins, I. and J. H. Thomas. 1962. A Flora of the Alaskan Arctic Slope. Toronto: University of Toronto Press. Willis, J. c. 1973. A Dictionary of the Flowering Plants and Ferns. 8th Ed. Cambridge, England: Cambridge University Press. Young, E. G. and c. F. Conway. 1942. On the Estimation of Allantion by the Rimini-Schryver Reaction. J. Biol. Chem. 142:839-853. Zacharius, R. M·.; Cathey, H. M.; and F. c. Steward. 1957. Nitrogenous Compounds and Nitrogen Metabolism in the Liliaceae 111. Changes in the Soluble Nitro- gen Compounds of the Tulip and their Relation to Flower Formation in the Bulbs. Ann. Bot. 21:193. Zhukova, P. G. 1969. Chromosome Numbers in Certain Plant Species Indegenous to the Northeast of the U.S.S.R. lV. Bot. Zurn. 54:1985-1990. THE GENUS HEDYSARUM IN NORTH AMERICA

Terry Edwin Northstrom Department of Botany and Range Science Ph. D. Degree, August 1974

ABSTRACT

In North America, Hedysarum taxa extend from southern Arizona to Banks Island and from northern Alaska to Newfound- land. They exhibit a high degree of polymorphism which is usually expressed in the form of a gradient in morphological characteristics. Four species, one subspecies and four varieties are recognized, including one new combination, H. boreale ssp. boreale var. rivulare (Williams) Northstrom. Comparative anatomical and morphological studies revealed two basic patterns. Chromosome numbers of all taxa were determined. The basic chromosome number.of!:!· alpinum, H. sulphurescens and H. occidentale is n=7. The basic chromosome number of H. boreale is n=8. Seed protein electrophoresis revealed two basic patterns which correspond to the two sections of the genus now delimited by classical methods. Total seed protein was estimated for each species. Tissue and seed extracts were tested for various ureides, uncommon amino acids and phytohemagglu- tinins. All taxa were found to be nonhemagglutinating and positive for canavanine, mangiferin, arginine and uric acid. Leaf element accumulation characters for each species were determined. COMMITTEE APPROVAL: