UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Systematics of Arnica, subgenus Austromontana and a new subgenus, Calarnica (Asteraceae : Senecioneae) Straley, Gerald Bane 1980

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Notice for Google Chrome users:
If you are having trouble viewing or searching the PDF with Google Chrome, please download it here instead.

Item Metadata

Download

Media
831-UBC_1980_A1 S87_3.pdf [ 14.69MB ]
Metadata
JSON: 831-1.0094954.json
JSON-LD: 831-1.0094954-ld.json
RDF/XML (Pretty): 831-1.0094954-rdf.xml
RDF/JSON: 831-1.0094954-rdf.json
Turtle: 831-1.0094954-turtle.txt
N-Triples: 831-1.0094954-rdf-ntriples.txt
Original Record: 831-1.0094954-source.json
Full Text
831-1.0094954-fulltext.txt
Citation
831-1.0094954.ris

Full Text

SYSTEMATICA OF ARNICA, SUBGENUS AUSTROMONTANA AND A NEW SUBGENUS, CALARNICA (ASTERACEAE:SENECIONEAE) by GERALD BANE STRALEY B.Sc, V i r g i n i a Polytechnic I n s t i t u t e , 1968 M.Sc, Ohio University, 1974 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS OF THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES (Department of Botany) We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA March 1980 © Gerald Bane Straley, 1980 In presenting this thesis in partial fulf i lment of the requirements for an advanced degree at the University of Bri t ish Columbia, I agree that the Library shall make i t freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. I t is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department nf Botany  The University of Brit ish Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 26 March 1980 ABSTRACT Seven species are recognized i n Arnica subgenus Austromontana and two species i n a new subgenus Calarnica based on a c r i t i c a l review and conserva-t i v e r e v i s i o n of the species. Chromosome numbers are given f o r 91 populations representing a l l species, including the f i r s t reports f o r Arnica nevadensis. Results of apomixis, vegetative reproduction, breeding studies, and a r t i f i -c i a l h y b r i d i z a t i o n s are given. I n t e r r e l a t i o n s h i p s of i n s e c t p o l l i n a t o r s , l e a f miners, achene feeders, and f l o r e t feeders are presented. Arnica  c o r d i f o l i a , the a n c e s t r a l species c o n s i s t s l a r g e l y of t e t r a p l o i d populations, which are e i t h e r autonomous or pseudogamous apomicts, and to a l e s s e r degree d i p l o i d , t r i p l o i d , pentaploid, and hexaploid populations. I t has given r i s e to Arnica nevadensis which i s t e t r a p l o i d and autonomously apomictic, and to a complex of d i s c o i d populations treated here as a s i n g l e polymorphic species, Arnica discoidea, with both d i p l o i d and t e t r a p l o i d , sexual and pseudogamous apomictic races. Arnica spathulata, derived from Arnica d i s c o i d e a , i s l a r g e -l y r e s t r i c t e d to serpentine s o i l s and i s a sexual d i p l o i d , with a few pseudo-gamous t e t r a p l o i d apomictic populations. Arnica l a t i f o l i a , the most poly-morphic species, has a r i s e n from Arnica c o r d i f o l i a , although i t has remained l a r g e l y sexual and d i p l o i d . Arnica l a t i f o l i a has given r i s e to Arnica cernua, the serpentine sexual, d i p l o i d endemic, and to Arnica g r a c i l i s , probably through h y b r i d i z a t i o n with other species. Arnica g r a c i l i s now e x i s t s l a r g e l y as a s e r i e s of autonomously apomictic t r i p l o i d races. The subgenus Calarnica was p o s s i b l y derived from subgenus Austromontana by way of Arnica spathulata through Arnica venosa, the l a t t e r g i v ing r i s e to Arnica v i s c o s a . These l a t t e r two species are rare sexual d i p l o i d s . i i i i i TABLE OF CONTENTS page Abstract • i i Frontispiece i i i L i s t of Tables v i L i s t of Figures v i i Acknowledgement x Introduction . 1 H i s t o r i c a l Background 2 Arnica - T r i b a l P o s i t i o n 2 Taxonomic History 3 Methods and Materials - General 6 Discussion of Characters 7 Habit and Branching Pattern 7 Underground Parts - Roots, Rhizomes, Caudices 8 Leaves 15 Odors of Leaves 16 Vesti t u r e 21 Capitula 32 P h y l l a r i e s 32 F l o r e t s - Ray, Disc, and Ampliate 35 Pappus 44 Achenes 45 D i s t r i b u t i o n and Ecology • 56 Reproductive Biology * 56 Phenology 56 iv page Cytology 58 Apomlxis 60 Pollen S t a i n a b i l i t y 89 Seed Production 95 Seed Germination 96 Seedlings 98 Vegetative Reproduction 101 C u l t i v a t i o n of Species I l l Hybridization 112 Sympatric Occurrence and Natural Hybridization " . . . 112 A r t i f i c i a l H ybridization 115 Entomological I n t e r r e l a t i o n s h i p s 120 Leaf Miners 121 F l o r e t Feeders 121 Achene Feeders 126 UV Patterns 127 P o l l i n a t o r s 130 Systematic Treatment 135 A r t i f i c i a l Key to Subgenera 135 Species Recognized 136 A r t i f i c i a l Key to Austromontana and Calarnica Species 136 F i e l d Key to Austromontana and Calarnica Species 138 Taxonomic Treatment 140 Phylogenetic Relationships 250 Summary 258 L i t e r a t u r e Cited 261 Appendices 265 v LIST OF TABLES pages I. Comparison of mature achenes. 55 I I . Previously reported chromosome numbers 63,64 I I I . New Chromosome numbers. 65-70 IV. Compatibility and Apomictic Experiments 86-88 V. Rhizome Growth. 110 VI. Sympatric Occurrence of Species 114 VII. A r t i f i c i a l Hybridizations 116,117 VIII. Leaf Miners. 122 IX. Floret Feeders 125 X. P o l l i n a t o r s . 132-134 v i LIST OF FIGURES pages 1-9. Schematic diagrams of habit and average branching. 9-14 10-25. Average leaves. 17-20 26-126. Comparison of v e s t i t u r e types on four vegetative parts. 23-31 127-136. Average i n v o l u c r a l bracts. 33,34 137-142. T y p i c a l ray f l o r e t s . 36,37 143-152. Ty p i c a l d i s c f l o r e t s . 38-41 153-156. Ampliate f l o r e t s of Arnica discoidea 42,43 157-165. Pappus. 45-48 166-186. Achene surfaces. 49-54 187-194. Meiotic chromosomes, microsporocytes, and po l l e n grains. 61,62 195-207. Chromosomes. 71-82 208-211. Experimental methods. 84,85 212-221. Pollen s t a i n a b i l i t y . 91-94 222-227. Seedlings. 99,100 228-231. Rhizome growth. 102-109 232. Summary of a r t i f i c i a l crosses. 118 233-240. Leaf Mines of Phytomyza species i n leaves of Arnica. 123,124 241-247. UV patterns i n Arnica heads and t y p i c a l heads of 3 species.128,129 248. D i s t r i b u t i o n map of Arnica c o r d i f o l i a . 145,146 249. Habit of Arnica c o r d i f o l i a . 147,148 250-259. V a r i a t i o n i n leaves of Arnica c o r d i f o l i a , large-leaved form. 149-151 260-267. Var i a t i o n i n leaves of Arnica c o r d i f o l i a , alpine form. 153,154 268-276. V a r i a t i o n i n leaves of Arnica c o r d i f o l i a , disturbed roadside form. 155,156 v i i L i s t of Figures, cont'd. pages 277-281. Habitats and habit of Arnica c o r d i f o l i a and Arnica nevadensis. 158,159 282. Habit of putative Arnica c o r d i f o l i a X Arnica l a t i f o l i a hybrid. 161,162 283. D i s t r i b u t i o n map of Arnica nevadensis. 165,166 284. Habit of Arnica nevadensis. 167,168 285. D i s t r i b u t i o n map of Arnica l a t i f o l i a . 174,175 286. Habit of Arnica l a t i f o l i a . 176,177 287-298. V a r i a t i o n i n leaves of Arnica l a t i f o l i a , small form. 179,180 299-304. V a r i a t i o n i n shortened ray f l o r e t s i n Arnica l a t i f o l i a . 181,182 305. D i s t r i b u t i o n map of Arnica g r a c i l i s and Arnica discoidea. 186,187 306. Habit of Arnica g r a c i l i s . 188,189 307-318. V a r i a t i o n i n leaves of Arnica g r a c i l i s , small form. 192,193 319-332. V a r i a t i o n i n leaves of Arnica g r a c i l i s , l a r g e r form. 194,195 333. D i s t r i b u t i o n map of Arnica cernua and Arnica spathulata. 198,199 334. Habit of Arnica cernua. 200,201 335-338. Habitats and habit of Arnica cernua. 203,204 339-355. Vari a t i o n i n leaves of Arnica cernua, small form. 205,206 356-365. V a r i a t i o n i n leaves of Arnica cernua, large form, sun form.207,208 366-370. Vari a t i o n i n leaves of Arnica cernua, large form, shade form. 209,210 371. Habit of Arnica discoidea. 214,215 372-375. Habitats and habit of Arnica discoidea and Arnica spathulata 217,218 376-385. V a r i a t i o n i n leaves of Arnica discoidea. 219,220 386. Habit of Arnica spathulata. 224,225 387-398. Va r i a t i o n i n leaves of Arnica spathulata, small form. 226,227 399-407. V a r i a t i o n i n leaves of Arnica spathulata, large form. 228,229 v i i i L i s t of Figures, cont'd. 408. D i s t r i b u t i o n of Arnica viscosa. 233,234 409. Habit of Arnica viscosa. 236,237 410- 414. Habitat and habit of Arnica viscosa. 238,239 415. D i s t r i b u t i o n map of Arnica venosa. 242,243 416. Habit of Arnica venosa. 244,245 417- 421. Habitat and habit of Arnica venosa. 246,247 422. Phylogeny of Arnica Subgenera. 254,255 423. Phylogeny of Arnica species i n subgenera Austromontana and Calarnica. 256,257 i x ACKNOWLEDGEMENTS I would l i k e to thank my advisory committee, Bruce A. Bohm, Fred R. Ganders, Christopher J . Marchant, Judith A. Myers, and my major advisor, Roy L. Taylor, for t h e i r h e l p f u l comments and c r i t i c i s m s during t h i s study. For c o l l e c t i o n s of seeds and plants I thank Bruce A. Bohm, Richard A. Gornall, Geraldine A. Guppy, Mary F. Jackson, Timothy A. Johns, Christopher J . Marchant, Kevin W. N i c h o l l s , and A l l a n A. Rose. For assistance i n l o c a t i n g rarer species I thank Susan Horner and numerous other members of the U. S. National Forest Service, Michigan Natural Resource Department, and U. S. and Canadian National Park Services f o r t h e i r cooperation i n allowing me to c o l l e c t i n t h e i r areas. Many hours of pleasurable c o l l e c t i n g were spent with Claude E. Garton, R. Patrick Harrison, A l l a n A. Rose and Roy L. Taylor. I was a s s i s t e d with SEM micrographs and other photographic techniques by Laszlo Veto and S y l v i a Taylor. Insects were determined by Graham C. D. G r i f f i t h s , F. C. Thompson, Robbin W. Thorp, Richard L. Westcott, and the L a t i n descrip-ti o n was done by P a t r i c i a M. Eckel. I thank my c a r e f u l t y p i s t , Betty Reubart, for her many hours of work. L a s t l y and most importantly, I would l i k e to thank Roy L. Taylor f o r h i s continued encouragement and f i n a n c i a l support through Natural Sciences and Engineering Research Council of Canada operating grant number 67-5705. I would l i k e to thank the curators of the following herbaria f o r allowing me to examine specimens of Arnica under t h e i r care: Brigham Young University (BRY) C a l i f o r n i a Academy of Sciences (CAS) Dudley Herbarium, Stanford University (DS) x Central Washington State University (ELRG) Humboldt State University (HSC) University of Idaho (ID) Jepson Herbarium, University of C a l i f o r n i a (JEPS) Kathryn Kalmbach Herbarium, Denver Botanic Gardens (KHD) University of C a l i f o r n i a , Los Angeles (LA) Lakehead University (LKHD) Museum of Northern Arizona (MNA) Missouri Botanical Garden (MO) University of Montana (MONTU) Northern Arizona University (NAU) C a l i f o r n i a Polytechnic State University (OBI) University of Oregon (ORE) Pomona College (POM) P a c i f i c Union College (PUA) Rancho Santa Ana Botanical Garden (RSA) Santa Barbara County Museum (SBM) Southern Oregon College (SOC) Unive r s i t y of B r i t i s h Columbia (UBC) University of C a l i f o r n i a , Berkeley (UC) University of C a l i f o r n i a , Santa Barbara (UCSB) Utah State University (UTC) University of V i c t o r i a (UVIC) B r i t i s h Columbia P r o v i n c i a l Museum (V) Willamette University (WILLU) x i Washington State University (WS) University of Washington (WTU) Crater Lake National Park x i i 1 INTRODUCTION Arnica i s a circumpolar genus of questionable t r i b a l p o s i t i o n i n the Asteraceae, occurring from the northern l i m i t s of vascular plant growth at 85° north l a t i t u d e , with four major southern r a d i i of d i s t r i b u t i o n ; south i n A s i a to Japan; i n Europe to Northern Jugoslavia and Northern Spain; i n Eastern North America along the A t l a n t i c Coast and Piedmont to Northern F l o r i d a ; and i n Western North America, where i t reaches i t s greatest d i v e r -s i t y , to Northern New Mexico and Southern C a l i f o r n i a . In the e a r l y years of Western North American exploration a vast number of species names were proposed, culminating i n the F l o r a of North America i n which Rydberg (1927) recognized 107 species i n North America i n 17 subgeneric groups. Rydberg presumably did not intend these groups to denote subgenera or s e c t i o n s , but rather convenient groups as an a i d i n i d e n t i f y i n g species. A much more conservative treatment, and the only comprehensive worldwide monograph, based l a r g e l y on morphology, was proposed by Maguire (1943), i n which he recognized 32 species (Appendix I) of.which 27 are from Western North America. Maguire was the f i r s t . t o d i v i d e the genus in t o 5 subgenera, A r c t i c a (7 s p e c i e s ) , Andropurpurea (3 s p e c i e s ) , Montana (2 s p e c i e s ) , Chamis-sonis (7 s p e c i e s ) , and Austromontana (13 s p e c i e s ) . Since Maguire's works (1943, 1947), the genus has recieved l i t t l e f u r -ther study other than a few reports of chromosome numbers (Ornduff , et a l . , 1963,1967; Strother, 1973; S t r a l e y , 1979), and an e x c e l l e n t survey of apo-mixis i n the genus by Barker (1966, unpublished). A recent monograph of the North American species (Ediger and Barkley, 1978) i s based l a r g e l y on Maguire (1943) with the reproductive information.of Barker (1966). Ediger and Barkley (1978) recognize 23 species and 26 i n f r a s p e c i f i c taxa (Appendix I I ) . There remain few taxonomic problems i n the subgenera Andropurpurea, Montana, and Chamissonis, although a re-evaluation of i n f r a s p e c i f i c taxa within the l a t t e r subgenus seems worthwhile. The greatest d i f f i c u l t i e s seem to be in the subgenus A r c t i c a . However, owing to the circumpolar d i s t r i b u -t i o n of these species and t h e i r r e l a t i v e i n a c c e s s i b i l i t y , p a r t i c u l a r l y of the Asian taxa, a comprehensive systematic study of subgenus A r c t i c a remains to be done. The subgenus Austromontana was selected for the present study be-cause of unanswered taxonomic problems i n t h i s subgenus following Maguire's monograph and the number of taxonomic changes proposed since then (Maguire, 1947; Cronquist, 1955, 1960; Ediger and Barkley, 1978). Furthermore, par-t i a l l y due to c u l t u r a l problems, Barker (1966) di d l e s s i n v e s t i g a t i o n of the reproductive behavior of Austromontana species than other subgenera. He states (Barker, 1966, p. 66), "The species belonging to the subgenus Austro-montana have proved to be l e s s amenable to i n v e s t i g a t i o n than those of other subgenera." L a s t l y , a l l the species i n t h i s subgenus are r e l a t i v e l y e a s i l y a c c e s s i b l e i n Western North America. HISTORICAL BACKGROUND T r i b a l P o s i t i o n of Arnica Although i t has long been placed i n the Senecioneae, Arnica, with i t s broad rays, opposite leaves, setose receptacle, base chromosome number of x=19, and apomictic element, does not f i t conveniently i n t h i s t r i b e . Nordenstam (1977) suggests e i t h e r a c l o s e a f f i n i t y to the Heliantheae or that possibly a new t r i b e , the "Arniceae", should be erected for Arnica and some eleven other genera. Baag^e (1977) presents evidence of l i g u l e micro-characters for i n c l u s i o n i n the Heliantheae. Chemical data ( S i e g l e r et a l . , 3 1974; Robins, 1977) supports the anomalous p o s i t i o n i n the Senecioneae. However, Turner and Powell (1977) believe that Arnica should be l e f t i n the Senecioneae. The pappus of c a p i l l a r y b r i s t l e s i s p a r t i c u l a r l y out of place i n the Heliantheae. The s i t u a t i o n i s well summarized by Cronquist (1977, p. 148), " I t seems l i k e l y that these two genera Arnica and Doronicum w i l l continue to be an embarrassment to future as well as present t r i b a l organi-zations i n the family." Further r e l a t i o n s h i p s of Arnica to Whitneya ( S i e r r a Nevada), Arnicastrum (Northwest Mexico), Jamesianthus (Southeast United States), and Mallopatopus (Japan) and possibly other genera need more c r i t i -c a l evaluation before they may be assigned with confidence to an e x i s t i n g t r i b e or placed i n a new t r i b e . Taxonomic History Maguire (1943) gives an excellent h i s t o r i c a l account of the genus as a whole. While l i t t l e more w i l l be discussed concerning the remaining four subgenera ( A r c t i c a , Montana, Andropurpurea, and Chamissonis) a more de t a i l e d account of the taxa i n Maguire's subgenus Austromontana, treated here as two subgenera, Austromontana and Calarnica, seems appropriate. The f i r s t name proposed f o r a species i n Austromontana was Arnica l a t i -f o l i a by Bongard (1832) which he described from material c o l l e c t e d by Mertens at S i t k a , Alaska. This proved to be the f i r s t i n a long series of names applied to what are now interpreted as a l l v ariants or even t y p i c a l specimens of the most polymorphic species i n the subgenus, i f not i n the whole genus. Hooker (1834) i n the F l o r a Boreali-Americana named Arnica c o r d i f o l i a from the Rocky Mountains, again the f i r s t name of many to be applied to a polymorphic, and the most widespread species i n t h i s subgenus. He also named 4 Arnica menziesii from the "Northwest Coast of America", the f i r s t of those species now considered as synonyms of Arnica l a t i f o l i a . N u t t a l l (1841) described a large-leaved plant from the Blue Mountains of eastern Oregon, Arnica macrophylla, now considered merely a large-leaved form of t y p i c a l Arnica c o r d i f o l i a . The f i r s t r a y l e s s taxon i n the group was named Arnica discoidea by Bentham (1849) from m a t e r i a l c o l l e c t e d by Theodor Hartweg " i n s y l v i s prope Monterey". Herder (1867) recognized two v a r i e t i e s of Arnica  l a t i f o l i a , ot genuina and 3 a n g u s t i f o l i a , and synonymized Hooker's Arnica  menziesii under Arnica l a t i f o l i a . In the Synoptical F l o r a of North America, Asa Gray (1884) proposed three names which are now considered synonymous with other names, Arnica p a r v i f l o r a and Arnica c o r d i f o l i a var. eradiata (now both = Arnica discoidea) and Arnica  l a t i f o l i a var. v i s c i d u l a (= Arnica d i v e r s i f o l i a i n subgenus Chamissonis). However, he d i d name the very d i s t i n c t i v e and rare Arnica v i s c o s a from Mt. Shasta, C a l i f o r n i a (Gray, 1878) and Arnica nevadensis from Lassen's Peak, C a l i f o r n i a (Gray, 1884), both of which are considered good species today. The three decades from 1897 to 1927 saw the greatest number of new names proposed, most of which are now relegated to synonymy under other species. Rydberg (1897, 1900, 1917, 1927), i n a s e r i e s of papers, proposed 18 new names, 17 of which are now considered synonymous with 5 older names. Only A r n i c a . g r a c i l i s from the Rocky Mountains of Western Montana was retained as a good species by Maguire (1943) and i n the present study, although others (Cronquist, 1955; Ediger and Barkley, 1978) considered i t a v a r i e t y of Arnica l a t i f o l i a . Greene (1900, 1901, 1902, 1910) likewise proposed 16 new names, with only A r n i c a spathulata from Southwestern Oregon retained today, although Greene's Arnica tomentella s t i l l remains a questionably v a l i d e n t i t y 5 (see discussion under Arnica nevadensis). Howell (1900) named the serpentine s o i l endemic which he c o l l e c t e d i n Josephine County, Oregon, Arnica cernua. I t i s one of the most d i s t i n c t i v e of the radiate species, as well as one of the r a r e s t . Nelson (1901, 1909) proposed the names Arnica p l a t y p h y l l a (= Arnica l a t i f o l i a ) , Arnica Columbiana, Arnica arcana (both = Arnica g r a c i l i s ) , and Arnica paniculata (? = Arnica  c o r d i f o l i a ; see discussion under that species). Jones (1910) proposed v a r i e t y g r a c i l i s of Arnica b e t o n i c a e f o l i a Greene (= Arnica l a t i f o l i a ) . H a l l (1915) named the rare but d i s t i n c t i v e Arnica  venosa which he c o l l e c t e d i n Shasta County, C a l i f o r n i a . Gandoger (1918) named Arnica eripoda (= Arnica l a t i f o l i a ) . Piper (1920) named Arnica andersonii from Skeena, B r i t i s h Columbia, another of the large-leaved shade forms of Arnica c o r d i f o l i a , and Arnica aphanactis from Mt. Baker, Washington, which i s a form of Arnica l a t i f o l i a . Fernald (1935) proposed the name Arnica whitneyi for plants from the Keewenaw Peninsula of Michigan, f ar disjunct from Western North American Austromontana species. I t was considered a subspecies of Arnica c o r d i f o l i a by Maguire (1943) and i s not given taxonomic status by Ediger and Barkley (1978) or i n the present treatment. Williams (1935) placed Greene's Arnica t e u c r i i f o l i a as a v a r i e t y of Arnica l a t i f o l i a and St. John (1937) named Arnica hardinae (= Arnica cordi-_ f o l i a ) from Benewah County, Idaho, the l a s t new name proposed i n the subgenus. Maguire (1943), i n his worldwide monograph of the genus, presented the most conservative treatment of the genus to that date, recognizing 13 species and 6 i n f r a s p e c i f i c taxa i n Austromontana (see Appendix I ) . His treatment has been l a r g e l y accepted by l a t e r workers and for the most part the findings of 6 the present study support his work. Although his work was based almost e n t i r e l y on morphological characters, he seems to have had an excellent understanding of the genus. However, the greatest shortcoming of his work i s that his keys are long and often very d i f f i c u l t to use. L i t t l e taxonomic work has been done since Maguire's monograph other than some new combinations and arbitrary lumping of species. In a study of Great Basin plants Maguire (1947) recognized four v a r i e t i e s of Arnica c o r d i f o l i a subsp. genuina: v a r i e t i e s c o r d i f o l i a , maefophylla, pumila, and humilis. Cronquist (1955), i n the Flora of the P a c i f i c Northwest, considered Arnica g r a c i l i s a variety of Arnica l a t i f o l i a and he lumped Arnica p a r v i f l o r a and Arnica grayi under the new combined name Arnica discoidea var. eradiata (A. Gray) Cronquist. He l a t e r (Cronquist, 1958) changed the status of Arnica  p a r v i f l o r a subsp. alata (Rydb.) Maguire to Arnica discoidea var. alata (Rydb.) Cronquist. In the most recent c l a s s i c a l monograph of the genus for North American Flora, Ediger and Barkley (1978) b a s i c a l l y follow Maguire's work except that they use Cronquist's new combinations and that they treat Maguire's sub-species as v a r i e t i e s . They recognize 9 species with 8 i n f r a s p e c i f i c taxa i n Austromontana (see Appendix II) . METHODS AND MATERIALS - GENERAL F i e l d collections were made from natural populations of Arnica species from B r i t i s h Columbia and Alberta to Ontario and Michigan and south to Colo-rado and Southern C a l i f o r n i a during several extensive c o l l e c t i n g t r i p s from the f a l l of 1976 through the summer of 1979. Some plants or seeds were also obtained from other collectors and from botanical gardens. 7 Small, whole heads were f i x e d i n the f i e l d i n Carnoy so l u t i o n (6 parts 95% ethanol : 3 parts chloroform : 1 part g l a c i a l a c e t i c acid) and l a t e r transferred to 95% ethanol and r e f r i g e r a t e d f o r c y t o l o g i c a l study. Live plants or seeds of many populations were c o l l e c t e d f o r further study i n the experimental garden, greenhouse, and growth chamber. Observations and records were made i n the f i e l d of p o l l i n a t o r s and other insect v i s i t o r s and feeders on the Arnicas. Insect specimens were c o l l e c t e d f o r i d e n t i f i c a t i o n and further study. Rhizome growth was measured for a number of populations. Specimens from more than 30 herbaria were studied to supplement f i e l d work. A morphological re-evaluation was made r e s u l t i n g i n the conservative taxonomic treatment presented herein. Voucher specimens of plants and insects are deposited at UBC. DISCUSSION OF CHARACTERS Among the morphological and other features used i n e l u c i d a t i n g taxa i n the subgenera Austromontana and C a l a r n i c a and i n determining phylogenetic r e l a t i o n s h i p s in these subgenera were the f o l l o w i n g : Habit and Branching Patterns A l l species i n Austromontana are herbaceous perennials a r i s i n g from rhizomes near the s o i l surface to several centimeters below the surface. The two species i n Calarnica are also herbaceous perennials but they lack rhizomes. These have a thick woody caudex a few to several centimeters below the s o i l surface. Only the rhizomatous species produce s t e r i l e basal rosettes of leaves (innovations). Plants may be unbranched above ground to much-branched near ground l e v e l or higher on the plant. In the non-rhizomatous species, and others with short rhizomes (as Arnica g r a c i l i s ) , the plant may be branched from below the ground l e v e l producing a clump. Heads may be erect or nodding and from one to many per stem. Figures 1 - 9 show a schematic representation of habit and average branching patterns. Underground Parts - Rhizomes, Caudices, and"Roots A l l species i n Austromontana are rhizomatous. The rhizomes are u s u a l l y slender (1-2 mm t h i c k ) , dark brown, and unbranched or with a few predomi-nantly terminal branches. They are naked or sparsely clothed with p a i r s of usually i n p e r s i s t e n t brown scales. In some of the short-rhizomed species ( e s p e c i a l l y Arnica g r a c i l i s ) and ecotypes of others (Arnica c o r d i f o l i a , Arnica l a t i f o l i a , and A r n i c a cernua) scales are more numerous or appear to be. These scales, i n a d d i t i o n to p e r s i s t e n t l e a f bases and stem bases from pre-vious years, give the rhizome a much more thickened appearance. Rhizome length i s r e l a t i v e l y consistent within a majority of populations of most species, being markedly greater i n some species (e.g., Arnica c o r d i -f o l i a ) than i n others (e.g., Arnica g r a c i l i s and Arnica l a t i f o l i a ) . (See Table V). within a species or population rhizomes are u s u a l l y longer i n shady than i n sunny h a b i t a t s . However, length seems to be l a r g e l y g e n e t i c a l l y c o n t r o l l e d . With few exceptions average yearly rhizome production under var-ious a r t i f i c i a l c u l t u r a l regimes va r i e d l i t t l e from yearly average production i n the same population i n the f i e l d . Rhizomes spread h o r i z o n t a l l y a few centimeters below the s o i l or duff survace i n most species and i n most f o r e s t and meadow h a b i t a t s . However, i n dry habitats, some populations, e s p e c i a l l y of Arnica spathulata, tend to have deeper rhizomes (6-10 cm) with v e r t i c a l extensions which branch g i v i n g r i s e 9 Figures 1-3. Schematic diagrams of habit and average branching pattern of Arnica species. 1. Arnica g r a c i l i s 2. Arnica l a t i f o l i a 3. Arnica cernua 11 Figures 4-7. Schematic diagrams of habit and average branching pattern of Arnica species. 4. Arnica c o r d i f o l i a 5. Arnica nevadensis 6. Arnica discoidea 7. Arnica spathulata 13 Figures 8, 9. Schematic diagrams of habit and average branching pattern of Arnica species. 8. Arnica viscosa 9. Arnica venosa 15 to h o r i z o n t a l rhizomes (Figure 7) c l o s e r to the ground surface. Roots are sparsely produced along the rhizome, except toward the branching terminal o r i g i n of a e r i a l portions. There are more roots produced on the shorter-rhizomed species and ecotypes. Primary roots are usually 0.5-1 mm thick and 20-30 cm long, rather v e r t i c a l and simple or sparsely branched. In d r i e r habitats and e s p e c i a l l y i n serpentine s o i l s the roots tend to be thicker (1-2 mm). Rhizomes generally remain a l i v e f or about 3 years with the oldest growth dying as new rhizomes grow i n the spring. In Calarnica both species lack rhizomes but rather possess a t h i c k woody perennial caudex, 4-5 cm i n diameter from which usually many flowering stems are produced each year. The bases of old flowering stems remain on the caudi-ces for several years giving them a rough character. The roots are quite t h i c k (5-6 mm) and long (to 40 cm or more) and tend to be quite v e r t i c a l and unbranched. Leaves A l l species of Austromontana produce basal rosettes of 4-6 p a i r s of u s u a l l y long-petiolate leaves. Flowering stems have opposite p a i r s of brown membraneous scales at the base, grading gradually, or more often abruptly, into small lower cauline leaves. There are t y p i c a l l y 2-4 p a i r s of opposite cauline leaves, the l a r g e s t produced toward the lower or middle portion of the stem, becoming smaller and b r a c t - l i k e and sometimes alternate below the i n -florescence. Cauline leaves are s e s s i l e to long p e t i o l a t e , with the p e t i o l e s often winged, r a r e l y wider than the blades. There i s much v a r i a t i o n i n d i s -p o s i t i o n of the leaves on the flowering stem as well as the angle at which the leaves are held. 16 Leaf o u t l i n e ranges from lanceolate to broadly ovate, with acute to cordate bases and acute to rounded t i p s . Margins may be e n t i r e to crenate, serrate, dentate, or doubly serrate, dentate, or crenate, often undulate, and often combinations of these characters. Leaves are one of the most r e l i a b l e characters i n separating taxa (Figures 10-25), although they are one of the most v a r i a b l e characters within some taxa. There i s usually much interpopulational consistency, although there may be a great deal of phenotypic v a r i a t i o n within a single population. Thick, r e l a t i v e l y narrow sun-leaves or t h i n , r e l a t i v e l y broad shade-leaves are often found within populations growing i n open f o r e s t habitats, e s p e c i a l l y i n Arnica l a t i f o l i a and Arnica cernua. Leaves, e s p e c i a l l y i n sunny locations and from plants growing i n serpentine s o i l s , are often flushed with red or purple. There are no basal rosettes i n subgenus Calarnica. Scales at the base of the flowering stems are very prominent and often several (6-12) pai r s which grade gradually into cauline leaves. Stems of t h i s subgenus are much l e a f i e r , with 7-15 or more pai r s of s e s s i l e ovate leaves on the primary axis and smaller leaves and/or bracts on numerous l a t e r a l branches and in the l e a f y inflorescence. Upper bracts are usually sub-opposite or alternate. Odors of Herbage Although d i f f i c u l t to describe q u a l i t a t i v e l y , pungent odors associated with those species bearing glandular hairs are excellent characters f o r sepa-r a t i n g some species. While odors are p a r t i c u l a r l y useful i n the f i e l d , herbarium specimens of c e r t a i n species r e t a i n t h e i r odor f o r many years, nota-bly Arnica v i s c o s a . 17 Figures 10-17. Average Basal and Cauline Leaves of Arnica Species. 10. Basal l e a f of Arnica l a t i f o l i a S-1676, WA, Clallam County. 11. Cauline l e a f of Arnica l a t i f o l i a S-1654, BC, Tod Mountain. 12. Basal leaf of Arnica c o r d i f o l i a S-1819, OR, Wasco County. 13. Cauline l e a f of Arnica c o r d i f o l i a S-1354, BC, Botanie V a l l e y . 14. Basal l e a f of Arnica cernua S-1743, OR, Josephine County. 15. Cauline leaf of Arnica cernua Sweetser, _s._n. (ORE), OR, Josephine County. 16. Basal l e a f of Arnica g r a c i l i s S-1603, AB, Lake Louise. 17. Cauline l e a f of Arnica g r a c i l i s C h r i s t 19463 (ID), Elmore County. 18 19 Figures 18-25. Average basal and cauline leaves of Arnica species. 18. Basal l e a f of Arnica spathulata S-1381, OR, Josephine County. 19. Cauline l e a f of Arnica spathulata Detling 4688 (ORE), OR, Josephine County. 20. Basal leaf of Arnica discoidea S-1789, CA, Marin County. 21. Cauline leaf of Arnica discoidea Copeland s.n. (ORE), CA, Butte County. 22. Basal l e a f of Arnica nevadensis S-1910, NV, Washoe County. 23. Cauline l e a f of Arnica nevadensis S-1910, NV, Washoe County. 24. Cauline l e a f of Arnica viscosa Muth 733 (PUA), CA, Siskiyou County. 25. Cauline leaf of Arnica venosa S-1793, CA, Shasta County. 20 A l l species have odors when the leaves are bruised except Arnica  l a t i f o l i a and Arnica cernua. Rarely Arnica l a t i f o l i a , or suspected hybrids of i t and Arnica c o r d i f o l i a are s l i g h t l y fragrant, e s p e c i a l l y i n younger, ra p i d l y growing leaves. The odor i s l o s t i n older leaves. Among the f r a -grant-leaved species there are consistent differences i n q u a l i t y and degree of odors. Arnica viscosa has by f a r the strongest odor, while the cl o s e l y r e l a t e d species, Arnica venosa, has only a s l i g h t fragrance, at le a s t i n the few populations studied i n the f i e l d . There also seem to be consistent d i f -ferences i n the odors of c l o s e l y r e l a t e d species, as i n Arnica c o r d i f o l i a and Arnica nevadensis, although there i s some v a r i a t i o n i n odor i n d i f f e r e n t populations of each of these species. Vestit u r e Maguire (1943, p. 403) comments on the use of pubescence as a c r i t i c a l character i n work on Doronicum by C a v i l l i e r (1907) and the Russian Arnicas by I l j i n (1926), but he cautions against r e l y i n g too heavily on th i s character alone. Pubescence, or the lack thereof, i s a good morphological character, e s p e c i a l l y i n the f i e l d , when used i n conjunction with other characters f o r determining Arnica taxa. With few exceptions a l l parts of a l l species are to some degree pubescent, most often densely so. There are two basic hair types i n Arnica, septate glandular and non-glandular. Each of these types commonly occur i n two types, long (1-2 mm) and short (0.1-0.2 mm), although there are numerous intermediate examples. The s t i p i t a t e glandular hairs have septate stalks usually two c e l l s thick except j u s t beneath the gland where they are one-celled. The gland i s com-posed of two to many c e l l s . A majority of the s t i p i t a t e glandular hairs are about 1 mm long on stal k s composed of 6-8 c e l l s , or they are quite short 22 (0.1-0.2 mm) with s t a l k s 2-3 c e l l s long. The degree of glandular pubescence imparts a more or l e s s v i s c i d or "clammy" and aromatic character to the leaves and stems of c e r t a i n species, e s p e c i a l l y Arnica c o r d i f o l i a , Arnica nevadensis, and Arnica v i s c o s a . This character alone makes them r e a d i l y separable i n the f i e l d from those species l a r g e l y lacking glandular h a i r s , most notably Arnica cernua and Arnica l a t i -f o l i a . In Arnica g r a c i l i s the predominant h a i r type i s the short, nearly s e s s i l e , glandular h a i r l e s s than 0.1 mm long g i v i n g the leaves a rough appearance under 10X magnification. In the other species bearing glandular h a i r s there are mixtures of both the long and short h a i r s , except i n Arnica  v i s c o s a which has only long glandular h a i r s . Non-glandular septate h a i r s are always one c e l l t h i ck except at t h e i r bases and they are u s u a l l y s t r a i g h t , the exceptions being on the lower part of the stems of some species, e s p e c i a l l y A r n i c a cernua. (Figure 62). Again two s i z e s of non-glandular h a i r s are seen, long ones 1-2 (-3) mm long and short ones 0.3-0.5 mm long, with some intermediates. The number of non-glandular h a i r s imparts a softness to the leaves of Arnica c o r d i f o l i a , Arnica  discoidea, and Arnica venosa. In c e r t a i n species or populations the glandular hairs are colored, notably the yellow-tipped h a i r s of Arn i c a v i s c o s a and the purple-tipped or based h a i r s of Arnica spathulata and to a l e s s e r degree Arnica discoidea. Figures 26 to 126 show the v e s t i t u r e types of a l l vegetative parts of the species i n these two subgenera, but with no i n d i c a t i o n of r e l a t i v e abun-dance of v e s t i t u r e . 23 Figures 26-126. Comparison of Vestiture Types on Four Vegetative Parts of Arnica Species. 24 25 Leaf, adaxial Leaf, abaxial Taxon cordifolia- l a t i f o l i a natural hybrid 26 Lower Cauline Upper cauline, Periclinum, Phyllaries Taxon gracilis nevadensis cernua 27 Leaf, adaxial Leaf, abaxial Taxon cernua none none 28 29 spathulata 30 31 Taxori viscosa Leaf, adaxial Leaf, abaxial 1mm venosa (§) 32 Capitula Heads vary from turbinate to campanulate i n the few-flowered species ( i . e . , Arnica viscosa and Arnica g r a c i l i s ) to broadly hemispheric i n most of the remaining species. They are l e s s than 1 cm broad i n the d i s c o i d Arnica  viscosa to nearly 6 cm broad i n terminal radiate heads of Arnica c o r d i f o l i a . Heads i n d i f f e r e n t species may be radiate with a single row of marginal ray f l o r e t s or d i s c o i d , lacking ray f l o r e t s . In the normally d i s c o i d species there may be elongated r a y - l i k e (ampliate) f l o r e t s (Figures 154-156) and i n the normally radiate species the ray f l o r e t s may be absent. There are nor-mally 1-3 heads per stem, with a larger terminal one and a p a i r or several p a i r s of l a t e r a l heads, with r a r e l y as many as 30 or more heads per stem as i n some populations of Arnica l a t i f o l i a , Arnica discoidea, and Arnica spathu-l a t a . Heads are held erect at anthesis i n most species. They are consis-t e n t l y nodding i n bud i n Arnica cernua and Arnica venosa and are often held at a 45-60° angle at anthesis i n Arnica cernua, Arnica c o r d i f o l i a , and Arnica  venosa. P h y l l a r i e s Average number, size and shape of i n v o l u c r a l bracts ( p h y l l a r i e s ) are shown i n Figures 127-136. These are v a r i a b l e characters within species and within populations, but o f f e r some consistent differences i n some taxa. A l l are lanceolate to ovate with acute t i p s , except Arnica cernua, Arnica spathu- l a t a , and Arnica discoidea which often have rounded t i p s . P h y l l a r i e s are c o n s i s t e n t l y smaller and narrower i n Arnica v i s c o s a and Arnica g r a c i l i s than i n other species. Putative natural hybrids between Arnica c o r d i f o l i a and Arnica l a t i f o l i a c o n s i s t e n t l y have more p h y l l a r i e s than e i t h e r parent. 33 Figures 127-136. Average Involucral Bracts of Arnica Species. 127. Arnica nevadensis S-1870, CA, S i e r r a County. 128. Arnica c o r d i f o l i a S-1343, BC, Botanie V a l l e y . 129. Arnica c o r d i f o l i a X Arnica l a t i f o l i a (putative natural hybrid) S-1475, MT, R a v a l l i County. 130. Arnica l a t i f o l i a S-1500, WA, Chelan County. 131. Arnica g r a c i l i s S-1605, AB, Moraine Lake. 132. Arnica cernua S-1386, OR, Josephine County. 133. Arnica discoidea S-1825, WA, K l i c k i t a t County. 134. Arnica spathulata S-1380, OR, Josephine County. 135. Arnica venosa S-1940, CA, Shasta County. 136. Arnica venosa S-1411, CA, Siskiyou County. 34 i l l l l l h l l l l I U 11111) 11 | iMIIIIIlUli® l l l l l l l t l l l l l ( S ) i l i l i i i l ® 35 Margins of p h y l l a r i e s are usually e n t i r e , although i n rare cases they have a few i r r e g u l a r teeth. Bases of the p h y l l a r i e s and the periclinum (area of attachment of p h y l l a r i e s to the peduncle) are usually the most densely pubes-cent part of most plants and often have the longest h a i r s on the plant. Ray F l o r e t s A l l ray f l o r e t s consist of a short tube and large f l a t t e n e d limb (Figures 137-142). The tube and lower surface of the limb are v a r i a b l y pubescent. The general o u t l i n e of the limb i s l i n e a r oblong to ovate with a rounded t i p or more frequently one to several c h a r a c t e r i s t i c teeth. The sizes and shapes of these teeth are used i n d i s t i n g u i s h i n g between some species. These are es-p e c i a l l y good f i e l d characters, not always apparent i n pressed herbarium specimens. The number of rays varies from t y p i c a l l y f i v e or r a r e l y l e s s i n depauperate heads to occasionally twenty or more per head. Rays are always bright yellow and p i s t i l l a t e with an elongated s t y l e extending beyond the tube. Disc F l o r e t s Heads contain as few as 9-10 to more than 180 disc f l o r e t s . These are mostly tubular or goblet-shaped (Figures 143-153). There i s often a d i s t i n c -t i v e ring at the base where the tube attaches to the ovary and another r i n g about midway up the tube where the filaments are attached within. At t h i s point the limb f l a r e s out, abruptly i n some cases, very gradually i n others. At the d i s t a l end of the limb are f i v e recurved lobes. Disc f l o r e t s are per-f e c t with f i v e anthers united around the p i s t i l . Nectar i s produced and the f l o r e t s are fragrant. They are always bright yellow except i n Arnica viscosa i n which the'y are white or cream-colored. Walls of the tube and limb are often nearly translucent. Disc f l o r e t s are sparsely to densely hairy on the 36 Figures 137-142. Typical ray f l o r e t s of Arnica subgenus Austromontana. 137. Arnica c o r d i f o l i a S-1353, BC, Botanie V a l l e y . 138. Arnica c o r d i f o l i a X Arnica l a t i f o l i a hybrid S-1474, MT, R a v a l l i County. 139. Arnica l a t i f o l i a S-1680, BC, Manning Park. 140. Arnica cernua S-1386, OR, Josephine County. 141. Arnica nevadensis S-1870, CA, S i e r r a County. 142. Arnica g r a c i l i s S-1605, AB, Moraine Lake. 37 38 Figures 143-148. T y p i c a l Disc F l o r e t s of Arnica Species i n Subgenus Austromontana. 143. Arnica c o r d i f o l i a S-1353, BC, Botanie V a l l e y . 144. Arnica c o r d i f o l i a X Arnica l a t i f o l i a S-1474 (putative natural hybrid), MT, R a v a l l i County. 145. Arnica l a t i f o l i a S-1476, MT, Missoula County. 146. Arnica nevadensis S-1870, CA, S i e r r a County. 147. Arnica g r a c i l i s S-1605, AB, Moraine Lake. 148. Arnica cernua S-1386, OR, Josephine County. 39 40 Figures 149-152. Typical Disc F l o r e t s of Arnica Species i n Subgenera Austromontana and Calarnica. 149. Arnica discoidea S-1789, CA, Marin County. 150. Arnica spathulata S-1742, OR, Josephine County. 151. Arnica viscosa S-1411, CA, Siskiyou County. 152. Arnica venosa S-1793, CA, Shasta County. 41 42 Figures 153-156. F l o r e t s of Arnica discoidea. 153. T y p i c a l d i s c f l o r e t , WA, K l i c k i t a t County, S-1825. 154. Marginal ampliate f l o r e t from same head. 155. Elongated marginal f l o r e t with abortive anthers, CA, Santa Barbara County, Smith 8330 (SBM). 156. Ray-like marginal f l o r e t with abortive anthers, CA, Mendocino County, Nelson & Anderson 843 (HSC). 43 44 tube or throughout ( e s p e c i a l l y i n Arnica venosa). There are always a few hairs on the t i p s of the lobes as well as d i s t i n c t i v e p a p i l l a e on t h e i r inner surface. The f l o r e t s are usually uniform i n s i z e and shape and symmetrical throughout the head. In many of the d i s c o i d heads of Arnica discoidea the marginal f l o r e t s are asymmetrical, known as ampliate f l o r e t s (Figure 155) or much elongated and r a y - l i k e with abortive anthers (Figures 154, 156). Pappus The pappus i s composed of a rin g or rings of about 27-65 m u l t i c e l l u l a r b r i s t l e s with l a t e r a l setae. The length of b r i s t l e s v a r i e s according to the length of the f l o r e t , but the length of l a t e r a l setae i s r e l a t i v e l y consis-tent within some species or species groups. Maguire (1943, p. 402) treats these setae i n three groups according to length: b a r b e l l a t e , with setae 0.1-0.2 mm; subplumose, 0.2-0.35 mm; and plumose, 0.35-0.6 mm. Most taxa i n these two subgenera f a l l within the f i r s t two groups (Figures 157-165, 186). The pappus of the whole head forms a round b a l l 1-3 cm i n diameter upon matur-i t y of the achenes. Color of the mature pappus i s usually pure white i n most species or often tawny i n Arnica nevadensis, Arnica viscosa, and Arnica  g r a c i l i s . Achenes The best s i n g l e reproductive character f o r separating some of the cl o s e -l y r e l a t e d species, i s the v e s t i t u r e of the achene surface (Figures 166-185). Table I gives a comparison of mature achenes. They are usually 5-8 mm long and 1 mm wide at t h e i r widest point, the d i s t a l end, l i n e a r oblong to clavate, weakly to strongly 5-10 angled. The surface i s dark grey or brown to black 45 Figures 157-161. Photographs with l i g h t microscope of near mid-length of pappus. 157. Arnica g r a c i l i s S-1603, AB, Lake Louise. 158. Arnica l a t i f o l i a S-1162, WA, Whatcom County. 159. Arnica cernua S-1772, CA, Humboldt County. 160. Arnica c o r d i f o l i a S-1939, CA, Mono County. 161. Arnica nevadensis S-1870, CA, S i e r r a County. 47 Figures 162-165. Photographs with l i g h t microscope of near mid-length of pappus. 162. Arnica discoidea Detling 6050 (ORE), CA, Siskiyou County. 163. Arnica spathulata S-1381, OR, Josephine County. 164. Arnica venosa S-1940, CA, Shasta County. 165. Arnica viscosa S-1411, CA, Siskiyou County. 48 49 Figures 166-171. Scanning Electron Micrographs of Achene Surf (mid-length) of Arnica species. 166. Arnica c o r d i f o l i a S-1353, BC, Botanie Valley, X65. 167. same, X130. 168. Arnica nevadensis S-1870, CA, S i e r r a County, X65. 169. same, X130. 170. Arnica discoidea S-1825, WA, K l i c k i t a t County, X65. 171. same, X130. 50 51 Figures 172-177. Scanning Electron Micrographs of Achene Surf (mid-length) of Arnica species. 172. Arnica cernua S-1772, CA, Humboldt County, X65. 173. same, X130. 174. Arnica l a t i f o l i a S-1476, MT, Missoula County, X65. 175. same, X130. 176. Arnica g r a c i l i s S-1562, BC, Monroe Lake, X65. 177. same, X130. 52 53 Figures 178-184. Scanning Electron Micrographs of Achene Surfaces (mid-length) of Arnica species. 178. Arnica spathulata S-1380, OR, Josephine County, X65. 179. same, X130. 180. Arnica venosa S-1940, CA, Shasta County, X65. 181. same, X130. 182. Arnica viscosa S-1411, CA, Siskiyou County, X65. 183. same, X130. 184. Arnica g r a c i l i s S-1562, BC, Monroe Lake, X250. 185. Photomicrograph with l i g h t microscope, achene surface of Arnica  c o r d i f o l i a S-1487, WA, Okanogan County, X500. 186. Scanning electron micrograph of pappus b r i s t l e (mid-length) of Arnica c o r d i f o l i a S-1353, BC, Botanie Valley, X575. TABLE I COMPARISON OF MATURE ACHENES OF ARNICA SPECIES Taxon Color c o r d i f o l i a dark grey l a t i f o l i a grey-brown g r a c i l i s black cernua nevadensis discoidea spathulata venosa viscosa dark grey dark grey to black dark grey black grey dark grey Length (mm) (6-)7-9(-10) (5-)6-8(-10) (4-)5-6(-7) 6-8 (4-)5-8 6- 8 7- 9 . 6-8 5-6 Width D i s t a l l y (mm) 1+ 1-1 1+ 1+ 1+ 1 1.5-2 1 Double Hairs uniform throughout or es p e c i a l l y d i s t a l l y none or few d i s t a l l y none or very few d i s t a l l y many throughout or d i s t a l l y few throughout or d i s t a l l y few to many throughout none to few throughout many throughout none Glandular Hairs uniform throughout none or few d i s t a l l y none or few through-out or many d i s t a l l y none uniform throughout few throughout few throughout or d i s t a l l y none uniform l o n g - s t i p i t a t e throughout 56 and minutely crested and undulate (Figure 184), although t h i s i s usually obscured by the surface h a i r s . There i s a thick white r i n g , the annulus, at the base where the achene was attached to the receptacle and another ri n g located d i s t a l l y to which the pappus i s attached. Achenes of ray f l o r e t s are shorter and broader than those of the di s c f l o r e t s . Most achene coats have few to many double-pronged or "duplex" hairs (Figure 185), e s p e c i a l l y d i s t a l -l y , i n addition to long s t i p i t a t e (Figure 183) to nearly s e s s i l e (Figure 184) glandular h a i r s . The p o s i t i o n and r e l a t i v e abundance or s p a r s i t y of these h a i r s i s used i n t h i s study as an important diagnostic character. DISTRIBUTION AND ECOLOGY Both of these subgenera are l a r g e l y Western C o r d i l l e r a n ( i n the broad sense) with eastern extensions or disjunct populations into the Cypress H i l l s of Saskatchewan, Black H i l l s of South Dakota, and around Lake Superior. None of these species i s t r u l y a r c t i c as are some of the species in other subgenera. They occur i n a v a r i e t y of habitats from open alpine or mountain meadows, for e s t edges, open f o r e s t s , disturbed roadside banks, s t a b i l i z e d rock s l i d e s , and s t a b i l i z e d burned-over areas, but mostly cool, mesic habitats. Only Arnica venosa i s a plant of the hot, dry f o o t h i l l s of the upper Sacramento River Valley of Northern C a l i f o r n i a . Two species, Arnica cernua and Arnica  spathulata are r e s t r i c t e d to or usually found on serpentine s o i l s . Arnica  viscosa i s l i m i t e d to high pumice slopes. REPRODUCTIVE BIOLOGY Phenology Being montane or alpine plants, a l l species of Austromontana begin new 57 innovation growth and flowering stem elongation as soon as spring or summer snows have melted. In coastal low mountains of C a l i f o r n i a and Oregon t h i s may occur i n A p r i l , while at higher altitudes and latitudes growth may begin as late as September or October (or probably not at a l l some years where snow cover i s excessive). High alpine populations often have elongated flowering stems under the melting snow or have stems emerging through the snow. In areas where there i s t y p i c a l l y dry weather following the spring snow-melt, as occurs east of the Cascade Mountains and i n the Siskiyou and Klamath Mountains, there i s usually a brie f flowering period of a few weeks with no more l a t e r a l flower buds being produced after the monocephalous or few-headed stems have flowered. This i s the common condition i n most species. In wetter habitats populations of the same species or other species may continue to branch and flower for months. Other species, especially Arnica l a t i f o l i a , produce only one " f l u s h " of flowers regardless of the weather conditions thereafter. In a growth chamber under conditions of 16 hour days, 25°C, and adequate moisture and f e r t i l i z e r , populations of several species, Arnica c o r d i f o l i a , Arnica discoidea, Arnica venosa, and especially Arnica viscosa continued to produce l a t e r a l branches and new growth from rhizomes and flowered continuous-l y for more than a year. Other experimental populations of Arnica c o r d i f o l i a , Arnica discoidea, Arnica spathulata, Arnica g r a c i l i s , Arnica nevadensis, and especially Arnica l a t i f o l i a , produced only one " f l u s h " of heads. Although leaves remained green for months and new vegetative growth was produced, no more flower buds were produced regardless of the growth regime. Only after several weeks of vernalization and then increasingly warmer temperatures and increased l i g h t were new flowering stems produced. 58 Actual flowering time from the opening of the f i r s t d i s c or ray f l o r e t to the opening of the l a s t d i s c f l o r e t may be as b r i e f as two days i n the few-flowered species (e.g., Arnica g r a c i l i s ) to more than a week i n the l a r g e r flowered species. During dry weather l a t e r a l buds often abort and at l e a s t during some years a l l the flower buds abort, as observed with numerous populations of Arnica c o r d i f o l i a i n c e n t r a l Wyoming during June 1978. Here, very large popu-l a t i o n s had flowering stems but no flowers. Achenes ripen within a few weeks of flowering and are blown away. New rhizome growth begins about the time of flowering with the new innovations emerging during or j u s t a f t e r flowering. A l l above-ground growth becomes dry and soon disappears except i n wetter habitats where plants may remain green u n t i l f a l l f r o s t . In Calarnica the stems become much elongated and often branched before flowers are produced. Several weeks or months may elapse from emergence to flowering. Flowering may be rather b r i e f with few heads per stem i n Arnica  venosa or longer for the much-branched Arnica viscosa. Cytology For chromosomal study, f i x e d anthers from small buds were excised on a glass s l i d e , a drop of aceto-carmine was added, and the anthers were macerated with iron needles f o r about 1-3 minutes u n t i l the aceto-carmine became dark purple. A drop of Hoyer's medium (Beeks, 1955) was added to make the s l i d e permanent, a cover s l i p was added, and the preparation was warmed b r i e f l y over a flame and squashed. Cover s l i p s were l a t e r ringed with diaphane. Observa-tions of chromosome number and meiotic behavior of microsporocytes were made with a p o s i t i v e phase-contrast microscope. Tapetal c e l l s were not used f o r chromosome number determinations since numerous observations of high p o l y p l o i d numbers were made i n tapetal c e l l s surrounding d i p l o i d or t e t r a p l o i d micro-sporocytes. Some chromosome number determinations were made from r o o t - t i p s from potted plants grown i n the greenhouse or growth chamber or from achenes germinated on moist f i l t e r paper. The r o o t - t i p s were soaked i n a supersatu-rated aqueous sol u t i o n of paradichlorobenzene f o r 1-2 hours (Strother, 1969) before f i x i n g and sta i n i n g as f o r flower bud material. Arnica chromosomes are small (ca. 5 Um), they do not f i x w e l l , and meiotic associations are often d i f f i c u l t to i n t e r p r e t , r e s u l t i n g i n many of the previous reports being approximations or probable e r r o r s . Often many microsporocytes from a number of squashes had to be observed before accurate counts or p a i r i n g was determined. Although d i p l o i d s usually form regular bivalents during metaphase I of meiosis (Figure 194), i n the higher ploidy l e v e l s there i s usually i r r e g u l a r behavior, with v a r i a b l e numbers of univa-l e n t s , b i v a l e n t s , multivalents, and chains present at metaphase I and lagging chromosomes and bridge formation during anaphase I (Figure 187) , as has been >. noted previously by Ornduff, et a l . (1963, 1967) and Barker (1966). Table II l i s t s a l l previously reported chromosome numbers f or these two subgenera, a number of which are approximations. There i s only one previous report f o r Arnica cernua and Arnica venosa and two each f or Arnica g r a c i l i s , Arnica spathulata, and Arnica v i s c o s a . Only f o r the two widespread species, Arnica c o r d i f o l i a and Arnica l a t i f o l i a , are there s u f f i c i e n t previous reports to show trends i n ploidy l e v e l s . Table III shows new chromosome numbers ob-served during t h i s study, more than twice the number of a l l previous reports. Counts f o r Arnica nevadensis are presented f o r the f i r s t time. Figures 187-60 189 and 193, 194 show photographs of meiotic chromosomes of Arnica g r a c i l i s , Arnica c o r d i f o l i a , and Arnica discoidea. Figures 195-207 show camera l u c i d a drawings of a l l ploidy l e v e l s observed during t h i s study, of a l l species recognized. To summarize known chromosome numbers, f i v e species in these subgenera are wholly d i p l o i d . These are Arnica viscosa, Arnica venosa, Arnica cernua, Arnica l a t i f o l i a (one t e t r a p l o i d reported), and Arnica spathulata (one t e t r a -p l o i d reported). Both d i p l o i d and t e t r a p l o i d populations are known for Arnica discoidea. Arnica g r a c i l i s i s predominately t r i p l o i d , with one t e t r a -p l o i d and one hexaploid previously reported, although voucher specimens from these reports have not been seen. Arnica nevadensis i s t e t r a p l o i d , based on counts from three populations. Arnica c o r d i f o l i a i s the most v a r i a b l e , being predominately t e t r a p l o i d , with a few d i p l o i d , t r i p l o i d , pentaploid, and hexa-p l o i d populations known. Apomixis To determine i f plants were apomictic, heads were bagged with c l o t h bags (Figure 211) before the rays opened on radiate species, or before the outer d i s c s opened on d i s c o i d species. Barker (1966) showed that sexual d i p l o i d s are self-incompatible, and there i s a d d i t i o n a l data to support h i s f i n d i n g s . However, i t was assumed that known polyploid populations producing v i a b l e achenes from bagged heads were indeed apomicts rather than self-compatible amphimicts (sexual p l a n t s ) . To insure that pollen was being transferred from anthers to stigmas, heads were gently rubbed on several successive days, as the f l o r e t s matured and stigmas emerged from the surrounding anthers. When the achenes matured each was checked for contents. Arnica achenes are 61 Figures 187-194. Meiotic Chromosomes, Microsporocytes, and Pollen Grains of Arnica Species. 187. Arnica g r a c i l i s S-1613, BC, Peyto Lake, anaphase I (upper c e l l ) , t r i p l o i d with lagging chromosomes. 188. Late anaphase I of same. 189. Late anaphase II of same. 190. "Quintad" and "Sextad" of same. 191. Arnica l a t i f o l i a S-1961, BC, Nancy Greene P r o v i n c i a l Park, pollen grains stained with cotton blue i n lactophenol, showing two unstained grains. 192. Arnica nevadensis S-1910, NV, Washoe County, SEM of pollen grains showing large and small grains, X1200. 193. Arnica c o r d i f o l i a S-1416, CA, S i e r r a County, t e t r a p l o i d (2n=76) with 37 chromosomes at each pole of the c e l l and one lagging p a i r . 194. Arnica discoidea S-1787, CA, Napa County, d i p l o i d (2n=38) with 19 pa i r s of chromosomes at metaphase I. Bars = ca 50 Jim. 63 TABLE I I . Previously Reported Chromosome Numbers for Arnica Species i n the Subgenera Austromontana and Calarnica. Taxon Sporophytic Number L o c a l i t y Reference cernua 38 OR, Josephine Co. Barker, 1966 c o r d i f o l i a 76 BC, Lambly Creek Taylor & Taylor, 1977 c o r d i f o l i a 76 AB Ornduff, et a l . , 1967 c o r d i f o l i a 76 SK, Cypress H i l l s Taylor & Brockman, 19( P r o v i n c i a l Park c o r d i f o l i a ca76 WA, Stevens Co. Barker, 1966 c o r d i f o l i a 72-76 OR, Lake Co. Strother, 1972 c o r d i f o l i a ca76 ID, Lewis Co. Ornduff, et a l . , 1967 c o r d i f o l i a 38 MT, Missoula Co. Schaack, et a l . , 1974 c o r d i f o l i a ca90+6 CO, LaPlata Co. Ornduff, et a l . , 1967 c o r d i f o l i a ca95+4 CO, Huerfana Co. Ornduff, et a l . , 1967 c o r d i f o l i a ca95 CO, Larimer Co. Love & Kapoor, 1968 c o r d i f o l i a 60 Cult., USSR Arcto-Alpine Bot. Gdn. Zhukova, 1967 discoidea ca72-76 OR, Josephine Co. Ornduff, et a l . , 1963 (as p a r v i f l o r a ) discoidea 38 CA, T r i n i t y Co. Barker, 1966 (as p a r v i f l o r a ) discoidea 38 CA, Santa Cruz Co. Ornduff, et a l . , 1967 (as p a r v i f l o r a ) discoidea 76 CA, Lake Co. Ornduff, et a l . , 1963 discoidea 76 CA, Orange Co. Ornduff, et a l . , 1963 g r a c i l i s c a l l 4 AB, Mt. Temple Love & Love, 1964 64 Table II cont'd. Taxon g r a c i l i s  l a t i f o l i a l a t i f o l i a l a t i f o l i a l a t i f o l i a l a t i f o l i a l a t i f o l i a l a t i f o l i a l a t i f o l i a spathulata spathulata venosa viscosa viscosa Sporophytic Number ca76 38 38 38 38 38 38 38 ca76 ca76 38 38 38 38 L o c a l i t y ID, Valley Co. BC, Queen Charlotte Is., Graham I s . BC, Queen Charlotte Is., Moresby I s . AB WA, K i t t i t a s Co. WA, Whatcom Co. WA, Lewis Co. MT, R a v a l l i Co. MT, Lake Co. OR, Josephine Co. CA, Del Norte Co. CA, Shasta Co. CA, Siskiyou Co. CA, T r i n i t y Co. Reference Ornduff, et a l . , 1967 Taylor & Mulligan, 1968 Taylor & Mulligan, 1968 Ornduff, et a l . , 1967 Ornduff, et a l . , 1963 Ornduff, et a l . , 1963 Barker, 1966 Schaack, et a l . , 1974 Powell, et a l . , 1974 Barker, 1966 Ornduff, et a l . , 1967 Barker, 1966 Barker, 1966 Strother, 1973 65 TABLE I I I . New Chromosome Numbers for Arnica Species i n the Subgenera Austromontana and Calarnica. Taxon Sporophytic Number L o c a l i t y C o l l e c t i o n '. cernua 38 CA, Humboldt Co. S-1396 Horse Mtn. cernua 38 CA, Humboldt Co. S-1772 west of Hoopa c o r d i f o l i a 76 YT, Rancheria S-1853 c o r d i f o l i a 76 BC, Botanie Valley S-1342 c o r d i f o l i a 76 BC, Botanie Va l l e y S-1353 c o r d i f o l i a 76 BC, Fountain Valley S-1360 c o r d i f o l i a 76 BC, Bonanza Pass S-1549 c o r d i f o l i a 57 BC, southeast of S-1594 Windermere c o r d i f o l i a 57 BC, Tod Mtn. S-1652 c o r d i f o l i a 76 BC, Hat Creek S-2009 c o r d i f o l i a 76 BC, Sunday Summit S-2014 c o r d i f o l i a 57 BC, O l a l l a S-2022 c o r d i f o l i a 76 BC, Apex Mountain S-2025 c o r d i f o l i a 76 BC, north of Kamloops S-2028 c o r d i f o l i a i 76 WA, Okanogan Co., S-1494 Slate Lake c o r d i f o l i a 114 WA, Okanogan Co., S-1497 Slate Lake c o r d i f o l i a 76 WA, Okanogan Co., S-1498 Slate Lake 66 Table III cont'd. Taxon c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a c o r d i f o l i a Sporophytic Number 76 76 76 76 76 76 76 76 57 114 76 76 76 76 L o c a l i t y WA, K l i c k i t a t Co. , north of Lyle WA, K l i c k i t a t Co., Appleton OR, Wasco Co., Pine Grove OR, Wasco Co., Pine Grove CA, S i e r r a Co., Calpine CA, Placer Co., Lake Tahoe CA, Siskiyou Co., west of Meiss Lake CA, Mono Co. , Saddlebag Lake CA, Mono Co., Saddlebag Lake AB, V e r m i l l i o n Pass AB, Lake Louise AB, Moraine Lake AB, Peyto Lake ID, Bonneville Co., west of Aspen, WY Co l l e c t i o n Number S-1822 S-1823 S-1819 S-1820 S-1416 S-1425 S-1797 S-1937 S-1939 S-1601 S-1604 S-1610 S-1616 S-1836 67 T a b l e . I l l cont'd. Taxon Sporophytic Number L o c a l i t y C o l l e c t i o n '. c o r d i f o l i a 76 MT, Missoula Co., Pattee Canyon S-1477 c o r d i f o l i a 57 WY, Teton Co., Grand Teton Nat. Park S-1464 c o r d i f o l i a 57 WY, Teton Co., Flagg Ranch S-1465 c o r d i f o l i a 76 WY, Park Co., Yellowstone River, Upper F a l l s S-1837 c o r d i f o l i a 76 SD, Lawrence Co., Deadwood Gulch S-1842 c o r d i f o l i a 76 CO, Routt Co., Rabbit Ears Pass S-1437 c o r d i f o l i a 76 CO, Routt Co., Rabbit Ears Pass S-1438 c o r d i f o l i a 76 CO, Grand Co., Muddy Pass S-1440 c o r d i f o l i a 76 CO, Grand Co., Winter Park S-1443 c o r d i f o l i a 57 CO, Grand Co., Grand Lake S-2035 c o r d i f o l i a 76 CO, Clear Creek Co., Berthoud Pass S-1458 c o r d i f o l i a 76 CO, Boulder Co., Nederland S-1460 68 T a b l e . I l l cont'd. Taxon Sporophytic L o c a l i t y C o l l e c t i o n Number Number c o r d i f o l i a 76 MI, Keweenaw Co., S-1846 Copper Harbor c o r d i f o l i a 76 MI, Keweenaw Co. S-2005 c o r d i f o l i a 57 Cult., BC, UBC Bot. Gdn., S-1512 o r i g . from BC, Kootenay D i s t . c o r d i f o l i a 57 Cult., BC, UBC Bot. Gdn. S-1513 discoidea 76 WA, K l i c k i t a t Co., Glenwood S-1825 discoidea 76 OR, Wasco Co., Warm Springs S-1821 Indian Reservation discoidea 38 CA, Mendocino Co., S-1775 L i t t l e River discoidea 38 CA, Napa Co., south of S-1787 Napa-Lake Co. l i n e discoidea 38 CA, Marin Co., S-1789 Mt. Tamalpias g r a c i l i s 57 BC, Monroe Lake S-1562 g r a c i l i s 57 AB, Lake Louise S-1603 g r a c i l i s 57 AB, Moraine Lake S-1605 g r a c i l i s 57 AB, Peyto Lake S-1613 g r a c i l i s 57 Cult., BC, UBC Bot. Gdn. S-1762 l a t i f o l i a 38 BC, Mt. Seymour S-1503 l a t i f o l i a 38 BC, Mt. Seymour S-1504 l a t i f o l i a 38 BC, Vancouver Island S-1529 Forbidden Plateau 6.9 Table III cont'd. Taxon Sporophytic Number L o c a l i t y C o l l e c t i o n 1 l a t i f o l i a 38 BC, Vancouver Island S-1538 Forbidden Plateau l a t i f o l i a 38 BC, Bonanza Pass S-1548 l a t i f o l i a 38 BC, Kootenay Pass S-1561 l a t i f o l i a 38 BC, Manning Park S-1680 l a t i f o l i a 38 BC, Nancy Greene Prov. Pk. S-1961 l a t i f o l i a 38 BC, Black Tusk S-2031 l a t i f o l i a 38 BC, Cypress Bowl S-2041 l a t i f o l i a 38 WA, Mt. Whatcom Co., Baker S-1161 l a t i f o l i a 38 WA, Chelan Co., S-1499 Washington Pass l a t i f o l i a 38 WA, Chelan Co., S-1500 Rainy Pass l a t i f o l i a 38 AB, Peyto Lake S-1612 l a t i f o l i a 38 AB, south of Grand P r a i r i e S-1855 l a t i f o l i a 38 MT, Missoula Co., S-1476 Pattee Canyon nevadensis 76 CA, S i e r r a Co., S-1870 Si e r r a Buttes nevadensis 76 CA, Mono Co., Saddlebag Lk. S-1938 nevadensis 76 NV, Washoe Co., Mt. Rose S-1910 spathulata 38 OR, Josephine Co., Kirby S-1380 spathulata 38 OR, Josephine Co., east of S-1381 Cave Junction 70 Table I I I cont'd. Taxon Sporophytic L o c a l i t y C o l l e c t i o n Number Number spathulata 38 OR, Josephine Co., S-1385 O'Brien spathulata 38 OR, Josephine Co., Selma S-1740 spathulata 38 OR, Josephine Co., north S-1742 of O'Brien spathulata 38 CA, Del Norte Co., S-1390 northwest of Gasquet spathulata 38 CA, Del Norte Co., east S-1751 of Gasquet spathulata 38 CA, Del Norte Co., S-1392 northwest of Gasquet venosa 38 CA, Shasta Co., S-1791 Shasta B a l l y venosa 38 CA, Shasta Co., South Fork S-1792 Lookout Mtn. Road venosa 38 CA, Shasta Co., Gibson S-1793 viscosa 38 OR, Crater Lake Nat. Park, S-1946 G a r f i e l d Peak viscosa 38 CA, Siskiyou Co., Mt. Shasta S-1411 Putative Natural Hybrids c o r d i f o l i a - 38 MT, R a v a l l i Co., Charles S-1474 l a t i f o l i a Waters Campground c o r d i f o l i a - 38 MT, R a v a l l i Co., Charles S-1475 l a t i f o l i a Waters Campground 71 Figure 195. Arnica cernua S-1772, CA, Humboldt Co. Microsporocyte, Metaphase I plate, d i p l o i d with 19 pairs of chromosomes. Figure 196. Arnica l a t i f o l i a S-1529, BC, Vancouver Island, Forbidden Plateau. Microsporocyte, metaphase I pl a t e , d i p l o i d with 19 pairs of chromosomes. 72 73 Figure 197. Arnica c o r d i f o l i a S-1819, OR, Wasco County. Microsporocyte, l a t e anaphase I, t e t r a p l o i d with 38 chromosomes at each pole. Figure 198. Arnica c o r d i f o l i a S-1601, AB, Vermil l i o n Pass. Micro-sporocyte, metaphase I pl a t e , hexaploid with v a r i a b l e univalents, b i v a l e n t s , and t r i v a l e n t s . 74 ® 75 Figure 199. Arnica c o r d i f o l i a X Arnica l a t i f o l i a S-1475, putative natural hybrid, MT, R a v a l l i County. Microsporocyte, d i a k i n e s i s , d i p l o i d with 19 pairs of chromosomes. Figure 200. Arnica c o r d i f o l i a S-1594, BC, E. of Windermere. Microsporocyte, l a t e anaphase I, t r i p l o i d with 30 chromosomes at one pole of c e l l and 27 at the other. 76 77 Figure 201. Arnica g r a c i l i s S-1613, AB, Peyto Lake. Microsporocyte, metaphase I plate, t r i p l o i d , with v a r i a b l e univalents, b i v a l e n t s , and t r i v a l e n t s . Figure 202. Arnica nevadensis S-1870, CA, S i e r r a County. Microsporocyte, anaphase I, t e t r a p l o i d , with 76 chromosomes. 79 Figure 203. Arnica spathulata S-1392, OR, Josephine County. Microsporocyte, metaphase I pl a t e , d i p l o i d with 19 pai r s of chromosomes. Figure 204. Arnica discoidea S-1787, CA, Napa County. Microsporocyte, metaphase I plate, d i p l o i d with 19 pairs of chromosomes. Figure 205. Arnica discoidea S-1821, OR, Wasco County. Microsporocyte, metaphase I plate, t e t r a p l o i d with 38 pai r s of chromosomes. 80 81 Figure 206. Arnica viscosa S-1411, CA, Siskiyou County. Microsporocyte, d i a k i n e s i s , d i p l o i d with 19 pai r s of chromosomes. Figure 207. Arnica venosa S-1793, CA, Shasta County. Microsporocyte, metaphase I palte, d i p l o i d with 19 pai r s of chromosomes. 82 83 produced regardless of the presence or absence of a v i a b l e seed within. These empty achenes are often as large as v i a b l e achenes and appear to be v i a b l e u n t i l examined c l o s e l y or u n t i l the achene coat i s removed. According to Barker (1966) the two common types of apomixis found i n Arnica are: Autonomous apomixis - Seed production without p o l l i n a t i o n or f e r t i l i z a t i o n . Pseudogamous apomixis - Seed production does not occur unless the polar n u c l e i are f e r t i l i z e d to produce endosperm. Embryo development i s parthenogenetic. In addition to the bagging experiments, two emasculation methods were used to determine autonomous apomicts. In radiate populations disc f l o r e t s were c a r e f u l l y removed from heads before the discs opened and before the rays had elongated (Figure 209) . The head was then bagged, leaving the p i s t i l l a t e ray f l o r e t s to mature in the absence of p o l l e n . On other heads, either radiate or d i s c o i d , at the same stage of development, the whole top of the head from j u s t above the ovary was removed with a razor blade, thus removing a l l anthers, stigmas, and the upper portion of the s t y l e s . The heads were then bagged and allowed to mature. Although severe, t h i s treatment seems to have no detrimental e f f e c t s on v i a b l e achene production i n autonomous apo-micts. The r e s u l t s of these bagging and emasculation procedures are shown i n Table IV. The breeding systems of the nine species are: Sexual (self-incompatible) Arnica cernua (Barker, 1966) Arnica discoidea populations (Barker, 1966, as Arnica p a r v i f l o r a ) 84 Figures 208-211. Experimental methods. Figure 208. Arnicas i n c u l t i v a t i o n i n experimental f i e l d , UBC Botanical Garden Nursery. Figure 209. Head of Arnica c o r d i f o l i a S-1497, WA, Okanogan County, with di s c f l o r e t s removed. Figure 210. Growth Chamber with plants i n flower, seedlings i n pots, and seeds germinating i n p e t r i dishes. Figure 211. Arnica head bagged. 85 TABLE IV. Compatibility and Apomixis Experiments with Arnica species. Taxon & Diploid C o l l . // Chromosome // Pollen S t a i n a b i l i t y it of Heads Selfed // of Heads Emasculated it of Achenes Produced // of Achenes With Seeds % of Ac With S. c o r d i f o l i a 1342 76 69% 4 - 111 5 4.5 c o r d i f o l i a 1343 1 89 4 - 221 38 17.2 c o r d i f o l i a 1343 1 89 - 3 101 0 0 c o r d i f o l i a 1353 76 83 2 - 112 7 6.2 c o r d i f o l i a 1353 76 83 - 2 55 8 14.5 c o r d i f o l i a 1359 1 38 2 - 96 13 13.5 c o r d i f o l i a 1359 1 38 - 1 7 0 0 c o r d i f o l i a 1360 76 42 4 - 157 2 1.3 c o r d i f o l i a 1416 76 24 3 - 272 67 24.6 c o r d i f o l i a 1458 76 82 1 - 189 0 0 c o r d i f o l i a 1458 76 82 - 1 71 0 0 c o r d i f o l i a 1477 76 32 6 - 371 24 6.5 c o r d i f o l i a 1477 76 32 - 2 46 0 0 c o r d i f o l i a 1487 ? 49 1 - 52 9 17.3 c o r d i f o l i a 1494 76 94 3 - 218 0 0 c o r d i f o l i a 1494 76 94 - 2 52 0 0 c o r d i f o l i a 1497 114 84 8 — 485 100 20.6 Table IV cont'd • Taxon & C o l l . # Diploid Chromosome # Pollen S t a i n a b i l i t y // of Heads Selfed // of Heads Emasculated // of Achenes Produced // of Achenes With Seeds % of Achenes With Seeds c o r d i f o l i a 2005 76 81 3 - 314 4 1.3 c o r d i f o l i a 2005 76 81 - 1 7 0 0 discoidea 1821 76 76 1 - 49 0 0 discoidea 1825 76 77 4 - 129 15 11.6 discoidea 1825 76 77 - 1 21 0 0 g r a c i l i s 1562 57 0 10 - 222 187 84.2 g r a c i l i s . 1562 57 0 - 4 72 50 69.4 g r a c i l i s '. 1605 57 3 4 - 93 64 68.8 g r a c i l i s 1605 57 3 - 2 54 18 33.8 l a t i f o l i a 1499 38 87 1 - 19 0 0 l a t i f o l i a 1499 38 87 - 1 36 0 0 l a t i f o l i a 1503 38 97 10 - 301 0 0 l a t i f o l i a 1504 38 97 2 - 69 0 0 l a t i f o l i a 1504 38 97 - 3 178 0 0 l a t i f o l i a 1561 38 75 2 - 57 0 0 l a t i f o l i a 1961 38 90 1 • _ 28 0 0 Table IV cont'd. Taxon & Diploid C o l l . // Chromosome # l a t i f o l i a 1961 38 nevadensis 1870 76 nevadensis 1938 76 nevadensis 1938 76 spathulata 1380 38 venosa 1940 38 viscosa 1411 38 Hybrids co r d . - l a t . 1474 38 cor d . - l a t . 1475 38 Pollen # of Heads // of Heads # of Achenes // of Achenes % of Achenes S t a i n a b i l i t y Selfed Emasculated Produced With Seeds With Seeds 90 - 1 8 0 0 43 2 - 122 59 48.4 3 3 - 155 65 41.9 3 - 3 74 23 31.1 99 3 - 113 0 0 100 4 108 0 0 100 20 - 302 0 0 7 7 12 - 1570 0 0 34 20 - 2051 0 0 89 Arnica l a t i f o l i a Arnica spathulata populations Arnica venosa Arnica viscosa Autonomous Apomicts Arnica c o r d i f o l i a populations Arnica g r a c i l i s Arnica nevadensis Pseudogamous Apomicts Arnica c o r d i f o l i a populations Arnica discoidea populations Arnica spathulata populations (Barker, 1966) P o l l e n S t a i n a b i l i t y For pollen s t a i n a b i l i t y t e s t s , contents of f r e s h l y dehisced anthers of one or more f l o r e t s of a plant from a w i l d - c o l l e c t e d population or herbarium specimen was tapped out on a glass s l i d e . Pollen grains were stained with a drop of cotton blue-lactophenol and a cover s l i p was added. The s t a i n a b i l i t y of a 200-grain random sample f o r each population was scored a f t e r 24 hours. Large, well-formed grains stained dark blue were scored as "good". Those that did not s t a i n or were small and misshapened were scored as "bad" (Figure 191). The range of p o l l e n s t a i n a b i l i t y f o r each species i s shown i n Figures 212-221. Several of the species are so rare in nature and i n herbaria that few specimens could be sampled. Barker (1966) discussed the c o r r e l a t i o n between ploidy l e v e l , apomixis, and pollen s t a i n a b i l i t y . The data presented here support his f i n d i n g s . 90 There are some d i s t i n c t trends among c e r t a i n species. The ex c l u s i v e l y or predominately d i p l o i d , sexual species including Arnica l a t i f o l i a , Arnica  cernua, Arnica spathulata, Arnica venosa, and Arnica viscosa a l l show very high p o l l e n s t a i n a b i l i t y , mostly greater than 90%. A second group of species includes Arnica g r a c i l i s and Arnica nevadensis which have mostly l e s s than 10% stainable p o l l e n . These two are known to be or presumed to be autonomous apomicts and mostly or e n t i r e l y p o l y p l o i d s . A number of populations of Arnica g r a c i l i s sampled had no pollen produced i n any f l o r e t s . In known t r i p l o i d populations of Arnica g r a c i l i s the production of small and unstainable normal sized p o l l e n grains (Figures 191, 192) can be followed from i r r e g u l a r p a i r i n g of chromosomes to the production of more than four groups of chromosomes a f t e r anaphase I I , the normal tetrad stage (Figure 189). Walls often form around these "quintads" or "sextads" r e s u l t i n g i n micro pollen grains and normal sized grains (Figures 187-192). Even these large grains apparently lack a f u l l complement of chromosomes and often f a i l to develop into a v i a b l e , or at l e a s t stainable, p o l l e n grain. A t h i r d group includes Arnica c o r d i f o l i a and the c l o s e l y related Arnica  discoidea i n which pollen s t a i n a b i l i t y i s e r r a t i c from 0-100% and does not seem to indi c a t e anything s p e c i f i c a l l y about the ploidy l e v e l or apomixis. In Arnica discoidea there are both d i p l o i d populations with high pollen s t a i n -a b i l i t y and t e t r a p l o i d populations with lower pollen s t a i n a b i l i t y . In Arnica  c o r d i f o l i a no d e f i n i t e sexual populations are known and f o r the known poly-p l o i d populations pollen s t a i n a b i l i t y i s v a r i a b l e . The best assumption f o r Arnica c o r d i f o l i a i s that lower pollen s t a i n a b i l i t y probably represents auto-nomous apomictic populations and the higher p o l l e n s t a i n a b i l i t y represents pseudogamous apomictic populations. The few d i p l o i d populations which are 91 jures 212-221. Range of pollen s t a i n a b i l i t y f o r Arnica populations sampled. cordifolia . A t : . •••• • ••••• • t • •••• • • • ••••••• Number of populations latifolia ••••• • ••••• ••••••• cordifolia-latifolia hybrids @ ) 10 2 0 ^ 0 " 4 0 % Pollen Stainability 5 0 6 0 70 8 0 9 0 100 nevadensis •••• • Number of populations cernua (216 - i 1 r-^ r-~l r- ^ "I 1— T grac ilis © : :. » » M • ( 1 r r- -i r-10 2 0 3 0 4 0 5 0 - i 1 r-6 0 7 0 8 0 9 0 100 % Pollen Stainability Number of populations T 1-- r — 1 r- -i r-discoidea © 9 9 Q QQ • •• e e ~1 r- • ••••••• • spathulata @ ) • •• • •••• venosa -i r--i 1 r- —i 1 1 viscosa 10 2 0 3 0 4 0 ' 5 0 ~i 1— 6 0 7 0 8 0 9 0 100 < Pollen Stainability 95 known to exi s t may be sexual, although they have not been investigated experimentally. Seed Production Seed production both i n nature and i n c u l t i v a t i o n i s often quite low even i n apomictic races. Although several dozen to a hundred or more achenes are produced i n each head, many of these are empty. Also, i n many populations of most or a l l species the heads are infested with one to several larvae of achene-feeding dipterans which often feed on most or a l l of the achenes i n a head before the pappus expands and the seeds are dispersed. (See further discussion under Entomological Relationships). Seed production i n c u l t i v a t i o n where there were no dipteran i n f e s t a t i o n was s t i l l low except i n the autonomous apomictic populations of Arnica g r a c i - l i s and Arnica nevadensis. Of achenes examined from 21 heads of Arnica  g r a c i l i s 54-100% (mostly greater than 80%) of the achenes contained seed. Eight heads of Arnica nevadensis had 0-70% (mostly greater than 60%) f u l l achenes. In the sexual populations of Arnica l a t i f o l i a 0-72% (mostly l e s s than 20%) of achenes from 25 heads examined contained seeds. Only wild c o l -l e c t e d achenes of Arnica cernua have been observed, with 6-14% f u l l achenes from 5 heads from one population. Of the sexual, self-incompatible popula-tions of Arnica spathulata, Arnica viscosa, and Arnica venosa, only one each has been grown s u c c e s s f u l l y to the flowering and f r u i t i n g stage. Therefore, no data of seed production are a v a i l a b l e . One head of Arnica venosa c o l l e c t e d i n the wild had 47% f u l l achenes. There i s poor seed set i n c u l t i v a t i o n of the predominately pseudogamous apomictic populations of Arnica c o r d i f o l i a and Arnica discoidea. Of more than 96 50 heads of Arnica c o r d i f o l i a examined, seed production was 0-64% (mostly les s than 30%) of achene production. Wild c o l l e c t e d achenes had 0-29% f u l l achenes. In Arnica discoidea 0-20% of the achenes from 16 heads examined were f u l l . Seed Germination Young seedlings i n t h e i r alpine or montaine environment would be at a disadvantage i f the achenes germinated immediately i n l a t e summer or f a l l a f t e r f a l l i n g or being c a r r i e d from the parent plant. I t was suspected that the achenes needed a v e r n a l i z a t i o n or s c a r i f i c a t i o n of the achene coat before the seeds would germinate. Germination probably occurs i n nature a f t e r the spring snow-melt. Achenes containing seeds were subjected to numerous schemes i n an e f f o r t to induce germination, including the following: 1. Planted i n pots of s o i l out-of-doors i n the f a l l and allowed to overwinter under "natural" ( i . e . , Vancouver winter) conditions. 2. Stored dry or i n moist sphagnum i n the r e f r i g e r a t o r (4°C) or freezer (0°C) or dry at room temperature (ca 20°C) f o r six weeks to several months before planting. 3. Achenes placed immediately a f t e r d i s p e r s a l or a f t e r storage on moist f i l t e r paper i n p e t r i dishes i n l i g h t (16 hours daylight) at 25°C. (Figure 210) . 4. Same as #3 except t o t a l darkness. 5. Achene coats c a r e f u l l y removed a f t e r they had been soaked i n d i s t i l l e d water f o r 24 hours, then placed on moist f i l t e r paper i n 16 hours daylight at 25°C. 97 The r e s u l t s of these various tests have been mixed and there are s t i l l some unanswered questions concerning seed germination. The greatest success was had with soaked achenes, with achene coats removed, and then placing the seeds on moist f i l t e r paper under long days at 25°C. In no cases did any seed germinate on moist f i l t e r paper before the achene coat was removed. Afte r removal, the hypocotyl often began to elongate within 12 hours. Using the above method there was nearly 100% germination of seeds from some populations of some species, e s p e c i a l l y Arnica c o r d i f o l i a , Arnica g r a c i -l i s , and Arnica v i s c o s a . With other populations of the same species or other species, e s p e c i a l l y Arnica c o r d i f o l i a , Arnica nevadensis, Arnica discoidea, and Arnica spathulata, none of the seeds germinated with t h i s , or any other method. In some cases a few days a f t e r the achene coat was removed, the cotyledons reflexed, but there was no further sign of growth, even a f t e r months. In yet other populations of Arnica c o r d i f o l i a , and e s p e c i a l l y Arnica  venosa, a few seeds germinated immediately, while others remained plump and apparently capable of germination f o r up to si x months, yet there were no signs of hypocotyl or e p i c o t y l elongation. Often during the longer tests the dishes became infested with mold or algae, so that the seeds had to be d i s -carded . Ve r n a l i z a t i o n i s not needed f or germination. Under a r t i f i c i a l conditions germination was not enhanced by cooling or freezing the achenes. However, s c a r i f i c a t i o n of the achene coats seems to be e s s e n t i a l f o r germination. Achenes planted d i r e c t l y i n s o i l out-of-doors did not germinate the following spring. The achenes were very d i f f i c u l t to locate to determine i f they s t i l l were v i a b l e or i f they had decomposed during the winter. 98 Seedlings Of more than 200 populations studied i n the f i e l d , i n c l u d i n g a l l taxa recognized i n t h i s study, no seedlings were observed. Seedlings grown from seeds germinated i n the growth chamber were trans-planted into a s t e r i l i z e d standard potting medium as soon as the f i r s t p a i r of true leaves s t a r t e d to develop. These were grown i n the growth chamber under the same growing conditions as f o r germination. M o r t a l i t y rate of the young seedlings was r e l a t i v e l y high f o r the f i r s t few weeks. Most healthy seedlings were very slow growing, taking 1-3 months to develop three p a i r s of leaves. Seedlings of the serpentine endemic Arnica cernua grown both i n potting medium and in serpentine s o i l grew equally slowly, the basal rosette reaching only 2-3 cm i n diameter and with 3-4 pai r s of leaves a f t e r 6-8 months. Others, e s p e c i a l l y A r n i c a c o r d i f o l i a , Arnica v i s c o s a , and Arnica  venosa grew qui c k l y a f t e r the t h i r d or fou r t h month and flowered by the s i x t h month (Figure 278) . One seedling of Arnica v i s c o s a had become much branched and produced more than 100 flowers by the eighth month. Seedlings of Arnica  l a t i f o l i a and Arnica g r a c i l i s produced robust basal rosettes and some rhizome growth within 6 months but no flowering stems were produced Two-month o l d seedlings grown i n a growth chamber are shown i n Figures 222-227. Even when quite young (by the f i r s t p a i r of true leaves) many of the seedlings assume the t y p i c a l characters of the more mature plant, i n lea f shape, roots, and e s p e c i a l l y i n pubescence, although the seedlings are often more pubescent when young than they are l a t e r . A f t e r a few weeks the Austromontana species remain as a basal rosette with no stem elongation, while the Calar n i c a species form a d e f i n i t e stem by the time the second or t h i r d p a i r of true leaves have developed. 99 Figures 222-227. Arnica Seedlings ca 2 months a f t e r germination, grown i n growth chamber. Figure 222. Arnica c o r d i f o l i a S-1497, WA, Okanogan County. Figure 223. Arnica l a t i f o l i a S-1855, AB, south of Grande P r a i r i e . Figure 224. Arnica g r a c i l i s S-1562, BC, Monroe Lake. Figure 225. Arnica cernua S-1772, OR, Josephine County. Figure 226. Arnica viscosa S-1411, CA, Siskiyou County. Figure 227. Arnica venosa S-1940, CA, Shasta County. 100 101 Vegetative Reproduction Considering the low seed production, i n f e s t a t i o n s of achene feeding Diptera, and poor germination of seeds, vegetative reproduction by rhizomes i n species of subgenus Austromontana i s c r i t i c a l f o r t h e i r successful c o l o n i -zation. L i t t l e or no information e x i s t s on vegetative reproduction p o t e n t i a l i n Arnicas. Herbarium specimens are of l i t t l e value. Although Arnicas make nice herbarium specimens ( i . e . , they f i t conveniently on a standard herbarium sheet) and they are frequently c o l l e c t e d and are well-represented i n herbaria, plants are usually pulled from the ground rather than dug, r e s u l t i n g i n the rhizomes being l e f t behind. Specimens with rhizomes usually show only one season's growth or a portion thereof. No known previously c o l l e c t e d s p e c i -mens of the two species i n subgenus Calarnica had enough of the underground parts present to determine i f they were or were not rhizomatous. Records of annual rhizome growth have been kept on populations of a l l species seen i n the f i e l d i n addition to several years of records of growth of some populations i n c u l t i v a t i o n (Figures 228-231, Table V). These studies have been supplemented by measurements from herbarium specimens. In most i n -stances the average growth i s lower than the median. A few populations grow-ing i n r e l a t i v e l y dense shade or i n very loose s o i l s tend to have the ex-tremely long rhizomes. Examples are the low elevation populations of Arnica  g r a c i l i s from f o r e s t s at Monroe Lake, BC (Straley 1562) with rhizomes up to 6 cm long, whereas a majority of the populations as represented by those from high alpine rock scree slopes of Banff and Jasper National Parks, AB (Straley  1603, 1605, 1613) average only 1-2 cm of growth per year. 102 Figure 228. Rhizome growth measured over four years, Arnica c o r d i f o l i a  S-1512, UBC Botanical Garden, Native Garden. Plant o r i g i n a l l y from Kootenay D i s t r i c t , BC. 2 0 c m Original Plant-1976 Growth -1977 1978 1979 104 Figure 229. Rhizome growth measured over 4 years, Arnica l a t i f o l i a  S-1162, WA, Whatcom County, Mt. Baker. 105 106 Figure 230. Rhizome growth measured over four years, Arnica cernua  S-1772, CA, Humboldt County, Hoopa Mountain. 2cm Original Plant - 1975 Growth -1976 1977 1978 108 Figure 231. Rhizome growth measured over 3 years, Arnica g r a c i l i s  S-1605, AB, Moraine Lake. 109 • - Original Plant -1977 O - 1978 A - 1979 110 TABLE V. Yearly Rhizome Growth i n Arnica, Subgenus Austromontana Species. Taxon Annual Growth (cm) Annual Average (cm) Breeding System Average Seed V i a b i l i t y cernua (l-)4-15(-20) 6 Sexual 14% c o r d i f o l i a 2-20(-60) 10 Apomictic 30% discoidea 5-15(-22) 7 Sexual and Apomictic 20% g r a c i l i s (0.5-)l-2(-6) 1.5 Apomictic 80% l a t i f o l i a (0.5-)l-7(-10) 4 Sexual 20% nevadensis (l-)3-12 6 Apomictic 60% spathulata (1-)3-7(-14) 5 Sexual and Apomictic ? I l l There seems to be no d i s t i n c t c o r r e l a t i o n between rhizome production and successful seed set or between the rhizome production of predominately sexual compared to apomictic species. However, there are some consistent differences between average rhizome growth of some of the c l o s e l y r e l a t e d species. The average growth i n Arnica c o r d i f o l i a , f o r instance, i s 2.5 times that of Arnica l a t i f o l i a and the growth of Arnica g r a c i l i s i s l e s s than ha l f as great as that of Arnica l a t i f o l i a . The derived species (Arnica cernua and Arnica g r a c i l i s from Arnica l a t i f o l i a , and Arnica discoidea, Arnica spathulata and Arnica nevadensis from Arnica c o r d i f o l i a ) a l l have lower rhizome produc-t i o n than do t h e i r presumed ancestral species. Several of these species frequently invade l a n d s l i d e s , burned-over areas, and other disturbed habitats. P a r t i c u l a r l y i n Arnica c o r d i f o l i a the yearly rhizome growth rates shown here could e a s i l y explain the vast coverage of an area by a single population a f t e r hundreds or thousands of years, from only one successful achene introduction. CULTIVATION OF ARNICA SPECIES Arnica populations have been transplanted from nature or grown from seed i n the growth chamber, greenhouse, and experimental f i e l d at UBC with mixed r e s u l t s . Some species, e s p e c i a l l y Arnica c o r d i f o l i a , Arnica nevadensis, and Arnica  g r a c i l i s have been most e a s i l y transplanted, with near 100% success, and with most populations flowering f r e e l y a f t e r transplanting. Arnica spathulata, Arnica discoidea, and Arnica viscosa have given mixed r e s u l t s . Some popula-tions transplanted e a s i l y and flowered seasonally for several years. Others e i t h e r did not survive or have l i v e d f o r only a few months and flowered poorly 112 or not at a l l . The widespread Arnica l a t i f o l i a u s ually survives moving, but flowers very poorly i n future years, although i t reproduces abundantly by rhizomes i n c u l t i v a t i o n . The serpentine endemic Arnica cernua has not been su c c e s s f u l l y transplanted. A few plants have survived f o r a year or two i n the experimental garden but have become progressively smaller and have never flowered i n c u l t i v a t i o n . A few achenes have germinated and the r e s u l t i n g seedlings have grown very slowly both i n serpentine s o i l s and in a standard s o i l - l e s s potting mixture. Two plants from d i f f e r e n t populations of the rare Arnica venosa have been unsuccessfully transplanted from the wild. However, un l i k e Arnica cernua, seedlings of Arnica venosa from achenes germinated on f i l t e r paper and transplanted to a s o i l - l e s s potting mixture have grown quickly and flowered f r e e l y i n c u l t i v a t i o n . In a l l cases, the tops died back to the rhizomes when plants were dug from the wild while i n flower or j u s t before or a f t e r . However, i n most cases, there were new sprouts produced from the rhizomes within a few weeks of transplanting, often producing flowering stems several weeks l a t e r . HYBRIDIZATION Sympatric Occurrence and Natural Hybridization There i s very l i t t l e evidence of recent h y b r i d i z a t i o n occurring i n nature between any of the species i n Austromontana and Calarnica, or with species i n other subgenera. There are a number of populations e x h i b i t i n g intermediate characters between Arnica c o r d i f o l i a and Arnica l a t i f o l i a , usually not growing with one or either parent. These are considered old s t a b i l i z e d hybrids, usu-a l l y apomicts, and w i l l be discussed under Arnica c o r d i f o l i a . 113 Because of d i f f e r e n t e c o l o g i c a l preferences, most of the nine species i n these two subgenera do not occur sympatrically. Where two or more species do grow i n close proximity or at l e a s t close enough f o r pollen exchange by insect p o l l i n a t o r s (see Table VI), they often have l i t t l e or no overlap i n flowering times. This was observed i n populations of Arnica discoidea and Arnica c o r d i f o l i a growing sympatrically i n Wasco County, OR (Straley 1820, 1821) and K l i c k i t a t County, WA (Straley 1825, 1826). In both of these popu-l a t i o n s the Arnica c o r d i f o l i a was f i n i s h i n g flowering as the Arnica discoidea was j u s t beginning to flower. A mixed population of Arnica cernua and Arnica  spathulata (Straley 1385, 1386) i n Josephine County, OR was observed i n May of 1977 and 1978. The Arnica cernua was j u s t f i n i s h i n g flowering as the Arnica spathulata was j u s t beginning to flower. These two species have also been c o l l e c t e d together i n Del Norte County, CA ( K l i p f e l 770, HSC), with no mention of any evidence of h y b r i d i z a t i o n . In most of the mixed populations there are also differences i n ploidy l e v e l s , breeding systems, and v i a b l e p o l l e n production, acting singly or usu-a l l y i n combination as b a r r i e r s to natural h y b r i d i z a t i o n . A specimen from Custer County, ID (Cronquist 3375, UTC) represents an apparent hybrid population growing with both parents, Arnica g r a c i l i s and Arnica l o n g i f o l i a . The specimen i s quite intermediate between the two paren-t a l types. This i s the only herbarium specimen seen of what appears to be a hybrid population between two d i s t i n c t species i n two d i f f e r e n t subgenera (Austromontana and Chamissonis). 114 TABLE VI. Sympatric populations of Arnica subgenus Austromontana species with no evidence of natural hybrids. C o l l e c t i o n L o c a l i t y Pollen D i p l o i d Breeding Number S t a i n a b i l i t y Chromosome System Number spathulata S -1385 OR, Josephine Co. 100% 38 sexual cernua s. -1386 OR, Josephine Co. 98% - sexual c o r d i f o l i a S--1573 BC, Yahk Mountain 30% - apomictic l a t i f o l i a s--1576 BC, Yahk Mountain 100% - sexual g r a c i l i s S--1603 AB, Lake Louise 0% 57 apomictic c o r d i f o l i a S--1604 AB, Lake Louise 52% 76 apomictic l a t i f o l i a S--1612 AB, Peyto Lake 99% 38 sexual g r a c i l i s S--1613 AB, Peyto Lake 2% 57 apomictic c o r d i f o l i a S--1616 AB, Peyto Lake 32% 76 apomictic c o r d i f o l i a S--1653 BC, Tod Mountain 74% - apomictic l a t i f o l i a S--1654 BC, Tod Mountain 95% - sexual c o r d i f o l i a S--1820 OR, Wasco County 85% 76 apomictic discoidea s--1821 OR, Wasco County 76% 76 apomictic discoidea s--1825 WA, K l i c k i t a t Co. 77% 76 apomictic c o r d i f o l i a s--1826 WA, K l i c k i t a t Co. 80% - apomictic nevadensis s--1938 CA, Mono County 3% 76 apomictic c o r d i f o l i a s--1938 CA, Mono County 8% 57 apomictic 115 A r t i f i c i a l H ybridization Reciprocal crosses between a l l the species i n these two subgenera are complicated by several f a c t o r s . The d i f f i c u l t y i n c u l t i v a t i o n of some species or populations has been previously discussed. Getting plants to flower con-currently i n c u l t i v a t i o n i s often a problem. In the experimental garden under natural spring conditions, most species produced only one set of flowering stems and the flowering duration was quite b r i e f , e s p e c i a l l y i n Arnica l a t i - f o l i a . While most of the montane and alpine species flower at the same time, Arnica viscosa takes several months f o r the flower stems to elongate, thus flowering much l a t e r than most other species. Under controlled conditions of the growth chamber Arnica viscosa, Arnica venosa, and some populations of other species could be kept flowering f o r many months, enabling experimental crosses with these and other more rap i d l y flowering species. In radiate forms, the ray f l o r e t s may be used as the female parent i n crosses, but the r e c i p r o c a l crosses cannot be made using the d i s c o i d forms with t h e i r complete f l o r e t s as the female parent, without f i r s t proving the dis c o i d form i s self-incompatible. Emasculation of the small d i s c o i d f l o r e t s before anthesis i s tedious and often r e s u l t s i n damage to the stigma or loss of the whole f l o r e t . Polyploidy and apomixis are further complications, with sexual populations not known i n several species, e s p e c i a l l y Arnica c o r d i f o l i a , Arnica nevadensis, and Arnica g r a c i l i s . Not only can these species not be used as the female parent i n r e c i p r o c a l crosses, but with very low p o l l e n f e r t i l i t y i n many of them, they cannot be used as a p o l l e n source f o r crosses. The r e s u l t s of a l l r e c i p r o c a l crosses are given i n Table VII and summarized i n Figure 232. TABLE VII. A r t i f i c i a l Hybridizations Using Sexual Populations (known or supposed) of Arnica Species. Taxon & Co l l e c t i o n # (tf Parent) Diploid Chromosome Taxon & // C o l l e c t i o n // ((^ Parent) Di p l o i d Chromosome # Heads Crossed Achenes Produced Achenes With Seeds l a t i f o l i a 1499 38 c o r d i f o l i a 1497 76 3 9 0 l a t i f o l i a 1504 38 c o r d i f o l i a 1512 57 2 13 7 l a t i f o l i a 1961 38 c o r d i f o l i a 2009 76 2 64 0 l a t i f o l i a 1961 38 discoidea 1821 76 1 56 0 l a t i f o l i a 1961 38 d i v e r s i f o l i a 1419 76 3 118 3 l a t i f o l i a 1961 38 g r a c i l i s 1605 57 1 28 0 l a t i f o l i a 1961 38 viscosa 1411 38 1 53 0 l a t i f o l i a 1503 38 mo l l i s 1515 ? 1 8 0 l a t i f o l i a 1504 38 mo l l i s 1515 1 1 9 0 spathulata 1380 38 c o r d i f o l i a 1477 76 2 69 3 spathulata 1380 38 c o r d i f o l i a 1497 76 O z. 37 0 venosa 1940 38 c o r d i f o l i a 2005 76 1 53 0 venosa 1940 38 cord.-lat. hyb. 1475 38 1 25 0 venosa 1940 38 l a t i f o l i a 2041 38 1 26 0 venosa 1940 38 nevadensis 1938 76 1 26 0 venosa 1940 38 viscosa 1411 38 1 27 0 Table VII cont'd. Taxon & D i p l o i d C o l l e c t i o n # Chromosome (0* Parent) viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 viscosa 1411 38 Hybrids cord.-lat. 1474 38 cord. - l a t . 1474 38 cord. - l a t . 1475 38 cord. - l a t . 1475 38 cord. - l a t . 1475 38 cord.' - l a t . 1475 38 Taxon & Diploid C o l l e c t i o n // Chromosome # (0_ Parent) c o r d i f o l i a 1343 ? c o r d i f o l i a 2005 76 l a t i f o l i a 1680 38 l a t i f o l i a 1961 38 l a t i f o l i a 2041 38 cord.-lat. hyb. 1474 38 cord.-lat. hyb. 1475 38 discoidea 1825 76 nevadensis 1870 76 venosa 1940 38 spathulata 1380 38 viscosa 1411 38 c o r d i f o l i a 1353 76 discoidea 1825 76 spathulata 1380 38 viscosa 1411 38 Heads Crossed Achenes Achenes Produced With Seeds 2 34 5 1 22 0 1 16 0 1 19 5 1 13 2 2 45 0 3 55 0 5 110 9 2 41 0 2 36 2 1 14 0 1 17 0 1 61 0 2 39 0 1 5 0 3 40 0 118 Figure 232. Summary of a l l a r t i f i c i a l crosses attempted between species i n subgenus Austromontana and Calarnica. S o l i d l i n e s indicate successful crosses. Dotted l i n e s , unsuccessful crosses. 119 120 Many of the seedlings from crosses were very weak and did not survive beyond a few days or weeks, or did not survive the transplanting from p e t r i dishes to s o i l . As of t h i s w r i t i n g several seedlings are s t i l l too young to determine very much about t h e i r characters except some vegetative ones. Seedlings of Arnica viscosa X Arnica venosa do ex h i b i t intermediate vegetative characters while quite young. The leaves are densely glandular as i n Arnica v i s c o s a , but with toothed margins as i n Arnica venosa. One seedling from a cross between Arnica l a t i f o l i a and Arnica v i s c o s a has grown r a p i d l y and has flowered. I t i s intermediate i n most respects. The leaves are la r g e , but nearly s e s s i l e and there are some glandular h a i r s and the odor of Arnica  v i s c o s a , yet greener than t h i s parent. The few heads were narrow but b r i g h t yellow and with a few short, twisted ray f l o r e t s . The pappus i s tawny and the achenes have a few glandular h a i r s . ENTOMOLOGICAL INTERRELATIONSHIPS Although many of the Arnicas have showy heads and are a conspicuous part of the montane-alpine-arctic f l o r a , very l i t t l e has been recorded on p o l l i n a -t i o n or any other p l a n t - i n s e c t r e l a t i o n s h i p s , other than work done by G r i f f i t h s (1974) on host s p e c i f i c Phytomyza f l y l e a f miners. Kunth (1908) has compiled records of a few insect v i s i t o r s to Arn i c a  montana (subgenus Montana), and some North American species c u l t i v a t e d i n bo-t a n i c a l gardens i n Europe. Many of these p o l l i n a t o r s were l i s t e d only as "bees" or " f l i e s " , etc. He records Arnica montana as being autogamous or geitonogamous ( p o l l i n a t e d by adjacent f l o r e t s i n the same head), although the fin d i n g s of Barker (1966) and i n t h i s current work suggest that a l l sexual 121 populations of Arnicas are probably self-incompatible. Leaf Miners G r i f f i t h s (1974) records seven species of Phytomyza (Diptera:Agromyzidae) which form mines i n the leaves of Arnica species. With only one exception a l l these are confined to the genus Arnica. However, there seems to be no species-s p e c i f i c Phytomyza. Some of the Phytomyza species can be i d e n t i f i e d from the pattern of the mine, as they tunnel below the epidermis. During t h i s study mines were seen on a l l species in.subgenus Austromon-tana, although none have been observed on e i t h e r species of C a l a r n i c a . In most instances the mines occupy h a l f or l e s s of the l e a f by the time the l a r -vae pupate. There are some severe i n f e s t a t i o n s observed i n which most of the leaves on the plants have the whole surface mined by several miners per l e a f . A l l c o l l e c t i o n s i d e n t i f i e d are given i n Table V I I I , and representative mines are shown i n Figures 233-240. No adult f l i e s have been bred from e i t h e r Arnica cernua or Arnica spathulata, so the species i n f e s t i n g those Arnicas are not known. F l o r e t Feeders There are occasional large h o s t - s p e c i f i c or omnivorous herbivores i n c l u d i n g inch worms (Geometriidae) or cutworms, e s p e c i a l l y Autographa simplex Guenee (Noctuidae), which may devour leaves, heads and f l o r e t s . There are also a number of smaller herbivores which feed on Arnica f l o r e t s . At l e a s t h a l f of the populations observed i n the f i e l d of both r a d i a t e and d i s c o i d species had one or many small beetles (Buprestidae, mostly i n the genus Anthaxia) feeding on the ray and/or d i s c f l o r e t s (Table IX). In some extreme 122 TABLE VIII. Arnica specimens c o l l e c t e d with l e a f mines of Phytomyza species. Phytomyza oreas G r i f . or Phytomyza a r h i c i v o r a Sehgal Arnica c o r d i f o l i a S-1472, MT, Lewis & Clark County Arnica c o r d i f o l i a S-1553, BC, Nancy Green P r o v i n c i a l Park Arnica c o r d i f o l i a S-1604, AB, Lake Louise Arnica c o r d i f o l i a S-1616, AB, Peyto Lake Arnica c o r d i f o l i a S-1819, OR, Wasco County Arnica c o r d i f o l i a S-1826, WA, K l i c k i t a t County Arnica c o r d i f o l i a Bohm 1109, BC, Osoyoos Arnica c o r d i f o l i a Guppy, _s.n_., BC, Botanie Valley Arnica l a t i f o l i a S-1676, WA, Clallam County Arnica l a t i f o l i a S-1680, BC, Manning Park Arnica nevadensis S-1870, CA, S i e r r a County Phytomyza saximontana G r i f . Arnica l a t i f o l i a S-1538, BC, Vancouver Island, Forbidden Plateau Phytomyza species Arnica cernua S-1396, CA, Humboldt County Arnica spathulata S-1385, OR, Josephine County 123 Figures 233-240. Leaf Mines of Phytomyza Species i n leaves of Arnica. Figures 233-237. Mines of P. oreas or P. a r n i c i v o r a . Figure 233. Arnica l a t i f o l i a S-1680, BC, Manning Park. Figure 234. Arnica l a t i f o l i a S-1676, WA, Clallam County. Figure 235. Arnica c o r d i f o l i a S-1819, OR, Wasco County. Figure 236. Arnica c o r d i f o l i a S-1604, AB, Lake Louise. Figure 237. Arnica nevadensis S-1870, CA, S i e r r a County. Figure 238. Mine of P_. saximontana on Arnica l a t i f o l i a S-1538, BC, Vancouver Island, Forbidden Plateau. Figure 239. Unidentified blotch mine on Arnica cernua S-1396, CA, Humboldt County. Figure 240. Unidentified blotch mine on Arnica spathulata S-1385, OR, Josephine County. 124 125 TABLE IX. Buprestid Beetles Collected Feeding on Arnica Species. Arnica c o r d i f o l i a S-1594, BC, Windermere Anthaxia expansa Lec. - 2 Arnica c o r d i f o l i a S-1819, OR, Wasco County, Pine Grove Anthaxia expansa Lec. - 1 Anthaxia aenescens Casey - 1 Arnica c o r d i f o l i a S-2022, BC, Apex Mountain Anthaxia expansa Lec. - 2 Arni c a c o r d i f o l i a S-1842, SC, Lawrence County, Deadwood Gulch Anthaxia expansa Lec. - 7 Arnica c o r d i f o l i a S-1797, CA, Siskiyou County, Meiss Lake Anthaxia sp. - 2 Arnica spathulata S-1381, OR, Josephine County, Cave Junction Acmaeodera connexa Lec. - 1 Anthaxia C a l i f o r n i a Obenb. -1 Arnica venosa S-1793, CA, Shasta County, Gibson Anthaxia C a l i f o r n i a Obenb. - 8 Anthaxia sp. - 1 126 cases, as i n Arnica c o r d i f o l i a near Windermere, BC, S-1594, and in Grand County, CO, S-2035, the i n f e s t a t i o n was so severe that a l l the rays had been eaten, giving the plants a close s u p e r f i c i a l resemblance to Arnica discoidea. Severe i n f e s t a t i o n s could a f f e c t p o l l i n a t o r v i s i t a t i o n due to changed UV reflectance-absorbance patterns (see further discussion under UV s e c t i o n ) . The Buprestid beetles are mostly too small and glabrous to transfer pollen from one plant to another, thus are not l i k e l y aiding i n p o l l i n a t i o n . Achene Feeders As discussed under "Seed Production" many populations of most or a l l species (observed i n a l l species except Arnica viscosa) are infested with the larvae of one or more small f l i e s of the genus Xenochaeta (Tephritidae). The young larvae begin feeding on the ovaries before the f i r s t f l o r e t s begin to expand. The larvae have usually pupated by the time the achenes are mature, occupying the remains of several achenes. In minor i n f e s t a t i o n s the larvae u s u a l l y feed upon only h a l f a dozen or fewer achenes i n the center of the head. The remaining achenes are undamaged. In more severe i n f e s t a t i o n s , which occur commonly, most or a l l of the achenes are either eaten or the pappus on the v i a b l e achenes i s damanged to the extent that i t does not open properly. The uninfested achenes may drop to the ground as a s o l i d unit rather than being c a r r i e d away i n d i v i d u a l l y by the pappus blown i n the wind. Frequently the larvae of the t e p h r i t i d s are themselves infested with larvae of Pteromalus wasps (Pteromalidae), so that the adult wasps, rather than adult t e p h r i t i d s , emerge from the pupal cases. The achenes are s t i l l u s ually i n f e s t e d enough that they are not v i a b l e . 127 U l t r a v i o l e t Reflectance-Absorbance of Arnica Heads Photographs of Arnica heads were taken out-of-doors on a sunny day with a tripod-mounted Minolta single-lense r e f l e x camera using Kodak Tri-X Pan black and white f i l m (ASA 400). At the same distance exposures were taken with a Wratten 18A f i l t e r attached and the camera stopped-down 3-4 f-stops to compensate f o r the lower l i g h t penetration through the opaque f i l t e r . The Wratten 18A f i l t e r allows only UV l i g h t i n the 350-400 nm range to pass through. Arnica heads are large enough that the v a r i a t i o n i n f o c a l length of the lens with and without the f i l t e r i s not c r i t i c a l , although some d i s t o r t i o n i s seen at t h i s distance (ca 30 cm). At distances closer than t h i s the d i f -ference i n f o c a l length must be compensated f o r ( H i l l , 1977; Kennedy and Ganders, 1979). The UV reflectance-absorbance pattern i n Arnica heads, which appear completely yellow to the human eye (Figure 241), appear l i k e those of Mono-p t i l o n b e l l i o i d e s (Kennedy and Ganders, 1979) i n which the dis c f l o r e t s absorb UV l i g h t and appear dark, while the ray f l o r e t s r e f l e c t UV and appear s o l i d l i g h t (Figure 242). This i s unlike the pattern i n many other genera of the Asteraceae i n which part of the ray f l o r e t s r e f l e c t UV and an inner portion of the ray absorbs UV as i n Rudbeckia spp. (Abrahamson and McCrea, 1977), Lasthenia chrysostoma (Kennedy and Ganders, 1979), or Coreopsis gigantea (Scogin et a l . , 1977). In Arnica, a f t e r the heads have been i n flower a few days and a l l the discs have opened and have probably been p o l l i n a t e d , the rays begin to absorb UV and the whole head appears dark (Figure 242). This p o s t - p o l l i n a t i o n change in UV pattern has been noted by Scogin et a l . (1977) f or Coreopsis gigantea and a s i m i l a r change was noted i n older herbarium specimens of that species. 128 Figures 241-247. UV Reflectance-Absorbance i n Arnica heads and Typ i c a l Heads of Three Species. Figure 241. Arnica c o r d i f o l i a X Arnica l a t i f o l i a (putative hybrid) S-1474, MT, R i v a l l i County, photographed i n natural sunlight. Figure 242. Same photographed with Wratten 18A f i l t e r showing dark, absorptive UV discs and l i g h t , r e f l e c t i v e UV rays i n two recently opened heads on l e f t and r i g h t , and dark, absorptive discs and rays of two p o s t - p o l l i n a t i o n center heads. Figure 243. Arnica discoidea S-1825, WA, K l i c k i t a t County, photographed i n natural sunlight. Figure 244. Same photographed with UV f i l t e r showing UV absorptive disc f l o r e t s (dark). Figure 245. Arnica c o r d i f o l i a S-1497, WA, Okanogan County, very wide-rayed population. Figure 246. Arnica l a t i f o l i a S-1961, BC, Nancy Greene P r o v i n c i a l Park, t y p i c a l head with "squared-off" rays. Figure 247. Arnica g r a c i l i s S-1603, AB, Lake Louise, t y p i c a l heads with few, narrow rays. 129 130 In the d i s c o i d species of Arnica the head absorbs UV and appears dark (Figures 243, 244). P o l l i n a t o r s An unexpedtedly small number of insect p o l l i n a t o r s has been observed v i s i t i n g Arnica populations i n these two subgenera, although many hours have been spent observing hundreds of populations at a l l hours of the day through-out most of the d i s t r i b u t i o n of the species. In many of the largest popula-tions with hundreds or thousands of heads i n flower no p o l l i n a t o r s were observed. There are several p o s s i b i l i t i e s f o r the lack of p o l l i n a t o r s i n some populations. In forest habitats where the shade i s r e l a t i v e l y dense there may be fewer insects than i n the more open habitats. In populations which are l a r g e l y monocephalous, most of the heads may open within a few days of one another. Few or no p o l l i n a t o r s may v i s i t a f t e r the heads have been p o l l i n a t e d because of the changing UV reflectance-absorbance patterns. In alpine meadows, where there are many d i f f e r e n t plants flowering at the same time, there may be a preference f o r other genera or species of flowers by the p o l l i n a t o r s . Many b u t t e r f l i e s and bees have been observed v i s i t i n g other flowers ( i . e . , Valeriana spp., Lupinus spp., and Eriogonum spp.) i n preference to Arnica spp. In a few instances as on Yahk Mountain, BC, and Moraine Lake, AB, there seemed to be a p o l l i n a t o r preference f o r Arnica species i n other subgenera. Euphydryas spp. (Nymphalidae) and Hesperia spp. (Hesperiidae) v i s i t e d Arnica r y d b e r g i i , Arnica m o l l i s , and Arnica d i v e r s i f o l i a , but r a r e l y v i s i t e d Arnica l a t i f o l i a , Arnica c o r d i f o l i a or Arnica g r a c i l i s . At s l i g h t l y lower elevations on Yahk Mountain and near the town of Yahk, BC, Arnica l a t i -f o l i a was commonly v i s i t e d by several species of Bombus. These two 131 populations are the only Arnicas i n these two subgenera where large numbers of insect p o l l i n a t o r s were observed. There seems to be no c o r r e l a t i o n between v i s i t o r s to apomictic compared to sexual populations of Arnicas or between d i s c o i d and radiate species, a l -though the UV patterns are d i f f e r e n t i n the two groups. Table X l i s t s the Diptera, Hymenoptera, and Lepidoptera p o l l i n a t o r s captured on Arnica heads. Nomenclature of b u t t e r f l y p o l l i n a t o r s follows Howe (1975), that of the other orders follows Borror et a l . (1976) above the generic l e v e l . 132 TABLE X. P o l l i n a t o r s Collected on Arnica Heads. Arnica cernua S-1772, CA, Humboldt County, Hoopa Mountain. Syrphidae sp. Arnica c o r d i f o l i a S-1464, WY, Teton County, Grand Teton National Park Lycaenidae Mitoura spinetorum (Hewitson) 0_ Arnica c o r d i f o l i a S-1648, BC, Sheridan Lake Nymphalidae Speyeria e l e c t a beani (Barnes & Benjamin) 0_ Arnica c o r d i f o l i a S-1797, CA, Siskiyou County, Meiss Lake Syrphidae Taxomerus o c c i d e n t a l i s Curran £ Arnica c o r d i f o l i a S-1819, OR, Wasco County, Pine Grove Syrphidae Syrphus opinator Osten Sachen 071  Arnica c o r d i f o l i a S-1823, WA, K l i c k i t a t County, Appleton Syrphidae Sphaerophoria sulphuripes (Thomson) 0_ Arnica c o r d i f o l i a S-1826, WA, K l i c k i t a t County, Glenwood Andrenidae Andrena sp. o" Arnica c o r d i f o l i a S-1836, ID, Bonneville County, Irwin Apidae Bombus b i f a r i u s Cresson 0_ Andrenidae Andrena sp. 133 Table.X cont'd. Arnica c o r d i f o l i a S-2022, BC, Apex Mountain H a l i c t i d a e Halictus sp. Arnica c o r d i f o l i a S-2028, BC, Kamloops Megachilidae Megachile sp. Arnica c o r d i f o l i a S-2029, BC, Sorrento Syrphidae E r i s t a l i s tenax (L.) ^  Arnica discoidea S-1777, CA, Napa County, S. of Lake/Napa Co. l i n e Hesperiidae Erynnis propertius (Scudder & Burgess) tf Arnica l a t i f o l i a S-1567, BC, Yahk Apidae Bombus mixtus Cresson 4 0^  7 \j (=worker cast) Bombus b i f a r i u s Cresson 2 \*j Psithyrus fernaldae Franklin 1 (f, 1 0_ Anthophoridae Nomada sp. 0_ Syrphidae E r i s t a l i s tenax (L.) cf Arnica l a t i f o l i a S-1574, BC, Yahk Mountain Apidae Bombus b i f a r i u s Cresson ^ 134 Table X cont'd. Arnica l a t i f o l i a S-1576, BC, Yahk Mountain Apidae Bombus melanopygus Nylander tj" Bombus mixtus Cresson 2 cf Nymphalidae Euphydryas sp. 5 0^  2 J Arnica l a t i f o l i a S-2031, BC, Black Tusk Sessidae Vespamina sp. ? Arnica spathulata S-1380, OR, Josephine County, Kirby Apidae Apis m e l l i f e r a L. Ha l i c t i d a e H a lictus sp. Arnica, spathulata S-1740, OR, Josephine County, Selma Apidae Bombus caliginosus (Frison) 2 0^  Arnica venosa S-1793, CA, Shasta County, Gibson H a l i c t i d a e sp. 135 SYSTEMATIC TREATMENT Genus Arnica L. Sp. Plant. 884. 1753. A r t i f i c i a l Key to the Subgenera of Arnica. A. Anthers purple; i n v o l u c r a l bracts callous-tipped; plants of n. B r i t i s h Columbia and Yukon to Alaska and Japan...Subg. Andropurpurea Maguire AA. Anthers yellow; i n v o l u c r a l bracts not callous-tipped B B. Leaves a l l s e s s i l e ; plants from woody caudex C C. Cauline leaves strongly b a s a l l y disposed on flowering stem; heads radiate; plants of se United States and cen t r a l Europe ...Subg. Montana Maguire CC. Cauline leaves not strongly b a s a l l y disposed, stem le a f y throughout; heads d i s c o i d ; rare plants of sw Oregon and nw C a l i f o r n i a Subg. Calarnica Straley BB. Leaves mostly p e t i o l a t e ; plants rhizomatous, producing basal rosettes C C. Pappus brown or tawny, subplumose Subg. Chamissonis Maguire CC. Pappus white ( r a r e l y tawny), barbellate D D. Leaf blades r e l a t i v e l y narrow, mostly 3-10 times as long as wide; plants mostly densely tufted Subg. A r c t i c a Maguire DD. Leaf blades r e l a t i v e l y broad, mostly 1-2.5 times as long as broad; plants mostly not densely tufted Subg. Austromontana Maguire 136 Species Recognized Subgenus Austromontana Maguire. B r i t t o n i a 4:432. 1943. Lectotype: Arnica l a t i f o l i a Bongard Sect. E u l a t i f o l i a e Maguire. B r i t t o n i a 4:432. 1943. Lectotype: Arnica l a t i f o l i a Bongard 1. Arnica c o r d i f o l i a Hooker 2. Arnica nevadensis A. Gray 3. Arnica l a t i f o l i a Bongard 4. Arnica g r a c i l i s Rydberg 5. Arnica cernua T. Howell Sect. Eradiatae Maguire. B r i t t o n i a 4:452. 1943. Lectotype: Arnica d i s c o i d e a Bentham 6. Arnica discoidea Bentham 7. Arnica spathulata Greene Subgenus Calarnica subgenus nov. Type Species: A r n i c a v i s c o s a A. Gray 8. Arnica v i s c o s a A. Gray 9. Arnica venosa H.M. H a l l The following keys are designed to be us e f u l f o r a majority of specimens of these two subgenera. An a r t i f i c i a l key i s provided f o r both l i v e and pressed specimens, and a f i e l d key. A r t i f i c i a l Key to the Species of Arnica, Subgenera Austromontana and Ca l a r n i c a . A. Leaves a l l s e s s i l e ; non-rhizomatous perennials l a c k i n g s t e r i l e basal rosettes of leaves; heads d i s c o i d (Subgenus Calarnica) B B. Margins of leaves toothed; 2° veins prominent; leaves rugose; heads nodding i n bud; f l o r e t s b r i g h t yellow; achenes densely 137 str i g o s e , non-glandular 9. Arnica venosa o BB. Margins of leaves e n t i r e ; 2 veins inconspicuous; leaves smooth; heads erect i n bud; f l o r e t s creamy-yellow or white; achenes s t i p i t a t e glandular 8. Arnica v i s c o s a AA. Leaves, at l e a s t lower ones, p e t i o l a t e ; rhizomatous perennials producing s t e r i l e basal rosettes; heads radiate or d i s c o i d (Subgenus Austromontana) B B. Heads radiate (Section E u l a t i f o l i a e ) C C. Leaves mostly glabrous ( i f pubescent then sparsely scabrous and non-glandular) D D. Heads erect i n bud; achenes hirsute from middle to top; leaves t h i n ; veins green; of widespread d i s -t r i b u t i o n i n w. N.A 3. Arnica l a t i f o l i a DD. Heads nodding i n bud; achenes sparsely to densely pubescent throughout; leaves thick, leathery, veins reddish; r e s t r i c t e d to serpentine s o i l s of sw Oregon and nw C a l i f o r n i a 5. Arnica cernua CC. Leaves pubescent with glandular and non-glandular h a i r s . . . D D. Pubescence of leaves glandular-puberulent only; plants mostly much-branched and clumped; mostly less than 20 cm t a l l ; achenes black, glabrous to sparsely pubescent. 4. Arnica g r a c i l i s DD. Pubescence of leaves glandular s t i p i t a t e and non-glandular; plants mostly unbranched and not clumped; greater than 20 cm t a l l ; achenes grey, mostly densely h i r s u t e at top or throughout E 138 E. Leaves of innovations and lower cauline leaves with cordiate bases; margins mostly toothed; pappus pure white 1. Arnica c o r d i f o l i a EE. Leaves of innovations and lower cauline leaves with attenuated or rounded bases; margins mostly e n t i r e ; pappus off-white or tawny 2. Arnica nevadensis BB . Heads d i s c o i d (Section Eradiatae) C C. Leaves of innovations and lower cauline leaves spathulate; plants mostly r e s t r i c t e d to serpentine s o i l s of sw Oregon and nw C a l i f o r n i a 7. Arnica spathulata CC. Leaves of innovations and lower cauline leaves with cordate or rounded bases; plants usually not of serpentine s o i l s ; s. Washington to s. C a l i f o r n i a 6. Arnica discoidea F i e l d Key to the Species of Arnica, subgenera Austromontana and Calarnica. A. Leaves d i s t i n c t l y fragrant when bruised B B. Heads d i s c o i d . C C. Leaves broadly s e s s i l e ; cauline leaves and bracts greater than 6 p a i r s ; basal rosettes lacking D D. Margins of leaves toothed; plants predominately eglandular; leaves rugose; heads nodding in bud; f l o r e t s bright yellow; rare at low elevations i n Shasta Co., C a l i f o r n i a 9. Arnica venosa DD. Margins of leaves e n t i r e ; plants strongly glandular; leaves smooth; heads erect i n bud; f l o r e t s white to creamy-yellow; rare at high elevations i n n. C a l i f o r n i a 139 and sw Oregon 8. Arnica viscosa CC. Leaves of basal rosettes and lower cauline leaves d i s t i n c t l y p e t i o l a t e ; cauline leaves and bracts l e s s than 6 p a i r s ; basal rosettes present D D. Lower cauline leaves spathulate; glandular h a i r s p u r p l i s h or reddish; of serpentine s o i l s i n sw Oregon and nw C a l i f o r n i a 7. Arnica spathulata DD, Lower cauline leaves not spathulate; leaves grey to green, lacking p u r p l i s h or reddish h a i r s ; usually not of serpentine s o i l s , s Washington to s C a l i f o r n i a . . . 6. Arnica discoidea BB. Heads radiate C C. Leaves appearing dark green, lacking non-glandular h a i r s ; plants mostly densely clumped, from short rhizomes 4. Arnica g r a c i l i s CC. Leaves appearing greyish due to many non-glandular h a i r s ; plants mostly not clumped, from long rhizomes D D. Basal rosette leaves with cordate bases; plants with many long non-glandular h a i r s (1-2 mm); widespread i n w. N.A. e s p e c i a l l y i n open Pinus ponderosa and Pseudotsuga menziesii f o r e s t s . . . 1 . Arnica c o r d i f o l i a DD. Basal rosette leaves with attenuated bases; plants lacking or with few long non-glandular h a i r s ; rare species, mostly at high elevations i n the S i e r r a Nevada 2. Arnica nevadensis AA. Leaves not at a l l or very f a i n t l y fragrant when bruised B B. Heads nodding i n bud; plants dark reddish or purp l i s h throughout, 140 e s p e c i a l l y along veins; rare species of serpentine s o i l s i n sw Oregon and nw C a l i f o r n i a ...5. Arnica cernua BB. Heads erect i n bud; plants mostly greenish througout except at base of stem; common widespread species not of serpentine s o i l s . . 3. 'Arnica l a t i f o l i a TAXONOMIC TREATMENT Subgenus Austromontana Maguire Perennial herbs from naked or scaly rhizomes or r a r e l y a short woody caudex and rhizomes; roots slender or fleshy; s t e r i l e rosettes of leaves (innovations) produced; rosette leaves and lower cauline leaves usually the l a r g e s t , mostly long p e t i o l e d , decreasing i n s i z e upward, becoming b r a c t - l i k e and s e s s i l e below inflorescence; cauline leaves opposite, blades ovate to lanceolate, bases cordate to attenuate, t i p s acute to rounded, margins e n t i r e to crenate, serrate, dentate and undulate; stems simple to much-branched; v e s t i t u r e of stems and leaves subglabrate to strigose, p i l o s e , and s t i p i t a t e glandular; inflorescence a s o l i t a r y head or corymb; heads radiate or d i s c o i d ; i n v o l u c r a l bracts narrowly to broadly lanceolate, acute to acuminate; ray f l o r e t s l i n e a r e l l i p t i c to oblong; disc f l o r e t s tubular or goblet-shaped; achenes ribbed, subglabrate to h i r sute and s t i p i t a t e glandular; pappus white ( r a r e l y tawny), ba r b e l l a t e to subplumose. Section E u l a t i f o l i a e Maguire Heads radiate; otherwise, as with the subgenus. 1. Arnica c o r d i f o l i a Hooker. F l . Bor. Am. 1:33. 1834. Arnica macrophylla Nutt. Trans. Am. P h i l . Soc. 7:407. 1841. TYPE: Oregon, Blue Mountains, N u t t a l l (Herb. Hook., KEW). 141 Arnica c o r d i f o l i a Hook, var. macrophylla (Nutt.) Maguire. Am. Midi. Nat. 37:137. 1947. Arnica grandifolia Greene. P i t t o n i a 4:173. 1900. TYPE: Montana, Bridger Mountains, July 28, 1896, J.H. Flodman 896 (holotype, ND; isotype, MO) (probable hybrid, also l i s t e d under synonymy of Arnica l a t i f o l i a ) . Arnica subcordata Greene. P i t t o n i a 4:173. 1900. TYPE: Alberta, Athabasca River, June 26, 1989, Spreadborough, Geol. Surv. Can. 19644 (UC). Arnica pumila Rydb. Mem. N.Y. Bot. Gdn. 1:433. 1900. TYPE: Colorado, Gray's Peak, 1872, J. Torrey, s.n. (NY). Arnica c o r d i f o l i a Hook. var. pumila (Rydb.) Maguire. Madrono 6:154. 1942. Arnica p a r v i f o l i a Greene. PI. Baker. 3:28. 1901. TYPE: Colorado, Marshall Pass, 3,000 m, July 19, 1901. C.F. Baker 515 (holotype, ND; isotypes, POM, WS). Arnica paniculata A. Nels. Man. Bot. Rocky Mts. 572. 1909. TYPE: Wyoming, Carbon Co., Bridger Peak, moist timber, August 24, 1903, L.N. Gooding 1974 holotype, RM; isotype, MO). Arnica evermannii Greene. Ottawa Nat. 23:215. 1910. TYPE: Idaho, P i t t i t Lake, 2160 m, August 13, 1895, B.W. Evermann 318 (US). Arnica abortiva Greene. Leaflets 11:47. 1910. TYPE: Wyoming, Wind River Mountains, July 23, 1882, W.H. Forwood, _s.n. (US). Arnica granulifera Rydb. F l . Rocky Mts. 978. 1917. TYPE: Montana, L i t t l e Belt Mountains, J.H. Flodman 896 (NY). Arnica andersonii Piper. Proc. B i o l . Soc. Wash. 33:106. 1902. TYPE: B r i t i s h Columbia, Skeene, E l l i s o n , damp woods, September 11, 1910, j;.R. Anderson 677 (US). Arnica ovalis Rydb. N. Am. F l . 34:338. 1927. TYPE: Alberta, Crow Nest Pass 142 J u l y 3 1 , 1897, Macoun, G e o l . S u r v . Can. 72719 (CAN?) (not s e e n ) . A r n i c a a u s t i n a e Rydb. N. Am. F I . 3 4 : 3 4 0 . 1927. TYPE: C a l i f o r n i a , Lake C o u n t y , June 1898, R . M . A u s t i n & Bruce 2165 (NY). A r n i c a h u m i l i s Rydb. N. Am. F I . 3 4 : 3 4 1 . 1927. TYPE: A l b e r t a , Lake L o u i s e , " t h e s a d d l e " , 2100 m, Macoun, G e o l . S u r v . Can. 65504 (NY). A r n i c a c o r d i f o l i a Hook, v a r . h u m i l i s ( R y d b . ) M a g u i r e . Am. M i d i . N a t . 3 7 : 1 3 8 . 1947. A r n i c a w h i t n e y i F e r n . Rhodora 3 7 : 3 3 4 . 1935. TYPE: M i c h i g a n , Keweenaw C o u n t y , Copper H a r b o r , d r y d e c i d u o u s woods, J u l y 4 , 1934, F e r n a l d & Pease 3579 ( G ) . A r n i c a c o r d i f o l i a Hook, s u b s p . w h i t n e y i ( F e r n . ) M a g u i r e . B r i t t o n i a 4 : 4 5 2 . 1943. A r n i c a h a r d i n a e S t . J o h n . F I . SW. Wash. 419. 1937. TYPE: I d a h o , Benewah C o u n t y , Lake C h a t c o l e t , open woods, (5. Weitman 226 ( h o l o t y p e , WASH; i s o t y p e , WS). S t r o n g l y r h i z o m a t o u s p e r e n n i a l h e r b ; rhizomes 2 - 1 0 ( - 6 0 ) cm g r o w t h / y e a r , s l e n d e r 1-2 mm t h i c k , naked e x c e p t f o r few s c a l e s and o l d l e a f b a s e s toward summit , unbranched or few b r a n c h e s n e a r summit , f o r m i n g l a r g e l o o s e p o p u l a -t i o n s of s t e r i l e b a s a l r o s e t t e s and f l o w e r i n g s t e m s ; stems s i n g l e or r a r e l y f e w , s i m p l e o r b r a n c h e d n e a r base or t h r o u g h o u t , ( 1 0 - ) 1 5 - 4 0 ( - 5 5 ) cm h i g h , s p a r s e l y t o d e n s e l y v i l l o u s and s t i p i t a t e g l a n d u l a r b a s a l l y , more d e n s e l y so h i g h e r , sometimes p u r p l i s h o r r e d d i s h a t b a s e ; i n n o v a t i o n s w i t h 2 - 6 ( - 1 4 ) l o n g - p e t i o l a t e l e a v e s , p e t i o l e s ( 2 - ) 6 - 1 0 ( - 2 0 ) cm l o n g , sometimes n a r r o w l y w i n g e d , b l a d e s o v a t e t o b r o a d l y o v a t e or l a n c e o l a t e , ( l - ) 4 - 8 ( - 1 2 ) cm l o n g , ( l - ) 3 - 6 ( - 9 ) cm b r o a d , bases c o r d a t e , s u b c o r d a t e , t o sometimes r o u n d e d , t i p s a c u t e t o r o u n d e d , m a r g i n s e n t i r e , s u b e n t i r e , t o u s u a l l y r e g u l a r l y d e n t a t e , 143 serrate, crenate, or r a r e l y doubly dentate or serrate; cauline leaves 2-3(-5) p a i r s , often reduced basa l l y and becoming b r a c t - l i k e toward summit, lower cauline leaves usually long-petiolate (l-)4-8(-10) cm long, blades (2-)5-10 (-13) cm long (2-)4-6(-9) cm broad, upper pa i r s of cauline leaves with pro-gre s s i v e l y shorter p e t i o l e s , uppermost p a i r usually b r a c t - l i k e and s e s s i l e ; shape and margins of lower cauline leaves as i n that of innovations, upper bracts usually ovate to lanceolate with subentire to e n t i r e margins; v e s t i -ture of herbage sparsely to densely strigose or v i l l o u s on under surfaces, sparsely to usually densely strigose or v i l l o u s and s t i p i t a t e glandular, leaf margins c i l i a t e ; inflorescence usually a s i n g l e head or open corymb of 3-7 (-35) heads; peduncles (4-)8-15(-20) cm long, peduncle and periclinum p i l o s e and s t i p i t a t e glandular; heads radiate, 15-25 cm high; i n v o l u c r a l bracts (7-)10-15(-21) + u n i s e r i a t e , 10-18 mm long, 2-4 mm broad, narrowly lanceolate, acute to acuminate; ray f l o r e t s , (5-)8-ll(-16) pale to dark gold-en yellow, tube 3-7 mm long, limb 10-25(-35) mm long, 4-7(-10) mm broad, l i n e a r e l l i p t i c , 1-3 teeth at t i p , tube and base of limb sparsely v i l l o u s and s t i p i t a t e glandular; disc f l o r e t s (15-)38-57(-94), tubular 8-12 mm long, sparsely to densely v i l l o u s and s t i p i t a t e glandular at base and sparsely glandular upward and on outer surface of lobes; pollen diameter 40-55 um; achenes dark grey, 4-10 mm long, 1 mm broad, sparsely to densely hirsute with b i f u r c a t e hairs and few s t i p i t a t e glandular h a i r s , e s p e c i a l l y d i s t a l l y ; pappus white, ba r b e l l a t e , the l a t e r a l setae .07-.10 mm long, 38-52 b r i s t l e s ; apomictic; 2n=38,57,76,95,114. Type: "Alpine woods of the Rocky Mountains", Drummond (Herb. Hook., KEW). D i s t r i b u t i o n : Arnica c o r d i f o l i a i s common i n many moderately dry habitats, e s p e c i a l l y i n open Pinus ponderosa, Pinus contorta, and Pseudotsuga menziesii 144 f o r e s t s or alpine meadows at many elevations from c e n t r a l Yukon and Eastern Alaska south to Northern New Mexico and Southern C a l i f o r n i a , and east to the Black H i l l s of South Dakota and d i s j u n c t i n Abies balsamea, Acer spp., and Betula spp. f o r e s t s i n Keweenaw Co., Michigan and Thunder Bay, Ontario. Figure 248. I l l u s t r a t i o n : Figure 249. This, the most widespread Ar n i c a i n Western North America, i s usually abundant where i t occurs, covering vast areas. I t i s often the dominant herbaceous plant i n open f o r e s t s i n the Western C o r d i l l e r a . In i t s more t y p i c a l form, where i t grows i n open pine or Douglas f i r f o r e s t s , i t i s d i s t i n c t i v e with i t s deeply cordate, toothed leaves, large s o l i t a r y heads, broad rays, and h i r s u t e achenes. In other habitats there i s great v a r i a b i l i t y i n form, which has l e d to the naming of a number of taxa. These are now generally accepted as f a l l i n g within the range of v a r i a b i l i t y of A r n i c a c o r d i f o l i a . In more shaded l o c a t i o n s very large leaved forms often occur. These have been given taxonomic r e c o g n i t i o n i n the past, as Arnica macrophylla Nutt. from the Blue Mtns. of Oregon, and Arnica andersonii Piper from Skeena, BC. However, t h i s v a r i a t i o n with large t h i n leaves can be seen from s i n g l e c o l l e c -t i o n s or populations growing i n sun, f i l t e r e d shade, and dense shade, as i n c o l l e c t i o n s from Siskiyou Co., CA, S-1797 (Figures 250-259). At the other extreme i s a d i s t i n c t s e r i e s of ecotypes from high elevations and disturbed roadsides to which the names A r n i c a pumila Rydb. and Arnica  humilis Rydb. have been applied. They are quite short, with small, usually e n t i r e leaves. They often resemble Ar n i c a nevadensis, from which they can be 145 Figure 248. D i s t r i b u t i o n of Arnica c o r d i f o l i a i n Western North America. Populations i n Michigan and Ontario are not shown. 146 147 ;ure 249. Arnica c o r d i f o l i a S-1494, WA, Okanogan County, hab X two-thirds. 148 149 Figures 250-259. V a r i a t i o n i n basal leaves from single population of Arnica c o r d i f o l i a S-1797, CA, Siskiyou Co. 150 151 152 distinguished, often only with d i f f i c u l t y , by the whiter, le s s plumose nature of the pappus, and usually a greater degree of p i l o s i t y of the leaves and stems than i n Arnica nevadensis populations. Typical of these small ecotypes are c o l l e c t i o n s from Slate Lake T r a i l , Okanogan Co., WA, S-1498 (Figures 260-267), and roadside i n Pinus contorta f o r e s t , Pine Grove, Wasco Co., OR, S-1820 (Figures 268-276). Many herbarium specimens from Siskiyou Co., CA, Oettinger 1082, Sawyer 941, 1236 (HSC) combine many characters of both Arnica nevadensis and Arnica c o r d i f o l i a . Only where there are several specimens or where they can be studied i n the f i e l d can they be more e a s i l y assigned to one or the other of these species. Larger, more t y p i c a l Arnica c o r d i f o l i a i s separable from t y p i c a l Arnica  nevadensis i n the generally narrower, paler rays, more densely p i l o s e stems and leaves, more toothed margins, whiter, shorter l a t e r a l setae of the pappus, and more densely hirsute achenes of the former. There are some populations of Arnica c o r d i f o l i a with broader rays (Figure 245) as i n a population from Siskiyou Co., CA, S-1797. These rays are as broad as t y p i c a l Arnica neva-densis rays. The two species are also usually e c o l o g i c a l l y separate. Arnica  c o r d i f o l i a usually grows at lower elevations and in more protected habitats of open f o r e s t s . The two have been observed growing together at Saddlebag Lake, Mono Co., CA, S-1937, 1938, with Arnica c o r d i f o l i a i n the f o r e s t edge and Arnica nevadensis on more exposed rock outcrops. Although Arnica c o r d i f o l i a generally has only one or three heads per stem, some populations or i n d i v i d u a l s e s p e c i a l l y i n wetter habitats, have much branched stems with many heads terminating each branch. The greatest extreme of t h i s was seen i n one plant with 35 heads from a c o l l e c t i o n from Tod Mtn., BC, S-1652. 153 Figures 260-267. Var i a t i o n i n basal leaves of high elevation population of Arnica c o r d i f o l i a S-1498, WA, Okanogan Co. 154 155 Figures 268-276. V a r i a t i o n i n basal leaves from population on disturbed roadside bank of Arnica c o r d i f o l i a S-1820, OR, Wasco Co. 156 157 Other c o l l e c t i o n s with several heads have been named Arnica paniculata by Nelson, a species retained by Maguire (1943), but considered a variant of Arnica c o r d i f o l i a by Ediger & Barkley (1978). Maguire (1943) points out the p o s s i b i l i t y of t h i s being a hybrid between Arnica c o r d i f o l i a and Arnica  p a r r y i (subgenus Chamissonis), although both species are l a r g e l y or wholly apomictic (Barker, 1966). A recent hybrid between the two i s u n l i k e l y . I t seems that the characters c i t e d could f a l l within the range of v a r i a t i o n of an apomictic race of Arnica c o r d i f o l i a , and i t i s treated i n t h i s work as a variant thereof. Maguire (1943) also gave taxonomic recognition to a population of Arnica  c o r d i f o l i a from Keweenaw Co., Michigan, treating i t as subsp. whitneyi. Since 1943 a d d i t i o n a l populations have been discovered on the north shore of Lake Superior i n Sibley P r o v i n c i a l Park, Thunder Bay D i s t r i c t , Ontario Garton  15164, 15486 (LKHD). Both of these populations have been studied i n the f i e l d (S-1846, 1849). They are on the average t a l l e r , have more heads (3/ stem), narrower rays, and le s s pubescence than average Arnica c o r d i f o l i a popu-l a t i o n s i n the Western mountains. However, there are numerous populations i n the West which cannot be distinguished from the Michigan and Ontario popula-tions. The population i n Michigan i s very rare and i s on the endangered species l i s t i n that state. I t i s found as one large widely dispersed popu-l a t i o n (or several small populations) between Copper Harbor and Eagle Harbor, growing i n open Abies balsamea, Acer spicatum, Acer saccharum, and Betula spp. f o r e s t . The dominant herbaceous cover i n the area i s Aster macrophyllus. When neither the Aster nor the Arnica are i n flower, the basal rosettes of leaves are separable only with great d i f f i c u l t y . The Arnica leaves tend to be smaller, s l i g h t l y greyer, and have a stronger odor. The Arnica may be 158 Figures 277-281. Habitats and Habits of Arnica c o r d i f o l i a and Arnica  nevadensis. Figure 277. Habitat of Arnica c o r d i f o l i a , WA, K l i c k i t a t Co., Appleton, (S-1823), open fo r e s t of Pseudotsuga menziesii and Pinus  ponderosa. Figure 278. Arnica c o r d i f o l i a , 6-month old seedling, from seed c o l l e c t e d at Hat Creek, BC, (Johns 648) (S-2009). Figure 279. Arnica c o r d i f o l i a X Arnica l a t i f o l i a (putative hybrid), MT, R a v a l l i Co., Charles Waters Campground, (S-1475), normal radiate head and d i s c o i d head on same stem. Figure 280. Habitat of Arnica nevadensis, CA, S i e r r a Co., S i e r r a Buttes, (S-1870), open forest of Pinus monticola, Tsuga mertensiana, and Abies sp. Figure 281. Arnica nevadensis, NV, Washoe Co., Mt. Rose, (S-1910). 159 160 more common i n the area than now presumed, but has been overlooked when not in flower because of the strong resemblance to the Aster. The plants from Sibley P r o v i n c i a l Park are e c o l o g i c a l l y and morphologically quite s i m i l a r to the Michigan ones. This plant should be sought on I s l e Royale in Lake Superior between the known populations i n Michigan and Ontario. Arnica  whitneyi Fern, i s not given s p e c i f i c status or recognized as a subspecies of Arnica c o r d i f o l i a i n t h i s treatment, despite the disjunct d i s t r i b u t i o n . Chromosomally they are l i k e the dominant Western po l y p l o i d complex (2n=76) and morphologically they are le s s d i f f e r e n t than some of the other extremes, e.g., the dwarf forms discussed above, or the southern extremes i n Arizona (Schreiber, _s.n. , Hemmingway, _s.n. MNA) . There are numerous i n t e r e s t i n g populations which are i n many morpho-l o g i c a l aspects intermediate between Arnica c o r d i f o l i a and Arnica l a t i f o l i a . Most notable are several c o l l e c t i o n s from Josephine Co., OR, Lang 798 (SOC), Savage, _s.n. (ORE), and Curry Co., OR, Leach 1544, 1545 (ORE), R a v a l l i Co., MT, S-1474, 1475 (Figure 282). These populations usually do not grow with one or either of the parents, and are considered to be old hybrids which have probably become apomictic a f t e r h y b r i d i z a t i o n and have retained t h e i r i n t e r -mediate characters. Pollen s t a i n a b i l i t y ranges from 33%-99%. D i p l o i d chromosome numbers have been obtained from two Montana c o l l e c t i o n s . They appear to be s t e r i l e having never produced any v i a b l e achenes i n c u l t i v a t i o n , e i t h e r when se l f e d or crossed with other species or s i m i l a r populations. These putative hybrids are usually much larger and more robust i n a l l characters than either parent. They are strongly rhizomatous l i k e Arnica  c o r d i f o l i a . The basal leaves and lower cauline leaves are usually + cordate, but the upper cauline leaves are s e s s i l e . The leaves are sparsely pubescent 161 Figure 282. Arnica c o r d i f o l i a X Arnica l a t i f o l i a S-1475, putative natural hybrid, MT, R a v a l l i County. X two-thirds. 162 163 and appear dark green as i n Arnica l a t i f o l i a . However, glandular hairs are present throughout giving the plants as strong an odor as i n Arnica c o r d i - f o l i a . Heads are usually larger than i n either parent, with more rays (up to 25), more disc f l o r e t s (nearly 200) and more i n v o l u c r a l bracts (up to 21). The shape of the rays (Figure 138) approaches that of Arnica l a t i f o l i a , with more truncate t i p s . Disc f l o r e t s (Figure 144) are more densely pubescent throughout than i n e i t h e r parent. There are numerous other populations or i n d i v i d u a l s closer to one or the other parent. A l l these are considered apo-m i c t i c races and are not given taxonomic status. 2. Arnica nevadensis A. Gray. Proc. Am. Acad. 19:55. 1883. Arnica chionophila Greene. P i t t o n i a 4:171. 1900. TYPE: Nevada, Ruby Mts., July 20, 1896, E.L. Greene (ND?) (not seen). Rhizomatous perennial herb; rhizomes (l-)3-12 cm growth/year, slender, 1-2 mm thick, naked except pairs of brown scales at nodes and crowded scales and old leaf bases toward the summit, unbranched or few branches a p i c a l l y , forming small to large populations of s t e r i l e basal rosettes and flowering stems; stems sin g l e or few, simple or branched from base or above, (10-)15-30 cm t a l l , very sparsely p i l o s e and + densely short glandular throughout; inno-vations with (2-)4-6 long p e t i o l a t e leaves, p e t i o l e s 3-5 cm long, blades ovate, rotund, or e l l i p t i c to lanceolate, 4-7 cm long, 2-4 cm broad, bases rounded to attenuate, t i p s acute to rounded, margins e n t i r e , subentire, to r a r e l y coarsely serrate or dentate; cauline leaves 2-3 p a i r s , lowest p a i r u s u a l l y with longest p e t i o l e s , (l-)2-3(-4) cm long, sometimes winged to 4 mm, next p a i r usually with largest blade 3-5.5 cm long, (l-)2-4 cm broad, upper p a i r u s u a l l y reduced and b r a c t - l i k e , lanceolate, shape and margins of cauline 164 leaves l i k e that of innovations; v e s t i t u r e of herbage sparsely scabrous, p i l o s e and short s t i p i t a t e glandular throughout, margins c i l i a t e ; i n f l o r e s -cence usually of a single head or open corymb of 3(-7) heads; peduncles 4-12 cm long; peduncles and periclinum densely short and long s t i p i t a t e glandular and sparsely p i l o s e ; heads radiate 15-20 mm high; i n v o l u c r a l bracts 10-15(-18) mm long, 2-4 mm wide, narrowly lanceolate, acute to acuminate; ray f l o r e t s 5-13, dark golden yellow, tube 2-7 mm long, limb 15-20 mm long, 6-7 mm broad, v i l l o u s and s t i p i t a t e glandular on tube and lower back of limb, l i n e a r e l l i p -t i c to broadly e l l i p t i c , 1-3 teeth at t i p ; disc f l o r e t s 21-51, tubular 7-10 mm long, sparsely to densely v i l l o u s and s t i p i t a t e glandular at base and sparing-l y d i s t a l l y and on outer surface of lobes; pollen diameter 35-45 um, achenes dark grey to black, 4-9 mm long, to 1 mm wide, sparsely hirsute with b i f u r -cate hairs e s p e c i a l l y d i s t a l l y , short to long s t i p i t a t e glandular throughout; pappus d u l l white to tawny, of 27-50 b r i s t l e s , barbellate to subplumose, the l a t e r a l setae .15-.20 mm; apomictic; 2n=76. TYPE: C a l i f o r n i a , S i e r r a Nevada, 2700 m, P r i n g l e s.n. (G). D i s t r i b u t i o n : Mostly at high elevations (1500-3500 m) i n open f o r e s t s of Tsuga mertensiana, Pinus monticola, and Pinus contorta i n the S i e r r a Nevada of C a l i f o r n i a and Nevada and sparingly east to the San Juan Mountains of Utah and north i n the Cascade Mountains to north-central Washington. (Figure 283). I l l u s t r a t i o n : Figure 284. Arnica nevadensis i s a d i s t i n c t species i n the S i e r r a Nevada, where i t can be r e a d i l y separated from Arnica c o r d i f o l i a most of the time. In much of i t s other d i s t r i b u t i o n i t i s only separable from high elevation forms of Arnica c o r d i f o l i a with d i f f i c u l t y , e s p e c i a l l y from herbarium specimens, as 165 Figure 283. D i s t r i b u t i o n of Arnica nevadensis. 166 167 gure 284. Arnica nevadensis S-1910, NV, Washoe Co. X two-thirds 168 169 further discussed under that species. Occasional populations of Arnica r y d b e r g i i growing i n shaded l o c a t i o n s , as from Hurrican Ridge, Clallam County, WA (S-1677) , are very a t y p i c a l of that species and could e a s i l y be i d e n t i f i e d as Arnica nevadensis. Many i n d i -v iduals have broad rounded basal and lower cauline leaves. However, the narrower heads, pure white pappus, and densely hirsute achenes r e a d i l y sepa-rate i t from Arnica nevadensis. In the S i e r r a Nevada, Arnica nevadensis i s not infrequently found growing with Whitneya dealbata, with which i t has often been confused. The p e r s i s t e n t , more-papery rays on mature achenes and absence of pappus i n Whitneya are d i s t i n c t i v e . Arnica tomentella Greene i s a very dubious species retained as v a l i d by both Maguire (1943) and Ediger & Barkley (1978). I t i s very rare i n c o l l e c -tions and has not been seen i n nature by any of the recent workers on the genus. Most of the specimens have very i n s u f f i c i e n t data, including the Type, Purpus 5625, "open woods, middle Tule River, southeastern C a l i f o r n i a " (holotype, US; isotype MO), so that r e l o c a t i n g i t i n the f i e l d seems a matter of luck. Maguire (1943) considered Arnica tomentella related to Arnica c o r d i f o l i a or Arnica nevadensis. The plants are t a l l e r than average Arnica nevadensis and the i n v o l u c r a l bracts are more pubescent, e s p e c i a l l y near the t i p s . The l a t e r a l setae are closer to those of Arnica nevadensis, although i n old herbarium specimens the color of the pappus cannot be determined and the shape of the rays i s d i f f i c u l t to determine. Ediger & Barkley (1978) consider Arnica tomentella a possible hybrid between Arnica c o r d i f o l i a and Arnica chamissonis var. f o l i o s a , probably based 170 on the attenuate leaf bases and pubescent i n v o l u c r a l bract t i p s . However, the general character of the pubsecence i s close to that of Arnica nevadensis, as are the leaves, and i t i s t e n t a t i v e l y placed here as a probable apomictic race of Arnica nevadensis, u n t i l i t can be found in the f i e l d again. 3. Arnica l a t i f o l i a Bongard. Mem. Acad. St. Petersb. VI. 2:147. 1832. Arnica l a t i f o l i a a genuina Herder. B u l l . Soc. Nat. Mosc. 40:424. 1867. Arnica l a t i f o l i a 8 a n g u s t i f o l i a Herder. B u l l . Soc. Nat. Mosc. 40:424. 1867. Arnica l a t i f o l i a A. Gray. Syn. FI. N. Am. 1:381. 1884. Arnica menziesii Hooker. FI. Bor. Am. 1:331. 1834. TYPE: North West Coast of America, A. Menzies, _s.n. (KEW). Arnica b e t o n i c a e f o l i a Greene. P i t t o n i a 4:163. 1900. TYPE: Washington, Olympic Mountains, Mt. Steele, 1800 m, August 1895, _C.V. Piper 2002 (ND). Arnica t e u c r i i f o l i a Greene. P i t t o n i a 4:164. 1900. TYPE: Idaho, Coeur d'Alene Mountains, divide between St. Joe and Clearwater Rivers, 1820 m, grassy mountain slopes, July 10, 1895, _J.I3. Leiberg 1229 (holotype, US; isotypes, MO, POM). Arnica l a t i f o l i a Bong. var. t e u c r i i f o l i a (Greene) L. Williams. L e a f l . West. Bot. 1:171. 1935. Arnica ventorum Greene. P i t t o n i a 4:173. 1900. TYPE: Wyoming, Wind River Mountains, Union Pass, August 1 1 , 1894, A. Nelson 836 (holotype, ND; isotypes, MO, WS). Arnica g r a n d i f o l i a Greene. P i t t o n i a 4:173. 1900. TYPE: Montana, Bridger Mountains, July 28, 1896, J^.H. Floodman 896 (holotype, ND; isotype, MO). Arnica p l a t y p h y l l a A. Nelson. Bot. Gaz. 31:407. 1901. TYPE: Oregon, Cascade Mountains, Hood River, f i r fo r e s t s , July 18, 1896, IJ.F. Henderson, s.n. (RM) . 171 Arnica l a e v i g a t a Greene. Ottawa Nat. 15:279. 1902. TYPE: B r i t i s h Columbia, Chilliwack V a l l e y , 900 m, by springs i n woods, August 15, 1901, J..M. Macoun, Geol. Surv. Can. 26926 (ND). Arnica aprica Greene. Ottawa Nat. 15:279. 1902. TYPE: B r i t i s h Columbia, Chilliwack Valley, 1050 m, open ground along streamlets, July 10, 1901, _J.M. Macoun, Geol. Surv. Can. 26284 (holotype, ND; isotype, MO) . Arnica j o n e s i i Rydb. F l . Rocky Mts. 978. 1917. TYPE: Utah, A l t a , Wasatch Mountains, 2700 m, July 31, 1879, M.E. Jones 119 (holotype, NY; isotypes, POM, UTC) . Arnica puberula Rydb. F l . Rocky Mts. 979. 1917. TYPE: Alberta, head of Lake Louise, July 22, 1904, J_.M. Macoun, Geol. Surv. Can. 65523 (NY). Arnica eriopoda Gandoger. B u l l . Soc. Bot. France 6538. 1918. TYPE: Oregon, Cascade Mountains, July 27, 1902, W._C. Cusick 2914, (holotype, US; isotypes, POM, MO). Arnica aphanactis Piper. Proc. B i o l . Soc. Wash. 33:105. 1920. TYPE: Washington, Mt. Baker, 1915, C^.W. Turesson, _s.n. (NY). Arnica flodmanii Rydb. N.Am. F l . 34:334. 1927. TYPE: Montana, Madison Range, Spanish Peaks, 2100-2400 m, July 14, 1896, j;.H. Flodman 898, (holotype, NY; isotype, MO). Arnica glabrata Rydb. N. Am. F l . 34:335. 1927. TYPE: Oregon, Crater Lake, August 10, 1897, Austin & Bruce 1627, (holotype, NY; isotypes, POM). Arnica o l i g o l e p i s Rydb. N. Am. F l . 34:336. 1927. TYPE: B r i t i s h Columbia, Skeena River, Hazel ton, June 23, 1917, J_.JA. Macoun, Geol. Surv. Can. 96048 (NY) . Arnica l e p t o c a u l i s Rydb. N. Am. F l . 34:336. 1927. TYPE: B r i t i s h Columbia, Vancouver Island, Mt. Mark, July 25, 1887, _J.M. Macoun, Geol. Surv. Can. 14570 (NY). 172 Arnica membranacea Rydb. N. Am. F l . 34:338. 1927. TYPE: Oregon, Jackson Co., Wimmer, June 13, 1892, E.W. Hammond 231, (NY). Rhizomatous perennial herb; rhizomes l-7(-10) cm/year, stout, 3-5(-7) mm thick, naked or densely clothed with old l e a f bases and dark brown scales, unbranched or few-branched near summit, forming loose or dense clumps of s t e r i l e basal rosettes and flowering stems; stems single or few, simple or branched above upper p a i r of leaves/bracts, r a r e l y branched lower, (10-)15-40 cm t a l l , glabrate to very sparsely v i l l o u s throughout, d i s t i n c t l y p u r p l i s h or reddish at base; innovations with 2-6(-12) long-petiolate leaves, p e t i o l e s (2-)4-8(-12) cm, blades narrowly e l l i p t i c to ovate or rotund, (l-)3-5(-6) cm long, (0.5-)2-4(-5) cm broad, bases attenuate to rounded, subcordate, or r a r e l y cordate, t i p s acute, margins e n t i r e to usually r e g u l a r l y dentate, serrate, or doubly dentate or serrate, or crenate; cauline leaves 2-4(-7) p a i r s , often reduced basa l l y and toward summit, where b r a c t - l i k e , lower cau-l i n e leaves s e s s i l e or with short p e t i o l e s l-3(-5) cm long, often winged 2-5 mm, blades 2-6(-9) cm long, 1-4(-7) cm wide, upper cauline leaves progressive-l y smaller, almost always s e s s i l e , shape and margins as i n leaves of innova-tions; v e s t i t u r e of herbage glabrate to sparsely ( r a r e l y densely) v i l l o u s , s t r i g o s e , and very r a r e l y short s t i p i t a t e glandular, margins often c i l i a t e ; i nflorescence of a single head or open corymb; peduncles (1-)3-10(-20) cm long, becoming more v i l l o u s upward; periclinum sparsely to densely v i l l o u s ; heads l-3(-13), radiate, 10-20 mm high; i n v o l u c r a l bracts (7-)ll-16(-23), + u n i s e r i a t e , 8-14 mm long, (l-)2-3 mm broad, narrowly lanceolate, acute to acuminate; ray f l o r e t s (5-)8-15(-18), usually pale yellow, tube 3-5 mm, limb (5-)10-25 mm long, 2-5 mm wide, l i n e a r e l l i p t i c , truncate at t i p , with 0-3 teeth, tube and base and back of limb sparsely v i l l o u s ; disc f l o r e t s 173 ( 1 5 - ) 5 0 - 9 0 ( - 1 2 4 ) , t u b u l a r , 5-10 mm l o n g , s p a r s e l y t o d e n s e l y v i l l o u s n e a r base and on o u t e r s u r f a c e of l o b e s ; p o l l e n d i a m e t e r 35-50 um; achenes d a r k g r e y - b r o w n , 5-10 mm l o n g , 1 mm w i d e , few b i f u r c a t e h a i r s from m i d - p o i n t d i s -t a l l y , r a r e l y t h r o u g h o u t , few s h o r t s t i p i t a t e g l a n d u l a r h a i r s d i s t a l l y o r t h r o u g h o u t , r a r e l y g l a b r o u s ; pappus w h i t e , b a r b e l l a t e , t h e l a t e r a l s e t a e . 0 5 -.07 mm l o n g , o f 43-64 b r i s t l e s ; s e l f - i n c o m p a t i b l e ; 2n=38. TYPE: A l a s k a , S i t k a , M e r t e n s , £.n_. (Herb. B o n g a r d , L e n i n g r a d ) . D i s t r i b u t i o n : A r n i c a l a t i f o l i a o c c u r s i n many h a b i t a t s a t v a r y i n g e l e v a t i o n s , m o s t l y i n c o o l mixed c o n i f e r f o r e s t s i n the mountains f r o m S o u t h e r n A l a s k a t o N o r t h e r n C a l i f o r n i a and C e n t r a l C o l o r a d o . ( F i g u r e 2 8 5 ) . I l l u s t r a t i o n : F i g u r e 2 8 6 . A r n i c a l a t i f o l i a i s one of t h e commonest and most p o l y m o r p h i c o f a l l s p e c i e s . I t u s u a l l y grows i n w e t t e r l o c a t i o n s than A r n i c a c o r d i f o l i a , e s p e c i a l l y i n c o o l f o r e s t s of P s e u d o t s u g a m e n z i e s i i , P i c e a s p p . , and A b i e s s p p . and n e a r s t r e a m m a r g i n s . There a r e so many e c o t y p e s t o w h i c h names have been a p p l i e d t h a t i t i s d i f f i c u l t to d e s c r i b e t h e " t y p i c a l " A r n i c a l a t i f o l i a . Most have s e s s i l e c a u -l i n e l e a v e s , the r a y s a r e u s u a l l y t r u n c a t e ( F i g u r e 1 3 9 ) , t h e l e a v e s and stems a r e u s u a l l y g l a b r o u s , and t h e mature achenes a r e u s u a l l y g l a b r o u s t h r o u g h o u t o r h i r s u t e o n l y on t h e d i s t a l h a l f . Among t h e v a r i a b l e e c o t y p e s a r e a group of b r o a d - l e a v e d , r o b u s t p o p u l a -t i o n s w i t h t h i n l e a v e s from shady l o c a t i o n s , and a n o t h e r group of s m a l l e r , n a r r o w e r - l e a v e d , o f t e n h i g h e l e v a t i o n , sun forms w i t h much t h i c k e r l e a v e s . The extreme c o n d i t i o n o f t h e shade form i s r e p r e s e n t e d i n c o l l e c t i o n s from M t . B a k e r , Whatcom C o . , WA, S - 1 1 5 8 , 1161. These are v e r y r o b u s t p l a n t s t o 174 ;ure 285. D i s t r i b u t i o n of Arnica l a t i f o l i a . Star indicates type l o c a l i t y . 175 176 Figure 286. Arnica l a t i f o l i a S-1500, WA, Chelan Co. X two-thirds. 177 178 40 cm t a l l , with t h i n broad leaves and coarsely serrate margins. This large form has received taxonomic recognition in the past with several names having been applied (Arnica p l a t y p h y l l a Nels., Arnica aprica Greene, Arnica l a e v i -gata Greene). At the opposite extreme are plants which s u p e r f i c i a l l y resemble Arnica g r a c i l i s . These usually alpine plants are short (as l i t t l e as 10 cm t a l l ) , clumped, and branched, with small heads, but lacking the glandular hairs (or lacking h a i r s of any type) of Arnica g r a c i l i s , and having paler, blunter rays than Arnica g r a c i l i s . Examples of these small ecotypes include c o l l e c t i o n s from Mt. Arrowsmith, Vancouver Is., BC, A l l e n , _s.ii. (UBC), (Figures 287-298), and Green Mtn., Vancouver Is., BC, Krajina et a l . 4987 (UBC). Records of Arnica g r a c i l i s from Vancouver Is. probably erroneously r e f e r to these small ecotypes of Arnica l a t i f o l i a . The name Arnica betonicae- f o l i a Greene has been applied to these ecotypes i n the past. One such form from Kootenay Pass, BC, S-1561 growing on a disturbed roadside, became much larger and more robust the following year when transplanted to the experi-mental garden i n Vancouver. Every possible combination of intermediate i n d i v i d u a l s and populations e x i s t s between the large and small ecotypes from much of the d i s t r i b u t i o n of the species. There i s a tendency i n the northern part of the range for populations to have a greater p i l o s i t y to the leaves and stems (Buttrick 779, A t l i n Lake, BC, and Taylor, et a l . 5993, Ilgachuz Mtns., BC, and e s p e c i a l l y some c o l l e c t i o n s from the Queen Charlotte Islands, BC, a l l UBC). A t r u l y d i s c o i d condition i n Arnica l a t i f o l i a has not been seen, although i n one population from Nancy Greene P r o v i n c i a l Park, BC, S-1961, some plants had rays which were var i o u s l y shortened (Figures 299-304), compared to t y p i c a l rays f o r t h i s species (Figure 139). 179 Figures 287-298. V a r i a t i o n i n basal leaves of small alpine form of Arnica l a t i f o l i a A l l e n s.n., BC, Mt. Arrowsmith. 180 181 Figures 299-304. Va r i a t i o n i n shortened ray f l o r e t s from several heads on one plant of Arnica l a t i f o l i a S-1961, BC, Nancy Greene Prov. Park. 183 The r e l a t i o n s h i p of Arnica l a t i f o l i a to Arnica m o l l i s and Arnica  d i v e r s i f o l i a (both i n Subgenus Chamissonis) deserves further i n v e s t i g a t i o n . Arnica d i v e r s i f o l i a has been treated i n previous works as a possible hybrid between the other two species. There are populations which strongly resemble Arnica l a t i f o l i a , including several c o l l e c t i o n s from Glacier National Park, MT, Harvey 5163, 5574 (MONTU), which have been determined as Arnica d i v e r s i -f o l i a . Further discussions of the r e l a t i o n s h i p of t h i s species to Arnica cernua, Arnica c o r d i f o l i a and Arnica g r a c i l i s may be found under the discussions of those species. 4. Arnica g r a c i l i s Rydberg, B u l l . Torr. Bot. Club 24:297. 1897. Arnica b e t o n i c a e f o l i a var. g r a c i l i s (Rydb.) M.E. Jones, B u l l . Univ. Mont. B i o l . 15:45. 1910. Arnica l a t i f o l i a Bong, var. g r a c i l i s (Rydb.) Cronq. Vase. PI. Pac. NW. 5:51. 1955. Arnica Columbiana A. Nels., Bot. Gaz. 30:200. 1900. TYPE: Montana, Columbia F a l l s , R._S. Williams 1049 (RM) . Arnica m u l t i f l o r a Greene. P i t t o n i a 4:162. 1900. TYPE: Idaho, Lake Pend O r i e l l e , Leiberg 234 (holotype, ND; paratypes, MONTU, NY, WS). Arnica o v a l i f o l i a Greene. P i t t o n i a 4:168. 1900. TYPE: Wyoming, Big Horn Mountains, Blankinship, _s._n. (ND) . Arnica arcana A. Nels., Bot. Gaz. 37:276. 1904. TYPE: Wyoming, Big Horn Mountains, Doyle Creek, N._L. Gooding 377 (holotype, RM; isotype, MO). Arnica l a c t u c i n a Greene, Ottawa Nat. 33:215. 1910. TYPE: Alberta, Banff, W.jC. McCalla 2014 (ND) . 184 Rhizomatous, often clumped perennial herb; rhizome growth 0.5-3(-6) cm per year, slender, 1-2 mm thick, naked on older parts or with old leaf bases and brown scales e s p e c i a l l y toward the summit, unbranched or usually several branched forming clumps of s t e r i l e basal rosettes and flowering stems; stems single or usually several and often much branched throughout or above upper leaves/bracts, 10-30 cm high; glabrate below to sparsely short s t i p i t a t e glandular, and sparsely scabrid or strigose, becoming densely short s t i p i t a t e and s e s s i l e glandular above with yellowish hairs; innovations with 4-6(-12) long p e t i o l a t e leaves, p e t i o l e s 0.5-2(-6) cm long, r a r e l y narrowly winged, blades e l l i p t i c to obovate, 2-3(-5) cm long, l-2(-3) cm broad, bases cordate to attenuate, often oblique, t i p s acute, margins subentire to serrate; lower cauline leaves, 2-3 p a i r s and one or more pairs of opposite or alternate bracts, p e t i o l e s 1.5-3(-4) cm long, often winged 2-3 mm, blades ovate to e l l i p t i c , 2-4(-5) cm long, 1-3.5 cm broad, bases usually attenuated, t i p s acute to rounded, subentire to i r r e g u l a r l y serrate; upper cauline leaves re-duced, ovate, usually s e s s i l e , often connate-perfoliate, 2-3(-6) cm long, 1-2(-3) cm broad, except uppermost b r a c t - l i k e , often alternate, 1-2 cm long, 1-2(-5) mm broad; v e s t i t u r e of leaves sparsely s t r i g u l o s e and very short s t i p i t a t e below, densely short s t i p i t a t e or s e s s i l e s t i p i t a t e above; i n f l o -rescence simple or open corymb; peduncles 2-8 cm; periclinum densely short and longer s t i p i t a t e glandular; heads ( l - ) 3 - l l , radiate, 10-15 mm high; i n -v o l u c r a l bracts 6-14, + u n i s e r i a t e 5-10(-12) mm long, 1-2 mm broad, narrowly lanceolate, acute to acuminate; ray f l o r e t s , 5-11, pale to bright yellow, tube 3-4 mm, limb 10-16(-20) mm long, 3-4(-6) mm broad, l i n e a r e l l i p t i c , t i p s acute with 1-3 blunt teeth at t i p , glandular and sparsely v i l l o u s on tube and lower back of limb; d i s c f l o r e t s (9-)16-23(-44) tubular, 5-6 mm long, glandu-l a r and sparsely v i l l o u s at base and on outer t i p s of lobes; pollen diameter 185 30-50 um; achenes dark brown to black, 4-7 mm long, 1 mm wide, glabrous to short s t i p i t a t e glandular throughout and few b i f u r c a t e hairs d i s t a l l y , pappus white, b a r b e l l a t e , the l a t e r a l setae .05-107 mm long, 37-48 b r i s t l e s . Apomictic. 2n=57. Type: Montana, Madison Range, Spanish Peaks, 1800 m, July 14, 1896, J.H. Flodman 901 (NY). D i s t r i b u t i o n : Arnica g r a c i l i s i s found on alpine scree slopes and rocky slopes i n open forests at mid to high elevations (1000-2400 m) i n Cascade to Rocky Mountains from southern B.C. and A l b e r t a to Wyoming and Oregon. Figure 305. I l l u s t r a t i o n : Figure 306. There has been much debate regarding the v a l i d i t y of Arnica g r a c i l i s . Maguire (1943) summarized his discussion of the species as "a loose e n t i t y which i s maintained as d i s t i n c t from Arnica l a t i f o l i a with some d i f f i c u l t y . " In more recent f l o r a s (Cronquist, 1955; Taylor & McBryde, 1978; Ediger & Barkley, 1978) the authors have treated i t as a v a r i e t y of the widespread Arnica l a t i f o l i a . In a subgenus (Austromontana) where there are no absolute-l y clear-cut species, and there are populations or i n d i v i d u a l s which are more or le s s intermediate or combine characters of two or more species, t h i s assemblage of populations seems as good as most of the other species. In the present treatment i t i s without h e s i t a t i o n retained as a separate species. Where Arnica g r a c i l i s grows i n i t s t y p i c a l form at high elevations on s t a b i l i z e d scree slopes, e s p e c i a l l y i n the Rocky Mountains of B r i t i s h Colum-b i a , Alberta, and Montana, i t i s densely clumped with l i t t l e rhizome growth 186 Figure 305. D i s t r i b u t i o n of Arnica g r a c i l i s (open c i r c l e s ) and Arnica discoidea (closed c i r c l e s ) . Stars indicate type l o c a l i t i e s . 187 188 Figure 306. Arnica g r a c i l i s S-1605. AB, Moraine Lake. X two-thirds. 189 190 each year (S-1603, 1605, 1613). Plants branch from near ground l e v e l and often above each p a i r of leaves, producing many heads per stem and per plant (Figures 307-318). At the other extreme, i n more shaded habitats often under Pinus ponderosa or Pinus contorta, as i n s t a b i l i z e d rock s l i d e s at Monroe Lake, BC (S-1562), the rhizomes are much longer and plants form open clumps more i n the nature of Arnica l a t i f o l i a or Arnica c o r d i f o l i a . These plants are r a r e l y branched, producing only one head ( r a r e l y 3) per stem. Yet the shape of the leaves, ray f l o r e t s , and the character of the short s t i p i t a t e glandular hairs i s t y p i c a l of those plants from higher elevations. (Figures 319-332). Arnica g r a c i l i s and Arnica l a t i f o l i a are e a s i l y distinguished i n the f i e l d . Arnica g r a c i l i s has a d i s t i n c t fragrance and v i s c i d nature to the leaves caused by the glandular hairs on the leaves. Arnica l a t i f o l i a i s usual-l y glabrous and when i t does have pubescence on the leaves ( e s p e c i a l l y from more northern l a t i t u d e s ) the pubescence i s non-glandular or predominately so, and i t i s usually not at a l l fragrant when bruised. Arnica g r a c i l i s leaves also look darker green, look and f e e l rough to the touch, and are noticeably rough under 10X magnification. Arnica g r a c i l i s also has fewer, narrower, more pointed rays (Figures 246, 247), long-petiolate ovate leaves, and broad ovate, usually s e s s i l e bracts. Very r a r e l y , and only at high elevations, does Arnica  l a t i f o l i a have the very short rhizomes t y p i c a l of Arnica g r a c i l i s . The predominance of t r i p l o i d apomictic populations has been demonstrated in t h i s species. I t probably represents a complex of apomictic races having o r i g i n a l l y arisen from h y b r i d i z a t i o n between the d i p l o i d Arnica l a t i f o l i a and the t e t r a p l o i d races of Arnica m o l l i s or Arnica d i v e r s i f o l i a (the l a t t e r two both i n Subgenus Chamissonis). The glandular pubescence of leaves and stems, 191 short scaly rhizomes, black, often glandular achenes, and often tawny pappus are a l l c h a r a c t e r i s t i c s s i m i l a r to the l a t t e r two species. Indeed, there are many populations which are separated only with d i f f i c u l t y from these species, e s p e c i a l l y when using herbarium specimens. Examples are Moraine Lake, Banff National Park, AB, where Arnica g r a c i l i s (S-1605) grows with a small form of Arnica d i v e r s i f o l i a (S-1606) on a scree slope. The l a t t e r i s di s t i n g u i s h e d i n the f i e l d by the more v i s c i d leaves, darker pappus, and l e s s branched habit . A r t i f i c i a l crosses between A r n i c a l a t i f o l i a and Arnica m o l l i s or Arnica  d i v e r s i f o l i a have been mostly unsuccessful. Arnica m o l l i s has grown very poorly i n c u l t i v a t i o n and a few crosses between i t as the male parent and Arnica l a t i f o l i a as the female parent have produced no achenes. Arnica d i v e r -s i f o l i a has grown well i n c u l t i v a t i o n . However only one cross between t h i s , and A r n i c a l a t i f o l i a has been s u c c e s s f u l , producing a few v i a b l e achenes. However, these did not survive beyond a few weeks. So, e f f o r t s to a r t i f i c i a l -l y produce an Arnica g r a c i l i s - l i k e hybrid have not been succ e s s f u l . There may also be confusion' between Arnica g r a c i l i s and Arnica r y d b e r g i i (Subgenus A r c t i c a ) , where they are growing i n s i m i l a r h a b i t a t s . However, Arnica r y d b e r g i i usually has more s e s s i l e c a u l i n e leaves which have a very d i s t i n c t p a i r of l a t e r a l veins nearly p a r a l l e l to the mid-rib, giving the leaves a t r i - n e r v e d appearance. The pappus i s pure white, the rays are darker orange-yellow, and the achenes are densely h i r s u t e throughout,in Arnica  r y d b e r g i i . 5. Arnica cernua T. Howell, FI. NW. Am. 373. 1903. Arnica c h a n d l e r i Rydb., N. Am. FI. 34:339. 1927. TYPE: C a l i f o r n i a , Humboldt Co., Hoopa Indian Reservation, June 1901, H.P. Chandler 1298 (holotype NY; isotypes MO, UC). 192 Figures 307-318. Vari a t i o n i n basal leaves of a single population t y p i c a l high-elevation Arnica g r a c i l i s S-1603, AB Lake Louise. 194 Figures 319-332. Va r i a t i o n i n basal leaves of a single population of lower-elevation Arnica g r a c i l i s S-1562, BC, Monroe Lake. 195 196 Rhizomatous perennial herb forming large open populations or t i g h t clumps of s t e r i l e basal rosettes and flowering stems; rhizomes (1-)4-15(-20) cm growth/year, (l-)2-3(-5) mm thick, naked except pairs of brown scales at nodes, and crowded scales and old leaf bases toward summit, simple or sparse-l y to much branched e s p e c i a l l y toward summit; roots 2-3 mm thick, mostly unbranched, succulent; stems single or few, simple or r a r e l y branched near base or higher, (5-)10-30(-45) cm high, glabrate to sparsely v i l l o u s with i n -curved h a i r s , p u r p l i s h or reddish at base or throughout; innovations with 4-8 long p e t i o l a t e leaves, p e t i o l e s (l-)4-6(-10) cm, blades 4-6(-7) cm long 2-4 (-5) cm broad, e l l i p t i c to ovate or rotund, bases rounded to subcordate, t i p s rounded to acute, margins e n t i r e or r e g u l a r l y or i r r e g u l a r l y serrate, dentate, crenate, or l a c i n i a t e and often undulate; cauline leaves 2-3(-5) p a i r s , the larges t p a i r usually near base of stem, long p e t i o l a t e , p e t i o l e s (-l)2-4(-6) cm long, blade (0.5-)2-4(-6) cm long, (l-)2-4(-5) cm broad, p e t i o l e s and blade siz e reduced upward, upper-most b r a c t - l i k e , often broadly s e s s i l e and often alternate; shape and margins of cauline leaves s i m i l a r to that of innovation leaves, although often more deeply i n c i s e d , leaves thick and succulent and often reddish or p u r p l i s h e s p e c i a l l y along veins of lower surface; herbage glabrate to scabrid e s p e c i a l l y on margins and along main veins, r a r e l y densely scabrid and p i l o s e on upper surface; inflorescence usually a s i n g l e head or an open corymb of 3-5 heads, nodding in bud, nodding or erect at anthesis, ped-uncles (2-)5-15(-22) cm long, upper peduncle and periclinum sparsely to densely p i l o s e and scabrid; heads 15-25 mm t a l l , radiate; i n v o l u c r a l bracts 7-12(-15), + u n i s e r i a t e , 10-15 mm long, 3-5 mm broad, broadly lanceolate, acute to obtuse; ray f l o r e t s (5-)7-9(-14), dark yellow, tube 4-6 mm long, limb 10-20 mm long, 4-8 mm broad, l i n e a r e l l i p t i c with 1-3 teeth at t i p , tubes and base of limbs densely short v i l l o u s ; d i s c f l o r e t s 24-57, tubular 10-15 mm 197 long, sparsely to densely v i l l o u s at base and sparingly toward summit and on lobes; pollen diameter 35-50 Um; achenes grey, 6-8 mm long, 1+ mm broad, sparsely to u s u a l l y densely h i r s u t e with b i f u r c a t e hairs throughout or from mid-point d i s t a l l y ; pappus pure white, 41-60 barbellate to subplumose b r i s t l e s , the l a t e r a l setae .10-.15 mm long; sexual d i p l o i d , 2n=38. Type: Oregon, Josephine County, near Waldo, on dry banks, base of Coast Mountains, June 1884, T. Howell 166 (ORE). D i s t r i b u t i o n : Arnica cernua i s r e s t r i c t e d to serpentine s o i l s at mid-elevations (400-1800 m) of the Coast Ranges and Siskiyou Mountains i n three counties of Southwestern Oregon (Coos, Curry, and Josephine) and three coun-t i e s of Northwestern C a l i f o r n i a (Del Norte, Siskiyou, and Humboldt). Figure 333. I l l u s t r a t i o n : Figure 334. This d i s t i n c t species i s one of the rarest Arnicas, although i t i s often common where i t does occur. Populations occupy rock crevices or rubble i n open exposed habitats in open f o r e s t s of Pinus j e f f r e y i , Calocedrus decurrens, Garrya fremontii, and Arctostaphylos spp., or at lower elevations are scat-tered i n open f o r e s t s of Pinus ponderosa, Pseudotsuga menziesii, and Calo-cedrus decurrens. I t i s not known to occur o f f serpentine s o i l s . Arnica cernua i s obviously derived from Arnica l a t i f o l i a , being most s i m i l a r to that species, but d i f f e r i n g from i t i n a number of characters other than t h e i r d i f f e r e n t e c o l o g i c a l niches. The thick, leathery leaves, which are usually purple or red are d i s t i n c t i v e i n Arnica cernua. The leaves of Arnica l a t i f o l i a are never as thick and fleshy nor are they often more 198 Figure 333. D i s t r i b u t i o n of Arnica spathulata (open c i r c l e s ) and Arnica cernua (closed c i r c l e s ) . Stars indicate type l o c a l i t i e s . 199 200 Figure 334. Habit of Arnica cernua S-1386. OR, Josephine X two-thirds. 201 202 than s l i g h t l y suffused with red or purple. The nodding or cernuous buds, from which Arnica cernua gets i t s name, may or may not become erect at anthesis. No specimens of Arnica l a t i f o l i a have been seen with nodding buds, Both the p h y l l a r i e s (Figures 130, 132) and the rays (Figures 139, 140) of Arnica cernua are larger and broader than those of Arnica l a t i f o l i a . The d i s t i n c t teeth at the t i p of the ray f l o r e t s of Arnica cernua d i f f e r from the more truncate t i p s of the Arnica l a t i f o l i a ray f l o r e t s . The achenes of Arnica cernua are much more densely hirsute d i s t a l l y or throughout than the smaller achenes of Arnica l a t i f o l i a . The species consists of two f a i r l y d i s t i n c t ecotypes, a small form (Figure 336) from exposed rocky slopes at higher elevations (750-1450 m) as i n c o l l e c t i o n s from Hoopa Indian Reservation and Horse Mountain, both from Humboldt Co., CA (Straley 1396, 1772) and a larger form (Figure 337) from lower elevations (400 m) i n open f o r e s t habitats as from near O'Brien, Josephine Co., OR (Straley 1386). The smaller ecotype i s barely rhizomatous, forming dense clumps with many stems, which often branch above producing 3-5 or more heads per stem. The leaves tend to be shorter-petioled and are r e l a -t i v e l y small (Figures 339-355). The la r g e r ecotype i s strongly rhizomatous forming large "loose" colonies. The stems are usually produced s i n g l y , are unbranched and consistently produce a single head per stem. Within popula-tions of the l a r g e r ecotype there i s a marked difference between plants growing i n nearly f u l l sun in that they have thicker , smaller, shorter p e t i o l e d leaves (Figures 356-365) compared to the thinner, l a r g e r , longer p e t i o l e d leaves of plants growing nearby i n rather dense shade (Figures 366-370). Although Rydberg (1927) considered the small ecotype a separate spe-c i e s , recognizing i t as Arnica chandleri, there are many c o l l e c t i o n s with 203 Figures 335-338. Habitat and habit of Arnica cernua. Figure 335. Habitat at summit of Hoopa-Redwood Valley Road, e l e . 750 m, Hoopa Indian Reservation, Humboldt Co., CA, Straley 1772. Serpentine s o i l s i n open fo r e s t of Pinus j e f f r e y i , Pinus lambertiana, Calocedrus decurrens, Arbutus menziesii, and Arctostaphylos spp. Figure 336. Dense clump with two open heads and many buds at above l o c a l i t y . Figure 337. Larger form with one head per stem i n dense forest of Pinus ponderosa, Calocedrus decurrens, and Arctostaphylos spp. at O'Brien, Josephine Co., OR, Straley 1386. Figure 338. Seed head from above population. 205 Figures 339-355. V a r i a t i o n i n basal rosette leaves of Arnica cernua S-1772, CA, Humboldt Co., small ecotype. 206 207 Figures 356-365. Va r i a t i o n i n basal rosette leaves of Arnica cernua S-1743, OR, Josephine Co., large ecotype, from plants growing i n f u l l sun. 209 Figures 366-370. V a r i a t i o n i n basal rosette leaves of Arnica cernua S-1743, OR, Josephine Co., large ecotype, from plants growing i n shade. 210 211 intermediate characters, so there seems to be no reason to give these populations separate taxonomic recognition. There are a number of exceptional c o l l e c t i o n s of t h i s species, including extremely narrow-leaved specimens from Red Mountain, Curry Co., OR, Leach  3442 (ORE), with the l a r g e s t cauline l e a f blades only 1.5 cm long and 0.5 cm broad and a c o l l e c t i o n from Tennessee Pass, Josephine Co., OR, Savage s.n. (ORE) with the lower cauline l e a f blades broader (5 cm) than long (3.5 cm) and deeply lobed. In an unusually pubescent specimen from Head S l i d e Creek, Curry Co., OR, Leach 2879 (ORE), a l l leaves and stems are scabrous throughout gi v i n g the plants a grey appearance. This population may represent a l o c a l ecotype or may r e f l e c t p ossible h y b r i d i z a t i o n with one of the pubescent species, l i k e l y Arnica c o r d i f o l i a . The Type specimen, Howell 166 (ORE) consists of three flowering stems and one basal rosette. Two of the plants are s i m i l a r to most of the l a t e r c o l l e c t i o n s of the species. However, the t h i r d specimen on the sheet has two sets of d i s t i n c t whorls of three leaves rather than opposite p a i r s , an unusual character f o r the genus. 6. Arnica discoidea Bentham. PL. Hartw. 319. 1849. Arnica p a r v i f l o r a A. Gray. Proc. Am. Acad. 7:363. 1867. TYPE: C a l i f o r n i a , Humboldt Co., chapparal, 1867, Geol. Surv. C a l i f . , H.H. Bolander 6051. (holotype, G; isotype, KEW). Arnica c o r d i f o l i a Hook, var. eradiata A. Gray. Syn. F l . N. Am. 1:381. 1884. TYPE: Oregon, Hood River, 1884, Barrett, s..n. (G) . Arnica discoidea Bentham var. eradiata (A. Gray) Cronquist. Vase. PI. Pac. NW. 5:49. 1955. 212 Arnica grayi H e l l e r . Muhlenbergia 1:5. 1900. TYPE: Oregon, Hood River, 1884, Barrett, _s.n. (G) . Arnica f a l c o n a r i a Greene. Ottawa Nat. 23:215. 1910. TYPE: Washington, K l i c k i t a t Co., Falcon Valley, June 27, 1892. W. Suksdorf 1617 (ND). Arnica a l a t a Rydb. N. Am. FI. 34:342. 1927. TYPE: C a l i f o r n i a , Yosemite, 1865, J . Torrey 258a (NY). Arnica p a r v i f l o r a A. Gray subsp. a l a t a (Rydb.) Maguire. B r i t t o n i a 4:455. 1943, Arnica discoidea Bentham var. a l a t a (Rydb.) Cronquist. Contr. Dudley Herb. 5:102. 1958. Arnica sanhedrensis Rydb. N. Am. FI. 34:342. 1927. TYPE: C a l i f o r n i a , Lake Co., south of Mt. Sanhedrin, July 25, 1902, H e l l e r 5985 (holotype, NY; isotype, POM). Rhizomatous perennial herb; rhizomes 5-15(-22) cm growth/year, 2-5 mm thick, naked except f o r pairs of brown scales at nodes and crowded scales and old leaf bases toward summit, forming large populations of s t e r i l e basal rosettes and flowering stems; roots + succulent, 2-3 mm thick, r e l a t i v e l y unbranched; stems single or few, simple or branched near base or throughout, (15-)20-45(-65) cm t a l l , sparsely to densely s t r i g o s e , v i l l o u s , and long s t i p i t a t e glandular throughout; innovations with 4-8(-12) long p e t i o l a t e leaves, p e t i o l e s (l-)4-6(-10) cm long, blades, ovate or broadly ovate to broadly lanceolate, 3-5(-9) cm long, 2-4(-5) cm broad, bases cordate, sub-cordate, to rounded, or r a r e l y attenuate, t i p s acute to rounded, margins subentire to usually r e g u l a r l y serrate, dentate, or crenate, or r a r e l y doubly serrate or dentate; cauline leaves (2-)3-5(-6) p a i r s , often reduced b a s a l l y and becoming b r a c t - l i k e toward summit, lower cauline leaves usually the l a r g e s t , long p e t i o l a t e , p e t i o l e s 2-6(-10) cm long, often winged 2-5(-14) mm, 213 blades (3-)4-7(-9) cm long, 2-6 cm wide, upper pairs progressively shorter p e t i o l e d , and often subalternate, shape and margins of cauline leaves l i k e that of innovations; v e s t i t u r e of herbage usually moderately to densely scabrous and v i l l o u s , e s p e c i a l l y along veins, and sparsely long s t i p i t a t e glandular throughout; inflorescences an open corymb of 3-7(-20) heads, r a r e l y subradiate, 12-15 mm high; i n v o l u c r a l bracts, 7-12 + u n i s e r i a t e , 10-15 mm long, 2-3(-5) mm wide, narrowly to broadly lanceolate, acute to acuminate; f l o r e t s 19-45, tubular, 8-12 mm long, marginal f l o r e t s often ampliate or r a r e l y + l i g u l a t e , to 20 mm long, 5 mm wide, sparsely to densely v i l l o u s and sparsely s t i p i t a t e glandular at base and sparingly upward, and on outer sur-face of lobes; pollen diameter 45-60 um; achenes dark grey, 6-8 mm long, 1 mm wide, sparsely to densely hirsute with b i f u r c a t e h a i r s , and s t i p i t a t e glandu-l a r throughout, e s p e c i a l l y d i s t a l l y ; pappus white, ba r b e l l a t e , of 42-64 b r i s t l e s , l a t e r a l setae .05-.15 mm long; apomictic; 2n=38, 76. Type: C a l i f o r n i a , Monterey Co., Monterey, i n f o r e s t , Hartweg 1805 (KEW). D i s t r i b u t i o n : This species occupies a wide var i e t y of habitats from open chapparal to conifer f o r e s t s , near sea l e v e l to 1800 m, from Southern Washing-ton ( K l i c k i t a t County) to Southern C a l i f o r n i a (Orange County). (Figure 305). I l l u s t r a t i o n : Figure 371. There has been much confusion and difference of opinion i n the treatments of Arnica discoidea, the most widespread and most v a r i a b l e of the d i s c o i d species. Arnica discoidea, as presented here, contains elements recognized by Maguire (1943) as three species, Arnica discoidea, Arnica p a r v i f l o r a (with 2 subspecies), and Arnica g r a y i . Ediger & Barkley (1978) tre a t these as one species, Arnica discoidea, with three v a r i e t i e s . 214 Figure 371. Arnica discoidea S-1775. CA, Mendocino Co. X two-thirds. 215 216 At the northern l i m i t of t h e i r d i s t r i b u t i o n there are populations formerly known as Arnica grayi H e l l e r , from K l i c k i t a t Co., WA (S-1825), Wasco Co., OR (S-1821) , and Hood River Co., OR, Barrett, s.xi. (G). , growing with Arnica c o r d i f o l i a . The leaves of these northern populations tend to be more densely strigose and with somewhat less cordate bases than i n most of the southern populations. However, there are numerous intermediates through-out the range of the species, as i n collections from San Mateo Co., CA, Keck  1775 (RSA). V a r i a b i l i t y i n leaf s i z e , shape, and especially i n margins can be seen i n many populations (Figures 376-385) and i s p a r t i c u l a r l y notable i n a c o l l e c -t i o n from Mt. Eddy, Siskiyou Co., CA, Copeland 3909 (POM). Many populations throughout the range of the species have marginal amp-l i a t e f l o r e t s (Figure 155) or elongated r a y - l i k e f l o r e t s (Figures 154, 156). This condition i s especially pronounced i n collections from Santa Barbara Co., CA, Smith 8330 (SBM), T r i n i t y Co., CA, Baker 328 (HSC), and Eldorado Co., CA, Peirson 9517 (RSA). S u p e r f i c i a l l y , these collections look very much l i k e Arnica c o r d i f o l i a and are often misidentified as such. However, none of the collections has true female ray f l o r e t s , although they are l i k e l y often func-tioning as female. Even the elongated marginal f l o r e t s are usually + tubular (Figures 154, 156) and have functional or abortive anthers. Arnica discoidea d i f f e r s from Arnica c o r d i f o l i a i n features other than an absence of ray f l o r e t s . Arnica discoidea i s usually more branched, has more heads (up to 20/stem), i s less strongly rhizomatous, and usually prefers drier habitats than Arnica c o r d i f o l i a . The discoid species also tends to be less pubescent, or at least the pubescence tends to be shorter than that of Arnica  c o r d i f o l i a . Trends toward, and differences between Arnica spathulata and 217 Figures 372-375. Habitat and habit of Arnica discoidea and Arnica spathulata. Figure 372. Habitat of Arnica discoidea S-1825, WA, K l i c k i t a t Co., K l i c k i t a t River Gorge, open forest of Pinus ponderosa and Pseudotsuga menziesii. Figure 373. same, flowering stems and basal rosettes. Figure 374. same, three flowering heads and two seed heads. Figure 375. Arnica spathulata S-1385, OR, Josephine Co., O'Brien. 218 219 Figures 376-385. V a r i a t i o n i n basal leaves of Arnica discoidea S-1789, CA, Marin County. 220 221 Arnica discoidea are discussed under Arnica spathulata. There does not seem to be any c o r r e l a t i o n between d i s t r i b u t i o n , chromosome number, and si z e of heads, whether leaves are b a s a l l y disposed or evenly d i s t r i b u t e d along the stem, or achene and pappus features i n any of the d i s c o i d populations. These features have been used to separate the d i s -coid species recognized i n previous treatments. I t seems best to treat these as one polymorphic species with both d i p l o i d , sexual and t e t r a p l o i d , apomictic races. 7. A m i ca. spathuXa tci Greens, Pittonia. 3 i l 0 3 . 1896. Arnica eastwoodiae Rydb. N. Am. F l . 34:343. 1927. TYPE: C a l i f o r n i a , Del Norte Co., Gasquet, French H i l l , September 14, 1912, A. Eastwood 2211 (NY). Arnica spathulata Greene subsp. eastwoodiae (Rydb.) Maguire. B r i t t o n i a 4:458. 1943. Arnica spathulata Greene var. eastwoodiae (Rydb.) Ediger & Barkley. N. Am. F l . I I . 10:43. 1978. Arnica c u s i c k i i Rydb. N. Am. F l . 34:343. 1927. TYPE: Oregon, Cascade Mountains, dry western slope, July 11, 1902, Cusick 2873 (holotype, NY; isotypes, MO, POM, US). Rhizomatous perennial herb often forming short woody caudex 0.5-1.5 cm broad, 1-2 cm high, 2-5(-7) cm below surface of ground, producing v e r t i c a l and h o r i z o n t a l rhizomes (l-)3-7(-14) cm/year, generally naked except p a i r s of brown scales at nodes and crowded scales and some old l e a f bases toward sum-mit, forming large open populations of s t e r i l e basal rosettes and flowering stems; roots r e l a t i v e l y few and succulent, 1-3 mm in diameter; stems single or few, simple to branched below and above, (15-)25-45(-55) cm t a l l , sparsely 222 to densely v i l l o u s and long s t i p i t a t e glandular throughout; innovations with 4-6(-10) long p e t i o l a t e leaves, p e t i o l e s 2-7 cm long, wingless or with wings 2-7(-12) mm broad, blades (2-)4-6(-8) cm long, 1-3.5 cm wide, spathulate to e l l i p t i c or obovate, bases attenuate or r a r e l y rounded, t i p s rounded to acute, margins e n t i r e to coarsely and evenly serrate or crenate; cauline leaves 3-6 p a i r s , basally disposed to + evenly d i s t r i b u t e d along stem, often subopposite, reduced and b r a c t - l i k e upward, where frequently alternate, with p e t i o l e s as wide as or wider than blade, l a r g e s t p a i r of cauline leaves with longest p e t i o l e s , usually near base, p e t i o l e s (l-)3-5(-8) cm. wings 2-10(-20) mm, blades (2-)3-6(-8) cm long, (l-)2-4(-5) cm wide, shape and margins as with leaves of innovations, veins usually prominent and reddish or p u r p l i s h e s p e c i a l l y on lower surface; v e s t i t u r e of herbage p i l o s e and long s t i p i t a t e glandular throughout, e s p e c i a l l y along veins, margins often c i l i a t e ; i n f l o -rescence a single head or an open corymb of 3-5(-25) heads, erect; peduncles (2-)4-8(-15) cm long, upper peduncles and periclinum long p i l o s e and s t i p i -tate glandular; heads 15-25 mm high, d i s c o i d , r a r e l y with ampliate marginal f l o r e t s ; i n v o l u c r a l bracts 7-13(-17), + u n i s e r i a t e , 10-15 mm long, 1-4 mm wide, lanceolate to broadly lanceolate, obtuse to acute; f l o r e t s 18-52, goblet-shaped to tubular, 8-11 mm long, dark yellow, sparsely to densely v i l l o u s and s t i p i t a t e glandular toward base and sparsely upward and on lobes; po l l e n diameter 35-55 um; achenes black, dark brown, or dark grey, short s t i p i t a t e throughout or from mid-point d i s t a l l y , few or no b i f u r c a t e h a i r s d i s t a l l y , 6-9 mm long, 1 mm wide, pappus pure white, 37-56 b r i s t l e s , barbel-l a t e , the l a t e r a l setae .07-.10 mm long; apomictic p o l y p l o i d and s e l f -incompatible sexual d i p l o i d ; 2n=38, 76. Type: Oregon, Douglas County, Glendale, June 3, 1887, T. Howell 1200 (US). 223 D i s t r i b u t i o n : Arnica spathulata occurs on open serpentine outcrops or open for e s t s at low to mid-elevations (180-1400 m) of Coast and Cascade Ranges from Douglas and Curry Counties i n SW Oregon to Siskiyou County i n NW C a l i f o r n i a . Figure 333. I l l u s t r a t i o n : Figure 386. This, the ra r e s t of the species i n section Eradiatae i s known from about two dozen populations l a r g e l y r e s t r i c t e d to serpentine s o i l s . Where i t does occur the populations often consist of several dozen to hundreds of i n d i v i d -u a ls. Populations or i n d i v i d u a l s are quite v a r i a b l e i n s i z e , branching habit, leaf shape, and pubescence. There i s a small ecotype to which the name Arnica  eastwoodiae Rydb. has been assigned. These populations, as i n c o l l e c t i o n s from French H i l l Rd., Gasquet, Del Norte Co., CA, (S-1751, probably the type l o c a l i t y of Arnica eastwoodiae) are smaller, have narrower leaves with l e s s -winged p e t i o l e s (Figures 387-398), narrower heads, and shorter pubescence than most other populations of the species. They are found on open serpentine slopes i n mixed f o r e s t s of Pinus attenuata, Calocedrus decurrens, and Pinus  ponderosa or Pinus j e f f r e y i . At the other extreme are a majority of popula-t i o n s , with t a l l e r stems, broader leaves, usually winged p e t i o l e s (Figures 399-407), and long glandular hairs throughout. Typical populations of t h i s l a rger ecotype are c o l l e c t i o n s from Kirby, Cave Junction, and Selma, a l l i n Josephine Co., OR, (S-1380, 1381, 1740). These populations are usually found on more densely forested h i l l s i d e s of Pinus ponderosa, Pseudotsuga menziesii, Arbutus menziesii, and Quercus k e l l o g g i i . There are numerous populations which are intermediate i n some or a l l of the above characters, including c o l -l e c t i o n s from Gasquet, Del Norte Co., CA, (S-1390) and O'Brien, Josephine Co., OR, (S-1385). 224 Figure 386. Arnica spathulata S-1740. OR, Josephine Co. X two-thirds. 225 226 Figures 387-398. V a r i a t i o n i n ba s a l r o s e t t e leaves of A r n i c a spathulata S-1751, CA, Del Norte Co., small ecotype. 227 228 Figures 399-407. Va r i a t i o n i n basal rosette leaves of Arnica spathulata S-1740, OR, Josephine Co., large ecotype. 229 2 3 0 O c c a s i o n a l l y i n t h e l a r g e - l e a v e d p o p u l a t i o n s , a s i n a s p e c i m e n f r o m P l e a s a n t C r e e k , J a c k s o n C o . , O R , Hammond 2 3 0 ( M O ) , t h e p e t i o l e s o f c a u l i n e l e a v e s a r e b r o a d a n d a r e o f t e n w i d e r t h a n t h e b l a d e s . T h i s c h a r a c t e r w a s n o t s e e n i n a n y o t h e r A r n i c a . A n u n u s u a l c o l l e c t i o n f r o m G o l d B a s i n , C u r r y C o . , OR , L e a c h 2 8 7 4 (ORE ) h a s s t r o n g l y b a s a l l y d i s p o s e d c a u l i n e l e a v e s , a n d u n -u s u a l l y d e n s e , s h o r t - p i l o s e v e s t i t u r e . A r n i c a s p a t h u l a t a m o s t c l o s e l y r e s e m b l e s A r n i c a d i s c o i d e a f r o m w h i c h i t c e r t a i n l y h a s b e e n d e r i v e d . T h e f o r m e r d i f f e r s i n i t s s p a t h u l a t e l e a v e s w h i c h a r e m o r e o f t e n p u r p l e o r r e d , i n i t s t a l l e r h e a d s , l a r g e r , d a r k e r a c h e n e s , a n d i n t h e c h a r a c t e r o f i t s r h i z o m e s . A s o m e w h a t i n t e r m e d i a t e p o p -u l a t i o n h a s b e e n c a l l e d A r n i c a c u s i c k i i R y d b . , C u s i c k 2 8 7 3 (NY, P O M , MO, U S ) . T h e e x a c t l o c a l i t y i s n o t k n o w n , o n l y " d r y w e s t e r n s l o p e o f t h e C a s c a d e M o u n t a i n s , S o u t h e r n O r e g o n . " T h e s e v e r a l s p e c i m e n s a r e v a r i a b l e , b u t t h e c h a r a c t e r o f t h e b a s a l l y d i s p o s e d c a u l i n e l e a v e s w i t h c o r d a t e b a s e s c l o s e l y r e s e m b l e s some A r n i c a d i s c o i d e a s p e c i m e n s , s o much s o t h a t i n t h e m o s t r e c e n t m o n o g r a p h o f t h e g e n u s ( E d i g e r a n d B a r k l e y , 1 9 7 8 ) , i t i s l i s t e d i n t h e s y n o n -ymy u n d e r A r n i c a d i s c o i d e a . H o w e v e r , i n t h e c h a r a c t e r s o t h e r t h a n t h e l e a v e s , e s p e c i a l l y i n t h e v e s t i t u r e , r h i z o m e s , a n d a c h e n e s , i t s e e m s t o f i t b e s t i n A r n i c a s p a t h u l a t a a n d i s h e r e i n r e t a i n e d a s a s y n o n y m o f t h a t s p e c i e s . A m p l i a t e o r e l o n g a t e d r a y - l i k e m a r g i n a l f l o r e t s s e e m t o b e much r a r e r i n t h i s s p e c i e s t h a n i n A r n i c a d i s c o i d e a . O n l y o n e s p e c i m e n , T r a c y 1 9 4 2 4 ( U C ) f r o m D e l N o r t e C o . , C A , n o r t h o f M o n u m e n t a l , h a s b e e n s e e n w i t h l o n g r a y - l i k e a m p l i a t e f l o r e t s . I t w a s r e p o r t e d a s g r o w i n g w i t h t y p i c a l d i s c o i d p l a n t s . 231 Subgenus Calarnica subgen. nov. Herbae perennes erectae a caudice ligno subterraneo; rhizomata carentes; radices succulentae, v e r t i c a l e s ; caules ramosi; f o l i a caulina ovata, s e s s i l e ; plantae dense v e l sparsim stipitato-glandulosae et pilosae; c a p i t u l a eradiata; pappus albusve subinfuscatus suffusus, barbellatusve subplumosus; chromosoma-tum numerus gamatae n=19. Erect perennial herbs from woody caudex, non-rhizomatous; roots f l e s h y , + v e r t i c a l and unbranched; lacking s t e r i l e basal rosettes (innovations); flow-ering stems branching from base, or sparingly to much-branched above; stems equably l e a f y , cauline leaves s e s s i l e , ovate, margins en t i r e to serrate, bases rounded, apex acute to rounded; stems and leaves p i l o s e and long s t i p i t a t e glandular; inflorescence a s i n g l e head or much-branched l e a f y corymb; heads d i s c o i d ; i n v o l u c r a l bracts narrowly to broadly lanceolate, acute; f l o r e t s tubular; achenes ribbed, hirsute and s t i p i t a t e glandular; pappus white to tawny, ba r b e l l a t e ; gametic chromosome number n=19. Type: Arnica v i s c o s a A. Gray 8. Arnica v i s c o s a A. Gray. Proc. Am. Acad. 13:374. 1878. R a i l l a r d e l l a paniculata Greene. Erythea 3:48. 1895. TYPE: C a l i f o r n i a , Siskiyou Co., Mt. Shasta, near the l i m i t of trees, August 4, 1894. W._L. Jepson, _s .n_. (not seen) . Chrysopsis shastensis Jeps. Man. FI. PI. Cal. 1037. 1927. TYPE: C a l i f o r n i a , Siskiyou Co., Mt. Shasta, Horse Camp, 2400 m, W.L. Jepson 51i (JEPS). Perennial herb from woody caudex 1-4 cm broad, covered with old stem bases and brown scales; rhizomes lacking; roots succulent, mostly v e r t i c a l 232 and unbranched to 40 cm long, 3-6 mm thick; stems s o l i t a r y or several, much branched throughout, 25-50 cm t a l l at anthesis, sparsely short s t i p i t a t e glandular and p i l o s e below, densely long s t i p i t a t e glandular and + p i l o s e above; innovations not produced; cauline leaves with 6-10(-12) p a i r s , l a r g e s t near middle of stem, reduced and s c a l e - l i k e toward base, reduced and bract-l i k e above, upper or a l l leaves/bracts often alternate or subopposite, l a r -gest cauline leaves 3-5 cm long, 1.5-3.5 cm broad, narrowly to broadly e l l i p t i c , broadly s e s s i l e , margins e n t i r e , bases rounded to acute, t i p s acute to obtuse, both surfaces sparsely to densely p i l o s e , e s p e c i a l l y along veins and margins, densely long s t i p i t a t e glandular, with yellow-tipped h a i r s ; heads s o l i t a r y terminating a l l upper branches; peduncles 0.5-5 cm long, peduncles and periclinum long s t i p i t a t e glandular; heads erect, d i s c o i d , 1.5-2 cm high; i n v o l u c r a l bracts, 8-14(-21), + u n i s e r i a t e , 6-12 mm long, 1-3 mm wide, narrowly lanceolate, acute; f l o r e t s 9-32(-54) tubular, 8-12 mm long, white to creamy-white, densely long s t i p i t a t e glandular below; po l l e n diameter 40-45 um; achenes grey to brown, sparsely to densely long s t i p i t a t e glandular throughout, shallowly ribbed, 4-6 mm long, to 1 mm wide, pappus off-white to tawny, 36-48 b r i s t l e s , b a r b e l l a t e , 5-9 mm long, l a t e r a l setae .05-.15 mm long; self-incompatible d i p l o i d ; 2n=38. TYPE: C a l i f o r n i a , Siskiyou County, Mt. Shasta, 2400 m, 1877, J.D. Hooker & A. Gray, _s._n. (G) . D i s t r i b u t i o n : This species occurs on loose pumice slopes at high elevations (1750-2500 m) in Crater Lake National Park, Klamath County, Oregon and i n Western Siskiyou and Northern T r i n i t y Counties, C a l i f o r n i a . Figure 408. 233 Figure 408. A l l known populations of Arnica v i s c o s a . @ ) 235 I l l u s t r a t i o n : Figure 409. Arnica viscosa i s the most d i s t i n c t i v e and one of the rarest Arnicas. Its much-branched habit, small, e n t i r e , s e s s i l e leaves, narrow, white, d i s -coid heads, and tawny pappus set i t apart from a l l other species. It i s the only species with white f l o r e t s , a character strangely never mentioned i n previous descriptions or monographs. The prominent yellow anthers give the heads a d e f i n i t e yellowish cast which may be the reason why the white f l o r e t s have not been observed previously. Arnica viscosa i s strongly v i s c i d throughout with long yellow-tipped glandular hairs on a l l vegetative parts and on f l o r e t s and achenes. I t s strong odor i s unlike that of any other Arnica. The odor i s retained i n herbarium specimens f o r many years. Described from Mt. Shasta, C a l i f o r n i a i n 1887 by Asa Gray, Arnica  viscosa was long known only from the type l o c a l i t y (Figures 410-412) and from G a r f i e l d Peak i n Crater Lake National Park, Oregon. Only i n the past 20 years has i t been noted from at l e a s t four other places i n Crater Lake, f i v e l o c a l i t i e s i n Western Siskiyou County, C a l i f o r n i a , and two l o c a l i t i e s i n Northern T r i n i t y County, C a l i f o r n i a , i n the Salmon Mountains and T r i n i t y Alps. Where Arnica viscosa has been observed on Mt. Shasta and G a r f i e l d Peak there i s v i r t u a l l y no other vascular plant l i f e . The very deep roots are able to grow almost v e r t i c a l l y among the loose pumice into the s o i l below. Both of these populations are quite large, composed of several hundred plants. A study of 25 herbarium specimens and two populations i n the f i e l d showed le s s v a r i a b i l i t y i n t h i s species than i n many of the others. There i s a great tendency f o r upper branches, leaves, and bracts to be subopposite to complete-l y a lternate. An extreme example of t h i s i s a specimen from Upper E n g l i s h 236 Figure 409. Arnica viscosa S-1411. CA, Siskiyou County, Mt. Shasta (Type L o c a l i t y ) . X two-thirds. 237 238 Figures 410-414. Habitats and habit of Arnica viscosa. Figures 410-412. Habitat of Arnica viscosa S-1411. CA, Siskiyou Co., Mt. Shasta (Topotype). Figure 413. Habitat of Arnica viscosa S-1946, OR, Klamath Co., Crater Lake. Figure 414. Flowering head of Arnica viscosa S-1411, CA, Siskiyou Co., Mt. Shasta. 2 3 9 240 Lake, Siskiyou County, CA, Oettinger 668 (HSC). One unusually p i l o s e specimen i n which the stem and veins are predominately long p i l o s e rather than predominately long s t i p i t a t e glandular as usual was c o l l e c t e d on Preston Peak, Siskiyou County, CA, Ground 820 (PUA). There i s a questionable c o l l e c t i o n by W.L. Jepson from Mt. Shasta, named R a i l l a r d e l l a paniculata by Greene. I t f i t s the description and exact l o c a l -i t y of Arnica viscosa. As mentioned e a r l i e r there i s v i r t u a l l y no other plant l i f e growing near the Arnica at that elevation on Mt. Shasta, so t h i s c o l l e c t i o n must be t h i s species and i s l i s t e d under the synonymy although a voucher specimen was not c i t e d i n the o r i g i n a l description and has not been located. 9. Arnica venosa H.M. H a l l . Univ. of Cal. Pubis. Bot. 6:174. 1915. ' Perennial herb from woody caudex 3-5 cm broad covered with o l d stem bases and dark brown scales; rhizomes lacking; roots succulent, mostly un-branched, to 30 or more cm long, 2-5 mm thick; stems s o l i t a r y or few, simple to r a r e l y branched above upper pa i r s of leaves/bracts, 30-50 cm high at anthesis, sparsely scabrous and p i l o s e below, becoming more densely p i l o s e and s t i p i t a t e glandular above; cauline leaves 7-10 p a i r s , largest near middle of stem, reduced and becoming s c a l e - l i k e toward base, and reduced and bract-l i k e above, upper leaves and bracts often subopposite or alternate, largest cauline leaves (3-)4-6(-7) cm long, (1.5-)2-3(-4) cm broad, narrowly e l l i p t i c to broadly so, broadly s e s s i l e , margins evenly coarsely serrate or r a r e l y doubly serrate, bases rounded, t i p s acute to obtuse, rugose, veins prominent, e s p e c i a l l y on lower surface, lower surface sparsely to densely p i l o s e and sparsely s t i p i t a t e glandular; inflorescence a single head terminating main 241 stem and l a t e r a l branches, i f any; peduncles 1.5-5 cm long, peduncles and periclinum densely p i l o s e ; heads nodding, e s p e c i a l l y i n bud, d i s c o i d , 2-2.5 cm high; i n v o l u c r a l bracts 8-14, + u n i s e r i a t e , 10-15 mm long, 4-6 mm broad, broadly lanceolate, acute to obtuse; f l o r e t s 32-56(-70), tubular, 8-10 mm long, dark yellow, densely v i l l o u s , e s p e c i a l l y below; pollen diameter 35-40 pm; achenes grey, strongly angled and ribbed, 6-8 mm long, 1.5 mm broad, densely hirsute with bifurcate h a i r s ; pappus white, of 45-55 b r i s t l e s , 6-10 mm long, b a r b e l l a t e , the l a t e r a l setae 0.1-0.2 mm long; self-incompatible; 2n=38. TYPE: C a l i f o r n i a , Shasta County, Salt Creek, 430 m, June 1903, H.M. H a l l & _E.F. Bab cock 4013 (UC). D i s t r i b u t i o n : Arnica venosa i s found i n f o o t h i l l s (430-530 m) of the Sacramento River Valley i n mixed conifer and Quercus spp. f o r e s t of Shasta County, C a l i f o r n i a . Figure 415. I l l u s t r a t i o n : Figure 416. This d i s t i n c t i v e species, c l o s e l y r e l a t e d to Arnica viscosa, has toothed leaves, nodding buds, yellow f l o r e t s , and a paucity of glandular h a i r s , set-t i n g i t apart from that species. This too, i s one of the rarer Arnica species. For many years i t was known from only two l o c a l i t i e s i n Shasta County, C a l i f o r n i a , Iron Mountain and S a l t Creek, the l a t t e r the type l o c a l -i t y . The exact l o c a t i o n of "Salt Creek" i s not known. There are several S a l t Creeks i n Shasta County, including one which i s now p a r t i a l l y under Lake Shasta. Iron Mountain i s now the s i t e of an active copper mining operation and the Arnica may have been exterminated from that l o c a l i t y . The mine owners w i l l not allow botanists to search the area. Having not been c o l l e c t e d 242 Figure 415. A l l known populations of Arnica venosa. 0 50 100 200k m 244 Figure 416. Arnica venosa S-1793. CA, Shasta County. X two-thirds. 245 246 Figures 417-421. Habitat and habit of Arnica venosa S-1793, CA, Shasta Co., Gibson. Figures 417-418. Habitat, roadside bank i n open mixed f o r e s t of Quercus spp. and c o n i f e r s . Figure 419. Habit of numerous branches from base. Figure 420. Strongly nodding bud. Figure 421. Head with f i r s t f l o r e t s open and nodding buds, and rugose leaves. 248 fo r more than 50 years, i t was thought to be possibly e x t i n c t . Only i n recent years has there been renewed i n t e r e s t by the National Forest Service personnel i n the area to f i n d i t , and at l e a s t 5 other l o c a l i t i e s have been found, a l l i n Shasta County. I t i s found i n d i s c r e t e populations of a few i n d i v i d u a l s to a few dozen plants, i n open f o r e s t s of Pinus ponderosa, Pinus lambertiana, Pinus atten-uata, Pinus sabiniana, Pseudotsuga men z i e s i i , Quercus k e l l o g g i i , Quercus  chrysolepis, Calocedrus decurrens, Arctostaphylos spp. and Ceanothus spp., and along open roadsides i n these f o r e s t s . Plants sparingly colonize the disturbed roadside banks. Plants begin to flower i n mid- to l a t e May and bloom f o r a few weeks before the h i l l s become very hot and dry, although a few s t r a g g l e r s may continue to flower throughout the early summer months. In c u l t i v a t i o n seedlings continued to flower f o r many months when grown under wetter conditions. Specimens are very rare i n herbaria. Of about 6 specimens seen from 3 l o c a l i t i e s plus 3 other populations studied i n the f i e l d during the spring and summer of 1978, there seems to be very l i t t l e v a r i a b i l i t y i n the plants. 249 PHYLOGENY Maguire (1943) suggests that the genus Arnica probably arose i n a r c t i c or subarctic Western North America and spread from there to the east and south. He recognized f i v e subgenera with A r c t i c a as the a n c e s t r a l subgenus. He considered the l a r g e , hemispheric or turbinate heads and densely pubescent periclinum as being p r i m i t i v e characters. If these morphological characters are considered p r i m i t i v e , then the p o s s i b l e phylogenetic scheme shown i n Figure 422 can be deduced f o r the other subgenera. This i s a m o d i f i c a t i o n of Maguire's phylogenetic scheme f o r h i s f i v e subgenera, plus my newly proposed subgenus Calarnica. Within subgenus Austromontana, Arnica c o r d i f o l i a i s the probable l i n k between the two subgenera, A r c t i c a and Austromontana. This species has large, u s u a l l y s o l i t a r y heads, and densely p i l o s e periclinum. From Arnica c o r d i f o l i a , the other species i n Austromontana and Calarnica could l o g i c a l l y have evolved as shown i n Figure 423. Arnica c o r d i f o l i a occurs today l a r g e l y as a s e r i e s of apomictic p o l y p l o i d populations, from t r i p l o i d to hexaploid, with t e t r a p l o i d predominating. How-ever, d i p l o i d populations do occur r a r e l y . Whether or not any of these d i p l o i d s are sexual has not been demonstrated. Based on p o l l e n q u a l i t y and the u s u a l l y strong c o r r e l a t i o n of d i p l o i d y with s e x u a l i t y i n the genus, there i s a strong p o s s i b i l i t y that some of these d i p l o i d populations are sexual. Barker (1966) hypothesized that apomixis i n the genus i s a r e l a t i v e l y recent development ( i n t e r - or p o s t - g l a c i a l ) . Sexual d i p l o i d s are assumed to have been more common and widespread i n the past. D i p l o i d Arnica c o r d i f o l i a probably gave r i s e to Arnica l a t i f o l i a i n generally wetter h a b i t a t s . A r n i c a l a t i f o l i a u s u a l l y has narrower heads, with 250 fewer f l o r e t s , and a l e s s - p i l o s e periclinum than Arnica c o r d i f o l i a . These characters are considered as being derived. Arnica l a t i f o l i a i s the most polymorphic species i n the subgenus, although i t e x i s t s almost e n t i r e l y as sexual d i p l o i d populations, with many ecotypes. The close r e l a t i o n s h i p of these two species can be seen i n numerous populations which combine many of the morphological characters of both species. Arnica l a t i f o l i a has i n turn probably given r i s e to two other species. Arnica cernua, also a sexual d i p l o i d , i s a rare serpentine s o i l endemic. I t combines many morphological characters of Arnica l a t i f o l i a but i s e c o l o g i c a l l y d i s t i n c t . A second species probably derived from Arnica l a t i f o l i a , p o s s i b l y through h y b r i d i z a t i o n , i s Arnica g r a c i l i s . I t i s considered a v a r i e t y of Arnica l a t i f o l i a by many authors. However, i n my opinion, most populations can be e a s i l y d i s t i n g u i s h e d from Arnica l a t i f o l i a . Arnica g r a c i l i s i s l a r g e -l y a t r i p l o i d , autonomous apomict, found mostly at higher elevations or i n rockier habitats than Arnica l a t i f o l i a . Arnica g r a c i l i s i s more branched and has much smaller heads, with fewer f l o r e t s than Arnica l a t i f o l i a . I i n t e r p r e t these characters as being advanced. Arnica g r a c i l i s has the glandular pubes-cence found i n Arnica d i v e r s i f o l i a and Arnica m o l l i s i n subgenus Chamissonis. Both of these l a t t e r species have been shown to have t e t r a p l o i d elements. The chromosome number and morphology of Arnica g r a c i l i s suggests i t may be of hybrid o r i g i n between d i p l o i d A r n i c a l a t i f o l i a and t e t r a p l o i d Arnica d i v e r s i - f o l i a or Arnica m o l l i s , although attempts to make t h i s cross a r t i f i c i a l l y have been unsuccessful. Arnica l a t i f o l i a often grows with e i t h e r or both Arnica d i v e r s i f o l i a and Arn i c a m o l l i s , but there are presently b a r r i e r s to hy b r i d i z a t i o n between the sexual d i p l o i d and the apomictic t e t r a p l o i d s . Both of the l a t t e r two species u s u a l l y have very low pollen f e r t i l i t y . 2 5 1 Arnica nevadensis c l o s e l y resembles Ar n i c a c o r d i f o l i a i n morphology and i n being a s e r i e s of t e t r a p l o i d autonomous apomictic races. The former i s u s u a l l y e c o l o g i c a l l y d i s t i n c t , being found at higher elevations and i n more exposed habitats. There are populations of these two species which are d i f -f i c u l t to assign to one species or the other. Arnica nevadensis may have a r i s e n as s e l e c t i o n s of apomictic races of A r n i c a c o r d i f o l i a , without hybrid-i z a t i o n with other species. However, the longer pappus setae, tawny pappus, shorter pubescence, and attenuated or rounded l e a f bases of Arnica nevadensis suggest that some other species may have been involved i n i t s early evolution. Based on morphology, Arnica nevadensis could have o r i g i n a t e d as the r e s u l t of h y b r i d i z a t i o n between Arn i c a c o r d i f o l i a and Arnica chamissonis i n the sub-genus Chamissonis. Arnica discoidea d i f f e r s l i t t l e from Arnica c o r d i f o l i a except i n i t s lack of ray f l o r e t s . The u s u a l l y , more-branched habit, more heads, and d i s -coid condition of Arnica discoidea are i n t e r p r e t e d as derived characters. A r n i c a discoidea i s a plant of chaparral and other d r i e r habitats than that of Arnica c o r d i f o l i a . Another d i s c o i d species, Arnica spathulata, seems to be a serpentine endemic d e r i v a t i v e of Arnica discoidea. Both of these d i s -c o i d species occur as both d i p l o i d and t e t r a p l o i d populations. The subgenus Calarnica with two species, Arnica v i s c o s a and Arnica  venosa i s considered as d i s t i n c t from subgenus Austromontana as the l a t t e r i s from any of the other subgenera. I consider subgenus Calarnica to be advanced with i t s numerous small heads, s e s s i l e leaves, lack of rhizomes, and adapta-t i o n to d r i e r h a b i t a t s . The r e l a t i o n s h i p of t h i s subgenus.to other subgenera i s unclear. I t i s p o s s i b l y derived from Arnica spathulata. The pappus, pubescence, and woody caudex of Arnica venosa are c l o s e r to that of Arnica 252 spathulata than any other species. Arnica v i s c o s a i s probably a high-e l e v a t i o n d e r i v a t i v e of Arnica venosa. They are both sexual d i p l o i d s . I consider Arnica v i s c o s a , with i t s very narrow heads and white f l o r e t s , the most advanced member of the genus. 253 Figure 422. P o s s i b l e Phylogenetic Relationships of Arnica Subgenera (modified a f t e r Maguire, 1943). 254 CALARNICA AUSTROMONTANA CHAMISSONIS ARCTICA ANCESTRAL SPECIES PHYLOGENETIC RELATIONSHIPS IN ARNICA SUBGENERA 255 Figure 423. P o s s i b l e Phylogenetic Relationships of Arnica Species i n Subgenera Austromontana and Calarnica. 256 VISCOSA VENOSA CALARNICA CERNUA GRACILIS SPATHULATA LATIFOLIA DISCOIDEA NEVADENSIS CORDIFOLIA AUSTROMONTANA 257 SUMMARY A conservative taxonomic treatment of Arnica subgenus Austromontana with seven species and a new subgenus, Ca l a r n i c a , with two species has been pre-sented. This treatment i s based on f i e l d studies, cytology, breeding systems, a r t i f i c i a l h y b r i d i z a t i o n s , i n s e c t i n t e r r e l a t i o n s h i p s , and a morphological r e -evaluation. In a monograph of the genus Arnica by Maguire (1943) these two subgenera were recognized as one, Austromontana, with 13 species and three i n f r a s p e c i f i c taxa. I recognize the following species i n subgenus Austro- montana: Sexual, incompatible d i p l o i d s Arnica l a t i f o l i a (one t e t r a p l o i d population reported) Arnica cernua Arnica spathulata (one t e t r a p l o i d population reported) Pseudogamous apomictic p o l y p l o i d s Arnica c o r d i f o l i a (some autonomous apomicts and r a r e l y , d i p l o i d s ) Pseudogamous apomictic p o l y p l o i d s and sexual, incompatible d i p l o i d s Arnica discoidea Autonomous apomictic p o l y p l o i d s Arnica g r a c i l i s Arnica nevadensis I recognize the f o l l o w i n g sexual, incompatible d i p l o i d s i n subgenus Calarnica : Arnica venosa Arnica v i s c o s a 258 No i n f r a s p e c i f i c taxa are recognized. Although several species contain two or more r e l a t i v e l y d i s t i n c t elements, there i s , i n my opinion, too much overlap i n characters to warrant g i v i n g them taxonomic rec o g n i t i o n . There also does not seem to be any consistent geographical or e c o l o g i c a l i s o l a t i o n of these populations. There i s a strong c o r r e l a t i o n between high p o l l e n s t a i n a b i l i t y , d i p l o i d y , and s e x u a l i t y . Autonomous apomictic polyploids have very low p o l l e n s t a i n -a b i l i t y , or p o l l e n i s not produced at a l l . Pseudogamous apomictic p o l y p l o i d s have higher p o l l e n s t a i n a b i l i t y than the autonomous apomictics, but u s u a l l y lower than the sexual d i p l o i d s . Seed set and v i a b i l i t y i s quite low i n most of the species, and those achenes which are produced are often fed upon by ins e c t predators. On the other hand, vegetative reproduction by rhizomes has been shown to be an impor-tant f a c t o r i n short distance d i s p e r s a l of several of the species i n subgenus Austromontana. There are no completely c l e a r - c u t species i n subgenus Austromontana. Every species, as I recognize them, intergrades into one or more other species. In the sexual d i p l o i d s there are a number of l o c a l ecotypes which are very p l a s t i c , responding to the l o c a l environmental conditions. I n d i v i d u a l s have been shown to exhibit morphological d i f f e r e n c e s from year to year under vary-ing environmental conditions i n c u l t i v a t i o n . Species i n these sexual d i p l o i d s are recognized as a s e r i e s of outbreeding populations, with high p o l l e n s t a i n -a b i l i t y , s i m i l a r morphology, and s i m i l a r e c o l o g i c a l niches. The two sexual, d i p l o i d species i n subgenus Calarnica are very c l e a r - c u t species. These two rare species show very l i t t l e i n t e r p o p u l a t i o n a l v a r i a t i o n . In the apomictic species, a.number of apomictic races have received taxonomic recognition i n the past. In r e a l i t y every population may be 259 considered a species. These are sel f - b r e e d i n g e n t i t i e s with b a r r i e r s to interbreeding with other morphologically s i m i l a r populations, even though they may have recently evolved from a common ancestor. To treat each of these populations as a species would be completely i m p r a c t i c a l . Other c r i -t e r i a such as p o l l e n s t a i n a b i l i t y or chromosome numbers are of l i t t l e help i n e l u c i d a t i n g species. It seems best to recognize groups of morphologically and e c o l o g i c a l l y s i m i l a r apomictic races as species. In one species, Arnica d i s c o i d e a , there are both sexual, d i p l o i d and apomictic, t e t r a p l o i d populations. The t e t r a p l o i d s are probably recent de-r i v a t i v e s , which have not become morphologically d i s t i n c t . I can see no characters which can be used to separate the t e t r a p l o i d s from t h e i r d i p l o i d counterparts, other than t h e i r d i f f e r e n t breeding systems. They are treated as a s i n g l e species with both sexual and apomictic elements. 260 LITERATURE CITED Abrahamson, W. & K.D. McCrea. 1977. U l t r a v i o l e t l i g h t r e f l e c t i o n and absorption patterns i n populations of Rudbeckia (Compositae). Rhodora 79:269-277. Baag^e, J . 1977. Microcharacters i n the l i g u l e s of the Compositae. I_n V.H. Heywood, J.B. Harborne, & B.L. Turner (eds.). The biology and chemistry of the Compositae ( v o l . I ) . Academic Press, New York, N.Y. Barker, W.W. 1966. Apomixis i n the genus Ar n i c a (Compositae). Ph.D. d i s s e r t a t i o n , U n i v e r s i t y of Washington, S e a t t l e , Wash. Bentham, G. 1849. Plantas Hartwegianas, London. Bongard, A.G.H. 1832. Observations sur l a vegetacion de l ' I s l e de S i t c h i a . Mem. Acad. St. Petersb. VI.2:119-177. Borror, D.J., D.M. DeLong, & C.A. T r i p l e h o r n . 1976. An introduction to the study of i n s e c t s , 4th ed. H o l t , Rinehart, & Wilson. New York, N.Y. C a v i l l i e r , F.G. 1907. Etude sur l e s Doronicum a f r u i t s homomorphes. Ann. Con. Jard. Geneve 10:277. Coulter, J.M. & A. Nelson. 1909. New manual of botany of the Central Rocky Mountains. American Book Company, New York, N.Y. Cronquist, A. 1955. Arnica. JEn Hitchcock, C L . , A. Cronquist, M. Ownbey, & J.W. Thompson. Vascular plants of the P a c i f i c Northwest. V. Compositae. Uni v e r s i t y of Washington Press, S e a t t l e , Wash. . 1958. Arnica. In F e r r i s , R. Taxonomic notes on Western plants. Cont. Dudley Herb. 5:102. . 1960. Arnica. In Abrams, L. & R. F e r r i s . I l l u s t r a t e d f l o r a of the P a c i f i c States. Stanford U n i v e r s i t y Press, Stanford, C a l i f o r n i a . . 1977. The Compositae r e v i s i t e d . B r i t t o n i a 29:137-153. Ediger, R.I. & T.M. Barkley. 1978. Compositae :Senecioneae. North Am. FI. I I . 10:1-245. Fernald, M.L. 1935. C r i t i c a l p l a nts of the upper Great Lakes region of Ontario and Michigan. Rhodora 37:324-341. Gandoger, M. 1918. Sertum plantarum novarum. Pars prima. B u l l . Soc. Bot. Fr. 65:24-69. Gray, A. 1878. XXV. Contributions to the botany of North America. Proc. Am. Acad. 13:361-374. 261 . 1 8 8 4 . A r n i c a . I n S y n o p t i c a l f l o r a o f N o r t h A m e r i c a . V o l . I , P a r t 2 . C a p r i f o l i a c e a e - C o m p o s i t a e . S m i t h s o n i a n I n s t i t u t u i o n , W a s h i n g t o n , D . C . G r e e n e , E . L . 1 9 0 0 . A s e r i e s o f p a p e r s r e l a t i n g t o b o t a n y a n d b o t a n i s t s . P i t t o n i a 4 : 1 0 3 - 2 2 6 . . 1 9 0 1 . A s e r i e s o f p a p e r s r e l a t i n g t o b o t a n y a n d b o t a n i s t s . P i t t o n i a 4 : 2 2 7 - 3 2 2 . . 1 9 0 2 . O t t a w a N a t . 1 5 : 2 7 9 , 2 8 0 . . 1 9 1 0 . Some w e s t e r n s p e c i e s o f A r n i c a . O t t a w a N a t . 2 3 : 2 1 3 - 2 1 5 . G r i f f i t h s , G . C . D . 1 9 7 4 . S t u d i e s o n b o r e a l A g r o m y z i d a e ( D i p t e r a ) V I . F u r t h e r P h y t o m y z a m i n e r s o n S e n e c i o n e a e ( C o m p o s i t a e ) . Q u a e s t . E n t o m o l . 1 0 : 1 0 3 - 1 2 9 . H a l l , H . M . 1 9 1 5 . New a n d n o t e w o r t h y C a l i f o r n i a n p l a n t s . U n i v e r s i t y o f C a l i f o r n i a P u b i s . B o t . 6 : 1 6 5 - 1 7 5 . H e r d e r , F . G . 1 8 6 5 . P l a n t a e r a d d e a n a e m o n o p e t a l a e . B u l l . S o c . N a t . M o s c . 3 8 : 3 6 9 - 4 2 1 . H i l l , R . J . 1 9 7 7 . T e c h n i c a l n o t e : U l a t r v i o l e t r e f l e c t a n c e - a b s o r b a n c e p h o t o g r a p h y ; a n e a s y , i n e x p e n s i v e r e s e a r c h t o o l . B r i t t o n i a 2 9 : 3 8 2 - 3 9 0 . H o o k e r , W . J . 1 8 3 3 - 1 8 4 0 ; F l o r a b o r e a l i - a m e r i c a n a . L o n d o n . H o w e , W . H . 1 9 7 5 . T h e b u t t e r f l i e s o f N o r t h A m e r i c a . D o u b l e d a y & C o . , New Y o r k , N . Y . H o w e l l , J . T . 1 8 9 7 - 1 9 0 3 . A f l o r a o f n o r t h w e s t A m e r i c a . P o r t l a n d , O r e g o n . I l j i n , M . M . 1 9 2 6 . A r n i q u e s d e l a f l o r e ^ r u s s e . T r u d y B o t . M u z . 1 9 : 1 0 7 - 1 2 0 . J o n e s , M . E . 1 9 1 0 . M o n t a n a b o t a n y n o t e s . B u l l . . U n i v . M o n t . B i o l . 1 5 : 1 - 7 5 . K e n n e d y , H . & F . R . G a n d e r s . 1 9 7 9 . T h e i n v i s i b l e p a t t e r n s o f f l o w e r s . D a v i d s o n i a 1 0 : 1 - 6 . K u n t h , P . 1 9 0 8 . H a n d b o o k o f f l o w e r p o l l i n a t i o n . O x f o r d , E n g l a n d . L o v e , A . & B . M . K a p o o r . 1 9 6 8 . I n I O P B c h r o m o s o m e n u m b e r r e p o r t s X V . T a x o n 1 7 : 9 1 - 1 0 4 . . & D . L o v e . 1 9 6 4 . I n I O P B c h r o m o s o m e n u m b e r r e p o r t s I . T a x o n 1 9 : 9 9 - 1 1 0 . M a g u i r e , B . 1 9 4 3 . A m o n o g r a p h o f t h e g e n u s A r n i c a . B r i t t o n i a 4 : 3 8 6 - 5 1 0 . ' . 1 9 4 7 . G r e a t B a s i n p l a n t s - I X . C o m p o s i t a e . A r n i c a , H y m e n o p a p p u s , H a p l o p a p p u s , a n d L y g o . d e s m i a . Am. M i d i . N a t . 3 7 : 1 3 6 - 1 4 5 . 2 6 2 N o r d e n s t a m , B . 1 9 7 7 . S e n e c i o n e a e a n d L i a b e a e - s y s t e m a t i c r e v i e s . I n H e y w o o d , W . H . , J . B . H a r b o r n e , & B . L . T u r n e r ( e d s . ) . T h e b i o l o g y a n d c h e m i s t r y o f t h e C o m p o s i t a e ( v o l . I I ) . A c a d e m i c P r e s s , New Y o r k , N . Y . N u t t a l l , T . 1 8 4 1 . D e s c r i p t i o n s o f n e w s p e c i e s a n d g e n e r a o f p l a n t s i n t h e n a t u r a l o r d e r o f t h e C o m p o s i t a e . T r a n s . Am . P h i l . S o c . I I . 7 : 2 8 3 - 4 5 3 . O r n d u f f , R . , P . H . R a v e n , D . W . K y h o s , & A . R . K r u c k e b e r g . 1 9 6 3 . C h r o m o s o m e n u m b e r s i n C o m p o s i t a e . I I I . S e n e c i o n e a e . A m . J . B o t . 5 0 : 1 3 1 - 1 3 9 . , T . M o s q u i n , D . W . K y h o s , & P . H . R a v e n . 1 9 6 7 . C h r o m o s o m e n u m b e r s i n C o m p o s i t a e . V I . S e n e c i o n e a e I I . Am. J . B o t . 5 4 : 2 0 5 - 2 1 3 P i p e r , C V . 1 9 2 0 . Some n e w p l a n t s f r o m t h e P a c i f i c N o r t h w e s t . P r o c . B i o l . S o c . W a s h . 3 3 : 1 0 3 - 1 0 6 . P o w e l l , A . M . , D .W . K y h o s , & P . H . R a v e n . 1 9 7 4 . C h r o m o s o m e n u m b e r s i n C o m p o s i t a e X . A m . J . B o t . 6 1 : 9 0 9 - 9 1 3 . R o b i n s , D . J . 1 9 7 7 . S e n e c i o n e a e - c h e m i c a l r e v i e w . I n H e y w o o d , W . H . , J . B . H a r b o r n e , & B . L . T u r n e r ( e d s . ) . T h e b i o l o g y a n d c h e m i s t r y o f t h e C o m p o s i t a e ( v o l . I I ) . A c a d e m i c P r e s s , New Y o r k , N . Y . R y d b e r g , P . A . 1 8 9 7 . R a r i t i e s f r o m M o n t a n a I I I . B u l l . T o r r e y B o t . C l u b 2 4 : 2 9 2 - 2 9 9 . 1 9 0 0 . S t u d i e s o n t h e R o c k y M o u n t a i n f l o r a - I . B u l l . T o r r e y B o t . C l u b 2 7 : 1 6 9 - 1 8 9 New Y o r k , N . Y . 1 9 1 7 . F l o r a o f t h e R o c k y M o u n t a i n s a n d a d j a c e n t p l a i n s . 1 9 2 7 . A r n i c a . ( C a r d u a l e s ) C a r d u a c e a e . L i a b e a e , N e u r o l a e n a e . S e n e c i o n e a e ( p a r s ) I n N . Am . F l . 3 4 : 2 8 9 - 3 6 0 S t . J o h n , H . 1937 . . F l o r a o f S o u t h e a s t e r n W a s h i n g t o n a n d o f a d j a c e n t I d a h o . P u l l m a n , W a s h . S c h a a c k , C . G . , J . T . W i t h e r s p o o n , . & T . J . W a t s o n , J r . 1 9 7 4 . I n I O P B c h r o m o s o m e n u m b e r r e p o r t s X L V . T a x o n 2 3 : 6 1 9 - 6 2 3 . S c o g i n , R . , D . A . Y o u n g , & C E . J o n e s . 1 9 7 7 . . A n t h o c h l o r p i g m e n t s a n d p o l l i n a t i o n b i o l o g y . I I . T h e u l t r a v i o l e t f l o r a l p a t t e r n s o f C o r e o p s i s  g l g a n t e a ( A s t e r a c e a e ) . B u l l . T o r r e y B o t . C l u b 1 0 4 : 1 5 5 - 1 5 9 . S i e g l e r , D . S . , D . H . W i l k e n , & J . J . J a k u p c a k . 1 9 7 4 . C h e m i c a l d a t a r e l a t i n g t o t h e t r i b a l a f f i n i t i e s o f H u l s e a a n d A r n i c a . B i o c h e m . - S y s t . E c o l . 2 : 2 1 - 2 4 . S t r a l e y , G . B . 1 9 7 9 . I n I O P B c h r o m o s o m e n u m b e r r e p o r t s L X I I I . T a x o n 2 8 : 2 7 8 . 263 Strother, J.L. 1972. Chromosome studies i n Western North American Compositae. Am. J. Bot. 59:242-247. Taylor, R.L. & R.P. Brockman. 1966. Chromosome numbers i n some Western Canadian plants. Can. J. Bot. 44:1093-1103. . & B. McBryde. 1977. Vascular plants of B r i t i s h Columbia. A d e s c r i p t i v e resource inventory. U n i v e r s i t y of B r i t i s h Columbia Press, Vancouver, Canada. . & G.A. Mulligan. 1968. F l o r a of the Queen Charlotte Islands. Part 2. C y t o l o g i c a l aspects of the vascular paints. Can. Dept. of Ag. , Research Branch, Monograph #4. part 2. Ottawa, Canada. . & S. Taylor. 1977. Chromosome numbers of vascular plants i n B r i t i s h Columbia. Syesis 10:125-138. Turner, B.L. & A.M. Powell. - 1977. Helenieae - systematic review. In Heywood, V.H., J.B. Harborne, & B.L. Turner (eds.). The biology and chemistry of the Compositae ( v o l . I I ) . Academic Press, New York, N.Y. Williams, L.O. 1935. Notes on Rocky Mountain plants I I . L e a f l . West. Bot. 1:171,172. Zukova, P.G. 1967. Karyology of some plants c u l t i v a t e d i n the A r c t i c - A l p i n e Botanical Garden. In A v r o r i n , N.A. (ed.): Plantarum i n Zonam Polarem Transplantatio I I . Leningrad. 139-149. 264 APPENDIX I Taxa recognized by Maguire (1943) Subgenus Aus tromontana 1. Arnica l a t i f o l i a Bongard 2. Arnica g r a c i l i s Rydb. 3. Arnica cernua Howell 4. Arnica tomentella Greene •5. Arnica nevadensis A. Gray 6. Arnica paniculata A. Nels. 7a. Arnica c o r d i f o l i a Hook, subsp. genuina Maguire 7b. Arnica c o r d i f o l i a Hook, subsp. whitneyi (Fernald) Maguire 8. Arnica g r a y i H e l l e r 9a. Arnica p a r v i f l o r a A. Gray subsp. genuina Maguire 9b. Arnica p a r v i f l o r a A. Gray subsp. a l a t a (Rydb.) Maguire 10. Arnica discoidea Bentham 11a. Arnica spathulata Greene subsp. genuina Maguire l i b . A rnica spathulata Greene subsp. eastwoodiae (Rydb.) Maguire 12. Arnica venosa H.M. H a l l -13. Arnica v i s c o s a A. Gray 265 APPENDIX II Taxa recognized by Ediger & Barkley (1978) Subgenus Austromontana l a . A r n i c a l a t i f o l i a Bongard var. l a t i f o l i a l b . A r n i c a l a t i f o l i a Bongard var. g r a c i l i s (Rydb.) Cronquist 2. Arn i c a cernua Howell 3. Arni c a tomentella Greene 4. Arni c a nevadensis A. Gray 5a. Ar n i c a c o r d i f o l i a Hooker var. c o r d i f o l i a 5b. Ar n i c a c o r d i f o l i a Hooker var. pumila (Rydb.) Maguire 6a. Ar n i c a discoidea Bentham var. a l a t a (Rydb.) Cronquist 6b. Arn i c a discoidea Bentham var. eradiata (A. Gray) Cronquist 6c. A r n i c a d i s c o i d e a Bentham var. discoidea 7a. A r n i c a spathulata Greene var. spathulata 7b. Ar n i c a spathulata Greene var. eastwoodiae (Rydb.) Ediger & 8. A r n i c a venosa H.M. H a l l 9. Arnica v i s c o s a A. Gray 

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            data-media="{[{embed.selectedMedia}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
https://iiif.library.ubc.ca/presentation/dsp.831.1-0094954/manifest

Comment

Related Items