Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Studies in the genus saxifraga in the Pacific Northwest : Part I: taxonomy of saxifraga occidentalis… Krause, David Larkin 1972

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

Item Metadata

Download

Media
831-UBC_1973_A6_7 K73.pdf [ 2.8MB ]
Metadata
JSON: 831-1.0101302.json
JSON-LD: 831-1.0101302-ld.json
RDF/XML (Pretty): 831-1.0101302-rdf.xml
RDF/JSON: 831-1.0101302-rdf.json
Turtle: 831-1.0101302-turtle.txt
N-Triples: 831-1.0101302-rdf-ntriples.txt
Original Record: 831-1.0101302-source.json
Full Text
831-1.0101302-fulltext.txt
Citation
831-1.0101302.ris

Full Text

c STUDIES IN THE GENUS SAXIFRAGA IN THE PACIFIC NORTHWEST PART I: TAXONOMY OF SAXIFRAGA OCCIDENTALS AND S. MARSHALLII. PART I I : NOTES ON SAXIFRAGA OCCIDENTALS AND CLOSELY RELATED SPECIES. by. DAVID L. KRAUSE B.Sc, University of B r i t i s h Columbia, 1964 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n the Department of Botany We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA December, 1972 < In presenting t h i s t h e s i s in p a r t i a l f u l f i l m e n t of the requirements f o r an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I f u r t h e r agree t h a t permission for extensive copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s r e p r e s e n t a t i v e s . I t i s understood that copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission. Department of The U n i v e r s i t y of B r i t i s h Columbia Vancouver 8, Canada Date £7 i i . ABSTRACT PART I. Taxonomy of Saxifraga o c c i d e n t a l i s and S_. marshal I i i . The taxonomy of Saxifraga m a r s h a l l i i and S_. o c c i d e n t a l i s i s reviewed. On morphological, c y t o l o g i c a l and d i s t r i b u t i o n a l evidence, the new combination S_. m a r s h a l l i i ssp. idahoensis i s proposed for the taxon o r i g i n a l l y referred to as S_. idahoensis. Keys to the taxa, synonomy, d i s t r i b u t i o n a l records, maps, and chromosome numbers are presented. PART I I . Notes on Saxifraga o c c i d e n t a l i s and c l o s e l y r e l a t e d species. A species key, d i s t r i b u t i o n maps and some chromosome numbers are presented f o r Saxifraga o c c i d e n t a l i s S. Wats., S_. rhomboidea Greene, S_. r u f i d u l a (Small) James Macoun, S^. r e f l e x a Hook., and the com-bined species S_. n i v a l i s L. - S_. tenuis (Wahl.) H. Sm. I t i s poss i b l e to separate these f i v e taxa u t i l i z i n g c e r t a i n q u a l i t a t i v e characters. Saxifraga n i v a l i s and S_. tenuis are two c l o s e l y r e l a t e d taxa whose re l a t i o n s h i p s are unknown. i i i . TABLE OF CONTENTS PART I. Taxonomy of Saxifraga o c c i d e n t a l i s and S_. m a r s h a l l i i . Abstract . i i . Introduction 1. Materials and Methods 3. Taxonomy 6. Discussion 20. L i t e r a t u r e Cited 24. Table 1 26. Figures 1-5 27. Figure 6 .28. Figure 7 29. PART I I . Notes on Saxifraga o c c i d e n t a l i s and c l o s e l y r e l a t e d species. Abstract i i . Introduction 30. Taxonomy 32. L i t e r a t u r e Cited 41. Table 1 43. Figure 1 44. Figure 2 45. Figure 3 .46. 1. PART I INTRODUCTION In the P a c i f i c Northwest the r e l a t i o n s h i p s i n the genus Saxifraga are often doubtful and the names d i f f i c u l t to apply. This statement i s c e r t a i n l y true for Section Boraphila subsection N i v a l i - v i r g i n i e n s e s established by Engler and Irmscher (1916) i n t h e i r comprehensive t r e a t -ment of Saxifragaceae. The marked disagreement over the i n t r a - and i n t e r - r e l a t i o n s h i p s of t h i s subsection are due p a r t l y to the morphological s i m i l a r i t i e s between e n t i t i e s and p a r t l y to the many s p e c i f i c and sub-s p e c i f i c descriptions which have resulted i n considerable synonomy. One of the problems within the group i s the r e l a t i o n s h i p of the taxa commonly re f e r r e d to as S_. o c c i d e n t a l i s S. Wats., Sj_ m a r s h a l l i i Greene, and S_. idahoensis Piper. In 1888 Greene published the name S^. m a r s h a l l i i f or a saxifrage found i n the Hoopa V a l l e y , Humboldt County, C a l i f o r n i a . Also i n 1888, Sereno Watson described a new species, jS. o c c i d e n t a l i s , "... from the Rocky Mountains of B r i t i s h America (Drummond) to B r i t i s h Columbia and Vancouver Island ( L y a l l , Macoun), Oregon (Cusick, Henderson, Howell), and the northern S i e r r a Nevada (Chico, Mrs. J . Bidwell, Gray)." No type specimen was designated and i t i s generally accepted that there i s more than one species w i t h i n the area c i t e d . The name S_. o c c i d e n t a l i s i s frequently used f o r the most common of the B r i t i s h Columbia s a x i -frages from the subsection N i v a l i - v i r g i n i e n s e s . In 1900 C. V. Piper described a new species from western Idaho, S^. idahoensis. He believed 2. i t to be re l a t e d to S_. o c c i d e n t a l i s and S_. r e f l e x a , but d i f f e r i n g by i t s "... nearly glabrous leaves, much branched inflorescence and small flowers ..." Engler and Irmscher (1916) reduced S_. idahoensis to a va r i e t y of m a r s h a l l i i . Abrams (1944) maintained S_. idahoensis as a species c l o s e l y r e l a t e d to S_. m a r s h a l l i i , but separated from the l a t t e r on the basis of smaller flowers and absence of yellow spots on the pe t a l s . In the l a t e s t treatment of the group, Hitchcock et a l . (1961) made S_. idahoensis a v a r i e t y of S_. o c c i d e n t a l i s because of i t s t r a n s i t i o n to other races of S_. o c c i d e n t a l i s , and because i t i s geographically i s o l a t e d from S_. m a r s h a l l i i . Small (1905) and Johnson (1923) published many new species belonging to subsection N i v a l i - v i r g i n i e n s e s , most of which have since been reduced to synonomy, mainly of S_. o c c i d e n t a l i s . The present paper i s an attempt to bring c y t o l o g i c a l , morphological, and d i s t r i b u t i o n a l evidence together to determine the relationships of S. o c c i d e n t a l i s , S. m a r s h a l l i i , and S. idahoensis. 3. MATERIAL AND METHODS F i e l d work has been done mainly across southern B r i t i s h Columbia and northwestern United States. Both l i v e material and buds were c o l l e c t e d whenever possible. The l i v e material was grown i n pots at The University of B r i t i s h Columbia. Live c o l l e c t i o n s and herbarium specimens were studied quantita-t i v e l y and q u a l i t a t i v e l y f o r the c h a r a c t e r i s t i c s used i n the taxa descriptions. The data from the quantitative c h a r a c t e r i s t i c s were processed f o r mean, midpoint, range, variance, standard deviation, standard e r r o r , and confidence l i m i t s at the 5% l e v e l . Scheffe's test for differences between groups using unequal sample si z e s was attempted between populations and between taxa. The quantitative measurements were taken of the l e a f , scape, and inflorescence. Of these measurements only the means and standard devia-tions were found to be u s e f u l . The means and standard deviations were used to de l i m i t the v a r i a b i l i t y of a s i n g l e morphological character; however, each character has such a wide range i n each taxon and over-laps so much the range i n other taxa that they are not useful as diagnostic characters. Graphic representation using computerized s c a t t e r diagrams confirmed t h i s high degree of overlap for the quantitative characters. The evaluation of the q u a l i t a t i v e characters was found to be most use f u l f o r the separation of the various taxa. A combination of q u a l i t a -t i v e characters i s usually needed as no s i n g l e character could be used to 4. separate a l l taxa. For example the presence of yellow spots on the petals i s diagnostic f o r S_. m a r s h a l l i i , but the absence of spots on herbarium specimens i s not diagnostic as the spots commonly fade upon drying. The buds c o l l e c t e d i n the f i e l d and taken from material grown i n pots were f i x e d i n 95% ethanol and g l a c i a l a c e t i c a c i d (3:1). Buds were stained using Snow's carmine technique (1963) and anthers dissected out and squashed i n haematoxylin (Whittman 1965). Voucher specimens f o r chromosome counts are kept i n the herbarium of The Univer-s i t y of B r i t i s h Columbia. C o l l e c t i o n data and chromosome counts are given.in Table 1. Herbarium material was borrowed from the herbaria l i s t e d below. ALTA Herbarium, Department of Botany, University of Alberta, Edmonton. UAC Herbarium, Department of Biology, University of Calgary, Alberta. CAN National Museum of Canada, Ottawa. COLO Herbarium, University of Colorado, Boulder, Colorado. DAO Plant Research I n s t i t u t e , Department of Agr i c u l t u r e , Ottawa. GH Gray Herbarium of Harvard University, Cambridge, Mass. ID Department of Botany, University of Idaho, Moscow, Idaho. JEPS Jepson Herbarium, Department of Botany, University of C a l i f o r n i a , Berkeley. MONTU University of Montana, Missoula, Montana. ND University of Notre Dame, Notre Dame, Indiana. NY The New York Botanical Garden, New York Cit y , New York. Herbarium of the University of Oregon, Eugene, Oregon. Herbarium, Department of Botany, Oregon State U n i v e r s i t y , C o r v a l l i s . Academy of Natural Sciences, Phil a d e l p h i a , Pennsylvania. Rocky Mountain Herbarium, University of Wyoming, Laramie, Wyoming. Herbarium of the University of C a l i f o r n i a , Department of Botany, University of C a l i f o r n i a , Berkeley. U.S. National Museum (Department of Botany), Smithsonian I n s t i -t u t i o n , Washington, D.C. Department of Plant Pathology, State College of Washington, Pullman, Washington. Herbarium, Department of Botany, University of Washington, Seat t l e . 6. TAXONOMY KEY TO THE SPECIES A. Inflorescence open, branches and peduncles usually d i v a r i c a t e , pedicels 2 to 4 times as long as flowers; flowers seldom grouped at ends of branches; filaments clavate, often showy a f t e r petals f a l l ; petals with 2 yellow spots at base (often inconspicuous on dried m a t e r i a l ) , deciduous; sepals reflexed; ovary le s s than 1/5 i n f e r i o r . B. Ovary almost completely superior; leaves oblong-ovate to e l l i p t i c -oblong; p e t i o l e s (0.5) 1 to 3 times as long as the blade length. Western Oregon and N.W. C a l i f o r n i a 1. S_. m a r s h a l l i i ssp. m a r s h a l l i i BB. Ovary from almost completely superior to 1/5 i n f e r i o r ; leaves ovate to rhombic; p e t i o l e s 0.5 to 1.5 (2) times as long as blade length. Chelan and K i t t i t a s Cos. of Washington; North Central Idaho and adjacent N.E. Oregon and western Montana 2. S_. m a r s h a l l i i ssp. idahoensis AA. Inflorescence open to congested, branches and peduncles spreading to ascending, pedicels equal to, to twice as long as, the flowers; flowers tending to be grouped at ends of branches; filaments subulate, l i n e a r or sometimes s l i g h t l y clavate, inconspicuous i f petals f a l l ; petals without basal yellow spots, often p e r s i s t e n t ; sepals spreading to p a r t i a l l y reflexed i n older flowers; ovary (1/10) 1/5 to 1/3 i n f e r i o r . B r i t i s h Columbia; Rocky Mountains of Alberta; Cypress H i l l s 7. of Saskatchewan; Mt. Rainier area i n Washington; northern Washington and Idaho; c e n t r a l Idaho; Rocky Mountains i n Montana, northeastern Oregon, and northern Wyoming; and occasionally i n the Rocky Mountains as f a r south as New Mexico... 3. S. o c c i d e n t a l i s 1. Saxifraga m a r s h a l l i i Greene ssp. m a r s h a l l i i , P i t t o n i a , 1: 159. 1888. TYPE C.C. Marshall s.n., Hoopa Val l e y , Humboldt Co., C a l i f o r n i a , Apr. 1887. ND! IS. p e t i o l a t a A.M. Johnson, Minn. Stud. B i o l . S c i . 4: 29. 1923. TYPE T. Howell s.n., Oregon, on wet banks, Woodville, May, 1889. WTU! Isotypes: NY I UC! (2) CAN! Micranthes m a r s h a l l i i (Greene) Small, N. Am. F l o r a 22(2): 140. 1905. S_. m a r s h a l l i i Greene var. m a r s h a l l i i f. dentata Engl. & Irmsch. Pflanzenr. IV, 117, 1: 36. 1916. S^. m a r s h a l l i i Greene var. m a r s h a l l i i f. crenata Engl. & Irmsch. Pflanzenr. IV, 117, 1: 36. 1916. S_. h a l l i i A.M. Johnson, Stud. PI. S c i . 4: 24. 1923. TYPE: E. H a l l 151, Oregon ann. 1871. MGB. S_. l a e v i c a u l i s A.M. Johnson, Minn. Stud. B i o l . S c i . 4: 26. 1923. TYPE: Piper 5061, Mt. Grayback, Oregon, June 15, 1904. _S. m a r s h a l l i i Greene var. d i v a r i c a t a Peck, L e a f l . West. Bot. 5: 59. 1947. TYPE: Peck 18158, Snake River Canyon, near mouth of Bat t l e Creek, Wallowa Co., Oregon, March 28, 1934. Stem frequently s o l i t a r y (2-3) from (0.5) 1 to 3 (4) dm long, covered with reddish purple glandular h a i r s with m u l t i c e l l u l a r heads; leaves oblong-ovate to e l l i p t i c - o b l o n g , upper surface glabrous, lower surface 8. often rusty-tomentose, e s p e c i a l l y smaller leaves, margins crenate with wide, blunt teeth at r i g h t angles to the margin, blade 2 to 6 cm long, (1) 1.5 to 3.5 cm broad, truncate to subtruncate at base, tapering i n t o the (0.5) 1 to 3 times longer p e t i o l e ; inflorescence an open d i f f u s e panicle, branches slender, d i v a r i c a t e , subtended by small, l i n e a r bracts which are occasionally l e a f l i k e and toothed, pedicels 3 to 4 times longer than flowers; sepals strongly reflexed, ovate-lanceolate to oblong-lanceolate, 2 to 3 times as long as broad; petals deciduous, oblong to ovate, 2.5 to 3.5 mm long, narrowed into a short d i s t i n c t claw, 2 yellow spots on base of p e t a l , mostly faded i n herbarium material; filaments p e r s i s t e n t , spreading, equal to or shorter than the pe t a l s , strongly clavate to p e t a l o i d , often showy a f t e r petals f a l l ; the 2 (3) carpels d i s t i n c t to base, f l u t e d nectary at base appearing as wavy ribbon as capsules mature, immature carpels long and t h i n , mature carpels erect, ti p s divergent; ovary almost completely superior, n_ = 10 (Table 1; F i g . 1). Moist banks beside streams i n north western C a l i f o r n i a and western Oregon (Fig. 6). Representative Specimens OREGON: BENTON Co.: Mary's Peak, C o r v a l l i s , Epling s.n. (ORE); COOS Co.: e. fork of Millicomes R., 4 mi from Allegany, Cronquist 6859 (WTU, WS, ID, OSC, CAN, UC, DAO, UBC, MONTU); Sitkum, Krause and Beamish  30-70 (UBC); Dora, Krause and Beamish 10-70 (UBC); Bear Ck., middle fork of Coquille R., Krause and Beamish 35-70 (UBC); CURRY Co.: Two Mile Creek, Ih mi s. of I l l a h e , Baker 5264 (WTU, WS, OSC, UC); Frye Place, 3 mi s. of Agness, Baker 3513 (OSC, UC); DOUGLAS Co.: Gardiner, Overlander s.n. (OSC); Mapleton, Krause and Beamish 36-70 (UBC); between North Bend and Roseburg, Mason 5290 (UC, OSC); JACKSON Co.: on wet banks, Woodville, Howell s.n. (NY, UC, CAN); Weimer, Hammond 125b (NY); JOSEPHINE Co.; 10 mi w. of Selma, Overlander s.n. (OSC); LANE Co.: Willamette Pass between Dexter and Oakridge, Packard and Gilkey s.n. (UBC, OSC); w. of Triangle Lake, Gilkey s.n. (WS, WTU); Scottsburg, Krause and Beamish  15-71 (UBC); LINCOLN Co.: Waldport-Yachats, Smith s.n. (OSC); wet c l i f f s s. side Oswego L., Nelson 2550 (WS); POLK Co.: M i l l C., 4 m sw. of B u e l l Peck 16213 (UC). CALIFORNIA: DEL NORTE Co.: Gasquet, Pack and Tracy 12351 (DAO, UC); HUMBOLDT Co.: Willow Creek, Tracy 6644 (UC, WTU); T r i n i t y Summit, 2 mi se. of Devil's Hole, Tracy 15511 (UC); TRINITY Co.: Salyer, Tracy  8011 (UC, JEPS) ; MENDOCINO Co.: Reel Mt. Creek b l u f f , on s. fork of E e l R., Tracy 16530 (UC) . 2. Saxifraga m a r s h a l l i i Greene ssp. idahoensis (Piper) Krause and Beamish, comb. nov. S_. idahoensis Piper, B u l l . Torrey Bot. Club, 27: 394. 1900. TYPE: Henderson 4588, Kendrick, Latah Co., Idaho, Apr. 26, 1896. GH! ISOTYPE: NYI' Photograph: WSPl S_. m a r s h a l l i i Piper var. idahoensis (Piper) Engl, and Irmsch., Pflanzenr. IV, 117, 1: 37. 1916. S_. microcarpa A.M. Johnson, Minn. Stud. B i o l . S c i . 4: 28. 1923. TYPE: M.F. Ehod 98a, Missoula v i c i n i t y . Mont. MONTU! S_. o c c i d e n t a l i s S. Wats. var. idahoensis (Piper) C.L. Hitch., Vascular Plants of the P a c i f i c Northwest., Univ. Wash. Press, Hitchcock et a l . IV. 49. 1961. 10. Caudex short and stout, rhizome short, oblique with 1 (2) flowering stem, 1.2-2 (2.5) dm t a l l with dense to sparse p u r p l i s h glandular h a i r s ; l e a f blades from ovate to e l l i p t i c to rhombic, glabrous on upper surface and sparsely rusty tomentose on lower surface, margin i r r e g u l a r l y dentate, blade from 2.3 to 3 cm long, 1.8 to 2.7 cm wide with an attenuate (obtuse) base; p e t i o l e from s l i g h t l y shorter than to twice as long as blade; inflorescence generally an open, d i f f u s e panicle, branches slender, d i v a r i c a t e , pedicels from 2 to 3 times as long as flowers, flowers seldom i n compact clusters at ends of branches; bracts small, l i n e a r , glandular-p i l o s e , occasionally leafy and toothed; sepals ovate-lanceolate, 2.5-3 times as long as broad, s l i g h t l y to strongly reflexed at anthesis; petals deciduous, oblong to ovate, 2-3.5 mm long, narrowed i n t o a claw, 2 yellow spots present at base (r a r e l y seen i n pressed m a t e r i a l ) ; filaments p e r s i s -tent, clavate to strongly clavate, about equalling the p e t a l s ; ovary from almost completely superior to 1/5 i n f e r i o r ; 2 (3) carpels, elongate t h i n ; nectary present at anthesis but r a r e l y v i s i b l e at maturity; n = 10 (Table 1; F i g . 2). Moist rocky slopes, K i t t i t a s and Chelan Cos., Washington; western Idaho, e s p e c i a l l y i n low l y i n g areas of the St. Joe and Clearwater River drainage systems; and adjacent northeast Oregon (Fig. 6). Type - A l l the specimens Piper examined i n e s t a b l i s h i n g h i s species were from western Idaho. He stated that the filaments were "subulate, p e r s i s t e n t , at length exceeding the p e t a l s . " In 1902 J.K. Small wrote asking him i f the d e s c r i p t i o n of "subulate filaments" was an error. He r e p l i e d " I t i s evident that the d e s c r i p t i o n of the filaments as 'subulate' was a blunder." (Letters on NY herbarium sheet.) In h i s type de s c r i p t i o n he also stated that S_. idahoensis was relat e d to r e f l e x a and o c c i - dentalis . As most, early S_. m a r s h a l l i i specimens from the United States with reflexed sepals were c a l l e d S_. ref lexa, this f i t s very w e l l with the hypothesis that S_. idahoensis i s c l o s e l y r e l a t e d to S_. m a r s h a l l i i but has some c h a r a c t e r i s t i c s of S^. o c c i d e n t a l i s . JS. microcarpa appears to be a smaller plant of ;S. m a r s h a l l i i ssp. idahoensis. Johnson states i t i s only found from the type l o c a l i t y , Missoula, Montana. He examined only two specimens. Representative Specimens WASHINGTON: CHELAN Co.: Moist draw above Camas Land Meadow, s. side, T23N, R18E S21, C L . Hitchcock and C.V. Muhlick 21542 (WTU); rocky moist slopes on Stuart fiidge above I n g a l l s , J.W. Thompson 8968 (NY, WTU); steep rocky h i l l s i d e , n. exposure above Swakane Cr., about 5 mi e. of divide to Nahukum Cr. , C L . Hitchcock 23440 (UC, WSP, WTU); moist rocky slopes on Lookout Mt., near Leavenworth, J.W. Thompson 6465 (OSC); above Peshastin Cr., ca. 6 mi n. of Blewett Pass, C L . Hitchcock and J.S. Martin  4738 (NY, WTU); wet granite slopes near Leavenworth, J.W. Thompson 6411 (OSC, WTU); moist slopes of high ridge above Thronson Cr., J.W. Thompson  8294 (NY, UC); 10 mi n. of Swaux Pass on Hy. 97, ca. 10 mi sw. of Cashmere, J. Maze et a l . 340 (CAN); KITTITAS Co.: moist rocks along Boulder Cr., J.W. Thompson 10422 (WTU); 1 mi below Camp Cr., upper Cle Elum River V a l l e y , A.R. Kruckeburg 4984 (UC); Sec. 4, T. 20N, R. 21E, on Ellensburg-Wenatchee Rd., ca. 5 mi n. of Colochum Pass, J . Maze et a l . 398 (CAN); on moist c l i f f s ca. 2 mi nw. of Mission Peak, Wenatchee Mts., C L . Hitchcock 20346 (WTU). 12. IDAHO: IDAHO Co.: along the Selway R. near Fenn Ranger Station, V. A l l e n s.n. (ID); 2 mi s. of Ahsahka, on c l i f f s along Clearwater R., J.H. Chr i s t and W.W. Ward 12107 (NY); Lewiston, moist h i l l s i d e , J.H. Small  6265 (NY, WTU); Hapwai, D. McDougal 93 (NY); moist s i t u a t i o n s i n granite gravel on south c l i f f at summit of John Beale Rd., Lolo Nat. Forest. M.J. Reed and F.A. Barkley 84 (MONTU); e. slope of mountain, 4000 f t s. of mouth of Sheep Cr., Snake River Canyon, S. Beinke 131 (WTU); rocky b l u f f s i n coniferous shade on Meadow Cr., above Selway F a l l s , L. Constance and  R.C. R o l l i n s 1638 (WS, WTU, ALTA); n. slope, s. of Selway R., Selway F a l l s , M. Ownbey and R.P. Ownbey 2040 (WTU, UC, WS, NY); 4 mi below Lowell, Middle fork of Clearwater R. at Three Devils Cr., L. Constance et a l . 1083 (WS, UC); moist, south-facing slope, 1 mi up Granite Cr., Snake R. Canyon, J . Packard  372 (UC); on rock outcropping near Selway R., 2 mi up from Selway F a l l s , J.L. McMullen 1487 (UC, ID); LATAH Co.: moist n. slope along Potlatch Cr. 4 mi s. of J u l i a e t t a , M. Ownbey and R.P. Ownbey 2032 (COLO, DAO, WTU, UC, WS, NY, OSC, ALTA); NEZ PERCE Co.: steep rocky h i l l s i d e , along the Clear-water R., near Myrtle, W.H. Baker 6606 (JEPS, WTU); moist rocky slope, near Culdesac, F.A. Warren s.n. (WS); moist rocky c l i f f , canyon of the Clearwater R., 4 mi sw. of Myrtle, W.H. Baker 6606 (UC); talus slopes, along Clearwater R., 1 mi e. of Spaulding, W.H. Baker 13909 (ID); Onfino^ on rd. to Nez Perce, J.H. Christ 7537 (NY); SHOSHONE Co.: moist s l a t e c l i f f w. of Twin L., C.L. Hitchcock and C.V. Muhlick 21745 (WTU, WS, NY); on c l i f f s along St. Joe R., 7 mi s. of Culdesac, J.H. Christ 10958 (ID, NY) OREGON: WALLOWA Co.: rocky slope, Snake R. Canyon near mouth of Ba t t l e Cr., M.E. Peck 18161 (NY); on basalt ledge at L i t t l e Creek, i n the Snake R. Canyon n. of the mouth of the Salmon R., W.A. Weber 2103 (COLO); moist rocky n. h i l l s i d e , mouth of Imnaha R., G.N. Jones 6295 (WS, UC). MONTANA: RAVALLI Co.: i n mossy cracks on bedrock above Glen L., B i t t e r Root Mts., K.H. Lackschewitz and R. Fageraas 1264 (MONTU); o f f the Lost Horse Cr. Rd., B i t t e r Root Mts. K.H. Lackschewitz and R. Fageraas  1215 (MONTU); MISSOULA Co.: on bedrock above Carlton L., Lolo Peak, B i t t e r Root Mts., K.H. Lackschewitz and R. Fageraas 477 (MONTU). 3. Saxifraga o c c i d e n t a l i s S. Wats., Proc. Am. Acad. 23: 264. 1888. LECTOTYPE: J. Macoun s.n., Yale Mt., B.C., May 17, 1875. Specimen A, CAN! S_. saximontana E. Nelson, Erythea, 7: 168. 1899. TYPE: A. and E. Nelson 5917, Yancey's Yellowstone Nat. Park, July 17, 1899. NY! ISOTYPES: WSP! RM! Micranthes o c c i d e n t a l i s (S. Wats.) Small, N. Am. F l o r a , 22(2): 144. 1905. M. a l l e n i i Small, N. Am. F l o r a , 22(2): 144. 1905. TYPE: O.D. A l l e n 242, Goat Mts., Washington, June 27, 1896. GH! ISOTYPES: NY! US! WS! WTU! M. l a t a Small, N. Am. F l o r a 22(2): 144. 1905. TYPE: John Macoun s_.n. , Lytton, B.C., Apr. 16, 1889. NY! ISOTYPES: GH! CAN! Photograph: DAO! M. saximontana (E. Nels.) Small, N. Am. F l o r a , 22(2): 145. 1905. S_. a l l e n i i (Small) Fedde, Just's Bot. Jahresber. 31(1): 613. 1906. S_. l a t a (Small) Fedde, Just's Bot. Jahresber. 31(1): 613. 1906. S_. o c c i d e n t a l i s S. Wats. var. wallowensis Peck, L e a f l . West. Bot. 4(4)60. 1947. TYPE: Peck 18542, moist slope above Ice Lake, Wallowa Mts., Wallowa Co., Oregon, July 4, 1934. 14. S_. o c c i d e n t a l i s S. Wats. var. a l l e n i i C.L. Hitchc. , Vascular Plants of the P a c i f i c Northwest. Univ. Wash. Press, H i t c h c . et a l . , I I I : 49. 1961. S_. o c c i d e n t a l i s S. Wats. var. l a t i p e t i o l a t a H i t c h c , Vascular Plants of the P a c i f i c Northwest. Univ. Wash. Press, Hitchc. et a l . I l l : £49. 1961. TYPE: M.W. Gorman 3561, Saddle Mt., Clatsop Co., Oregon, June 20, 1915. WTU! Stems 1 (2-3), from 0.8 to 2 (2.5) dm t a l l with dense to sparse pu r p l i s h glandular h a i r s ; leaf blades v a r i a b l e i n s i z e and shape, from oval-ovate to rhombic, upper surfaces glabrous, lower surfaces sometimes rusty-tomentose, margins serrate to crenate, blade from 1.5 to 3 cm long, 1.2 to 2.5 cm wide tapering into a p e t i o l e equal to, to twice as long as the blade; inflorescence an open p a n i c l e , from considerably c o n s t r i c t e d with short erect branches to open with spreading branches, glomerate to rounded or pyramidal; pedicels e q u a l l i n g twice the length of flowers, flowers grouped at ends of branches; bracts small and l i n e a r , often toothed and (or) rufose-tomentose; sepals ovate to oblong-oblanceolate, spreading to reflexed; petals white, immaculate, deciduous or p e r s i s t e n t , ovate to oblong, 1.5 to 3.5 cm long with a d i s t i n c t claw or clawless, filaments subulate, l i n e a r , or s l i g h t l y clavate, p e r s i s t e n t , equalling the petals; carpels 2 (3) narrowing gradually into long s t y l e s , s h arply divergent f o r 3/4 t h e i r length at maturity; ovary from 1/10 to 1/3 i n f e r i o r ; nectary v i s i b l e at anthesis, but drying a f t e r anthesis and d i f f i c u l t to see; n = 10, 19, 28, 29 (Table 1; Figs. 3 and 4). Moist banks and meadows to rocky alpine slopes. B r i t i s h Columbia; Rocky 15. Mountains i n Alberta; Cypress H i l l s i n Saskatchewan; area around Mt. Rainier, Washington; northern Washington and Idaho; c e n t r a l Idaho, Rocky Mountains i n Montana and northwest Wyoming and northeastern Oregon and occasionally the Rocky Mountains as f a r south as New Mexico (Fig. 7). Type - No type specimen has been designated f o r J3. o c c i d e n t a l i s . Watson (1888) c i t e d s e v e r a l specimens f o r the range of S_. oc c i d e n t a l i s "From the Rocky Mountains of B r i t i s h America (Drummond) to B r i t i s h Columbia and Vancouver Island ( L y a l l , Macoun), Oregon (Cusick, Henderson, Howell) and the northern S i e r r a Nevada (Chico, Mrs. J. Bidwell, Gray)." It i s generally accepted that the material from Vancouver Island and Oregon belong to other species. One of Macoun's specimens from Vancouver Island (Mt. Finlayson, V.I., Macoun s.n. DAOl NY! CAN!) has been desig-nated as the holotype of S_. r u f i d u l a ; the material c o l l e c t e d by Howell from the Columbia River Gorge i s most l i k e l y S_. r u f i d u l a (high c l i f f s along the Columbia River at Multnomah F a l l s , J . Howell s.n. NDl PHI), and a l l the material from southern Oregon i s most l i k e l y S_. m a r s h a l l i i as j3. o c c i d e n t a l i s does not grow as far south as southern Oregon (Grant's Pass, J. Howell s_.n. NY!). We have chosen one of Macoun's specimens as the lectotype f o r _S. o c c i d e n t a l i s : Yale Mountain, B.C., May 17, 1875, J. Macoun s.n. CAN! ( F i g . 5, specimen A). Representative Specimens BRITISH COLUMBIA: 2 mi ne. of Salvus on Prince Rupert - Terrace Highway, J.A. Calder, D.B.O. Savile and J.M. Ferguson 13117 (DAO); Paradise Mine 15 mi w. of Windermere, J.A. Calder and D.B.O. Savile 11232 (DAO); Skagit V a l l e y , B.C. - Washington boundary to 10 mi n., H. Rhodes 16. and C. Brayshaw 6641 (DAO); B l u f f Lake about 12 mi ssw. of T a t l a Lake P.O., 51°46'N 124°43'W, J.A. Calder, J.A. Parmelee, R.L. Taylor 17263 (DAO); Kimberley, F. Fodor 119 (UBC); along Hwy. 3, about 2 mi w. of Nelson, W.C. McCalla 8127 (UBC); Hope-Princeton Hwy., n. of mile 126, K.I. Beamish et a l . 7638 (UBC); Blackwall, Manning Park, K.I. Beamish  and F. Vrugtman 60792 (UBC, DAO); r.ocky mountain summit, Liumchen area, above cabin, Cultus L. area, K. Beamish 7910 (UBC); Tod Mt., Kamloops, T.M.C. Taylor and A.F. Szczawinski 784 (UBC); Corrie Ridge, G a r i b a l d i , A.E. Beggs s.n. (UBC); Empetrum Ridge, G a r i b a l d i P r o v i n c i a l Park, K.I.  Beamish 680176 (UBC); Mt. Brent, Penticton, J.W. Eastham s.n. (UBC); Cathedral Lakes, Ashnola D i s t . , T.M.C. Taylor 1346 (UBC, WS, UC); Princeton, T.M.C. Taylor 2053 (WS); Mile 19, Flathead Rd., K.I. Beamish  156 (UBC, DAO); Goat Mt., 49°58'N 114°42'W, R.L. Taylor and D.M. Fergu- son 3039 (DAO); Moosehorn L., 58°10'N 132°07'W, R.J. P i l f r e y 57 (DAO); 3 mi s. of Camp 7, 18 mi sw. of Anahim L., D.W. Hyndman and H.W. Tipper  27 (DAO); subalpine slopes near Lake Botanie, Marble Mts. 5000 f t , J.W. Thompson and E.M. Thompson 70 (UC, WS); crevices of rocks, Mt. Queert, B.C., Macoun 8375 (CAN); Skagit R., Macoun 79347 (CAN); Crow's Nest Pass, Rocky Mts., J.M. Macoun 20142 (CAN). The following f i v e specimens from B r i t i s h Columbia have open inflorescences, p a r t i a l l y reflexed sepals, and l e a f shapes which make them resemble some of the S. m a r s h a l l i i ssp. idahoensis plants from Idaho: 6 mi n. of Clear-water Station, North Thompson River at Bear Creek F a l l s , J.A. Calder, J.A. Parmelee and R.L. Taylor 17423 (DAO); 1 mi sw. o f Midway, J . Grant 38 (DAO); Goat River Canyon on Erickson Mt. near Creston, J.A. Calder 17. and D.B.O. Savile 7735 (DAO); 3 mi e. of South Slocan on Nelson-Trail Highway, J.A. Calder and K.W. Spicer 32877 (DAO). The following plant had excessively large broad leaves which give i t a strange appearance: 38 mi on road from Anahim L. to B e l l a Coola, 52°25'N 125°53'W, J.A.  Calder, J.A. Parmelee and R.L. Taylor 17338 (DAO); ALBERTA: Shoulder of Wilcox Mt., Jasper Nat. Park, E.H. Moss s.n. (CAN); Banff Nat. Park., near lookout point above Peyto L., W.C. McCalla  7073 (ALTA); head of L. Louise, Rocky Mts., J.M. Macoun 65316 (CAN); rocky slope above Bertha L., Waterton Lakes Nat. Park, CO. Rosendahl  3618 (ALTA); edge of woods above Red Rock Canyon, Waterton Lakes Nat. Park, W.C. McCalla 11673 (ALTA); Goat Cr., e. flank of Flathead Range, R.A. P r i c e 10 (DAO); Savannah Cr. j u s t w. of I s o l a t i o n (Isola) Peak 36 mi due n. of Coleman, E.W. Mountjoy 1 (DAO); SASKATCHEWAN: Cypress H i l l s Park., dry h i l l s i d e , A.J. Breitung  4212 (DAO); WASHINGTON: OKANOGAN Co.: moist rocky ledges, Sawtooth Ridge, Valley of War Cr., H. St. John, W.D. Courney and C S . Packer 3822 (WTU, WS, UC); sandy h i l l s i d e , Ridge, Ptarmigan Cr., Hidden Ls., O.T. Edwards  263 (WS, WTU); rocky outcrop below State Pk., at the middle fork of the Pasayten R., Cascade Mts., M. Ownbey and F.C Meyer 2299 (ALTA, WTU, UC, OSC, WS); exposed open summit of Muckamuck Lookout, T i f f a n y Range, J.W. Thompson 6985 (WTU); near Ruby Grade, on Salmon Cr., Ruby, C.B. Fiker 1183 (WS, WTU); SKAMANIA Co.: Wettenburg, Chequash Mts., W.N. Suksdorf 9671 (WSP); crevices of c l i f f s under w a t e r f a l l s , Paradise Park, Mt. Rainier, J.W. Thompson 7614 (WTU); SPOKANE Co., damp rocky 18. h i l l s i d e near Latah Cr., southeast of Spangle, W. Suksdorf 8616 (UC, WS, WTU); YAKIMA Co.: crags and moist slopes i n Chinook Pass, Cascade Mts., J.W. Thompson 9841 (WTU); IDAHO: BLAINE Co.: Boulder Ck. Canyon, Sawtooth Nat. Forest, Sawtooth Mts., J.W. Thompson 14056 (NY, WTU); Alpine Ck., nw. of Alturus L., Sawtooth Primitive Area, C.L. Hitchcock and C.V. Muhlick 10505 (NY, WS); BOISE Co.: on rocks 5 mi n. of Elk L., Sawtooth P r i m i t i v e Area, headwater of S. Fork Payette R. above Sacajawea Hot Spr., C.L. Hitchcock and C.V. Muhlick 9872 (NY); BONNER Co.: Continental Mt., 25 mi w. P o r t h i l l , J.H. Christ 1678 (NY): CUSTER Co.: moist, protected slopes, Josephus Ls., J.F. Macbride and E.B. Payson 3549 (NY, UC); saddle w. of Castle Pk., White Cloud Range, C h a l l i s Nat. For., C.L. Hitchcock and C.V. Muhlick 10869 (NY, WTU, WS); meadow on w. base of Ryan Pk., Boulder Mts., Sawtooth Nat. For., C.L. Hitchcock  and C.V. Muhlick 10647 (NY, WTU); i n semiswampy s o i l i n rocky canyon about 10 mi w. of C h a l l i s , C.L. Hitchcock and C.V. Muhlick 9514 (UC, NY, WTU, WS); Livingston Mine, R.J". Davis 674 (WS); DEERLODGE Co.: dry slope above Storm L., Anaconda Mts., C.L. Hitchcock and C.V. Muhlick  14820 (NY); FREMONT Co.: base of summit of mountains ne. of lake, Henry L., E.B. Payson and L.B. Payson 2059 (NY, UC); IDAHO Co.: on divide between head of Sheep Cr. and head of the East fork of Sheep Cr., J . Packard 344 (WS). MONTANA: BEAVERHEAD Co.: on top of Odell Peak, Pioneer Range, C.L. Hitchcock and C.V. Muhlick 14925 (WTU, NY, WS); Cirque above Ajax L., j u s t below summit of Continental Divide, on granite talus and 19. rock, B i t t e r Root Range, C L . Hitchcock and C.V. Muhlick 12679 (WS, NY,. WTU); East P i n t l a r Peak, head of P i n t l a r Cr., Anaconda Range, C L . Hitch- cock and C.V. Muhlick 12846 (WTU, NY, WS); talus slope above Torrey L., Pioneer Range, C L . Hitchcock and C.V. Muhlick 15056 (WTU, NY); DEERLODGE Co.: w. slope of Mt. Tiny, s. of Storm L., Anaconda Mts., C L . Hitchcock and C.V. Muhlick 14860 (WTU, WS) ; FLATHEAD Co.: on ridge, Mt. Aeneas, Swan Range, L.H. Harvey 3283 (MONTU); Mt. Cannon above McDonald Cr., G l a c i e r Nat. Park, L.H. Harvey 6050 (MONTU); i n seep below snow bank on talus slope, Logan Pass, G l a c i e r Nat. Park, L.H. Harvey 6689 (MONTU); high above and se. of McDonald L., Mission Range, C L . Hitchcock 18300 (WS, WTU, UC, COLO, NY, ID); MISSOULA Co.: talus s l i d e , granite rock, 5 mi above Bonner, C L . Hitchcock and C.V. Muhlick  11425 (WTU, NY, WS) ; h mi w. of Upper Holland L., Flathead Range, C L .  Hitchcock 18422 (WTU, WS); POWELL Co.: top of Gordon Mt., 6 mi s. Big P r a i r i e Ranger Station, Flathead Nat. For., C L . Hitchcock 18848 (WTU). WYOMING: rocky part of Canyon Cr., Grand Teton Nat. Park, W.C. McCalla  4836 (ALTA); on mossy rocks i n spray of f a l l s , Crazy Cr., on Red Lodge-Cooke City Highway, Park Co. , C L . Porter 5437 (DAO, WTU); moist mossy h i l l s i d e , Cascade Canyon, L. Williams 1135 (ORE, ID); Gl a c i e r Nat. Park, moist rocky c l i f f s near Cracker L. t r a i l , D.H. Brant 45 (WTU). 20. DISCUSSION The r e l a t i o n s h i p of the taxa previously referred to as Saxifraga  m a r s h a l l i i , S_. idahoensis, and S^. o c c i d e n t a l i s has been doubtful. We f e e l that this r e l a t i o n s h i p i s best expressed by the following taxonomy: _S. m a r s h a l l i i Greene ssp. m a r s h a l l i i , J3. m a r s h a l l i i Greene ssp. idahoen- s i s (Piper) D.L. Krause and K.I. Beamish, and S_. o c c i d e n t a l i s S. Wats. Saxifraga m a r s h a l l i i ssp. m a r s h a l l i i grows i n western Oregon and northwestern C a l i f o r n i a , S_. m a r s h a l l i i ssp. idahoensis i n Chelan and K i t t i t a s Counties of Washington and i n northcentral Idaho and adjacent areas of Oregon and Montana. Saxifraga O c c i d e n t a l i s grows i n B r i t i s h Columbia and adjacent Alberta, Cypress H i l l s of Saskatchewan, Washing-ton, Idaho, northeastern Oregon, the Rocky Mountains of Montana and Wyoming, and occasionally the Rocky Mountains as f a r south as New Mexico. Saxifraga m a r s h a l l i i ssp. m a r s h a l l i i can be separated quite r e a d i l y from J5. o c c i d e n t a l i s . However, ssp. idahoensis i s much more d i f f i c u l t to separate from S_. o c c i d e n t a l i s because, where t h e i r ranges meet, the two taxa have c h a r a c t e r i s t i c s which tend to approach each other. For example^ i n Idaho some ssp. idahoensis specimens are found which have p a r t i a l l y i n f e r i o r ovaries, filaments that are not strongly clavate, rhombic leaves and more congested inflorescences than t y p i c a l s s p . idahoensis. These ssp. idahoensis plants can be separated from S_. o c c i - dentalis by the former's reflexed sepals, yellow spots on the p e t a l s , deciduous nature of the petals, p e r s i s t e n t filaments, and the long pedicels of the flowers. We believe that ssp. idahoensis i s more c l o s e l y related to S_. m a r s h a l l i i than to S_. o c c i d e n t a l i s and therefore cannot be c l a s s i -f i e d as an i n f r a s p e c i f i c taxon of S_. oc c i d e n t a l i s (Hitchcock 1961) even though there may have been some intergradation of c h a r a c t e r i s t i c s with this species. Saxifraga idahoensis has been reduced to a subspecies of S_. m a r s h a l l i i because some specimens of ssp. idahoensis are almost i n d i s t i n -guishable from specimens of ssp. m a r s h a l l i i . The Washington and Idaho populations have been separated from the Oregon and C a l i f o r n i a popula-t i o n as a subspecies because the i n f e r i o r i t y of the ovary, open nature of the inflorescence, shape of the filaments, and the l e a f shape d i f f e r s l i g h t l y i n most specimens. The morphological characters i n ssp. idahoensis are skewed s l i g h t l y towards those i n S_. o c c i d e n t a l i s . The tendency toward morphological s i m i l a r i t y between the l a t t e r two taxa could be the r e s u l t of l i m i t e d gene flow between the two taxa at some previous stage i n t h e i r evolution. Present c y t o l o g i c a l information precludes gene flow at the present time. The presence or absence of yellow spots on the petals of fresh material appears to be a good c h a r a c t e r i s t i c f o r separating the sub-species of S_. m a r s h a l l i i from S_. o c c i d e n t a l i s . A l l the l i v e c o l l e c t i o n s of ssp. idahoensis and ssp. m a r s h a l l i i had yellow spots, whereas no yellow spots were observed on c o l l e c t i o n s of S_. o c c i d e n t a l i s . Unfor-tunately, this c h a r a c t e r i s t i c i s not us e f u l for most dried specimens as the yellow spots frequently fade upon drying. The q u a n t i t a t i v e measurements taken from both the l i v e populations and the herbarium specimens could not be used to separate the taxa 22. because the v a r i a b i l i t y w i t h i n each taxon was so great that the taxa overlapped f o r a l l measurements taken. The chromosome counts strengthen the hypothesis that ssp. idahoensis i s more c l o s e l y r e l a t e d to ssp. m a r s h a l l i i than i t i s to j>. o c c i d e n t a l i s . As seen i n Table 1, a l l the ssp. m a r s h a l l i i and ssp. idahoensis popula-tions counted had 10 pairs of chromosomes. Only one population of S_. o c c i d e n t a l i s with 10 pairs of chromosomes has been reported, and i t i s from Alberta, w e l l to the north of the range of ssp. idahoensis. The chromosome counts obtained for S_. o c c i d e n t a l i s i n d i c a t e that the cyto-genetics of t h i s species i s more complicated than previously suspected. The d i f f e r e n t counts of jS. o c c i d e n t a l i s could not be correlated with morphological d i f f e r e n c e s . More populations of S_. o c c i d e n t a l i s need to be counted to discover the extent of t h i s chromosome v a r i a b i l i t y . The p o s s i b i l i t y of combining S_. m a r s h a l l i i and S_. o c c i d e n t a l i s into one species as suggested by Piper (1906) and Hitchcock (1961) does not seem s a t i s f a c t o r y . The separation of these two taxa over most of t h e i r range i s quite s a t i s f a c t o r y and, as Engler and Irmscher (1916) point out, there i s abundant h y b r i d i z a t i o n between species i n the same se c t i o n , and even between sections. If species i n the genus Saxifraga were lumped together because they occasionally intergrade, then the genus would have one or two extremely complex superspecies. Saxifraga o c c i d e n t a l i s i s an extremely v a r i a b l e species. Many workers, p a r t i c u l a r l y Johnson (1923) and Small (1905), have described species which have subsequently been put into the synonomy of S_. o c c i - dentalis . The v a r i a b i l i t y within t h i s species i s so great that most 23. of the combinations of c h a r a c t e r i s t i c s used to d i s t i n g u i s h i t s sub-s p e c i f i c categories can be found throughout the range of the species. We f e e l that nothing would be gained by r e t a i n i n g any su b s p e c i f i c categories f o r S_. o c c i d e n t a l i s as we have been unable to co r r e l a t e d i s t r i b u t i o n with any of the proposed combinations of c h a r a c t e r i s t i c s . The possible exception i s the p o s i t i o n of S_. r u f i d u l a . This taxon has been c l a s s i f i e d as a species, and as a v a r i e t y and sub-species of S_. o c c i d e n t a l i s . The taxonomic p o s i t i o n of _S. r u f i d u l a i s s t i l l i n doubt as i n s u f f i c i e n t data have, as yet, been obtained. Some l i v e c o l l e c t i o n s as w e l l as a few herbarium specimens from southern Oregon appear to be s l i g h t l y d i f f e r e n t from S_. m a r s h a l l i i . Some of these specimens were labeled S_. h o w e l l i i . We have not seen enough material of t h i s taxon to reach any conclusions regarding i t s taxonomic p o s i t i o n . The subpopulation of S_. m a r s h a l l i i has been changed from a v a r i e t y (Engler and Irmscher 1916) to a subspecies as our views on the use of the rank subspecies i n l i e u of v a r i e t y are s i m i l a r to those of Raven (1962) as based on the views of Du Rietz (1930) and Pennell (1949). 24. LITERATURE CITED Abrams, L. 1944. Saxifraga. In 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. V o l . 2. Stanford Univ. Press, Stanford Univ., C a l i f , pp. 355-367. Beamish, K. I. 1961. Studies of meiosis i n the genus Saxifraga of the P a c i f i c Northwest. Can. J . Bot. 39: 567-580. Du Rietz, G. E. 1930. The fundamental units of b i o l o g i c a l taxonomy. Svensk. Bot. Tidskr. 2_4: 333-428. Engler, A., and E. Irmscher. 1916, 1919. Saxifragaceae - Saxifraga. In Das Pflanzenreich, 4, 117, H. 67 (1916); H. 69 (1919). Reprinted 1958. Greene, E. L. 1888. New or noteworthy species XXIII. P i t t o n i a , 1_: 159. Hitchcock, C. L., and A. Cronquist. 1961. Saxifraga. In Vascular plants of the P a c i f i c Northwest, Part 3. Saxifragaceae to E r i c a -ceae. Edited by C. L. Hitchcock. A. Cronquist, M. Ownbey, and J . W. Thompson. University of Washington Press, Seattle, pp. 31-55. t Johnson, A. M. 1923. A r e v i s i o n of the,North American species of the se c t i o n Boraphila Engler of the genus Saxifraga (Tourn.) L. Minn. Stud. B i o l . S c i . _4: 1-110. Packer, J . G. 1968. IOPB chromosome number reports. Taxon, 17(3) : 287. Pennell, F. W. 1949. Toward a simple and cl e a r nomenclature. Am. J . Bot. 36: 19-22. Piper, C. V. 1900. New and noteworthy northwestern plants. IV. B u l l . Torrey Bot. Club, 27: 394. . 1906. Saxifraga. In F l o r a of the state of Washington. Contrib. U.S. Nat. Herb. 11: 312-317. Raven, P. H. 1962. The systematics of Oenothera subgenus Chylismia. Univ. C a l i f . Publ. Bot. 34: 1-122. Small, J. K. 1905. Micranthes. N. Am. F l o r a , 22(2): 132-148. Snow, R. 1963. A l c o h o l i c hydrochloric acid-carmine as a s t a i n f o r chromosomes i n squash preparations. Stain Technol. 2_3: 9-13. 25. Watson, S. 1888. XVII. Contributions to American botany. Proc. Am. Acad. Arts, 2_3: 264. Whittman, W. 1965. Aceto-haematoxylin - chloral hydrate for chromosome staining. Stain Technol. 10_(3) : 161-162. TABLE I. CHROMOSOME COUNTS Collect o r C o l l e c t i o n Number Location Saxifraga occidentalis 1 Packer 2839(ALTA) Blakeston Mt., Waterton Nat. Park, Alberta Beamish, Vrugtroan and Sperrings 8224 2Beamish 7901 Krause and Beamish 31-71 Krause 69018 Krause 43-70 Krause 33-70 Beamish 57501 Krause 19-71 Krause 16-71 Krause and Maze 690001 Krause 42-70 Krause 40-70 Krause 20-70 Botanie Valley, near Lytton, B.C. Mt. Lihumitson, S. of Chilliwack, B.C. Yale, B.C. Blackwall, Manning Park,B.C. Mt. Apex, near Penticton, B.C. Dominion Mt., near Salir.o, B.C. Tchaikazan River Valley, B.C. Tweedsmuir park,B.C. Bluff L.•, B.C. Chehalis, Washington Boulder Mt., N. of Ketchum, Idaho Bayhorse L., near C h a l l i s , Idaho New Meadows, Idaho Saxifraga m a r s h a l l i i ssp. m a r s h a l l i i Krause and Beamish 30-70 Krause and Beamish 10-70 Krause and Beamish 35-70 Krause and Beamish 15-71 Krause and Beamish 36-70 Stikum, Oregon Dora, Oregon Bear Cr., Middle fork of Coquille R., Oregon Scottsburg, Oregon Mapleton, Oregon Saxifraga m a r s h a l l i i ssp. idahoensis Krause 38-70 Krause 5-71 Krause 6-71 Krause 41-70 Krause 32-70 Krause 32-71 Krause 33-71 Krause 39-70 1 Packer, Taxon 17(3):287. 1968. St. Joe River, Avery, Idaho Myrtl-e, Idaho O r i f i n o , Idaho Winchester, Idaho Cow C.r., Hamilton, Montana 7 mi N. of Leavenworth, Wash. I c i c l e Cr., S. of Leavenworth, Wash. Mt. Tiff a n y , Washington 2 6 . Chromosome numbers n - 10 n - 19 n - 19 n - 19 n - 19 n - 19 n - 19 n - 19 n - 28 n - 28 n - 29 n - ca 29 n - ca 29 n - ca 29 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 n - 10 Beamish, Can. J . Bot. 39:567-580. 1961. 2 7 V © • © *| N.rUir.il \\\<»x\ Swui "I' l'.ni:i'l;i nmn II i M — tt \ \ 1 © 1» t- » 4* • •© Figs. 1-4. The. black l i n e represents 2)i. Fig. 1. Saxifraga marshallii ssp. mar- s h a l l i i ; n = 10; Sitkum, Oregon; Krause and Beamish 30-70. Fig. 2. S_. inarshallii ssp. idahoensis; n = 10; Winchester, Idaho; D.L. Krause 41-70. . Fig. 3. _S. occi-dentalis ; n = 28; Tweedsmuir Pk., B.C., Krause 39-71. Fig. 4. S_. occidentalis ii - 29; Cbehalis, Washington; Krause & Maze 690001. Fig. 5. Photograph of lecto-type for Saxifraga occidentalis: specimen A.J. Macoun s.n.; Yale Mountain, B.C.; May 17, 1875, CAN. 28. FIGURE 6. D i s t r i b u t i o n of Saxifraga m a r s h a l l i i spp. mar_sh,aJJ;ii, S. m a r s h a l l i i ssp idahoensis, and S. o c c i d e n t a l i s i n Washington, Idaho, Montana and southern B r i t i s h Columbia. 29. FIGURE 7. D i s t r i b u t i o n of Saxifraga o c c i d e n t a l i s i n B r i t i s h Columbia. 30. PART II INTRODUCTION The d i s t r i b u t i o n and systematics of Saxifraga o c c i dentalis and i t s c l o s e l y r e l a t e d species i n B r i t i s h Columbia are poorly known. Fragmentary treatments have been included i n the works of Calder and Taylor (1968), Hulten (1968), Hitchcock et a l (1961), Rydberg (1917) and Henry (1915). However, no f l o r a has dealt with the B r i t i s h Columbia complex as a whole. Even the monographic works of Engler and Irmscher (1916) , Small (1905) and Johnson (1923) do not adequately treat this complex. In B r i t i s h Columbia the Saxifraga o c c i d e n t a l i s complex consists of the following species of Engler and Irmscher's (1916) Saxifraga section Boraphila subsection N i v a l i - v i r g i n i e n s e s : S_. o c c i d e n t a l i s S. * Wats., S_. r e f l e x a Hook.; S_. rhomboidea Greene , S_. r u f i d u l a (Small) James Macoun, jS. n i v a l i s L., and S_. tenuis (Wahl.) H. Sm. The l a s t two taxa are c l e a r l y d i s t i n g u i s h a b l e from the other four but, as w i l l be explained l a t e r , t h e i r r e l a t i o n s h i p to each other i s s t i l l not c l e a r . Throughout t h i s paper, therefore, they are referred to as the "S_. n i v a l i s - t e n u i s complex". In an attempt to c l a r i f y the systematic problems i n t h i s species complex, a program of f i e l d and herbarium studies was undertaken. The herbarium material and l i v i n g material c o l l e c t e d by the authors were the source of morphological data. Quantitative measurements were taken from Although j ^ . rhomboidea does not extend as far north as B r i t i s h Columbia, i t i s included here because most f l o r a s give B r i t i s h Columbia and Alberta, as i t s northern l i m i t . leaves, scapes and inflorescences and were analyzed s t a t i s t i c a l l y . However, they proved to be inadequate to delimit the taxa under study. A combination of q u a l i t a t i v e c h a r a c t e r i s t i c s was found to be the most useful c r i t e r i o n f o r the separation of various taxa. These were used to erect the diagnostic key to the species. As t h i s paper deals p r i m a r i l y with B r i t i s h Columbia, the d i s t r i -bution maps and l i s t s of representative specimens do not include, i n a l l cases, adjacent areas. Herbarium material was examined from the following h e r b a r i a : ALTA, Dept. of Botany, University of Alberta, Edmonton; UAC, Dept. of Biology, University of Calgary, Alberta; CAN, National Museum of Canada, Ottawa; COLO, University of Colorado, Boulder, Colorado; DAO, Plant Research I n s t i t u t e , Dept. of A g r i c u l t u r e , Ottawa; ID, Dept. of Botany, University of Idaho, Moscow, Idaho; MONTU, University of Montana, Missoula, Montana; ND, University of Notre Dame, Notre Dame, Indiana; NY, the New York Botanical Garden, New York C i t y , New York; ORE, University of Oregon, Eugene, Oregon; OSC, Dept. of Botany, Oregon State U n i v e r s i t y , C o r v a l l i s , Oregon; UC, University of C a l i f o r n i a , Berkeley, C a l i f o r n i a ; WS, Dept. of Plant Pathology, State College of Washington, Pullman, Washington; and WTU, Dept. of Botany, University of Washington, Sea t t l e . 32. TAXONOMY KEY TO THE SPECIES A. Ovary 1/2 i n f e r i o r ; filaments subulate, equalling the sepals; sepals about as broad as long, ascending; inflorescence a t i g h t glomerate head or a few-flowered cyme. B. Filaments not i n f l e x e d at t i p ; leaves rhombic to spatulate, teeth usually only on upper 1/2 of l e a f blade; inflorescence usually a few-flowered cyme; pubescence of scape sparse to dense, hairs white with purple glandular t i p s ; circumboreal, southward i n the high mountains of B r i t i s h Columbia and Alberta^ p a r t i c u l a r l y the Rocky Mountains as f a r south as Colorado 1. S_. n i v a l i s - t e n u i s complex BB. Filament t i p s i n f l e x e d ; leaves rhombic with w e l l developed crenate margins over most of l e a f blade; inflorescence a t i g h t , o ccasionally interrupted, glomerate head; pubescence of scape dense, hairs yellowish with yellow glandular t i p s ; from c e n t r a l Montana south 2. S_. rhomboidea AA. Ovary superior to 1/3 i n f e r i o r ; filaments subulate to clavate, longer than the sepals; sepals longer than broad, spreading to strongly reflexed; inflorescence open, occasionally tight-glomerate. C. Petals with 2 basal yellow spots; filaments strongly clavate to p e t a l o i d ; sepals strongly reflexed; ovary completely superior; leaves puberulent (seldom glabrate) on both upper and lower surface; from f a r northern B r i t i s h Columbia i n t o Alaska, Yukon and Northwest T e r r i t o r i e s 3. S_. r e f l e x a 33. CC. Petals immaculate; filaments subulate, l i n e a r or clavate; sepals spreading to s l i g h t l y reflexed; ovary 1/5 to 1/3 i n f e r i o r ; leaves glabrous to rusty-tomentose on lower surface, glabrous on upper surface. D. Inflorescence a corymb; leaves ovate to rhombic with crenate margins with small, well-formed teeth, often copiously rusty-tomentose on lower surfaces; filaments subulate; i n B r i t i s h Columbia r e s t r i c t e d to Vancouver Island, also found i n the Olympic Peninsula, the Columbia River Gorge, and the Santeam River V a l l e y , Oregon 4. S_. r u f i d u l a DD. Inflorescence various, seldom f l a t - t o p p e d ; leaves various, seldom rusty-tomentose on lower surface; filaments subulate to s l i g h t l y clavate; from northern B r i t i s h Columbia south to Washington, Idaho, Montana, northeast Oregon and occasionally i n the Rocky Mts. as f a r south as New Mexico 5. S_. o c c i d e n t a l i s 1. Saxifraga n i v a l i s - t e n u i s complex The taxonomy of the _S_. n i v a l i s - t e n u i s complex needs extensive biosystematic study to separate the taxa involved. Most workers (Savile 1961, Love & Love 1951 & Jorgensen et a l 1958) f e e l that S_. n i v a l i s var. tenuis Wahl. i s s u f f i c i e n t l y d i s t i n c t from S_. n i v a l i s to warrant s p e c i f i c rank, although others (Hult£n 1968, F l o v i c k 1940) f e e l i t should remain a subcategory of S_. n i v a l i s . Over much of t h e i r range they can be adequately separated using c r i t e r i a s i m i l a r to those proposed by Savile 34. (1961): number of flowers; shape of inflorescence; colour of pet a l s , calyx and scape; type of pubescence; and e s p e c i a l l y shape of leaves. In the Yukon, however, these d i s t i n c t i o n s seem to break down and the plants appear to intergrade. Cytotaxonomists i n Europe (see Love & Love 1961) and North America (Hedberg 1967, Johnson & Packer 1968) have reported S_. n i v a l i s to have either 58, 60 or about 60 chromosomes and J3. n i v a l i s var. tenuis to have 20 chromosomes. It i s d i f f i c u l t to c o r r e l a t e the d i f f e r e n t counts with morphological features as we have not obtained enough counts and the previous authors have not given the c r i t e r i a they used f o r i d e n t i f i c a -t i o n . We have obtained counts of 10, 20 and 30 pairs of chromosomes for the material c o l l e c t e d i n the Yukon (Table 1, F i g . 1A & IE). The v a r i a -b i l i t y i n the chromosome numbers wit h i n t h i s complex may be more prevalent than was previously suspected. The S_. n i v a l i s - t e n u i s complex i s circumboreal. It i s common i n the Rocky Mountains of B r i t i s h Columbia and A l b e r t a y ( F i g . 2) and occasionally southward as f a r as Colorado. The scattered d i s t r i b u t i o n i n B r i t i s h Columbia and southward i s probably due to c o l l e c t i n g anomalies. In B r i t i s h Columbia i t i s sympatric with S_^_ o c c i d e n t a l i s i n high mountains but i t can be r e a d i l y separated from S_. o c c i d e n t a l i s as shown i n the key to species. Some representative specimens of the S_. n i v a l i s - t e n u i s complex i n A B r i t i s h Columbia and Alberta follow: BRITISH COLUMBIA: On rocks below summit of O t t e r t a i l Pass, Rocky Mt. Park, Macoun s.n. (CAN); A l i c e Mt., A d d i t i o n a l c i t a t i o n s f o r a l l species appear with, chromosome counts i n Table 1. 35. n.e. of Turnagain L., Cassiar D i s t . A.F. Szczawinski s.n. (DAO); Murray Range at north end of Azouzetta L., Hart Highway, J.A. Calder,  D.B.O. Savile and J.M. Ferguson 14075 (DAO); Red Rose Tungsten Mine about 9 miles e. of Skeena Crossing, J.A. Calder, D.B.O. Savile and J.M. Fer- guson 15182 and 13369 (DAO); Mile 75, Haines Rd., T.M.C. Taylor, A.  Szczawinski and M. B e l l 824 (DAO, UBC); Mile 60, Haines Rd., T.M.C. Taylor,  A. Szczawinski and M. B e l l 1204 (UBC, DAO); Mile 68, Haines Rd., T.M.C.  Taylor, A. Szczawinski and M. B e l l 896 (DAO, UBC); Mile 61, Haines Rd., D. Krause, K.I. Beamish and J. Luitjens 681807 (UBC). ALBERTA: Snow L. Pass about 30 miles n. of Banff, J.A. Calder 23996 (DAO); Banff Nat. Pk., alpine, wet c l i f f s above Lake Agnes, A. Harmon 8 (DAO); Rocky slope, above timber l i n e , Jasper Nat. Pk., E.H. Moss 2736a (ALTA); top of rocky ridge about 5300 f t . above Cadomin, E.H. Moss 10339 (ALTA). The following sheets carry specimens of both S_. occ i d e n t a l i s and the j>. n i v a l i s - t e n u i s complex. ALBERTA: Mt. Edith C a v e l l , J.A.  Calder 37210 (DAO). BRITISH COLUMBIA: Mt. T h o r n h i l l , Terrace, J.A. Calder, D.B.O. Savile and J.M. Ferguson 14888 (DAO); Sylvan Mine, Hudson Bay Mt., about 9 miles w. of Smithers, J.A. Calder, D.B.O.  Savile and J.M. Ferguson 14612 (DAO). The l a s t possibly has hybrids. 2. Saxifraga rhomboidea Greene Most f l o r a s f o r the P a c i f i c Northwest i n c o r r e c t l y give southern B r i t i s h Columbia and Alberta as the northern l i m i t s of S. rhomboidea 36. d i s t r i b u t i o n . This i s probably due to the s u p e r f i c i a l resemblance of some _S. o c c i d e n t a l i s plants with t i g h t glomerate heads and densely pubescent scapes to S_. rhomboidea. As shown i n the key these two species can be separated r e a d i l y using filament shape, i n f e r i o r i t y of ovary, shape, and p o s i t i o n of sepals, and colour of the glandular hairs on the scape. Saxifraga rhomboidea extends only as f a r north as c e n t r a l Montana (Fig. 3). In southern Montana and Wyoming i t i s occasionally sympatric with S^ . o c c i d e n t a l i s . The following herbarium sheets were found with both _S. rhomboidea and S_. o c c i d e n t a l i s on them: H e l l Roaring t r a i l , Wyoming, L i l l i a n Winiecki 67 (WS); Mts. at 7000 f t , Sheridan, Montana, L.A. F i t c h s.n. (NY)f Columbia F a l l s , Montana, R.S. William s.n. (NY). The northern d i s t r i b u t i o n of S_. rhomboidea i s shown by the following representative specimens from Montana: MONTANA: CARBON Co.: 4 miles n.w. of Red Lodge, CH. Draper s.n. (UC); GALLATIN Co.: Bozeman, G a l l a t i n Canyon near Squaw Cr., W.E. Booth 1514 (WTU, WS, MONTU); Bozeman Bridge Canyon, J.W. Blankinship s.n. (UC); MEAGHER Co.: s.w. slope of King's H i l l , L i t t l e B elt Mts., C L . Hitchcock and C.V. Muhlick  12337a (WTU, WS); Yoho Pk., L i t t l e B elt Mts., south side, C L . Hitch- cock 16131 (WTU, WS, UC); MADISON Co.: . 1 mile s. of Crockett L. (Devil' s L.) , Gravelly Range, C L . Hitchcock and C.V.. Muhlick 12500 (WTU); s. side of Black Butte, Gravelly Range, C L . Hitchcock 16875 (WTU); PARK Co.: Cooke Guard Station, 3 miles e. of Cooke C i t y , J . C Witt  1161 (COL, WTU, WS); Wallrock Basin, n.w. of W i l l s a l l , W.N. Suksdorf  272 (WS); STILLWATER Co.: n. slope of Mt. Haystack, head of Boulder Cr., Absaroka Nat. For., C L . Hitchcock and C.V. Muhlick 13424a (WTU). 37. 3. Saxifraga r e f l e x a Hook. Saxifraga r e f l e x a extends southward only i n t o the most northerly portion of B r i t i s h Columbia ( F i g . 2). Its taxonomy i s r e l a t i v e l y c l e a r -cut. I t can be i d e n t i f i e d by i t s reflexed sepals, yellow spots on the pet a l s , and i t s puberulent leaves. Saxifraga r e f l e x a appears to be most c l o s e l y r e l a t e d to S^ . mar- s h a l l i i Greene growing i n Oregon, C a l i f o r n i a , Washington and Idaho. Both have yellow spots on the pe t a l s , reflexed sepals, superior ovaries, open inflorescences, and 10 pairs of chromosomes (Krause and Beamish, i n press), with the exception of one population of S_. r e f l e x a from Q u i l l Creek, Yukon. At the Q u i l l Creek s i t e counts were found of n = 10, n = 20, and (probably i n a hybrid) n = ca. 30 (Table 1). Representative specimens from B r i t i s h Columbia: F l o r a of Cassiar, T.M.C. Taylor, A. Szczawinski and M. B e l l 512, 623, and 498 (UBC); Mile 87, Haines Rd.,, T.M.C. Taylor, A. Szczawinski and M. B e l l 932 (UBC); Mile 81, Haines Rd., T.M.C. Taylor, A. Szczawinski and M. B e l l  1266 (UBC); B.C. - Yukon boundary, 59°58*W, M.P. and R.T. P o r s i l d 198 (CAN); Stonehouse Cr., Mile 61, Haines Rd., K.I. Beamish, D.L. Krause  and J . Luitjens 681801 (UBC). 4. Saxifraga r u f i d u l a (Small) James Macoun* Saxifraga r u f i d u l a i s found on Vancouver Island, the Olympic Peninsula, the Columbia River Gorge and i n the Santeam River Valley of Oregon (Fig. 2). In B r i t i s h Columbia i t i s confined to Vancouver Island. I t i s geographically d i s t i n c t from S_. o c c i d e n t a l i s as the l a t t e r i s found on the mainland. The name Saxifraga r u f i d u l a (Small) Macoun was i n c o r r e c t l y a t t r i b u t e d to John Macoun by Beamish (1967). 38. I n the Columbia River Gorge S_. r u f i d u l a appears to intergrade with S_. o c c i d e n t a l i s and perhaps with S_. m a r h s a l l i i . Much work remains to be done to determine the extent of the intergr^adation. Chromosome counts obtained for S_. ruf i d u l a are given i n Table 1. On Vancouver Island i t was found to have 10 and 29 pairs of chromosomes but d e t a i l e d study revealed no morphological difference between the 10-paired populations and the 29-paired population (Fig. ID). In the Columbia River Gorge, counts f o r S_. ruf i d u l a were obtained of n = 10, n = 19 ( F i g . IB), n = 28 ( F i g . IC & F) and n = ca. 28. The plants i n the Gorge are morphologically very v a r i a b l e but none of the morphological differences could be correlated with chromosome numbers. Representative specimens for B r i t i s h Columbia follow: Mt. Helmcken, n.w. of V i c t o r i a , L.C. Sherk 302 (DAO); along s. side of Cameron L. between Port A l b e r n i and P a r k s v i l l e , J.A. Calder, J.A. Parmelee, and R.L. Taylor 16414 (DAO); near C l i f f s i d e on east side of Shawnigan L., J.A. Calder, J.A. Parmelee, and R.L. Taylor 16245 (DAO); along Sooke R. about 3 miles n. of Milne's Landing, w. of V i c t o r i a , J.A. Calder 28420 (DAO); Castlecrag Mt., J.E. U n d e r h i l l 406 (DAO); Saanich Arm, J.R. Anderson 644^ (DAO); Mt. Douglas, V i c t o r i a , P.P. Henson s.n. (DAO); Mt. Arrowsmith, J.A. Calder, J.A. Parmelee and R.L. Taylor 19431 (DAO); Observatory H i l l , n. of V i c t o r i a , J.A. Calder and M.T. MacKay 28660 (DAO); above Boston Lk., below Mt. Becher, J.A. Calder and M.T.  MacKay 32557 (DAO); Nanoose H i l l , Nanoose Bay, J.A. Calder and  M.T. MacKay 429021 (DAO). 39. 5. Saxifraga o c c i d e n t a l i s S. Wats. Saxifraga o c c i d e n t a l i s can be s a t i s f a c t o r i l y separated from other species of sec t i o n Boraphila subsection N i v a l i - v i r g i n i e n s e s using the characters i n the key,although as mentioned e a r l i e r , some s p e c i -mens with t i g h t glomerate heads have been mistakenly i d e n t i f i e d as S_. rhomboidea (See under S_. rhomboidea) . The d i s t r i b u t i o n of S_. o c c i - dentalis i s given i n Figures 2 and 3. Representative specimens of S_. o c c i d e n t a l i s with t i g h t glomerate heads, many of which have been i d e n t i f i e d as S_. rhomboidea, follow: BRITISH COLUMBIA: Quiniscoe L., Ashnola d i s t r i c t , M.E. Barr PB69 (UBC); Mt. McLean, near L i l l o o e t , B.C., J.M. Macoun s.n. (CAN); head of M c G i l l i v r a y Cr., Cascade Range, J.M. Macoun s.n. (CAN); Slesse Mt., Chilliwack Valley, W. Spreadbourough s.n. (CAN); Blackwall Pk., Manning Park, L. Holm 428 (UBC). MONTANA: Rising Sun Meadows, G l a c i e r Nat. Park, L.H. Harvey 4773 (MONTU); Logan Pass, G l a c i e r Nat. Park, G.N. Jones  5525 (UC, WTU); Siyeh Pass, G l a c i e r Nat. Park, S. Bamberg 288 (COL); LAKE Co.: s.e. of McDonald L. , Mission, Range, C.L. Hitchcock 1830,0 (ID, COL, UC); MISSOULA Co.: 1 mile w. of Gordon Pass, Flathead Range, C.L. Hitchcock 18421 (UC, WS) . ALBERTA: Shoulder of Wilcox Mt.,, Jasper Nat. Park, E.H. Moss 4965 (ALTA); above Sunwapta Pass, Jasper Nat. Park, E.H. Moss 4887 (ALTA); near Blakiston Brook, Waterton L. Park, E.H. Moss 10174 (ALTA, CAN); 5 miles w. of Pincher Cr., S . S . Servey,  1193 (ALTA); Upper Spring Cr., between Livingston and Porcupine Range Stn., ca. 30 miles n. of Lindbreck, R.G.H. Cormack ,52 (UAC); Banff v i c i n i t y , E.S. Dowding s..n. (ALTA); Raspberry Ridge., Kananaskis Forest 40. Preserve, B. de Vries 2348-64 (DAO, UAC); Opal H i l l s , n.e. of Rainbow Lodge, Maligne L., L. Jenkins 7903 (DAO); Snow Ck. Pass, 20 miles n. of Banff, J.A. Calder 23976 (DAO); s.w. of d r i l l s i t e , Bighorn Mts., 4 miles n.w. of South Ram R., j e t . with Nordegg - Cochran Forestry Rd., C.D. Bir d  7538 (UAC). Not a l l S_. o c c i d e n t a l i s specimens from northern Montana have t i g h t glomerate heads. The following have open inflorescences: GLACIER Co.: 2.6 miles n. of St. Mary, D. Lynch 6284 (WS); L. St. Mary, L.W. Sayloc 43 (UC); BEZVERHEAD Co.: Top of Odell Pk., Pioneer Range, C.L. Hitchcock and C.V. Muhlick  14925 (WS); MISSOULA Co.: h mile w. of Upper Holland L., Flathead Range, C.L. Hitchcock 18422 (WS); 5 miles above Bonner, C.L. Hitch- cock and C.V. Muhlick 11425 (WS, WTU); DEERLODGE Co.: Dry slope above Storm L., Anaconda Mts., C.L. Hitchcock and C.V. Muhlick 14820 (WS, WTU); LAKE Co.: High above and s.e. of McDonald L., Mission Range, C.L. Hitchcock 18300 (WTU, WS). 41. LITERATURE CITED Beamish, K. I. 1967. A P a c i f i c Coast saxifrage with 10 pairs of chromosomes: Meiosis, development of the female gametophyte, and seed production. Can. J. Bot. 45: 1797-1801. Calder, James A. and Taylor, Roy L. 1968. Saxifraga. In, F l o r a of the  Queen Charlotte Islands. Systematics of the.Vascular Plants. Res. Branch, Canada Dept. A g r i c , Monogr. 4, Part 1. Queen's P r i n t e r , Ottawa, p. 380-389. Engler, A. and Irmscher, E. 1916. Reprint 1958. Saxifragaceae -Saxifraga, Sect. 1 Boraphila Engl. Tn, Das Pflanzenreich 4, 117, H67: 5-89. Flov i c k , K. 1940. Chromosome numbers and polyploidy within the f l o r a of Spitzbergen. Hereditas 2J>: 430-440. Hedberg, 0. 1967. Chromosome numbers of vascular plants from a r c t i c and s u b - a r c t i c North America. Ark. f. Bot. n.s. 6_: 309-326. Henry, J. K. 1915. Saxifraga. In, Flo r a of Southern B r i t i s h Columbia  and Vancouver Island. Gage, Toronto, p. 164-168. Hitchcock, C. L., Cronquist, A., Ownbey, M., and Thompson, J . W. 1961. Saxifraga. In, Vascular Plants of the P a c i f i c Northwest. Part 3. University of Washington Press, Seattle, p. 31-55. Hulten, E. 1968. Saxifraga. In, F l o r a of Alaska and Neighboring T e r r i t o r i e s . Stanford University Press, Stanford, p. 563-583. Johnson, A. M. 1923. A r e v i s i o n of the North American species of the sec t i o n Boraphila Engler of the genus Saxifraga (Tourn.) L. Minn. Stud. B i o l . S c i . 4_: 1-110. Johnson, A. W. and Packer, J. G. 1968. Chromosome numbers i n the f l o r a of Ogotoruk Creek, N.W. Alaska. Bot. Not. 121: 403-456. Jorgensen, C. A., Sorensen, T. H., and Westergaard, M. 1958. The flowering plants of Greenland. A taxonomical and c y t o l o g i c a l survey. • Danske Vidensk. Selsk. B i o l . Skr. 9(4): 1-172. Krause, D. L. and Beamish, K. I. Taxonomy of Saxifraga o c c i d e n t a l i s S. Wats, and S. m a r s h a l l i i Greene. Can. J . Bot. In press. Lbve, A. and Love, D. 1951. Studies on the o r i g i n of the I c e l a n d i c f l o r a . I I . Saxif ragaceae. Svensk Bot. Tidskr. 45_: 368-399. 42. LbVe, A. and Love, D. 1961. Chromosome numbers of c e n t r a l and north-west European Plant species. Opera Botahica 5_: 1-581. Rydberg, P. A. 1917. Saxifragaceae. In, F l o r a of the Rocky Mountains  and Adjacent P l a i n s . Hafner, New York, p.* 376-391. Sav i l e , D. B. 0. 1961. The botany of northwestern Queen Eliz a b e t h Islands. Can. J . Bot. 39: 909-942. Small, J . K. and Rydberg, P. A. 1905. Saxifragaceae. In, North Ameri-can F l o r a 22(2) : 81-158. T A B L E 1. CHROMOSOME COUNTS F R O M BRITISH C O L U M B I A AND A D J A C E N T A R E A S Coll e c t o r C o l l e c t i o n Number Locati o n Chromosome numbers S. n i v a l i s - t e n i u s complex Krause, B e a m i s h and Luitjens Krause, B e a m i s h and Luitjens Krause, B e a m i s h and Luitjens Krause 682000 M i l e 101, Lapie L., Yukon 681807 M i l e 61, Haines Rd., B.C. 681398 S.E. of L a p i e L., Yukon 68 Mt. S.S.W. of A l t u r u s L., Idaho *n = 20 2n = 60 n = 10 n = 10 S. r c f l e x a Krause, B e a m i s h and Luitjens Krause, Beamish and Luitjens Krause, B e a m i s h arid L u i t jens Krause, Beamish and Luitjens Krause, B e a m i s h and Luitjens 681768 Q u i l l C r . w. of Kluane L., Yukon 681768 Q u i l l C r . w. of Kluane L., Yukon 681768 Q u i l l C r . w. of Kluane L., Yukon 682001 Midnight Dome, Dawson City, Yukon 681436 N.E. of F i s h L., Whitehorse, Yukon n = 10 n = 20 2n ca. 30 n = 10 n = 10 Krause and Maze Krause and Maze Kr a u s e Krause Krause Krause. Krause Krause and Beamish Krause and Beamish Krause and Beamish 22 - 71 21 - 71 1 - 71 1 - 70 69000 5 7 - 71 690006 690007 9 - 71 11 - 71 1 2 - 7 1 4 - 71 2 - 71 27 - 70 24 - 70 28 - 70 Sooke, B.C. Mt. Finlayson, B.C. Shawnigan L., B.C. Campbell R i v e r , B.C. 1 rni w. of Cook at Dog Cr., Washington Beaver Cr. F a l l 10 mi e. of Clatskanie, Oregon n = 10 n = 10 n = 10 *n = 29 *n = 19 "n = 19 Viento State Park, 8 mi w. of Hood R., Oregon n = 19 W. of A i n s w o r t h State P a r k , near Oeneonta Gorge, Oregon n = 10 Elk Rock, Willamette R., Oregon n = 10 B i g C l i f f Dam, near Detroit, Oregon n = 10 5 mi e. of S. Santiam R., Linn Co., Oregon n - 10 Mosier, Oregon n = ca. 28 1 mi w. of Clark-Skamania Co. line, Columbia R. Gorge, Washington n = 10 The Dalles, Oregon n » 28 Troutdale, Oregon ' n - 10 White Salmon, Washington n - ca. 28 Gh r oino some numbers in bold fact! indicate new count!! fur the taxon. 44. Figure 1. A-D. The black l i n e represents 2u. Fig. 1A. Saxifraga nivalis-tenuis complex, n = 20, Mile 101, Lapie Ls., Yukon, Krause, Beamish and Luitjens  682000; Fig. IB. S. ruf idula, n = 19, 1 mi w. of Cook at Dog Cr., Washington, Krause & Maze 690005; Fig. IC. J3. rufidula, n = 28, The Dalles, Oregon, Krause & Beamish 27-70; Fig. ID. S. rufidula, n = 29, Campbell River, B.C., Krause 1-70; Fig. IE. Inter-pretive drawing of Fig. 1A; Fig. IF. Interpretive drawing of Fig. IC. 0 • O » V © . © © • . # occidentalis, j5. reflexa and j>. rufidula in British Columbia and some adjacent areas, derived from field work and herbarium specimens. I f I • Saxifraga rhomboid ea • S. occidentalis I i \ I I." Figure 3. D i s t r i b u t i o n of Saxifraga rhomboidea and S_. occidentalis i n B r i t i s h Columbia and adjacent areas, derived from f i e l d work and herbarium specimens. 

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}]}"
                            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:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0101302/manifest

Comment

Related Items