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Studies in the genus DODECATHEON of North-western America, with some reference to its use in floriculture Beamish, Katherine Isabel 1951

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STUDIES IN TBS GENUS DODEGATfiEON OF NORTHWESTERN AMERICA with, some reference to its use in floriculture by KATHERMS ISABEL BEAMISH A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF Master of SCIENCE IN AGRICULTURE in the Department of Biology and Botany We accept this thesis as conforming to the standard required from candidates for the degree of MASTER OF Members of the Department of THE UNIVERSITY OF BRITISH COLUMBIA April, 1951 Abstract A study of the genus Dodecatheon has been made with two pur-poses in view: 1* A reclassification, of the genus in northwestern America, in-cluding Alaska, Yukon, British Columbia, Alberta, Saskatchewan, Washing-ton, Oregon, Idaho and Montana. 2. An incidental survey of its horticultural possibilities. The method of approach has included collection of living mater-i a l , hybridization, cytological study, and examination of seven to eight hundred herbarium specimens. Throughout the progress of this work, a number of horticulturally valuable characteristics have been observed and noted. Chromosome counts have been obtained for a number of species and from a number of localities. These counts indicate an interesting pattern of polyploidy within the genus: diploids on the east of the Cas-cade Mountains, polyploids extending north to Alaska along the Pacific Coast. Similarity of diploids and polyploids suggest autoploidy. As a result of the work outlined above, the genus has been re-classified into ten species and one variety on the basis of morphology, cytology, and distribution. Though a number of these species, two part-icularly, are variable, further subdivision is considered unwise until much more can be learned about the cytogenetics of the genus. Finally, the suggestion is made that cytogenetic study is the next step in horticultural improvement and might provide valuable evi-dence regarding the course of evolution in the genus. ACKNCILDEGEMENTS S i n c e r e g r a t i t u d e i s e x t e n d e d t o D r . V . C . B r i n k and D r . T . M . O . T a y l o r f o r t h e i r h e l p f u l g u i d a n c e a n d e n c o u r a g e m e n t i n t h i s s t u d y . O t h e r s , t o o , h a v e g i v e n v a l u a b l e a s s i s t a n c e i n many p h a s e s o f t h e p r o j e c t , a n d , t h o u g h m e n t i o n o f e a c h one i s i m p o s s -i b l e , c o - o p e r a t i o n o f a l l i s g r a t e f u l l y a c k n o w l e d g e d . S p e c i a l m e n -t i o n s h o u l d be made o f l i b r a r i e s and h e r b a r i a w h i c h h a v e made a v a i l -a b l e v a l u a b l e b o o k s a n d p r e s s e d s p e c i m e n s f o r s t u d y . A f i n a l w o r d o f a p p r e c i a t i o n goes t o t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a i n r e c o g n i t i o n o f t h e g r a n t s w h i c h have made p o s s -i b l e two summers o f r e s e a r c h o n D o d e c a t h e o n . TABLE OS" CONTENTS Page I I n t r o d u c t i o n A . The p r o b l e m . 1 B . The f a m i l y and g e n u s , a g e n e r a l s u r v e y e 2 I I E x p e r i m e n t a l w o r k a n d o b s e r v a t i o n s . . . . . < > 5 A . A b b r e v i a t e d l i s t o f synonyms . . . . . . . . . . . . . 5 B . C r o s s i n g e x p e r i m e n t s . . . . . . . . . . . 6 1 . p u r p o s e s . . . . . . . . . . . . . . . . 5 2 . t e c h n i q u e s 7 3 . v i a b i l i t y t e s t 8 4 . r e s u l t s . « 9 C . C y t o l o g i o a l s t u d y . . . . . . . . . . . . 1 1 1 . t e c h n i q u e s . . . . . . . . . . . . . . . . . . . . 1 1 2 . chromosome c o u n t s . . . . . . . . . . . . . . . . 1 2 I I I ' T a x o n o m i c t r e a t m e n t . . . . » 1 3 A . D i s c u s s i o n o f s p e e l a t i o n . . . . . . . . . . . . . . . 1 4 B . D e f i n i t i o n o f t a x o n o m i c c a t e g o r i e s • • • 22 C . H i s t o r y o f t h e t a x o n o m i o t r e a t m e n t 2 3 D . D i a g n o s t i c c h a r a c t e r s . £4 E . H e r b a r i a c o n s u l t e d • • • • • . . . . . . 28 F . C l a s s i f i c a t i o n 28 1 . D o d e c a t h e o n , t h e genus . . . . . . . 28 2 . K e y t o t h e s p e c i e s 50 I l l T a x o n o m i c t r e a t m e n t ( c o n t ' d ) Page 3 . D o d e o a t h e o n f r i g i d u n u . . . . . . . . . . . . . . 3E 4 . D o d e o a t h e o n d e n t a t u m . . . . . . . . . . . . . . 35 5 . D o d e o a t h e o n J e f f r e y ! 37 6 . D o d e o a t h e o n a l p i n u m . . . . . 45 7 . D i s c u s s i o n o f D . f r i g i d u m , d e n t a t u m , J e f f r e y i a n d a l p i n u m . . . . • . . . . . . » • 47 8 . D o d e o a t h e o n p o e t i o u m . . . . . . . 5 2 9 . D i s c u s s i o n o f D . p o e t i c m . . . . . . . . . . . . 5 3 1 0 . D o d e o a t h e o n H e n d e r s o n ! . . . . . . . . . . . . . 54 1 1 . D i s c u s s i o n o f D . H e n d e r s o n ! . . . . . . . . . . . 58 1 2 . D o d e o a t h e o n c o n j u g e n s . . . . . . . . . . . . . . 60 1 3 . D o d e o a t h e o n co lumn i a n u m 65 1 4 . D i s c u s s i o n o f D . c o n j u g e n s a n d D . oo lumbianum . . 67 1 5 . D o d e o a t h e o n C u s i c k i i . . . . 68 1 6 . D o d e o a t h e o n p a u c i f l o r u m • 74 1 7 . K e y t o the v a r i e t i e s o f D . p a u c i f l o r u m . . . . . 75 1 8 . D o d e o a t h e o n p a u o i f l o r u m v a r . t y p i c u m . . . . . . 76 1 9 . D o d e o a t h e o n p a u c i f l o r u m v a r . m a c r o c a r p u a . . . . 8 2 2 0 . D i s c u s s i o n o f D . O u a i c k i i , D . p a u c i f l o r u m a n d D . p a u c i f l o r u m v a r . maorooarpum • • 8 6 I V C o n c l u s i o n 91 7 A p p e n d i x I • * 96 V I B i b l i o g r a p h y . « 1 0 0 i INDEX OF S P E C I E S Page 1* D o d e o a t h e o n a l p i n u m ( G r a y ) Greene . . . . . . . . . . . . 45 2 . c o l u m b i a n u m n . s p . . . . . . . . . . . . . 65 3 . Greene 60 4 . v a r . l e p t o p h y l l u m ( S u k s d . ) P i p e r 60 5 . v a r . v i s o i d u m ( P i p e r ) Mason e x S t . J o h n . • . . • 60 6 . c rena tu ta Greene . . . . . . • • . . • » • < > » 37 7 . C u a i c k i i Greene . . . . . . . . . . . . 68 8 . v a r . a l b u m S u k s d 68 9 . d e n t a t u m Hook 35 1 0 . e x i l i f o l i u m M a c b r i d e a n d P a y s o n . . . . . . . . . . . 38 1 1 . f r i g i d u m Cham, a n d S o h l e c h t 32 1 2 . v a r . d e n t a t u m G r a y • . » . e . . . . . . . . . e 35 1 3 . H e n d e r s o n ! G r a y 54 1 4 . v a r . l e p t o p h y l l u m S u k s d . . . . . . . . . . . . . 60 1 5 . i n t e g r i f o l i u m M i o h x v a r . l a t i f o l i u m Hoofc . . . . . . 54 1 6 . v a r . v u l g a r e Hook 76 1 7 . J e f f r e y ! M o o r e e x V a n H o u t t e 37 1 8 . v a r . a l p i n u m Gray . . . . . . . . . . . . . . o 45 1 9 . v a r . t e t r a n drum ( S u k s d . ) J e p s o n . . . . . . . . . 38 2 0 . l a t i f o l i u m (Hook) P i p e r . . . . . . « . » 55 2 1 . l a t i l o b u m ( G r a y ) E l m e r e x P a x a n d K n u t h • 35 8 2 . m a c r o c a r p u m ( G r a y ) K n u t h . . . . . . . . . . . » 0 . . 82 i i Pag© 2 3 . m a c r o c a r p u m v a r . a l a a k a n u m H u l t . . . . . . . . . . . 8 2 2 4 . M e a d i a L . v a r . a l p i n u m G r a y . . . . . . . . . . . . . . • 45 2 5 . v a r . b r e v i f o l i u m G r a y « 55 2 6 . v a r . f r i g i d u m Hook 32 2 7 . v a r . I a n c i f o l i u m G r a y 37 2 8 . v a r . l a t i l o b u m G r a y . . . . . . . . . . . . . . 35 2 9 . v a r . m a c r o o a r p u m G r a y • 8 2 3 0 . v a r . p a u c i f l o r a . . . • 76 3 1 . v a r . p u b e r u l a . . . » 68 3 2 . p a u c i f l o r u m ( D u r . ) G r e e n . . . . . . . . . . . . . . 74 3 3 . v a r . C u a i c k i i (Greene) M a s o n e x S t . J o h n . . . . 68 3 4 . v a r . C u s i c k i i (Greene) M a s o n f . a l b u m ( S u k s d . ) S t . J o h n . . . . , 69 3 5 . v a r . macro carpum ( G r a y ) n . comb. . . . . . . . . 8 2 3 6 . v a r . monanthum G r e e n e . . a . . . . . . . . . . . 76 3 7 . s u b s p . monanthum (Greene ) K n u t h . . . . . . . . . 76 3 8 . s u b s p . s a l i n u m ( N e l s . ) K n u t h . 5 • . . . . . . . 0 76 3 9 . v a r . s h o s h o n e n s i s N e l s . . . . . . . . . . . . . 76 4 0 . v a r . t y p i c u m n . n . . . . . . . . . . . . . . . . . 76 4 1 . p h i l o s o l a H e l s 76 4 2 . p o e t i c u m H a n d . . . . . . . . . . . . . . 52 4 3 . p u b e r u l e n t u m H e l l e r . . . . . . . . . . . . . . . . . 68 4 4 . p u b e r u l u m ( N u t t . ) P i p e r 68 i i i Page 4 5 . s a l Intra N e l s » . . . , . . 76 4 6 . t e t r a i l drum S u k s d . . . . . . . . . 38 4 7 . u n i f l o r u m R y d b . « . . . 76 4 8 . v i s o i d u m P i p e r . . . . . . 60 4 9 . v i v l p a r u m ( G r e e n e ) 37 5 0 . v u l g a r e (Hook) P i p e r 76 STUDIES IN THE GENUS DODECATHEON OF NORTHWESTERN AMERICA with some reference to its use in floriculture INTRODUCTION Many gardeners tend toward the belief that ornamentals, to be of much horticultural value, must be Imported, while members of our native flora, as beautiful as the exotics and often having the advantage of suitability to our sail and climate, they ignore as being "Just wild flowers." Among the more promising of such native plants are species of the genus Dodeoatheon, commonly known as shooting stars. Their gay colours, graceful habit of growth, and suitability to a variety of locations recommend them for a place among spring flowers In the gardens of the Middle West and Yfost Coast. To learn more of the horticultural possibilities of this genus, study was begun at the University of British Columbia In 1948. Three obvious approaches to the problem were: 1. a survey of the species, their ecological preferences, and their range of morphological variability. 2. cytological study to determine chromosome numbers within the genus. A knowledge of the presence of polyploidy would indicate the possibility of hybridization. It would also provide a basis for under-standing the inheritance patterns of genetic characters, patterns which are quite different in a polyploid from those in a diploid. 2 3 . hybridization to combine the desirable features of more than one species in a single plant. One of the great difficulties in carrying out such a study has been the lack of material in the university botanical garden. Only two species, both from southern Vancouver Island, were available at the outset. A l l others have had to be collected at the expense of considerable time, money, and effort from their native habitats, from co-operative gardeners or botanical gardens, or, in a few cases, from American commercial growers dealing in so-called "wild flowers." A second difficulty, even less easily overcome, arose when collections began* Attempts to identify the collections revealed among the western species a state of taxonamic confusion which prevented defin-ite classification of most of the plant material* Very few published works have done more than describe new species or provide a key to a small or local group. As a basis to any horticultural improvement, a thorough knowledge of the genus appeared necessary. Horticultural work was there-fore laid aside for the future while the problem became a study of speciation within the genus and a reclassification of the present north-western species based on cytology, morphology, and distribution. In secur-ing the necessary material for such a study, the willing coroperation of many individuals and institutions was obtained and is gratefully acknow-ledged; Both living material and herbarium specimens have been provided from many parts of the region covered. Dodeoatheon belongs to the family Primulaceae of which several 3 * other genera include horticulturally important species and varieties. Notable among such genera are Primula and Cyclamen. In general appear-ance, the latter, with its re flexed corolla, most closely resembles Dodecatheon, differing in that i t produces one comparatively large bloom on each scape whereas Dodecatheon produces an umbel of one to more than twenty smaller flowers. The species are al l perennial and, generally speaking, spring-blooming, though their actual month of flowering, March to August, depends on the advent of spring in a particular location. After bloom, many completely disappear from above ground while others retain their green leaves t i l l autumn. Propagation may be either vegetative or by seed. By reason of their type of root, discussed more fully later, single plants rapidly become clumps. Seeds of most species germinate easily and in good per-centages outdoors, but require at least two years to bloom. Dodecatheon appears to be essentially a North American genus. One species has been recorded (Gray 1886b.Hulten 1948, Anderson 1949) from the Asiatic Coast. The same or other species range from Alaska to California along both the Pacific Coast and the Cordilleran System. East of the Rocky Mountains, representatives are found in Alberta and Saskat-chewan, across United States to the east coast, and south to Texas. The species east of the 100th meridian have been dealt with systematically by Fassett (1944). The present paper is concerned with those of the Northwest found in Alaska, Yukon, British Columbia, Alberta, Saskatchewan, Washington, Oregon, Idaho, and Montana. From the physical nature of the Northwest, distributions are more or less discontinuous and related to the mountain barriers and climatic variation* For example, species of the Pacific Coast are general-ly distinct, at least genetically, from those of the dry Interior Plateaux, and the latter often, though less consistently, differ again from those of the Eastern Cordilleran Mountains* Where two species inhabit the same general area, separation is usually effected by altitude or moisture pre-ferences or, as in the case of two overlapping species on Vancouver Is-land, by physiological differences which vary the time of maturity. In certain sections, however, especially of the Interior Plateaux, hybridiza-tion appears to have occurred. Habitat varies from sea level to at least ten thousand feet altitude and from situations of extreme moisture throughout the year to those of extreme drought after blooming* Species of upper elevations are generally to be found along streams in highly organic soil, often with their roots in water. Coast species, those of the dry inter-mountain area and of the prairies grow and bloom in wet meadows or otherwise most loca-tions, but are subject to extreme drying later in the summer. Dodeoatheon seldom grows in other than well-drained soil or fresh water, but a prairie form of one species frequents saline sloughs and a coast form of the same species is found in the brackish water of tidal flats. Light preferences vary from ful l sun in most cases to the partial shade of open woods. More detailed discussion of both distribution and habitat appears with the taxonomic treatment of the genus. 5 EXPERIMENTAL WOIK AND OBSERVATIONS The ultimate object of this study, then, is the introduction of Dodeoatheon as an ornamental, but the immediate problem, as previously stated, is the reclassification, based on morphology, cytology, and distribution, of the genus as i t occurs in the Northwest, emphasizing always any information of possible use in horticultural improvement. Part of the work has been done during winter sessions but a great deal of field and laboratory study has been made possible by the generosity of the University of British Columbia in providing grants for two summers* re-search. The classification which has resulted from this study is given in detail later, but before proceeding with an account of the work done, a l i s t of the specific names accepted and their more commonly used synonyms seems desirable* The l i s t is as follows: 1* Dodeoatheon frigidum Cham. & Schlecht. 2. D. dentatum Hook. 3 . D. Jeffrey! Moore ex Van Houtte D. vlviparum Greene D. tetrandrum Suks. 4. D. alpinum Greene. 5. D. poeticim Hend. 6 . D. Henderson! Gray. 7. D. latifolium (Hook.) Piper 6 » 7. D. conjugens Greene. D. cylindrocarpum Gray. D. T i a c i d u m Piper 8. D. columbianum n«sp. 9. D. Cusickii Greene D. puberulum (Nutt.) Piper 10* D» pauciflorum (Dur.) Greene. D. vulgare Hook. As an indication of field study, collections made personally, either, alone or with a party, have been included under the author's name in the complete l i s t , Appendix 1* Unfortunately, as such study has been confined to southwestern and south central British Columbia and to one location in northern Washington, not a l l species have been observed in their native habitats. However, living material of most species has been collected and is now growing in the university botanical garden. Of three, D. conjugens, D. poetlcum, and D. alplntim, only pressed specimens have been available. 1. Crossing Experiments The live material has been brought to the university during three summers and was either planted directly outdoors or moved out before any experimental work was begun. In the first sunuer, 1949, sufficient collections were secured that the rollowing year some crosses could be undertaken. The hope was that such crosses would serve three purposes : 1. that they would indicate relationships between the various groups collected. 7 2. that selfs and intraspecific crosses would show the presence or absence of incompatibility factors within the species. This informa-tion would have an Important bearing on the methods used in horticultural Improvement. 3 . that interspecific crosses would indicate the possibilities in hybridization and provide some hybrid seed to be used as a basis for later work* Plants from such seed might yield information regarding the inher-itance of certain characteristics or, through their meiotic behaviour, might give a clearer picture of relationships, as here structural differ-ences should show up as well as differences in chromosome numbers. In the hope of accomplishing the three purposes stated above, a l l types of pollinations were made: selfs, crosses within collections, crosses between collections of possibly the same species, and crosses be-tween obviously different species. It might be noted here that in both crossing and cytological study, D. Meadia and D. Clevedlandi have been included,; not because they belong to the Northwest, but because they show particular promise horti-culturally* The former is a native of eastern United States, the latter of California* Both were obtained as plants from a California nursery* They are strong-growing and free-flowering species possessing, besides these two characteristics, a number of others which would be extremely valuable in a breeding programme. The technique used in pollination was determined by observation of the earliest blooms* Dodeoatheon is apparently a protogynous genus as the stigma and style often protrude from the staminal column before the cololla is re flexed and certainly before authesis. In pollinating, there-8 fore, flowers were emasculated before they opened and stigmas protected both before and after pollination by bagging* In every case an entire umbel was pollinated, as its flowers matured, from the same plant so that one bag could be used to cover the umbel* Seed was harvested when the capsules began to open* Results of pollination experiments are shown in Table 1. As a test of viability, seeds were planted from each cross which had set seed, as well as from selfed and open-pollinated flowers* Prev-iously, in obtaining root tips for cytological work, some difficulty had been experienced with germination, but this difficulty had been largely overcome by pre -chilling the seeds* Therefore, in the viability tests, one series of seeds was chilled for three weeks in moist petrie plates in o a 5 C. cold chamber, then planted in flats in the greenhouse* A duplicate series was planted directly into pots and put in a cold frame at the end of November to chill naturally until spring* Results of the viability tests appear in Table 2 * Comparison of the two tables shows that germination closely parallels seed-set. Open pollinations, selfs, and intraspecific crosses produced, in most cases, much seed which geminated in average to high percentages. Interspecific crosses produced no seed or a few seeds which germinated in low percentages or not at a l l . A number of desirable crosses had to be emitted or were lost. Omissions were for two reasons* First, there were differences in blooming time of the various species* To some degree these differences were no doubt, genetic, but they were also due in part to differences in environ-0 8 A TABLE 1 C; 1 6 8 Made. Se l f . 49- 49* 49* 49* 49 49., Open ed -3 100 101 102 » S 277 49 • 49 49 49» 50* 49 103 *12 #10 105 106 . 4 Source of Plant* 1 . Clevelandi. L*.%. C * l . 3 3 3 0 0 loat 0 0 0 0 © California 2. D. Hendersoni. 49*100. ¥ . T . 2 2 0 2 0 1 Vic in i ty of Victor ia Vancouver Island 3 . B, Baueiflerum. A9-101, V . I - 3 2 0 0 3 1 0 1 0 0 Vic in i ty of Victor ia Vancouver Island 4* naueiflorum. A9*102r L.T- 2 loat 0 2 3 3 leet loat Lulu Ialand, near Vancouver 5. J»t. Dauciflorum. L9*&. Alaaka lest 0 1 lest Bklutna Plata, near Anchorage. Alaska 6. Pf naueifloraa. A9«277.Ht- Mt. Arxfevemith, Van* oouver Ialand 7. £*. Meadia. 4 9 * 2 . Gal 3 3 0 3 1 California 8. PA. Cuaiekii . 49*103. Bot.V- 2 1 1 0 0 Botanic Valley, near Lytton, B«C # 9- feu Cuaicki i . 49*12. Clinton 2 2 Cairn Mt», near Clinton, B«C« 1 0 . Dt dentatum. A9»10. Pentleten 1 1 0 V ic in i ty of Pentiet* on, B.C. l i e *>.*.  Jeffrevi.49*105. Stevens P- 1 0 Stevene Paas, tfaeh. 1 2 . B t Jeffrey!. 49*106.Stevens P- 1 Stevena Paas, Waeh. 13* D t Jef frevi . 4 9 * 4 . P.R. 3 Prince Bupert, B.C, Notes Seed set, as determined by inspection of opflapsulee: 0 « none 2 * f« • 25 seeds 1 - poor, 1 * 10 aeeda 3 * g< s e « d » numerous * 8 B • Parent 1. »1 . Clevelandi. Cal . 2. B. Ca l . 3. D* Cleveland!. Ca l . 4. »t Cleveland!. Cal . 5. D t Cleveland!. Cal . 6. D, Headerseal. V . I . 7. Bt Henderson!. T . I . 8. fit Henderson!. T . I . f . C. Henderson!. T . I . 10. B t Pauciflorum. T . l . 11. D T Dane!florum. T . I . 12. D, p a u c i f l o r u m , T . I . 13. D. pauciflorum. T . I . 14. D. paud florum. T . I . 15. D. pauciflorum. T . I . 16. D, pauciflorum. L . I . 17. D f pauciflorum. L . I . 18. Dt paueiflariimr L . I . 19. D. pauciflorum. L . I , 20. D. pauciflorum. L . I . 21. D, pauciflorum. L . I . 22. D. Headia. Cal . 23. »• Meadia. Cal . 24. Dt Headia. Ca l . 25. Dt Meadia. Cal . 26. Dt, Meadia. Cal . 27. * i . Cus icki i . Bot i . T . 28. D,t„ Cusicki i . Bot . T . 29. D t Cusicki i . Bot.V. 30, B. Cusieki i . Bot.V. 31. D, Cueiekii . Bot • T . 32. D, Cusieki i . Bot . T . 33. D,t Jeffrevi . Stevens Parent B. Cleveland!. Cal. D. Meadia. Gal. selfed D. Henderson!. T . I , open poll ination open p o l l i n a t i o n selfed D. Henderson!. V . I . P. Cusieki i . Bot.V. epen poll ination D. Pauciflorum. V . I . D. pauelfloraa. Mt.A. D. Meadia. Cal . B. pauciflorum. V . I , s e l fed Do Meadia. Cal. B. pauciflorum. V . I , B. Meadia. Cal . open poll ination D. pauciflorum. Alaska B. pane!f lorurn. L . I , B. Meadia. Cal. D. pauciflorum. V , I . selfed open poll ination D. ntueiflorua. Alaska B. Henderson!. V . I , B. Cleveland!. Cal . B. Henderson!. V . I . open poll ination open poll ination selfed P. selfed TABLE 2 • Tiafel Test Seed Seeds per Germir % Set t r i a l at! oi srmin« (Greenhouse) it ion 3 5© 48 f6 3 5© 45 9© 3 5© 38 7© 0 2 0 0 3 5© 25 5© 2 15 1 6.6 2 25 1 4 2 25 0 0 1 2 0 © 3 5© 7 14 1 5 © 0 1 0 0 0 2 25 0 0 3 5© 0 0 2 45 © 0 3 5© 18 36 2 17 © © 3 1© 1 10 2 5© 5 1© 3 50 13 26 3 5© 5 10 2 20 10 50 3 5D 2 4 3 50 12 24 3 50 31 62 1 25 1 4 1 8 0 © 1 0 © © 1 5 © 0 2 10 © 0 2 1© © © 2 10 0 0 1 10 0 6 Seeds per Germin* % t r i a l ation Germina (garden) tion 50 Good *> 50 Good -50 Good -2 1 50 50 Good • 20 17 85 25 15 60 25 16 64 2 © 0 50 32 64 5 0 0 1 0 0 25 7 28 50 42 84 45 32 71 5© 21 42 17 6 35 10 7 70 50 38 76 5© 25 50 50 30 m 20 8 4© 50 17 34 5© 36 72 50 10 20 25 4 16 8 1 12,5 1 0 © 5 0 0 10 1 1© 10 © 0 10 1 1© 10 0 0 meat. For instance, species from wet habitats, such as D. Jeffrey! and D. dentatum, had been planted in a wetter part of the garden and therefore developed more slowly in the spring* Second, certain collections bloomed very sparingly or not at a l l . Among these were D. pauciflorum from Mon-tana and D. pauciflorum var. macrocarpum from Mt. Arrowsmith, Vancouver Island. As the scarcity of bloom was in collections from high elevations, the change in altitude may have produced the adverse effect. Losses were due to damage by slugs or wind. Control of slugs proved difficult, especially in the wetter location, in spite of constant baiting. Staking in most cases protected the scapes from wind, but two breaks occurred. In regard to the three aims previously outlined, the crossing ex-periments were in some respects indicative, in others inconclusive. 1. The lack of seed-set between entities here accepted as species might be considered as suggestive of genetic distinction. Such a conclu-sion, however, must be drawn with definite reservations since a number of other agents may have prevented seed-set. Moreover, crosses between collections here accepted as one species were not successful in a l l cases, a fact which detracts from the value of the above evidence. The group of four collections designated as D. pauciflorum var. macro carpum serves as an illustration. As shown in Table 1, none of these, crossed with other species, set any amount of seed other than with D. Meadia. On the other hand, among themselves seed-set was somewhat better, though even here, not good in a l l cases. 10 «• 2. The success of the selfs and crosses within collections indi-cate an absence, at least within the species used, of incompatibility factors. 3 . Very l i t t l e hybrid seed was obtained* This, too, is negative evidence but may indicate that hybridization within the genus will not occur readily. Whether or not such is the case can be proved only by further experimentation* At any rate, l i t t l e hybrid seed is at present available to provide plants for later study. Three other points in the results of the crossing experiments merit special notice: 1* Seed from crosses involving D. Cleveland!, the California species, germinated in high percentages and very quickly. In the garden series, they germinated before Christmas and were, unfortunately, frozen before a definite count was taken. 2. D. Meadla, though tetraploid, crossed freely with most species, whether used as male or female parent* Hybrid seed from these crosses germinated in al l cases, though sometimes in small percentages. 3 . D. Cuslckii from the dry Interior Plateaux set seed poorly and what l i t t l e seed was produced germinated practically not at a l l * On the whole then, this part of the experimental work can be used only as an indication of the desired information* A'mo re extensive crossing programme would have to be carried out before results could be called conclusive. 11 II. Oytologlcal Study Material used for cytological study represents a variety of loca-tions in the Northwest. From this material, chromosome counts have been made for many, though not a l l , species, some from several populations. Two types of material have proved satisfactory. Sporogenesis gave most accurate counts, preferably dlakinesis, but occasionally first anaphase. In some oases, however, where flowers were not available, root tips were substituted. By 2 hours' pre fixation treatment of these tips in paradichlorobenzene as outlined by Meyer (1945), mitotic chromosomes are contracted sufficiently to give closely approximate counts. In a l l cases aceto-carmine squashes were used, made from mater-ial fixed in acetic-alcohol (3:1). Added iron in the form of ferric chloride intensified the stain. Boot tip squashes were used as temporary mounts, but anther squashes have been made permanent according to "the technique of Bradley (1948) with the following modification: to avoid darkening of the cytoplasm during dehydration in a vapour jar, dehydration was carried out by drawing 95%, then absolute alcohol under the cover slip until a l l surplus stain had been removed. This process required only 5-10 minutes after which diaphane was put around the cover slip and allowed to infiltrate in a vapour jar. Chromosome counts obtained are listed below, those from meiotic divisions as Mn w, from mitotic as n2n H. Diploid meiotic counts are con-sidered accurate. Polyploid meiotic and a l l somatic counts are considered a minimum and, except where otherwise indicated, within 2 of accuracy. D* dentatum from Penticton, B.G ....2n « 44 D. Jeffrey! 1* seed from California 2n - 44 2. from Stevens Pass. Washington ...... n - 22 3. from Forbidden Plateau, Vancouver Island • 2n - 88 4. from Prince Bupert, B. C 2n 88 D. Henderson! 1. seed from California diploid 2. from Vancouver Island .............. n - 44 - 50 D. columblanum from Natal B.C.. n - 22 D. Cuaickii from Princeton, B.C....... n - 23 or 24 D. pauciflorum 1* from Martlach, Saak n - 22 2. from Fairmont Springs, B.C 2n «* 44 3. from Montana diploid D. pauciflorum var. macro carpum 1. from Vancouver Island at least tetraploid 2. from Lulu Island near Vancouver .... n «- 44 3. from Eklutna Flats, Anchorage, Alaska tetraploid Species from outside the area studied: D. Meadia, plants from California n * 46 D. Clevelandi, plants from California n «» 23 The small size and large numbers of chromosomes have made accur-acy difficult to obtain in anything but the most favorable material, usually diakinetic figures of meiosis. In four instances no actual draw-- 1 3 ings could be made but from an approximate count of chromosome figures seen, D. Hendersoni from California and D. pauciflorum from Montana are almost certainly diploid, D. pauciflorum var, macro carpum from Eklutna t Elats tetraploid, and D, pauciflorum from southern Vancouver Island at least tetraploid, possibly hexaploid. Small size has also made chromosome morphology and behaviour difficult to study. In somatic material, of course, distinctive morphol-ogy was lost in the excessive and unnatural shortening and thickening pro-duced by paradichlorobenzene• One general observation concerning the polyploids should be made. These polyploids, though they differ l i t t l e morphologically from their diploid counterparts, show no definite evidence of multivalent pairing unless, perhaps, in D, Hendersoni from Vancouver Island, This lack of multivalents -correlates with a lack of noticeable sterile pollen in the many anthers seen under the microscope. Where further cytological information has been obtained concern-ing individual species, i t appears with the taxonomic treatment of that species, III, Taxonomic Treatment Before presenting a classification of the genus Dodeoatheon some definition is necessary of the taxonomic categories into which the genus has been divided* Also a discussion would be helpful of the process-es by which the plant groups represented by these categories, arise. The latter is treated first as the definitions are based on this discussion. 14 • A. Speelation Four processes are generally accepted as Important agents in speelation (Dobzhansky, 1941} Huxley, 1940): mutation, recombination, selection, and isolation* Mutation provides new variability within a pop-ulation* Recombination through sexual reproduction, either alone or accompanied by hybridization, produces newly-combined genotypes which may or may not be able to establish themselves* Selection eliminates the un-successful variants* Successful variants may behave in either of two ways. By backcrossing they may merely spread their variation through the popula-tion or, i f isolation prevents backcrossing, they may accumulate differ-ences until they become varieties or, in time perhaps, new species. Further consideration of mutation makes clearer the source of variability* Taken in the broad sense, mutations are of three types: 1* gene mutations, that is, changes in the chemical make-up or molecular arrangement of the individual genes. 2* chromosomal rearrangements and loss or gain of chromosome parts or whole chromosomes* These include translocations, inversions, defic-iencies, duplications, etc. 3. polyploidy, that is, the duplication of whole sets of chromo-somes* Such duplication may be autoploidy or, i f combined with hybridizaw tion, alloploldy. Within a genus any one or any combination of these three types of mutation may be at work in speelation. The significance of the first two may be illustrated by examples from genera which have been thoroughly - 15 studied. The evolutionary value of the third is s t i l l a matter of debate and. therefore, will be considered in more detail. 1. Gene mutation is considered by Bab cock (1947) as one of two major genetic evolutionary processes in Grepis, producing three principal effects: a. morphological and physiological differentiation. b. the accumulation of intersterility in isolated populations. c. reduction in chromosome size. Harlsnd (1933) and Silow (1944) conclude that speciation in Gossypium is also brought about by gene mutation, principally in modifier complexes. 2. More recently Stebbins (1945) points out that genie changes could, in many cases, be indistinguishable from what he terms "cryptic structural differentiation". By this term he means structural dianges, (translocations* inversions, etc.)., so small as to be unrecognizable cytologically, which behave as mendelian units in fertile individuals. Stephens (1950) has applied this concept, rather than gene mutation, as the major explanation of breeding phenomena observed in Gossypium, and... considers gene changes to play a minor part. Babcock (1947) places structural changes beside genie changes as primary agents of speciation in Crepis. Speaking of translocations sufficiently large to cause meiotic abnormalities, he assigns them two roles: a. production of intersterility allowing accumulation of gene mutations. b. karyotype evolution in the form of reduction in chromo-some numbers, reduction in chromosome length, and increase in asymmetry. 16 3* Polyploidy and its significance in evolution seem best discussed with hybridization as the two so often operate together. Difficulties in the discussion arise from difficulties in definition of the terms. Polyploids are of two types, autoploids and allopoids, (or amphiploids)• The former are produced by the multiplication of a single genom and the latter by multiplication of two different genams. However, as pointed out by Stebbins (1947), a l l degrees of difference exist between genoms, from those producing variation within species to those producing distinct species and genera. By the above definition alone, most polyploids would be alloploids, unless, as Stebbins says, they were produced by somatic doubling of a homozygous3 diploid. The same problem arises in defining a hybrid* According to Darlington (1937), a hybrid is "a zygote produced by union of dissimilar gametes." By this standard most plants are hybrids, even those within species. Before proceeding further, then, the usage of the terms auto-ploid, alloploid, and hybrid must be made clear. Clausen, Keck, and Hiesey (1945) have somewhat clarified the polyploid situation and their classification has been accepted, with some modification, as the basis of Stebbins1 (1947) review of polyploids* In-stead of the usual systematic categories they use what what they term *biosystematic units* on which they base their classification of poly-ploids* Their biosystematic units are. a. ecotypesj ecologic races of one species which are capable of interbreeding freely i f they meet. b. ecospecies: species whose internal and external balance are so *• 17 different that exchange of genes is limited even where their required environments overlap* c* cenospecies: species unable to exchange genes except to pro-duce sterile hybrids* d* comparium: a group of cenospecies s t i l l able to produce sterile hybrids. Polyploids, then, are classified by Clausen et a l . according to their origin from these biosystematic units* Autoploids may either spring spontaneously from nonhybrid individuals of a given species, or arise from intra-ecospecific hybrids, which, of course, are fully fertile. Amphiploids, on the other hand, are derived only from inter-ecospecific and inter-cenospecific hybrids, and consequently the Fj_ hybrids giving rise to them are either partially or wholly - sterile. This statement means, in effect: a* Autoploids are derived from fertile individuals produced by crosses below the species level, though there may be minor differences in genoms. b. A l l opioids are derived from more or less sterile individuals produced by crosses between entities of species rank or higher. These two concepts, then, are here accepted as defining auto-ploid and allopioid. Heiser (1949), in his review of natural hybrids, limits his dis-cussion to the products of crosses between species* This, in the classi-fication of polyploids, would make hybrids,alloploids. It is in this sense then, as interspecific crosses, that the term 'hybrid* is here used. - 18 -With definition of the terms, at least as used in this paper, established, further discussion of polyploids is possible. Two considera-tions are important, their behaviour during meiosis and their signific-ance in evolution. a* The behaviour of polyploids during meiosis depends upon the type of polyploid. Allopioids are usually fertile by reason of preferential pairing between like genoms. Autoploids, on the other hand, have more than two chromosomes of one kind and "this circumstance often leads to formation of multivalents followed by unequal distribution of chromosomes to the poles, which in turn produces varying degrees of ster i l i t y 0 That large numbers of such multivalents are not necessarily the rule however, has been shown by Upcott (1935) in her work on autotetraploid Lycopersicum  esculentum, where she found the number of multivalents to vary from only one to eight per cell* Moreover, Stebfeins (1947) points out that sterility of auto-ploids is not always due to multivalent formation, but more often to gene-controlled irregularity of chromosome distribution, or to unexplained physiological incompatibility unassociated with irregular meiotic division. b. Difference of opinion exists on the evolutionary role of the two types of polyploidy. Alloploidy is generally accepted as being the more significant. Clausen et al* state that because of barriers to back-crossing set up by both chromosome numbers and genetic balance, an allo-ploid is a fully formed species from its beginning, combining the char-acters of its two parents* Stebbins (1947) states that about half the species of Angiosperms are polyploid, and he believes that most, i f not a l l , of these are alloploid* However, he maintains that the chief sig* 19 nificance of even alloploidy, since it provides no really new genetic material, is in combination with hybridization where i t may establish fer t i l i t y of hybrids between otherwise intersterile species. That autoploids exist in nature is agreed but their value in evo-lution is s t i l l uncertain* Clausen et al* (1945) l i s t ten species which they consider, after the study of extensive evidence, to be natural auto-ploids. They conclude that autoploids differ less than alloploids from the parental forms in ecological preference and that sometimes they are morpho-logically indistinguishable* However, they also state, Simple doubling of the chromosomes, with a l l their included genes, clearly is a process that may alter the physiologic balance of a plant, so that i t can occupy an ecologic niche to which its progenitor was not ad-apted* From then on, differential selection should eventually make diploid and tetraploid forms morpholog-ically distinct also, so that they would become distinct species* Again they say, There is no doubt that most of these autoploids units play the same roles in nature as do well re-cognized species, for most of them have distinct ecologic preferences and do not interbreed freely with their diploid counterparts* They are there-fore important units for the ecologist and the evo-lutionist. Wherever morphological differences are available, the autoploids should be recognized as taxonomic species, even though the differences are less distinct than those between species on one chromosome level* Stebbins (1947), on the other hand, believes that most or a l l of the oo-called autoploid species are in reality alloplold* He admits only one known naturally-occurring polyploid, Galax aphylla, as an undisputed autoploid species, and i t , he says, is so like its diploid parent as to have l i t t l e significance in evolution* so Stebbins* view seems somewhat extreme though, in part, the difference in viewpoint may be due to a narrower interpretation on his part of the term •autoploid*• In any case, this paper tends toward the view of Clausen et al« that doubling of even very similar genams must produce a new internal balance which, with isolation, could lead to diff -erentiation of the autoploid from its diploid ancestor. Certainly, as pointed out by Clausen et al», and as illustrated by many species, both auto- and alloploidy may enlarge the species dist-ribution* On this subject Stebbins (1940) makes three observations. a. In general, the range of diploid forms is restricted, that of polyploids widespread. In particular instances, such as Crepis, both diploids and autoploids are restricted, whereas in Tradesoantia the auto-ploids are widespread. b. The centre of distribution of a polyploid complex is the centre of variation and the spread may be from south to north, or from Arctic regions south, from inland to the sea coast, or from the coast inland. c. The situation in respect to these three points in any given genus depends on what genus is in question. These observations are borne out by the work of Flovik (194u) on Scandinavian floras. He found, in some instances, that the polyploids had spread north to Spitzbergen. In other eases, the diploids were in the north, the polyploid derivatives farther south. If i t does no more, then, than extend distribution, polyploidy may provide isolation which could act as an agent in speelation. * 21 Hybridization has been discussed in connection with polyploidy* Farther mention should be made of its action alone. Hybridization on the diploid level and its effect in speciation are discussed by Heiser (1950). He divides hybrids into two groups; 1. Those which are completely sterile. 2* Those which are not completely sterile and therefore are able to backcross to the parents. For the first group, alloploidy offers the only chance of fe r t i l i t y * In the second group occurs what is termed introgression, or, as Heiser de-fines i t , "the transfer of genes from one species to another by means of hybridization and backcrossing*tt As examples of genera in which intro-gression occurs, Heiser mentions Cistus, Helianthus, Salvia, Silene, etc. He concludes that introgression has at least two effeots in speciation: 1* It inoreases the variability of a species. 2. It possibly creates incipient species. He also points out that, though no new genetic material is directly creat-ed, the interaction of newly combined genes may produce new characters and also that hybridization is known in certain instances to stimulate muta-tion rate, thus creating new genetic material indirectly. To recapitulate then, and beyond doubt, to over-simplify the process of speciation as i t occurs in the course of evolution: variabil-ity within an existing species may be provided by any one or any combina-tion of three types of mutation; sexual reproduction and hybridization spread the variability, recombining mutated characters; selection acts on the variants, eliminating the unfit; successful variants, i f by some means isolated from the parent stock, may further differentiate to become 2 2 -new s p e c i e s » B . D e f i n i t i o n o f t a x o n o m i o c a t e g o r i e s . A f t e r a d i s c u s s i o n o f s p e c i a t i o n a s a c o n t i n u o u s p r o c e s s i n t h e l a r g e r p r o c e s s o f e v o l u t i o n , d i f f i c u l t i e s may be f o r e s e e n i n d e f i n i n g t h e s p e c i e s , t h e b a s i c t a x o n o m i c u n i t * H o w e v e r , numerous d e f i n i t i o n s have been o f f e r e d by b i o l o g i s t s ( D o b z h a n s k y , 1 9 4 1 ; H u x l e y , 1 9 4 0 ; a n d o t h e r s ) . D i f f e r e n t c r i t e r i a have b e e n a d o p t e d b y d i f f e r e n t w r i t e r s , b u t i n t h e many d e f i n i t i o n s c e r t a i n p o i n t s may b e c h o s e d w h i c h a p p e a r r e p e a t e d l y . 1 * S p e c i e s a r e n a t u r a l u n i t s , n o t a r t i f i c i a l g r o u p i n g s s e t up f o r c o n v e n i e n c e i n n a m i n g * 2 . S p e c i e s a r e s e l f - p e r p e t u a t i n g because t h e y m a i n t a i n a b a l a n c e i n t e r n a l l y among t h e i r genes a n d e x t e r n a l l y w i t h t h e i r e n v i r o n m e n t . 3 . They b r e e d u s u a l l y w i t h i n t h e i r l i m i t s a n d be tween s p e c i e s t h e r e i s u s u a l l y some d e g r e e o f g e n e t i c , e c o l o g i c a l , o r g e o g r a p h i c a l i s o l a t i o n . 4 * Some m o r p h o l o g i c a l d i f f e r e n c e i s g e n e r a l l y p r e s e n t b e t w e e n s p e c i e s . 5 * A g e n e r a l l a c k o f i n t e r g r a d e s e x i s t s be tween s p e c i e s * These f i v e p o i n t s , t h e n , p r o v i d e a s p e c i e s c o n c e p t s u f f i c i e n t l y c o m p l e t e t o s e r v e a s a d e f i n i t i o n o f t h e t e r m a s h e r e a c c e p t e d . A v a r i e t y i s c o n s i d e r e d as a s u b - d i v i s i o n o f a s p e c i e s , d e s i g n a t i n g a g r o u p w h i c h shows some degree o f g e n t i c a l l y - d e t e r m i n e d m o r p h o l o g i c a l d i f f e r e n c e a n d some d e g r e e o f i s o l a t i o n f r o m i t s p a r e n t g r o u p * The v a r i e t y t h e n , i s i n a p o s i t i o n t o d i f f e r e n t i a t e s t i l l f u r t h e r , and t h e r e f o r e , i s a s p e c i e s i n t h e m a k i n g . A f o r m i s a s t i l l l e s s e r s u b - d i v i s i o n o f a s p e c i e s , d i f f e r i n g f r o m 23 -its parent group by perhaps only one or a few genes. Though the differ-ence is genetically determined, lack of isolation permits spread of the mutant gene or genes within the species. A form, then, is a mere variant, but with the introduction of some isolating mechanism it might become a variety. C. History of the Taxonomic Treatment of the Genus. Dodeoatheon is a Linnaean genus described in Species Plantarum as having only the type species, D. Meadia, of eastern United States. The type locality is given as Virginia. Linnaeus includes three references to the previous use of Meadia as the generic name. None of these refer-ences has been available, but apparently the two earliest are both publica-tions of 1731: Mark Catesby in The Natural History of Carolina, Florida, and the Bahama Islands and Philip Miller in The Gardener's Dictionary. Because the genus has no economic value and, until now at least, its horticultural value has been unrecognized, except by a few gardeners, i t has not been extensively studied* From 1753 to 1876 several species were described but no attempt was made to treat the entire genus until Gray (1876) published a classification in which he recognized only one species, D. Meadia with six varieties. This classification he maintained in a second publication (1878), but, in a revision of the genus (1886b), he established five species and two varieties. From 1886 to 1900, many new species were described, so many that in the next revision, that of Pax and Knuth in Engler (1905), no less than thirty species appeared. At the present time, however, a large number of these, at least in the Northwest, seem to be synonymous. 24 o « In 1944, Fassett published a reclassification of Dodecatheon as i t appears east of the 100th meridian* He recognizes two species, D*  Meadia and D. amethystlnum, each with a number of varieties and forms* The present paper treats the genus in the Northwest. The com-plete area covered is shown on any one of Maps 1-7. After considerable field work and examination of seven to eight hundred herbarium specimens, it is felt that representatives of a l l species present in this area have been 3 e e n and studied* The material appears to f a l l into ten quite dis-tinct groups recognized as species* One of these contains one variety. Aside from these ten species, a small number, perhaps fifteen to twenty-five specimens, have been examined which f i t rather poorly into any group, but their variation from one another and their distribution are so wide as to preclude their being another species* Some combine characteristics of two recognized species and may well be hybrids; others, which vary in only a few minor respects from a recognized species, have been, at least for the present, included with that species. Within several groups there is considerable variation in those characteristics found not to be particu-larly useful in classification. Rather than further confuse the taxonomic state of the genus with uncertain varieties and forms, such variants have been mentioned in discussions but left unrecognized as distinct entities, at least until further work indicates their correct status. D. Diagnostic characteristics Possible diagnostic characters have been divided into three groups according to the value they have been found to possess in classi-fication. - 25 -Group A. type of root. length and colour of filaments, type of capsule dehiscence. Group B. leaf shape. degree and distribution of pubescence, seed wings. corolla colour and markings. anther length. Group C. number of parts. size of plant, presence of filament tube, characteristics of sepals, bracts, and petals. Characteristics in Group A have been found to be most constant throughout the genus and will be dealt with in some detail. 1. Roots are of two types and on this basis the genus falls into two subdivisions, those species haying a tap root with numerous fleshy rootlets along its length and those having a short crown with a whorl of fleshy-fibrous rootlets at its base. In general, the first type belongs to species which live in running water or locations otherwise wet through-out the growing season. The second type belongs to species which grow in areas moist only during blooming season, later becoming dry to extremely dry. Either type is an efficient means of vegetative reproduction as, after flowering, the fleshy rootlets form new crowns close to the mother plant, and in the next season send up new shoots. In species of wet habi-26 tats the rootlets seem often to remain attached, thus forming dense masses of intertwined roots as well as dense clumps of plants. In the species of arid locations, the rootlets break away during the dry season and the following year produce separate plants. In either case, the result is the eventual production of clumps of plants vegetatively from a single indiv-idual, a characteristic extremely useful in horticultural work. 2. Usually correlated with type of root, is length of filament. The species with tap roots have filaments extremely short or laoking, while those with a short crown have filaments 1 mm. long or more. The excep-tions to this are D. conjugens and D. columblanum, each with a short crown but filaments less than 1 mm. long. Filaments are either yellow or purple, and this characteristic is usually quite constant. However, variation has been found in both length and colour of filament in a few specimens from Oregon. 3. Capsule dehiscence, like root habit, is of two distinct types, but dehiscence and root habit are i£ no way correlated. In some species the capsules split from the apex by valves, in others the fruit is a pyxis whose cap may be relatively large and distinct or very small but nevertheless present. The distinction here is quite obvious, but not as useful as some others because, unfortunately, mature fruits are so often missing from herbarium material. Characteristics of Group B are useful to a limited extent and only in separating certain species. 1. Leaf shape varies considerably even within most species, but the sub-orbicular leaves of D. Hendersoni and sub«cordate of D. dent a turn and 87 -D . frigidum. are reasonably distinctive. 2. Likewise pubescence through a good deal of the genus is variable but the thickly glandular pubescent inflorescence of D . Cusickii sets i t apart from otherwise similar species. 3. Winged seeds are peculiar to D . Jeffrey! and possibly D . alpinum, but seeds unmistakably of D . alpinum have not been seen. 4. Corolla colour is generally limited to shades of rose or purple with occasional white forms occurring in some species. The only consist-ently white-flowered species is D . dentatum, of which apparently no colour-ed form has been collected. The only obvious corolla marking is the scallop on the re flexed tube and i t serves only as a guide in certain groups. 5 . Anther length is quite constant but varies so l i t t l e between species that only in a few cases is it helpful in distinguishing species from each other. Between D . columblanum and D . conjugens and between D . pauciflorum aid its variety, maorooarpum, anther length proves useful. Characteristics of Croup C are so variable even within species as to be of l i t t l e diagnostic value. I s Most species are D i m e r o u s but tetramery occurs in D . Henderson!, D . Jeffrey!, and D . alpinum. In the first of these its occurrence is rare but so variable that 4-, 5-merous, and even 6-merous flowers appear on the same scape. In the last two, tetramery is the more consistent but, at 1 least in D . Jeffreyi, variability is also common. 8.. Size of plant depends so much on water supply, soil conditions, elevation, etc. that it fails to be of use as a diagnostic character. 3. Presence or absence of a filament tube seems more closely related to age than to species. It has been used as a distinguishing feature of 28 D. conjugena but even here, on herbarium sheets examined, a l l degrees of fusion has been found. 4. Sepals, bracts, and petals vary in both quantitative and quali-tative characters so greatly that nothing has been found in any one of them to be of help in classification. Characteristics used, then, in the key are from Group A and Group B. Those of Group C have been included in the descriptions for completeness. E. Herbaria from which specimens were borrowed, showing abbrevia-tions used in citations: 1. National Museum of Canada, Ottawa .............. N.M. 2. Provincial Museum, Victoria, B.G 3. University of British Columbian Vancouver . 4. Unviersity of California, Berkeley ........ 5* University of Washington, Seattle ......... P. Classificat ion. Dodeoatheon L. . ••• P.M. « . .. U.B.C. . • .. U.C. .... U. w. Dodeoatheon L. Sp. PI., ed. 1, 144, 1753. Low perennial herbs with basal leaves; glabrous throughout or variously glandular pubescent; rootlets from a fleshy tap root or a short crown; herbage persistent or withering after flowering; leaves simple, spreading or erect; scape single, a few centimeters to over 40 cm. t a l l , bearing a bracteate umbel of 1-20 nodding flowers; pedicels 1-4 cm. long, elongating and becoming erect with maturity; flowers pentamerous, less commonly tetramerous; calyx and corolla reflexed, each with a short tube; «• 29 -c o r o l l a l o b e s i n s h a d e s o f r o s e o r p u r p l e , somet imes w h i t e , u s u a l l y w i t h a w h i t e zone a t t h e i r b a s e ; c o r o l l a t u b e y e l l o w i s h , b e a r i n g u s u a l l y a r e d -o r a n g e o r p u r p l e s c a l l o p ; s tamens 4 o r 5 , e x s e r t e d f r o m t h e c o r o l l a a t u b e o p p o s i t e t h e p e t a l s , a n t h e r s f r e e , f i l a m e n t s , i f v i s i b l e , y e l l o w o r p u r p l e , u s u a l l y f o r m i n g a. t u b e ; s t i g m a c a p i t a t e , s t y l e e x c e e d i n g t h e s t a m e n s , o v a r y u n i l o c u l a t e , c a r p e l s 5 , f u s e d , o v u l e s n u m e r o u s , p l a c e n t a t i o n f r e e c e n t r a l o n a c o l u m n ; f r u i t a c a p s u l e , c l r c u m s c i s s i l e o r d e h i s c i n g by t e e t h ; s e e d s s m a l l , 0 e 5 ~ 2 . 0 mm. l o n g , d a r k t o p a l e b r o w n , w i n g e d o r w i n g l e s s . Type s p e c i e s : D . M e a d i a Type l o c a l i t y : V i r g i n i a D i s t r i b u t i o n ; C o a s t , I n s u l a r and C o r d i l l e r a n M t s . f r o m A l a s k a a n d Y u k o n t o O r e g o n and I d a h o , d r y I n t e r i o r P l a t e a u x , e a s t a c r o s s M o n t a n a , A l b e r t a , S a s k a t c h e w a n , and i n t o M a n i t o b a . Above i s t h e d i s t r i b u t i o n i n t h e a r e a w i t h w h i c h t h i s p a p e r i s c o n c e r n e d . D o d e o a t h e o n e x t e n d s s o u t h t h r o u g h C a l i f o r n i a , a c r o s s t h e U n i t e d S t a t e s t o t h e e a s t c o a s t a n d s o u t h t o T e x a B . so K E Y TO THE SPECIES R o o t l e t s f r o m a f l e s h y t a p r o o t ; f i l a m e n t s e x t r e m e l y s h o r t o r l a c k i n g , ( e x c e p t r a r e l y i n D . a l p i n u m ) ; s p e c i e s o f wet h a b i t a t s . L e a v e s o v a t e w i t h s u b c o r d a t e o r s h a r p l y c o n t r a c t e d b a s e ; s eeds w i t h o u t w i n g s . F l o w e r s l i l a c p u r p l e ; c a p s u l e c i r c u m s c i s s i l e ; i n f l o r e s c e n c e g l a n d u l a r p u b e s c e n t . 1 . D . f r i g i d u m F l o w e r s w h i t e ; c a p s u l e d e h i s c i n g b y v a l v e s ; p l a n t s g l a b r o u s t h r o u g h o u t . 2 . D . d e n t a t u m L e a v e s o b l a n c e o l a t e , t a p e r i n g g r a d u a l l y t o a w i n g e d p e t i o l e ; s eeds w i n g e d on t h e a n g l e s . C a p s u l e c i r c u m s c i s s i l e , d e h i s c i n g b y a v e r y s m a l l c a p ; l e a v e s h a l f o r more a s l o n g as t h e s c a p e ; p l a x t s u s u a l l y g l a n d u l a r p u b e s c e n t , e s p e c i a l l y i n t h e i n f l o r e s c e n c e . 3 . D . J e f f r e y i C a p s u l e o p e n i n g b y v a l v e s ; l e a v e s l e s s t h a n h a l f a s l o n g a s t h e s c a p e ; p l a n t s g l a b r o u s . 4 . D . a l p i n u m R o o t l e t s f r o m a s m a l l c r o w n ; f i l a m e n t s e v i d e n t . u s u a l l y more t h a n 5 . mm. l o n g ; m o s t l y s p e c i e s o f h a b i t a t s b e c o m i n g d r y i n summer. F i l a m e n t s d a r k p u r p l e ; 1 mm. l o n g o r m o r e . G l a n d u l a r p u b e s c e n t t h r o u g h o u t ; c a p s u l e o p e n i n g b y v a l v e s . 5 . D . p o e t i c u m G l a b r o u s e x c e p t m o d e r a t e l y g l a n d u l a r p u b e s c e n t i n t h e i n f l o r e s c e n c e ; c a p s u l e c i r c u m s c i s s i l e . 6 . D . H e n d e r s o n i F i l a m e n t s y e l l o w . F i l a m e n t s u s u a l l y l e s s t h a n 1 mm. l o n g ; c a p s u l e c i r c u m -31 s c i s s i l e . A n t h e r s 6 -7 rata, l o n g , d a r k p u r p l e ; l e a v e s g l a b r o u s o r s p a r s e l y g l a n d u l a r p u b e s c e n t . 7 . D . c o n j u g e n s A n t h e r s 4 . 5 - 5 . 5 mm. l o n g ; l e a v e s t h i c k l y g l a n d u l a r p u b e s c e n t , i n f l o r e s c e n c e g l a b -r o u s . 8 . D . c o l u m b i a n -um F i l a m e n t s 1 mm. l o n g o r m o r e ; c a p s u l e o p e n i n g b y v a l v e s ; Whole p l a n t t h i c k l y g l a n d u l a r p u b e s c e n t , e s p e c i a l l y t h e i n f l o r e s c e n c e . 9 . D . C u s i c k i i A t l e a s t t h e i n f l o r e s c e n c e g l a b r o u s . 1 0 . D . p a u c i f l o r -urn 32 -1. Dodeoatheon frigidum Cham, and Schlecht. Dodeoatheon frigidum Cham, and Schlecht., Linn. 1:222, 1826. Dodeoatheon Meadia L. var. frigidum Hook., Curt. Bot. Mag. a. 3, t. 5371, 1870, pro parte. Plants of small stature, a few cm. to rarely 30 cm. or more high; herbage essentially glabrous, upper part of scape and inflorescence dis- tinctly glandular pubescent, often showing red-purple coloration even in  the calyx; rootlets red-brown, fie shy-fibrous, from a slim tap root; leaf  blades ovate, 1.5-5.0 cm. long, 1.5-3.0 cm. wide, margin entire with minute callus teeth terminating the veins or irregularly repand with calluses in the sinuses, apex rounded to obtuse, base abruptly contracted  or subcordate; petioles 1*5-7.0 cm. long, narrowly winged; scape 9.0-22.0 cm. long; bracts 1*8-8*0 mm* long, narrowly acuminate; flowers 2-6, pent-amerous; calyx tube 1*5-2.0 mm. long, lobes 3.0-3.5 mm. long, obtuse; corolla lilac-purple with a white zone at the base of the lobes and a broad regular, dark purple scallop on the reflexed tube; stamens dark purple, anthers 4*5^ 5.5 mm. long, filaments very short or lacking, connectives smooth, narrowly acuminate; capsule narrowly ovoid, about 1§ times the length of the calyx, sparsely glandular pubescent, circumscissile. walls delicate, almost transparent; seeds 1.5-1.9 mm. long, wingless. Type locality: St. Lawrence Bay on the Asiatic Coast, Bering St. Habitat: wet alpine meadows and rook crevices, stream banks, 2000-7000 f t . Distribution: Asiatic Coast, Alaska, Yukon, McKenzle Delta, north-ern British Columbia. Map 1* Maps 1 - 7 show t h e d i s t r i b u t i o n o f t h e s p e c i e s o f D o d e c a t h e o n i n A l a s k a , Y u k o n , B r i t i s h C o l u m b i a , A l b e r t a , S a s k a t c h e w a n , W a s h i n g t o n , O r e g o n , I d a h o , a n d M o n t a n a . M o u n t a i n a x e s shown o n t h e maps i n d i c a t e t h e f o l l o w i n g r a n g e s : W e s t e r n C o r d i l l e r a n C o a s t , I n s u l a r - S t . E l i a s M t s . , o f A l a s k a n d Y u k o n . I s l a n d M t s . o f V a n c o u v e r I s l a n d , Queen C h a r l o t t e I s l a n d s , and t h e A l a s k a n P a n h a n d l e . O l y m p i c M t s . o f W a s h i n g t o n . C o a s t Range o f O r e g o n . M a i n A x i s - M c K i n l e y M t s . o f A l a s k a C o a s t Range o f Y u k o n and B r i t i s h C o l u m b i a . Cascade M t s . o f W a s h i n g t o n and O r e g o n . C e n t r a l C o r d i l l e r a n - C e n t r a l Ranges o f Y u k o n , e . g . G l e n l y o n M t s . O m i n e c a S y s t e m i n c l u d i n g S t i k i n e M t s . o f B r i t i s h C o l u m b i a . C o l u m b i a n S y s t e m i n c l u d i n g S e l k i r k and Monashee M t s . C a b i n e t M t s * o f I d a h o a n d M o n t a n a . S a w t o o t h R a n g e , S a l m o n R i v e r M t s . o f I d a h o . B l u e M t s . o f Oregon and W a s h i n g t o n . W a l l o w a M t s . o f W a s h i n g t o n . E a s t e r n C o r d i l l e r a n - E n d i c o t t M t s . o f A l a s k a M c K e n z i e M t s . , o P e e l M t s . o f A l a s k a . N o r t h e r n and S o u t h e r n R o c k i e s o f C a n a d a . A m e r i c a n R o c k i e s i n c l u d i n g t h e o u t l i e r s , i e . , L i v -i n g s t o n e R a n g e , e t c . Map 1 D o d e o a t h e o n f r i g i d u m • . * • c i r c l e s D o d e o a t h e o n d e n t a t u m . • . . s q u a r e s 33 Chromosome count: no satisfactory material obtained. Specimens examined: ALASKA: Head of Savage Hiver, McKinley National Park, July 1, 19E8, Ynes Mexia, 8070 (U.W., U.G.); Nome Hiver, Nome, July 17, 1946, Christine Heller, s.n. (U.W.)j Nome, 1900-1904, Mrs. 0«N.Powers,  110, (U.C.); vicinity of Nome, Mrs. C.N. Powers, 44 (U.C»); Nome, 1900, Dr. ff.E. Blaiadell, 48 (U.C.); Kuskokwim River region, Russian Mt., Aug.6, 1944, A.J. Stewart, 656 (U.C.)j KuskokwimPRiver region, Horn Mt., July 8, 1944, A.J. Stewart, 609 (U.C.); Thompson Pass, Valdez, July 14, 1949, G. Frohne, 49845 (U.B.C.); southeast Alaska, head of Chitina River, June 26, 1925, H.M. Laing, 169 (N.M.); Kokrines Mts., near hot sulphur springs, 65°17'N. 154PZ0*^,, July 6, 1926, A.E. and R.T. Poraild, 680 (N.M.); Kokrines Mts., north side of divide towards Melozitna River, June 23-July 5, 1926, A.E. and R.T. Porsild, 785 (N.M.); Alaska Range, between Paxon and Sunmit, 63°to 63°20,N. 145°30*W., June 26, 1926, A.B. and R.T. Porsild, 562 (N.M.); Alaska Range, Paxon, south of divide, 63°N. 143°3o'w., June 27, 1926, A.E. and R.T. Porsild, 584 (N.M.)j Cantwell, Broad Pass in the Alaska Range* eS^'N. 149°W., June 8-11, 1926, A.E. and R.T. Porsild, 86 (N.M.); Takotna, July 27, 1941, J.P. Anderson and G.W. Gasser, 7459 (N.M.); YUKON: vicinity of Carcross, 1950, J.P. Aitken, s.n. (U.B.C.); Pelly River, 1949, J.P. Aitken, s.n. (U.B.C.); Mt. Decoeli, Dezadeash Mts., July 4, 1943, V.C. Brink, s.n. (U.B.C.)} Dawson Range, about 62°N. 138°W., south of Selkirk, west of Cairnacks, near source of KLaza River, July 12, 1933, H.S. Bos to ok, 28 (N.M.); Kluane Lake, Burwash Landing, above timer-line, July 3, 1943, C.H.D. Clarke, 279 (N.M.); Haines Rd., near Mile 85, B.C., July 17, 1944, C.H.D. Clarke, 466 (N.M.); McQueston area, east side of White Mt., June 22, 1948, J.P. Campbell, 154 (N.M.); Klotassin area, 34 * southwest of Yukon River, between Selkirk and White River, Aug., 1916, P.P. Cairnes, s.n. (N.M.); north of McQueston River, Aug. 15, 1947, R.L. Christie, 81 (N.M.); Keno, Mayo Pistrict, June 20-28, 1927, Mrs. George Black, s.n. (N.M.); 63°50'N. 141°W., July 5, 1917, C.E. Cairnes, s.n. (N.M.); Single Pt., MacKenzie Pelta, Aug. 4, 1914, J.R. Cox and J.J. O'Neill, 173 (N.M.); Herschel Island, 69°55fN. 139°Wi, Aug., 1914, Frits Johansen, 196 (N.M.); May Pistrict, between heads of Wedwards and Nelson Creeks, 6340'N. 134°20'w., July 5, 1939, Hugh Bostock, 108 (N.M.); 60-*iile River, June 2, 1939, R.T. Porsild, s.n. (N.M.); Shingle Pt., 69°N. 137°30'w., Sept. 13, 1933, A.E. Porsild, 6908 (N.M.); Shingle Pt., 69°N. 137o30'w., July 24, 1934, A.E. Porsild, 7111 (N.M.)j Canol R., Rose-Lapie Pass, Mile 105, near Lapie Lake, June 11, 1944, H.E. Porsild and A.J. Breitung, 9540 (U.W., N.M.); Canol R., pass between Teslin and Nisutlin Rivers, July 6, 1944, A.E. Porsild and A.J. Breitung,11006 (N.M.); Canol R., east slope of Rose River valley near pump station No. 10 at Mile 77, July 3, 1944, A.B. Porsild and A.J. Breitung, 10515 (N.M.); Canol Rd., Rose-iapie River Pass, Mile 105, near Lapie Lake, June 10, 1944, A.B. Por- sild and A.J. Breitung, 9325 (N.M.); Canol Rd., Rose-Lapie River Pass, Mile 105, schist mountain east of lake, June 11, 1944, A.E. Porsild and A.J. Breitung, 9408 (N.M.); Canol Rd., Rose-Lapie River Pass, east slope of granite^schist mountain west of Mile 118, June 29, 1944, A.E. Porsild and A.J. Breitung, 10143 (N.M.); Canol Rd., south and east slopes of Mt. Sheldon, Aug. 3, 1944, A.E. Porsild and A.J. Breitung, 11728 (N.M.)j Canol Rd., Mile 132, lower Lapie River Crossing, June 26, 1944, A.E. Por*  slid and A.J. Breitung, 9941 (N.M.); NORTH WEST TERRITORIES: eastern slope of Richardson Mts., west of Mackenzie River delta, 68°N. 136°W., - 35 July 1, 1931, 0. Bryant, 6616 (N.M.); shore of Mackenzie Bay, mouth of Mackenzie River, 1893, Rev. J. C. Stringer, 14424 (N.M.); BRITISH COLUMB-IA: Kechika River, 20 miles north of S i f ton Pass, July 20, 1940, N.C. Stewart, 13205 (P.M.); White Pass, June 24, 1936, E. & J. Lohbrunner, 119 68 (P.M.); Cambrey Mt., Northern B.C., July 9, 1945, A.G. Slocomb, 17883 (P.M.); near Yukon Boundary, sg^'N. Iseoss'w., June 29-30, 1947, M.P. and R.T. Pro slid, 224 (N.M.); Rabbit River, Liard region, 59°N. 127°W., 1913, E.B. Hart, s.n. (U.B.C.); / Rainy Hollow, Aug.14, E.S.  Wilkinson, 5 (U.B.C.). 2. Dodeoatheon dentatum Hook. Dodeoatheon dentatum Hook., El* Bor. Am. 2:119, 1940. Dodeoatheon Meadia L. var. latilobum Gray, In Geol. Sur. Cel. Bot., ed. 2, v. 1:467, 1880. This publication is considered the equivalent of ed. 1, 1876, as i t states, "In this edition of the present volume, no changes have been made excepting such as may properly be called corrections of slight verbal mis-takes or errors of the press. n Dodeoatheon frigidum Cham, and Schlecht. var. dentatum Gray, Bot. Gaz. 11:234, 1886b. Dodeoatheon latilobum (Gray) Elmer ex Pax and Knuth, In Daz Pflanz. 4:239, 1905. Plants generally taller than D. frigidum, commonly 15-30 cm. high but scapes and petioles slim; completely glabrous, lacking red- purple coloration; rootlets white, fleshy-fibrous from a slim tap root; leaf blades ovate, 2.0-9.5 cm. long, 1.5-4.5 cm. wide, margin repand with calluses in sinuses or Irregularly dentate with callus-tipped teeth, apex 36 « rounded to obtuse, base subcordate; petioles 3.0-12.5 on. long, narrowly winged; scape 13.0-35.0 an. long; bracts 1.0-9.0 mm. long, acute to acuminate, papery; flowers 3-5, pentamerous; calyx tube 1.5-2.0 mm. long, lobes 1.5-4.5 mm. long, lanceolate and aoute to ell i p t i c and abruptly acuminate; corolla white with a faint to very distinct dark scallop on or almost within the re flexed tube; stamens purple-brown, anthers 5.0-6.5 mm. long, filaments very short or lacking, connectives smooth, narrowly acumin-ate; capsules narrowly ovoid, about 1§ times the length of the calyx, de- hiscing by valves, walls firm, opaque; seeds about 1 mm. long, wingless. Type locality: N. W. interior. Habitat: stream banks, or boggy meadows along streams, 1000-6000 f t . Distribution: Northern Idaho, east slope of the Cascades through Washington and northern Oregon, Okanagan Valley of Southern British Columbia. Map 1. Chromosome count: somatic, 2n - approximately 44. plate 1. Specimens examined; along Tenas Creek near Apex Mt., Penticton, May 10, 1940, J.W. Eastham, s.n. (U.B. C.); between Penticton and Kereme-os, May 10, 1940, J.W. Eastham, 12322 (P.M.); west of Penticton along Tenas Creek, July 26, 1950, K.I. Beamish, 50108 (U.B.C.); Tenas Creek, near Apex Mt., Aug. 4, M.C. Stonor, s.n. (U.B.C.); Summerland, July 3, 1934, Miss Stonor, s.n. (U.B.C.); Okanagan, 1937, M.C. Stonor, s.n. (U.B. C ) ; WASHINGTON: Mt. Hamilton, May 24, 1919, M.W. Gorman, 4515 (U.C.); Chelan Co.; head of Falls Greek, near South Navarre Peak, north of Lake Chelan, July 19, 1936, Marvin Kelly, 22 (U.C.); Mt. Stuart, Wenatchee Mt., Aug. 20, 1930, J. Wm. Thompson, 5846 (U.W.); banks of Eagle Creek near Leavenworth, May 23, 1931, J. Wm. Thompson, 6447 (U.W.); banks of Nigger PLATE 1 Dodecatheon dentatum, vicinity of Penticton,B.C. Camera lucida drawing, anaphase of mitosis in a root tip pretreated with paradichlorobenzene, showing approximately 44 chromosomes. Ca. X 3400 37 Creek, Weatchee Mts., June 2, 1934, J. Wm. Thompson, 10552 (U.W.)j base of Bowlder Pk., June 23, 1935, J. Wm. Thompson, 11789 (U.Wo); Tronson Creek, Wenatehee Mts., June 1, 1940, J. Wm. Thompson, 14613 (U.W., U.C.); Kittitas Co.: Blewett Pass region, Cascade Mts., Aug. 29, J.M. Grant, s.n. (U.W.); stream bank near Blewett Pass, June 7, 1924, G.A. Warren, 558 (U.W.); Blewett Pass, west side, June 15, 1925, G.V. Copley, 45 (P.M.); Blewett Pass, June 11, 1925, G.V. Copley, s.n. (P.M.); 3 miles north of Liberty, June 12, 1938, C.L. Hitchcock et al. , 5646 (U.W.); Table Mt., Wenatehee Mts., July 4, 1940, J. Wm. Thompson, 14959 (U.W., U.C.); Table Mts., Aug. 9, 1935, J. Ufa. Thompson, 9745 (U.W.); Table Mtd., July 24, 1947, Fred & Lill i a n Meyer, 2255 (U.C.); Okanogan Co.: Salmon Creek to Much amuck Lookout, Tiffany Range, June 25, 1931, J. Wm. Thompson, 6955 (U.W.); OREGON: bluffs of Columbia River near Cascades, June 1886, Thomas Howell, s.n. (U.W.); Clackamas Col: c l i f f s of Willamette River, Milwaukie, May 17, 1884, L.F. Henderson, 645 (U.C.); Multnomah Co.: Mult-nomah Falls, Columbia Gorge, May 27, 1928, J. Wm. Thompson, 4552 (U.W.); IDAHO: Shoshone Co.: below forks of Fishhook Greek, May 19, 1940, Marion  Ownbey, 2048 (U.W., U.C.); 3. Dodecatheon Jeffrey! Moore ex Tan Houtte. Dodecatheon Jeffrey! Moore ex Van Houtte, F l . des Serres 16:99, 1865>1867. Dodecatheon Meadia L. var. lancifolium Gray, In Beol. Sur. Cal. Bot., 2, v. 1:467, 1880, equivalent to ed.l, 1876 (see D. frigidum). Dodecatheon crenatum Greene, Pitt. 2:74, 1890, not Raf. Atl. Joum. 180, 1853. Dodecatheon viviparum Greene, Eryth. 3:58, 1895. - 38 Dodeoatheon tetrandrum Buks* ex Greene, Eryth. 3 : 4 0 , 1 8 9 5 . Dodeoatheon exlllfolium Macbrlde and Payeon, Contrib. Gray Herb^.n.s* 4 9 : 6 3 - 6 4 , 1 9 1 7 . Dodeoatheon Jeffreyl Moore var. tetrandrum Jepson, Man* F l o PI, Cal. 7 5 3 , 1 9 2 5 . Plants large, scapes stout, commonly over 30 cm. long, leaves often large and coarse; Inflorescence and sometimes leaves and scape var-iably glandular pubescent, rarely, entirely glabrous; rootlets thick, fleshy-fibrous from a heavy tap root; leaf blades lanceolate, e l l i p t i c , or oblancedate, 3 , 0 s 1 8 0 0 cm* long, 1 . 0 - 3 . 0 mm* wide, entire with minute  callus teeth to crenate with calluses in the sinuses, apex rounded to ac-ute, base cuneate, tapering to a winged petiole; petiole 2 » 0 - 1 2 o 0 cm* long; scape 1 0 . 0 - 4 5 . 0 on. long; bracts 2 . 0 - 1 8 . 0 mm* long, narrowly acumin-ate; flowers 2 - 8 , 4 - or 5«<aerous; calyx tube 1 * 0 - 3 . 0 mm* long, lobes 3*0<* 8 * 0 mm. long, broadly rounded to narrowly acuminate; corolla pale mauve to li l a c purple, scallop on the re flexed tube lacking to distinct purple, sometimes almost within the tube; stames dark purple, anthers 5*0-8*0 mm. long, filaments very short or lacking, connective transversely rugose; capsule broadly ovoid, about 1 £ times the length of the calyx, circum- scissile by a very small cap, walls delicate, almost transparent; seeds about 2 mm* long, winged on the angles* Type locality: "Mont ague s-Rocheuses" but described from a horti-cultural specimen. Habitat: Marshy meadows and stream banks, 3000-10000 f t . in the southern mountains, descending to sea level in the northern coast islands and Alaska* 39 Distribution; Central and Western Cordilleran Mts. of Montana, Idaho and Washington, along the coast from Alaska to Oregon, south into California. Map 2. Chromosome count; Stevens Pass, Wash*, meiotic n - 22 (PI 2,3) Forbidden Plateau, Vancouver Island . somatic ................. .**.2n -approx. 88 (W.4J Prince Rupert, B.C., somatic *• 2n -approx. 88 Specimens examined: ALASKA: Juneau, June 2, 1940, J.P. Anderson, 63541 (N.M.); Thum Bay, Knight Island, Prince Wm. Sound, Sept. 20, 1940, W.J. Eyerdam, 5641 (N.M.); Hinchinbrook Id*, July 15, 1924, W.J* Eyerdam, s.n. (U.C.); Gravina Island, Ketchikan, Aug. 3, 1927, J.P. AnderBon,658 (N.M.); Port San Juan, Evans Island, Aug. 25, 1948, W.J.Eyerdam, 7111 (N.M.); Port San Juan, Evans liand, July 5, 1948, W.J. Eyerdam, 5823 (K.M.) Thum Bay, Knight Island, Prince William Sound, Oct. 2, 1939, W.J. Eyer-dam, 5288 (U.C.); vicinity of Sitka, July, 1891, W.G. Wright, 1547 (U.C); Alaska Range near Paxon*s, July-Aug., 1934, ff.W. Went, s.n.(U.C.h Mite Cove, Yakobi Island, June 22, 1944, Max cine Williams, s.n. (U.W.); Peters** burg, June 22, 1949, W*0. Frohne, 4994 (U.B.C.); Juneau, July 23, 1949, W»C. Frohne, 49317 (U.B.C); Douglas Island, Juneau, July 26, 1949 , W.C,  Frohne. 49322 (U.B.C.); BRITISH COLUMBIA: muskegs above Hague Point, Princess Royal Island, May 29, 1936, T.T. McCabe, 5579 (U.C*); Canoona Lake, Princess Royal Island, June 21, 1956, T.T. Mc Cabe, 5550 (U.C*); Goose Island, July 20, 1959, T.T. McCabe, 7288 (U.C); near Deadman Cove, Banks Island, Aug. 19, 1959, T.T. McCabe, 7556 (U.C); mainland opposite Kaien Island, June 19, 1957, T.T. McCabe, 4551 (U.C); Leroy Lake, mouth Map 2. D o d e o a t h e o n J e f f r e y ! . . „ . c i r c l e s Dodeoa theon a l p i n u m . . . * s q u a r e s PLATS 2. X 0 © Dodecatheon Jeffrey!, Stevens Pass, Washington Camera lucida drawing, first division prophase of meiosis in a developing anther, showing 22 pairs of chromosomes. Ca. X 1500 PLATE 3 . D o d e c a t h e o n J e f f r e y ! , S t e v e n s P a s s , W a s h i n g t o n P h o t o m i c r o g r a p h , f i r s t d i v i s i o n p r o p h a s e o f m e i o s i s i n a d e v e l o p i n g a n t h e r , s h o w i n g 22 p a i r s o f chromosomes . C a . X 1000 V P L A T E 4. Dodecatheon Jeffrey!, Forbidden Plateau, Vancouver Island. Camera lucida drawing, prophase of mitosis in a root tip pre treated with paradidi loro-benzene, showing approximately 88 chromo-somes. Ca. X 1500 ELATE 5 Dodeoatheon Jeffrey!, Prince Rupert, B3. C. Camera lucida drawing, prophase of mitosis in a root tip pre treated with pared iohloro-benzene, showing approximately 88 chromo-somes, Ca. X 1500 - 40 of Smith Inlet, Sept. 23, 1934, T.T. McCabe, 1807 (U.C.); Smyth Island, Bardswell Group, Sept. 22, 1935, T.T. McCabe, 5172 (U.C.); Borrowman Harb-or, Aristazabel Island, May 31, 1936, T.T. MoCabe, 5386 (U.C., U.W.); Borrowman Harbor, northwest part of Aristazabel Island, June 5, 1936, T.T. MoCabe 5410(U.C); Calvert Island, head of Kwakshua Pass, May 16, 1953, T.T. McCabe, 59 (U.C.); Mount Buxton, Calvert Island, June 14, 1957, T.T. McCabe, 4204 (UlC.); Mount Buxton, Calvert Island, June 14, 1937, T.T. McCabe, 4205 (TJ.C.)j Calvert Island, northwestern part, south of Kwakshua, Sept. 8, 1935, T.T. McCabe, 5079 (U.C.) i Calvert Island, May 50, 1957, T.T. McCabe, 4060 (U.C.); Nootka Sound, V.I., May 29, 1907, W. Spreadborough, s.n. (N.M.); vicinity of Ucluelet, May 14, 1909, J. Macoun, s.n. (N.M.); vicinity of Ucluelet, June 1, 1909, J. Macoun,s.n. (H.M.); Ucluelet, June 22, l9l7, J.K. Henry, s.n. (U.B.C.); Ucluelet, May 12, 1912, G. Fraser, s.n. (U.B.C.); Mackenzie Lake, Forbidden Plateau, 7.1., Aug. 17, 1950, K.I. Beamish, 50111 (U.B.C.); 2 miles north on the t r a i l from Croteau Camp, Forbidden Plateau, V.I., Aug. 15, 1932, C. Hand*  ford, s.n. (U.B.C.); Forbidden Plateau, 7.1. near Mt. Alma, Aug. 15,1952, C. Handford, s.n. (U.B.C.); Forbidden Plateau, 7.I., July 15, 1959, J. Bostock, s.n. (U.B.C.); Paradise Meadows, Forbidden Plateau, 7.1., Aug. 3, 1932, R. Connell, s.n. (P.M.); mountains, Ucluelet, 7.I., May 16, 1924, W.B. Anderson, s.n. (P.M.); Prince Rupert, May 23, 1923, W.B. Ander- son, s.n.(P.M.); Tle l l River, July, 1914, C. de B. Green, s.n. (U.B.C.); WASHINGTON: Baldy Peak, July 25, 1897, F.H. Lamb, 1357 (U.C.); Mt. Paddo, fruit from Chi quash Mts., June 14, Oct., 1890, W.N. Suksdorf, 988, isotype or type of D. tetrandrum, (U.C.)"; near the Upper Valley of the "Nesqually" Aug. 18,- 1896, P.P. Allen, 246, (U.C, U.W.); Chelan Co.* Stevens Pass, •» 41 Aug., 1893, J.H. Sandberg and J.B. Leiberg, s.n. (U.C); Stevens Pass, Cascades, Aug. 9, 1893, J.H. Sandberg and J.B. Leiberg, 715 (U.C. U.W.); Mt. Inglls, Mt. Stuart Dist., July 4, 1957, W.J. Eyerdam, s.n. (U.C.); Mt. Stuart, Wenatehee Mts., July 27-31, 1931, J. Ufa. Thompson, 7710 (U.W.); Stevens Pass, June 6, 1949, K.I. Beamish, 49105 (U.B.C); Stevens Pass, June 23, 1950, K.I. Beamish, 50106 (U.B.C.); Ice Creek, Aug. 22, 1935, G.B. Morrill, 312 (U.W.); Clallam Co.: Olympic Mts., July, 1900, A.D.E.  Elmer, 2802 (U.W.); Grays Harbor Co.: near summit of Col. Bob Lookout, July 9, 1931, J. Wm. Thompson, 7267 (U.C); mountain meadows of Mt. Col. Bob. Aug. 23, 1953 J. Wm. Thompson, 9982 (U.C, U.W.,)j King Co.: Hester L., Cascades, July 18, 1955, John Broadbent, s.n. (U.W.); Kittitas Col: head of Beverly Creek, July 23, 1955, J. Wm. Thompson 9562 (U.C); Mt. Daniels, 4 miles west of Hyas L«, July 12, 1942, C.L. Hitchcock, 8085, (U.W.); stream, Scatter Creek Trail, July 6, 1942, C.L. Hitchcock, 7998, (U.W.); Fish L., 12 miles northeast of "Salmon La sac", June 21, 1942, C.L. Hitchcock, 7945 (U.W.); summit of Snoqualmie Pass, June 12, 1938, C.L. Hitchcock et al. , 5656 (U.W.); Klickitat Co.: Satus Pass region, May 14, 1958, J. Win. Thompson, 14289 (U.W.); Okanogan Co.: Chelan Nation-al Forest, valley of War Creek, July 22, 1921, H. St.John et a l . , 5256 (U.C, U.W.); banks of Lewis Creek, west of Twisp, Aug. 8, 1933, C B. F i -ker, 1347 (U.W.); Pierce Co.: Tan Trump Pk«, Mt. Rainier, July 31, 1929, A.A. Heller, s.n. (U.W.); slopes above Paradise Inn, Rainier National Pk., Aug. 11, 1926, A.A. Heller, 14777 (U.W.); near Spray Pk., Mt. Rainier, July 27, 1950, J. Wm. Thompson, 5456 (U.W.) J J.B. Flett, s.n. (U.C); Paradise Talley, Mt. Rainier, Aug. 16, 1955, G.N.. Jones, 8100 (U.C.).; Paradise Talley, Rainier National Pk., Aug. 16, 1928, H.E. and S.T. Parks, 0295£21051, (U.C); Paradise Talley, Mt. Rainier, J.B. Tarleton, 29 (U.C.);' - 42 -head of Mazsma Ridge, Mt. Rainier, July 22, 1930, Lyman Benson, 2503 (U.C.); Mt. Rainier, July 51, 1924, H.M. Hall, 12045 (U.C.); Paradise Pk., Mt. Rainier, Aug. 9, 1926, E.I. Applegate, 4888 (U.C.); Snohomish Co.; Silver-lake, Monte Cristo Dis., July 27, 1955, W.J. Eyerdam, s.n. (U.C); Yakima Co.: near Jennie's Butte Ranger Station, Yakima Indian Reservation, July 15 1932, V.T. Heldenreioh, 221 (U.W.); OREGON: W.C.Cuslck, 1526 (U.C); Rogue River Mts., head of EL at Creek, near Rabbit Ears, July 15, 1929, E.I. Applegate, 6015 (U.C.); subalpine bogs of the Blue Mts., July 20, Sept. 12, 1899, Wm. C. Cusiek, 2252 (U.C); Baker Co.: Imperial mine, headwaters of Powder River, about 10 miles above Sumpter, June 14, 1950, E.I. Applegate, 6429 (U.C.); Clackamus Co.: Mt. Hood, May 26, 1941, J.W.  Mcffarland, s.n. (U.W.); Paradise Park, Mt. Hood, Aug. 11, 1926, J. Wm.  Thompson, 1655 (U.W.); Coos Co.: Bandon-on-the-sea, May 5, 1954, M.W*  Gorman, s.n. (U.C); Deschutes Co.: McKenzie Pass, June 29, 1956, Helen M.  Gilkey, s.n. (U.C); McKenzie Pass, June 29, 1956, G.M. Powell, s.n.(U.W«); 6 miles west of MacKenzie Pass, June 22, 1939, CL. Hitchcock and J.S.  Martin, 4854 (U.C, U.W.); just east of Little Cult us Lake, June 25, 1939 CL. Hitchcock and J.S. Martin, 4919 (U.C, U.W.); Douglas Co.: Silent Creek, south end of Diamond Lake, Cascade Mts., July 16, 1926 E.I. Apple- gate, 4811 (U.C, U.W.); Silent Creek, Diamond Lake, Cascade Mts., July 16, 1926, E.I. Applegate, 4809 (U.C); Silent Creek, Diamond Lake region, Cas-cade Mts., July 16, 1926, B.I. Applegate, 4806 (U.C); Hood River Co.: Mt. Hood, July 51, 1950, Lyman Benson, 2507, (U.C); margin of Lost Lake June 23, 1938, Sister Mary Milburge, 1457 (U.C, U.W.); Hood River Meadows, Mt. Hood, July 8, 1925, M.W. Gorman, s.n. (U.C); Lane Co.: high McKenzie - 43 River, June 26, 1926, M.W. Gorman, 7709 (U.C*); Wasco Co*: Fifteen Mile Meadow, Mt. Hood National Forest, July 19, 1933, G.N. Jones, 4126 (U.C, U.W.); IDAHO: Loop Creek, ^  mile below mouth of Ward Creek, St. Joe National Forest, T.46N. R.7E. June 29, 1939, W.R. Moore, 455 (U.C); Loop Creek, below mouth of Ward Creek, St. Joe National Forest, T.46N. R.7E* June 29, 1939, W.R. Moore, 456 (U.C.); divide between St. Joe and Clear-water River, July U, 1895, J.B. Leiberg, 1252 (U.C); Priest Lake, May 14, 1938, CL. Hitchcock, 2914 (U.W.); Adams Co.: Black Lake T.21N. R.2W., July 3, 1940, R.J. Davis, 2545 (U.C); § mile from Valley*Adams County line, about 5 miles west of McCall, May 29, 1958, F.G. Meyer, 1476 (U.C.) J Blaine Co.: Alturas Lake, June 29, 1958, Ray J. Davis, 461 (U.C); Smoky Mts., Aug. 15, 1916, J.F. Macbride and E.B. Pay son, 3744, type or isotype of D. exilifolium, (U.C*); Alpine Creek, northwest of Alturas Lake, Sawtooth Primitive Area, July 7, 1944, CL* Hitchcock and CV. Muhlick, 10461 (U.8T.)j upper Norton Lake, vicinity of Norton Peak, Smoky Mt., Aug. 3, 1944, CL.  Hitchcock and C.V. Muhlick, 10716a (U.W.); Boise Co.: head of Clear Creek, about 15 miles north of Lowman, July 8, 1944, CL. Hitchcock and C.V.  Muhlick,9729 (U.W.); margin of Elk Lake, Sawtooth Primitive Area, head-waters of South Fork of Payette River above Sacajawea Hot Springs, July 11-12, 1944, CL. Hitchcock and C.V. Muhlick, 9879 (U.W.); Clear Creek, 16 miles north of Lowman, Sawtooth Mt. region, Payette River watershed, June 18, 1930, B..I* Applegate, 6503 (U.C.); Custer Co.: Bonanza, July 28, 1916, J.F. Macbride and E.B. Pay son, 5496 (U.C); Josephus Lakes, Aug. 3, 1916, (U.C.); J.F. Macbride and E.B. Pay son, 3583 (U.C.).; meadow midway from Stanley Lake to Cape Horn, July 7, 1944, CL* Hitchcock and C.V. Muhlick, - 44 9669 (U.W.); Elmore Co.: near Big Roaring River Lake, 20 miles north, of Pine, Aug. 25, 1947, F.G. and L.B. Meyer, 2559 (U.C); Trinity Lakes, T.3N. R.9E., R.J. Davis, 2875 (U.C.); above Trinity Lake, about 10 miles west of Featherville, July 25, 1944, C.L. Hitchcock and C. 7. Muhlick, 10579, (U.W.); Sawtooth Primitive Area, July 19, 1944, C.L. Hitchcock and C.T.  Muhlick, 10152 (U.W.); vicinity of Bald Mt. about 10 miles south of Atlan-ta, July 25, 1944, CL. Hitchcock and C.7. Muhlick, 10285 (U.W.); Idaho Co.: "The Swamps" near Lily Pad Lake, T.23N. R.2W., June 15-50, 1957, Jacque- line Packard, 525 (U.C); Warren Summit, T.22N. R.7E., July 5, 1940, R.J. Davis, 2559 (U.C); Bernard Lakes, T. 23N. R.2W., June 26, 1940, R.J. Davis  2518 (U.C); headwaters of Obia Creek, Rocky Ridge, Lolo Trail, Bitterroot Mts., July 15, 1957, L. Constance and F.W. PenneH, 2017 (U.C); f mile be-low Pilot Knob Lookout, S.33 T.30N. R.6E., July 2, 1947, W.N. Elwood, 55 (U.C, U.W.); Lemhi Co.: Mt. Baldy, Salmon, July 1, 1920 E.B. and L.B. Pay son, 1845 (U.C); west slope of Quart ziteMt., Yellowjacket Mts., June 50, 1946, C.L. Hitchcock and C.7. Muhlick, 14222 (U.C, U.W.); Shoshone Co.: main fork of St. Joe River, 5 or 4 miles east of Avery, S.18 T.45N. R.6E., July 11, 1941, CB. Wilson, 122 (U.C); Freezeout Saddle, S.35 T.45N. R.3E., July 18, 1941, CB. Wilson, 122A (U.C); Talley Co.: head of Deadwood Riv-er, June 20, 1939, R.J. Davis, 925 (U.C); Profile Summit, T.20N. R.9E., July 7, 1940, R.J. Davis, 2686 (U.C); Gold Fork Lookout, Payette National Forest, Sawtooth Mts., July 9, 1937, J. Wm. Thompson, 15784 (U.C, U.W.); east side of Brudage Mt., July 5, 1957, L. Constance and F.W. Fennel!, 1967 (U.C); 1 mile south of Landmark, June 27, 1946, C.L. Hitchcock and C T.  Muhlick, I4g66 (U.C.U.W.); MONTANA: Beaverhead Co.: June, 1889, Frank  Tweedy, s.n. (U.C); near Granite Peak near Melrose, June 50, 1956, Bert T. 45 -Bradley, 21 (U.W.); Torrey Lake, Pioneer Range, July 27, 1946, CJ,. Hitch- cock and C.V. Muhlick, 15029 (U.W.); Oreamnos Lake, head of Pintlar Creek, Anaconda Range, July 27, 1945, CL. Hitchcock and C.7. Muhlick, 12755 (U.C U.W.); 1% miles east of Lake Waukena, Pioneer Mts., Aug. 1, 1945, CL.  Hitchcock and C.V. Muhlick, 15152 (U.C.) J above AJax Lake, Bitter Root Range, July 25, 1945, CL. Hitchcock and CV. Muhlick, 12667 (U.C, U.W.); Granite Co,: Skalkaho Pass, July 7, 1946, CL. Hitchcock and C.V. Muhlick,  14466 (U.C, U.W.); along skalkaho road, west slope of the Continental Divide near summit, 8 miles west of Rock Creek Ranger Station, June 27, 1950, E.I. Applegate, 6424 (U.C.); Mineral Co.: Lee Creek Camp, Lolo Hot Springs, July 12, 1946, CL. Hitchcock and C.V. Muhlick, 14608A (U.C,UW.); Missoula Co.: top of Mt. Stuart, July 9, 1946, CL. Hitchcock and C.V.  Muhlick, 14556 (U.C, U.W.); Ravalli Co.: Hamilton, "Skalkaho" Rd., July 4, 1924, J.E. Klrkwood, 1780 (U.C); 1 mile west of St. Mary's Peak, Aug. 5, 1947, CL. Hitchcock, 17111 (U.C, U.W.); St. Mary's Lake, Bitterroot Mts., Aug.10, 1946 CL. Hitchcock and C.V. Muhlick, 15514 (U.W.); 4. Dodeoatheon alpinum (Gray) Greene. Dodeoatheon alpinum (Gray) Greene, Eryth. 5:59, 1895. Dodeoatheon Meadia L. var. alpinum Gray, In Geol. Sur. Cel. Bot., ed. 2, v. 1:467, 1880, equivalent to ed. 1, 1876 (see D.  frigidum). Dodeoatheon Jeffrey! Moore var. alpinum Gray, Bot. Gaz. 11:252, 1886b. Plants generally smaller than D. Jeffrey!, entirely glabrous; - 46 rootlets fieshy-fibrous from a short, slim tap root; leaf blades linear-oblanceolate, 1,8-6.0 cm. long, 0.2-0.9 an. wide, margin entire, apex rounded to acute, base narrowly cuneate, tapering to a winged petiole; petiole 1.5-4.0 an. long; scape 11.0-30.0 cm. long; bracts 3.5»8.5 mm. long, acute to acuminate; flowers 2-6, usually 4-, occasionally 5-enerous; calyx tube 1.0-2.0 mm. long, lobes 3.0-5.0 mm.long, acute to acuminate; corolla rose-purple with a distinct dark purple scallop on the re flexed tube; stamens dark purple, anthers 5.5-7.0 mm. long, filaments 0.5-0.8 mm. (rarely 1.0 mm) long, connective transversely rugose; capsule broadly ovoid, dehiscing by valves, walls delicate, almost transparent; seeds un-known when completly mature, apparently winged; Type locality: 'on the higher mountains, 9,500-12,000 f t . Habitat: stream banks and wet mountain meadows, 5000*8000 f t . in the northern part of its range. Distribution: southern and eastern Oregon, south to California. Map. 3-Chromosome count: no satisfactory material obtained. Specimens examined: OREGON: Wallowa Mts., Joseph-Cornucopia t r a i l , Aug.5, Eagle Creek meadows, fr. , Sept. 2, 1899, Wm. C. Cusick, 2316 (U.C.); sub-alpine Powder River Mts., Aug., 1896, Wm. C. Cusick, s.n. (U.C); Baker Co.: rocky slopes of Wallowa Mts., near Cornucopia, July 18, 1936, J. Wm. Thomp- son, 15334 (U.W.); Harney Co.: meadow above Pish Lake, Steen Mts., June 14, 1946, B. Maguire and A.H. Holmgren, 26449 (U.C, U.W.); canyon of Wild Horse Creek, Steen Mts.,- June 29, 1925, M.E. Peck, 14134, (U.W.); Josephine Co.: Illinois River 3 miles west of Selma, May 18, 1942, L. Constance et. al . , 2979 (U.C, U.W*); north slope of Grayback Mt., Siskiyou Mts., July 17, 1933, E.I* Apple gate, 8753 (U.C); Klamath Co.: near Admin-istration Bldg., Crater Lake National Park, June 29, 1926, F.W. Peirson,  6927 (U.C); region of Crater Lake, expedition to Fossil Lake, Aug. 1*16, 1901, H.W. Furlong et a l . , s.n. (U.C); Four Mile Lake, Cascade Mts., June 2, 1926, E.I* Apple gate, 4681 (U*C); Crater Lake National Park, July 20, 1918, A.A. Heller, 15056 (U.W.); Grater Lake, July 21, 1935, J. Wm* Thomp-son, 12250^ (U.W.); Lake Co.: Forest Camp on Dairy Creek, 35 miles north-west of Lakeview, July 3, 1927, M.E. Peck,15415 (U.W.); slopes of Crane Mt*, near Lakeview, July 11, 1956, J. Wm. Thompson, 15226 (U.W.); Willow Creek, east base of Warner Mts., July 6, 1952, B.I. Applegate 7438 (U.C); head of Deep Creek, eastern slope of Crane Mt., Warner Mts., July 6, 1932, S.I. Apple gate, 7459 (U.C); Wasco Co.: north side of Mirror Lake, Lake Basin, Wallowa Mts., Aug. 3, 1935, L. Constance and CD. Jacobs, 1355 (U.C.) i DISCUSSION OF D. FRIGIDUM, D. DENT ATOM, D. JEFFREYI, AND D. ALPINUM Dodecatheon frigidum appears to be a well differentiated northern species, most closely related to D. Jeffrey! and D. dent at um. Like D. Jeff- reyl i t possesses a tap root, circumscissile capsule, and glandular pubes-cent inflorescence. It differs markedly, however, In possessing: 1* generally smaller size. 2* red-brown rootlets and red pigmentation in the upper scape, bracts, calyx, and even capsule. 5. wingless seeds. 4. sharply contracted or sub cor date leaves. — 48 «• D* dentatum also has subcordate leaves, short filament tube, and tap root, but lacks any glandular pubescence, has only white flowers, no red colora-tion, and its capsule dehisces by teeth* Habitats of a l l three are much alike but distribution is very different* D» frigidum is entirely a northern species centred in Alaska and Yukon but reported also from the Asiatic Coast (Gray 1886, Hulten 1948, Anderson 1949) and northern British Columbia* (Map 1). D. dentatum is found on the east side of the Cascade Mountains in southern British Columb-ia, and Washington* One specimen has been seen from Idaho (Map 1)* No chromosome count has been obtained for D« frigidum, but D.  dentatum, from somatic counts, is diploid (Plate 1)* Dodecatheon Jeffrey! ranges from California to Alaska* In the territory covered by this paper, i t extends across the area south of the American border, in the mountains, reaching into Canada apparently only along the coast and islands, and by this route north to Alaska (Map 2). D» alpinum, on the other hand, though closely related, appears to be essentially a Californian species, coming north into Oregon only to a lim-ited extent (Map 2), and therefore largely beyond the scope of this paper* The two species appear to be distinct, though from Oregon very few speci-mens have been available on which to base conclusions* Californian mat-erial might establish the distinction with greater certainty. In general, the two seem to he separated by: 1* the glandular pubescent inflorescence of D. Jeffreyi, lacking in P. alpinume 2* the generally larger stature of D. Jeffrey!, in eluding the heav-ier tap root. This, of course, could be environmental. 49 3. the much smaller ratio of scape to leaves in D. Jeffrey!* 4. the clrcumscissile capsule of D. Jeffrey! as compared to the valvular dehiscence of D* alpinum* 5. the preference of D . Jeffrey! for lower altitudes than D. alpin* urn, though in Oregon this preference is less evident than i t apparently is farther south* Both species, however, vary in many of these characteristics. Four speoimens of D. Jeffrey! from southern Oregon, Applegate 6015, 4806, 4811, and 4809, are a l l glabrous and yet in other respects they belong with D. .Jeffrey! and have been included there* On the other hand, three speci-mens, Constance et al* 8979, and Applegate 7458 and 7459, also from south-ern Oregon, have scape to leaf ratios which hardly f i t D* alpinum. but they are glabrous and their capsules dehisce by valves* As capsule dehis-cence has been found to be one of the most constant characters in the genus, these three intermediates are included with D. alpinum with the reservation that they may be hybrids possibly. One other unstable characteristic in the alpinum group is the length of filament a and this is unusual as length of filaments, like mode of dehiscence, has proved generally very constant within a species* inform-ation regarding this trait in the California alpinum would perhaps throw more light on its value as a diagnostic characteristic for the species* P.. alpinum, then, as i t occurs in Oregon, is not too well defined* However, it is being maintained because its chief area of distribution is California, outside the present limits, and there i t is generally consider-ed a distinct species* The one attempt to subdivide each of the two entities, D. Jeff-50 -reyi and D. alpinum, has been made by Hall (1912), who recognizes, in Cal-ifornia, one variety end one form of D. Jeffrey! and one form of D. alpinum. His division into forms is based largely on stature and leaf length and he states that the differences may be due to ecological responses. Unless such differences could be shown by experiment to be gpnetic they seem scarcely adequate as a basis of speciation. His variety reddens, he says, is scarcely more redolent than typical D. Jeffrey!, but can be easily dis-tinguished from the latter by the degree of rolling of the corolla and by . the lack of a purple ring on the corolla tbroat. In the more northern specimens examined, both of these characters are variable. The lack of a purple scallop, combined with pentamery, has been used by Macbride and Payson as a basis for their D. exilifolium described from Idahoo Neither character, after examination of many specimens, seems constant enough to be of any specific value. Plants from both Idaho and Montana show a l l combinations of numbers of parts and distinctness of scallop, D. exilifolium has, then, been included under D. Jeffrey!. D. tetrandrum and D. viviparum, as described by Greene (1895), are apparently synonymous with D. Jeffrey!, there being nothing in their original descriptions which could not come within the range of variability of that speciese They are, therefore, here both treated as synonyms. Admittedly D. Jeffrey! is not, in many characteristics, a homo* geneous spe cies, but on those morphological points found most stable wl thin the genus, the many specimens examined agree very well* Minor characterist-ics, such as pubescence, size, number of stamens, and scallop on the cor-olla tube are so scattered throughout the extensive range of the species as to suggest mere variability rattier than incipient spe cies. Either the 51 variations are due to environment, or lack of isolation has permitted the spread of mutated genes* One possible isolating mechanism, chromosome number, which might have set up a barrier behind which speciation could proceed, has had, at least as yet, l i t t l e visible effects As indicated previously, counts made in material from California and Stevens Pass, Washington, show 22 pairs of chromosomes, the diploid number for the genus as so far ascertained, (Plate 2, 3)• Material from the Forbidden Plateau, Vancouver Island, and from Prince Rupert on an island near the British Columbia mainland, both show approximately 88 mitotic chromosomes, or the tetraploid number (Plate 4, 5)« Tet morphologically the difference is so l i t t l e that plants of the two groups are indistinguishableo The northern members are more constantly tetramerous, but in stature, flower characteristics, and habi-tat there is no differentiation to justify separating them taxonomioally* This similarity suggests autoploidy, possibly derived from the product of an intraspeciflc cross, since the first anaphase of meiosis in the only diploid studied, shows chromatin bridges and lagging chromosomes (Plate 6), though each pole appears to receive 22 chromosomes (Plate 7)* Such behavi-our during meiosis is characteristic of unlike gen cms, which are,however, not too unlike for bivalent pairing and regular separation to occur* With the morphological variation observed in the species, such intraspeciflc crosses could be expected* As the counts of tetraploid populations have been from somatic material only, they give no clue as to the type of poly-ploidy* An insufficient range of material has been studied to generalize too broadly, but the indication is that the southern representatives of PLATE 6. Dodeoatheon Jeffrey!, Stevens Pass, Washington Camera lucida drawing, first division anaphase of meiosis in a developing anther, showing ohxmatin bridges and lagging chromosomes* Ca. X 1700. PLATE 7 Dodeoatheon Jeffreyi. Stevens Pass. Washington Camera lucida drawing, fi r s t division late ana-phase of meiosis in a developing anther, showing regular segregation of chromosomes, 22 at each pole. Ca. X 2000 - 52 the species are diploid and the more northern polyploid. This fact sug-gests that polyploidy, i f i t has made l i t t l e morphologioal difference, has at least greatly increased the range of the species into the northern climates* Doubtless also the tetraploids are genetically isolated from the diploids, but being tetraploid, and therefore having four chromosomes of each kind, in this case a l l very much alike, mutated genes would take relatively longer in the former to become fixed. Therefore accumulation of morphological differences must be very slow. 5. Dodecatheon poeticurn Rend. Dodecatheon poetloum Hend*, Shod. 32:27, 1930. Plants glandular pubescent throughout, especially in the inflor- escence; rootlets fleshy-fibrous from a short crown; leaf blades elliptic to oblanceolate, 1.0-9.0 cm. long, 0.5-2.3 cm. wide, margins entire to dentate, apex narrowly rounded to acute, bass sharply contracted or narrow-ly cuneate and tapering to a winged petiole; petioles 2*0-9.0 cm. long; scape 12.0-28.0 cm. long; bracts 3.0-7.0 sm» long, tapering or abruptly contracted to rounded, acute, or acuminate apices; flowers 2-10, pen ta-mer ous; calyx tube 1.5-2.5 mm. long, lobes 4.0-6.0 mm. long, acute to acu-minate; corolla lobes rose-purple with a broad white area at their bases; reflexed tube bearing a distinct purple scallop; stamens dark purple, anthers 4.5-7.0 mm. long, filaments 1*0-2.0 mm. long, connectives smooth, abruptly contracted near their bases, tapering to acuminate apices; cap- sule narrowly ovoid, about equalling the calyx, glandular pubescent, de- hiscing by teeth, walls very firm, opaque; seeds unknown. 53 -Type locality: bluffs of the Columbia River hear the east line of Hood River County, Oregon. Habitat: open grassy slopes and moist stream banks. Distribution: Deschutes and Columbia River Valleys near their Junction, extending north into Klickitat and Yakima Counties of Washington. Map 3. Chromosome Counts no material available. Specimens examined: WASHINGTON: Klickitat Co.: Satus Pass region, May 14, 1938, J. Wm. Thompson, 14289 (B.C.); near Bingen, April 15, 1932, J. Wm. Thompson, 8154 (U.C, U.W.); Yakima Co.: h i l l s south of White Swan, May 16, 1942, B.F. Hoover, 5845 (U.C.); h i l l s south of White Swan, April 4, 1942, R.ff. Hoover, 5656 (U.C.); north fork of Ahtanum Creek, May 2, 1942, R.F. Hoover, 5755 (U.C.); 10 miles north of Satus Pass, April 50, 1953, CL. Hitchcook, V.L. Marsh, 5509 (U.C, U.W.) J Toppenisn Creek, Yakima Indian Reservation, April 8, 1952, V.T. Heidenrelch, 272 (U.W.); OREGON: Hood River Co.: between Ortley and Mosier, Columbia River, April 25, 1952, L.F. Henderson, 14505 (U.C); slopes east of Hood River, April 5, 1926, J. Wm. Thompson, 794 (U.W.); Wasco Co.; south1 side of the Columbia River, |f mile west of Mosier, April 12, 1936, L. Constance et al. , 1497 (U.C, U.W.); Mosier, April, 1891, M.W. Gorman, s.n. (U.W.); along the Columbia River west of the Dalles, Aprill 22, 1937, Sister Mary Mllburge, 1551, (U.W.); Siwash Flats, 2 miles west of Mosier, Columbia Gorge, April 7, 1928, J. Wm. Thompson, 4061 (U.W.); DISCUSSION OF D. POETICUM Dodeoatheon poeticum is a very small species centred around the junction of the Deschutes and Columbia Rivers, spreading north into Map 3. D o d e o a t h e o n p o e t i c u m 54 -Washington along the east slope of the Cascade Mountains* It resembles Dodecatheon Cusickii, especially in its glandular pubescence, and occurs on the western edge of the range of that species (Maps 3 and 6)» On the other hand, in its dark purple filaments, i t resembles Dodecatheon Hender- son! , and occurs on the eastern edge of the range of that species (Maps 3 and 4)* Hybridization is a possibility but, since, owing to lack of mat-erial, no cytologlcal work has been done on D» poeticum, no information is at hand regarding its genetic constitution. At any rate, i t appears to be maintaining its identity and extending its range, and i s , therefore, worthy of recognition* The question of whether i t should be an independ-ent species or a variety of D. Cusickii, which i t most nearly resembles, has been considered. The decision to consider i t of speoific rank has been made on the basis of the stability of filament colour within the genus. D» Henderson! and D. Cusickii, in the remainder of their range, show no variation; D. oonjigen3 and D. columbianum are likewise constant. In fact, the only evidence of variability in this characteristic has been found in southern Oregon in a half dozen specimens placed with D. pauciflorum. From this evidence, i t would seem that filament colour is determined by some mechanism more complicated than one or a few genes and that a variant in this respect, with some supporting differences In leaves, flower colour, and distribution, deserves more than varietal rank. D. poetlcum has, therefore, been maintained as a species. 6. Dodecatheon Henderson! Gray Dodecatheon Henderson! Gray, Bot. Gaz. 11:233, 1886. Dodecatheon lntegrlfollum Michx. var. latifolium Hook., F l . Bor. 55 -Am. 2:119, 1840. Dodeoatheon Meadia L. var. brevifolium Gray, In Geol. Sur. Cel. Bot., ed. 2, v. 1, 467, 1880, equivalent to ed. 1, 1876, see D. dentatum. Dodeoatheon latifolium (Hook.) Piper, Contr. U.S. Nat. Herb. 11:446, 1906. Herbage glabrous, inflorescence glandular pubescent; rootlets  fieahy-fibrous, often only a few millimeters long, from a short crown;  leaf blades sub-orbicular to broadly elliptic, 1.5-6.0 cm. long, 1.0-6.0 cm. wide, margins entire, apex rounded to broadly obtuse, base sharply contracted or broadly cuneate; petioles 0.5-7.5 cm. long; scape 13.5-40.0 cm. long; bracts 3.5*17.0 mm. long acuminate; flowers 2-7, 5-, occasionally 4-merous; calyx tube l.O^S.O mm. long, lobes 3.5-6.5 mm. long, tapering or sometimes sharply contracted to acuminate apices; corolla rose-purple with a distinct dark purple scallop on the re flexed tube; stamens  dark purple, anthers 3.0-5.5 mm. long, filaments 1.0-3.0 mm. long, connsot-tive narrowly acuminate; capsule broadly cylindrical, about twice the length of the calyx, glandular pubescent, circumscissile, walls firm, opaque; seeds 1.5-2.0 mm. long, wingless. Type locality: Tualatin Plains, Oregon. Habitat: light, open woodland, moist meadows, becoming dry in summer, sea level to 2000 f t . Distribution: from California north through Oregon and Washington west of the Cascade Mts.; Gulf Islands of Washingt-on, also southern Vancouver Island. Map 4. Chromosome count: from southern Vancouver Island, meiotic, n - 44-50 (Plate 8). Map 4 . D o d e c a t h e o n H e n d e r s o n ! PLATE 8. Dodaoatheon Hendersoni, southern Vancouver Island. Camera lucida drawing, first division prophase of meiosis in a developing anther, showing 44-50 pairs of chromosomes. Ca. X 2000 - 56 Specimens examined: BRITISH COLUMBIA: Victoria, March 29, 1915, C.F.N., s.n. (P.M.); Victoria district, April 21, 1930, W.A. Newcombe. 8* n* (P.M.)j Viotoria, April 19, 1900, A.J. Plneo, s.n. (U.C.); Oak Bay, Victoria, April U, 1895, A.J. Pineo, s.n. (U.B.C.)} Victoria, islands south May 1, 1917, Mrs. J.T. Higglns, s.n. (U.B.C.) ; Victoria, April 9, 1912, J.K. Henry, s.n. (U.B.C); Shawnigan Lake, J.K. Henry, s.n. (U.B.C.); Victoria, June 12, 1913, W. Taylor, s.n.(U*B.C.h near Victoria, April , 1938, J. Bnckland, a.n. (U.B.C.); Cadboro Bay, V.I., April 11, 1913, Bn.  Taylor, s.n. (U.B.C.); Beacon H i l l Park, Victoria, April 10, 1938, J.W.  Bastham, s.n. (U.B.C); Beacon H i l l Park, Victoria, April 7, 1938, J.W.  Bastham, s.n. (U.B.C.); Quarantine Lake, V.I. April 28, 1938, T.T. Mo Cable, 5696, (U.C.); Colwood, V.I., April 28, 1938, T.T. MoCabe, 5737 (U.C.); 1 mile south of Sooke, V.I., April 27, 1938, T.T. McCabe, 5650 (U.C); Uplands, Victoria, May 11, 1950, K.I. Beamish, 50100 (U.B.C.); Mt. Douglas near Victoria, May 3, 1949, K.I. Beamish, 49100 (U.B.C.); Duncan, V.I. May 14, 1950, D. Charter 210 (U.B.C.); Mt. Tzouhalam, near Duncan, V.I. June 4, 1950, D. Charter s.n. (U.B.C.); Point Gonzales, V.I., April 7, 1900, J.R. Anderson, s.n. (P.M.); Point Gonzales, V.I., April 22, 1899, J.R. Anderson, s.n. (P.M.); Thetis Lake, V.I., May 12, 1901, J.R. Ander-son, s.n. (P.M.); Thetis Lake, V.I., May 2, 1903, J.R. Anderson, s.n. (P.M.); Saanich Aim, V.I., April 9, 1899, J.B. Anderson, 512 (P.M.); Cedar H i l l , V.I., April 7, 1895, J.R. Anderson, s.n. (P.M.); Baauimalt, V.I., May 10, 1896, J.R. Anderson, s.n. (P.M.); Seymour H i l l , V.I., April 6, 1924, C.F. Newcombe, s.n. (P.M.); Skirt Mt., V.I., May 24, 1896, C J .  Newcombe, s.n. (P.M.); Mt. Newton, V.I., May 19, 1940, W.V. Hardy, 12536 (P.M.); Lost Lake, Saanich, V.I., April 13, 1946, G.A. Hardy, 19508 (P.M.); 57 Langford, V.I., May 10, 1929, Mrs, Donald, s,n. (P.M.); Baaver Lake, Mt, Newton, V.I,, April 9, 1914, John Maooun, s.n. (P.M.); Victoria, Burnaide Rd., May 8, 1893, Maooun, s.n. (N.M.); Victoria, Christmas H i l l , April 27, 1908, John Maooun, s.n. (N.M.); Victoria, April 13, 1908, John Maooun, s.n. (N.M.); Victoria, May U, 1906, John Maooun, s.n. (H.M.); Cedar H i l l , V.I., May 24, 1887, Maooun, s.n. (N.M.); Yale, May 17, 1875, Maooun, s.n. pro parte (N.M.); Mt. Newton, Sydney, V.I., May 25, 1912, J.M. Maooun, a»n* (N.M.); Cedar H i l l , V.I., March 16, 1921, G.V. Copley, s.n. (P.M.)} Cedar H i l l , V.I., March 17, 1915, C.F.N. & F.K., s.n. (P.M.)} Mt. Tolmie V.I., May 16, 1933, F. Kermode, s.n. (P.M.); "Fowl*1 Bay, Victoria, April 7, 1918, W.R. Carter, s.n. (P.M.); near Victoria, June 1, 1872, Cowley, a.n. pro parte (N.M.). WASHINGTON. Cowlitz Co.: along the Columbia River near Kalama, April 7, 1934, J. Wm. Thompson, 10121 (U.W.); Island Co.: Whidby Island, NJ.. Gardner, s.n. (U.C.)} Whidby Island, April 17, 1897, N.L. Gardner, s.n. (U.W.); Lewis Co.: Centralia, April 23, 1928, R.M. Owen,a. n. (U.W.); Mason Co.: Scott's Prairie near She It on, May 11, 1935, George  Neville Jones, 6493 (U.W.); near Shelton, May 4, 1935, J. Wn. Thompson.  11435 (U.C.)} Pierce Co.: Nisqually Indian Reservation, May 16, 1937, W.J. Syerdam, a.n. (U.0«); Mt. Rainier, Aug. U , 1894, J.B. ZLott, s.n. (U.W*); Skamania Co.: Hood, 5 miles west of Bingen, April 15, May 21, July 9, 1909, Wllhelm Suksdorf, 6446 (U.C, U.W.); Thurston Co.; Granger's Prairie, vicinty of Olympia, May 2, 1905, B.C. Townaend a.n. (U.C.)} Baker'a Prairie, April 8, 1934, I.C Otis, 1850 (U.W.); OREGON: Benton Co.; -Cemetery H i l l , Corvallls, April 22, 1955, Helen Maw, a.n. (U.C.); Curry Co.: about 5 miles south of Agness, Rogue River, May 10, 1952, E.I. Apple- gate, 7128 (U.C)} Rogue River Canyon, south of Illahe, March 19, 1948, Tttn. H. Baker, 5194 (U.C.}; Douglas Co.; summit of ridge west of Drain. May 11, 1929, H«L. Maaon, S294 (U.C.); Roseburg, Urapqua Valley, April, 1887, Thomas Howell, s.n. (U.C.); west of Hoseburg, April 7, 1927, J. Wm. Thompson, 2029 (U.W.); Jackson Co.: headwaters of Emigrant Creek, north slope of Siskiyou Mts., south of Ashland, March 28, 1926, Elmer I. Applegate, 4580 (U.C, U.W.); Bun comb, Little Applegate River at the mouth of Sterling Creek, April 16, 1926, B.I. Applegate, 4598 (U.C); Jacksonville, April 7, 1925, S.I. Apple- gate, 4150 (U.C); Tolman Springs, upper Emigrant Creek, March 28, 1926, B.I. Applegate, 4579 (U.C, U.W.); ridge between Emigrant and Tyler Creeks, 12 miles s.e. of Ashland, March 29, 1926, B.I. Applegate, 4585 (U.C.); east of Brownsboro, April 8, 1927, J.Ttta. Thompson, 2155 (U.W.); Josephine Co.: near Waldo, April 10, 1925, Elmer I. Applegate, 4183 (U.C); west fork of William's Creek, April 13, 1940, L.E. Pet ling, 4005 (U.C.); I l l i n -ois Valley, April 17, 1959, E.A. Meola, 24 (U.W.); Grant's Pass, April, Thomas Howell, 1210 (U.C); Lane Co.: Willamette River, 1 mile above Lowell, April 7, 1956, L. Constance et al. , 1485 (U.C, U.W.); meadows along Crow River, April 16, 1932, L.F. Henderson, 14257 (U.C); Marion Co.: Salem, O.B. Johnson, s.n. (U.W.); Washington Co.: near Forest Grove, March 9, 1926, J. Win. Thompson, 532 (U.W.); Tualatin Plains, May 1883, L.F. Hend- erson, 642, isotype (U.C.); Yamhill Co»: May 1874-9 Rev. R.W. Summers, 2165 (U.C); DISCUSSION OF D. HENDERSONI Dodeoatheon Hendersoni in the Northwest is a well-differentiated group, maintaining its identity even where its range overlaps that of D.  pauciflorum var. macro carpum on Vancouver Island. It is apparently a - 59 common California species but is found also north through Oregon and Wash-ington west of the Cascade Mountains, on the Gulf Islands of Washington, and on southern Vancouver Island. No specimen has been seen from farther north than Duncan, Vancouver Island, nor from east of the Cascades. One, Macoun s.n., May 17, 1875, is said to be from Tale, B.C. Yale is some 185 miles up the Eraser Hiver (Map 4) from Vancouver. The specimen is authen-ti c , but, as D. Hendersoni has appeared nowhere else on the southern B.C. mainland, the location may be wrong. In at least two respects D. Henderson! has been found to be un-like any other northern member of the genus. 1. Its roots are of the short crown type with fleshy rootlets which break away each year to form new plants. However, many of these rootlets are extremely short, much shorter than those of other species of the seme group. 2. Meiosis occurs much earlier in the floral development than in other species. In a l l others from which meiotic counts have been obtained, satisfactory stages could be found in the younger flowers of an umbel after the scape was fully grown, or, in a large umbel, after some flowers had opened. In D. Henderson!, meiosis is completed before the umbel is well above ground level. This early maturation is probably one means by which D. Henderson! is Isolated from D. pauciflorum var. macro carpum where the two grow almost side by side in the vicinity of Victoria, Vancouver Island. Another point of interest, in which D. Henderson! resembles D.  Jeffrey!, i s the apparent distribution of polyploidy. Again, generaliza-tion is unwise, but an indication of the situation can be obtained from chromosome counts. Seeds from California show the diploid number (approx-- 60 -imately 44 chromosomes). Unfortunately, the material was not sufficiently clear for drawings to be made, but the count is considered reasonably accurate. From meiotio material obtained from southern Vancouver Island, the count is tetraploid, n - 44-50 pairs. (Plate 6)» The uncertainty is due to difficulty in resolving certain associations in the cells. The complexity of these groupings suggests the possible presence of t r i - or tetravalents. Such a condition may well be expected, for as in D. Jeff- rey!, the diploids (presuming the California representatives of the species to be diploid) and tetraploids look very much alike. D. Henderson!, then like D. Jeffrey!, suggests autoploidy. 7. Dodecatheon conjugens Greene. Dodecatheon conjugens Greene, Eryth. 3:40, 1895. Dodecatheon Henderson! Gray var. leptophyllum Suks., Deutsch. Bot. Monatss. 18:132, 1900. Dodecatheon viseidurn Piper, Bull. Torr. Bot. Club 28:43, 1901. Dodecatheon conjugens Greene var. leptophyllum (Suks.) Piper, Oontrib. U.S. Nat. Herb. 11:446, 1906. Dodecatheon conjugens Greene var. viscidum (Piper) Mason ex St. John, F l . S.E. Wash. Adj. Ida. 311, 1937. Plants of moderate stature, usually not over 20 cm. t a l l , glabrous throughout except leaves rarely glandular pubescent; rootlets  fleshy-fibrous from a small crown; leaf blade elliptical to oblanceolate, rarely linear, 1.0-5.5 cm. long, 0.6-1.8 cm. wide, entire, apex rounded to acute, b a B e sharply contracted to the petiole, less commonly narrowly cuneate and tapering; petioles 1.5-4.0 cm. long; scape 6.5-17.0 cm. long; - 61 -bracts 1.5-9.0 mm. long, broadly rounded to acuminate, papery; flowers ]3*6 5- (rarely 4- or 6-Omerous; calyx tube 1.5-2.8 mm. long, lobes 2.5-6.5 mm. long, broadly acute to narrowly acuminate; corolla lilac-purple or, less commonly, white, with a narrow dark purple scallop on the re flexed tube; anthers 4.5-8.5 (usually 5.0-;7.0)mm. long, purple, filaments under 1.0 mm.  long, yellow, free or fused, connective transversely rugose; capsule cyl-indrical, about twice the length of the calyx, circumscissile, walls firm, opaque; seeds 1.0-2.0 mm. long, wingless. Type locality: Helena, Montana. Habitat; open meadow or light woodland, 4000-5500 ft . in Washington, up to 8000 f t . in southern Oregon. Distribution: dry areas east of the Cascade Mts., across central and southern Washington, Oregon, Idaho, and into Montana. Map 5. Chromosome count; no material obtained. A few glandular plants, otherwise resembling D. conjugens, appear in western Washington. They have been described by Piper (1901) from Spangle as D. viscldum and reclassified by Mason ex St. John (1937) as D.  conjugens var. viscidum. However, in the herbarium material examined, very few such specimens were found. Those present appear to be typical D. con- jugens except that they are glandular pubescent especially on the upper part of the scape and in the inflorescence. In this respect they seem id^ » entloal with D. Cusickii. Since both D. conjugens and D. Qusjckiisare found in the vicinity of Spangle, the viscid plants may well be hybrids. For this reason, and since they most resemble D. conjugens, they have been, for the present at least, included here. Specimens examined, showing this glandu* Map 5 . Dodecatheon conjugens . . , . circles Dodecatheon columbiaT"1™ » . . squares 62 lar pubescence, are listed by themselves after those of typical D. conjug- ens. A second variant within D. conjugens .is the narrow leaved plant described as D. conjugens Greene var. leptophyllum (Suks.) Piper. It appears along the east slope of the Cascade Mountains in southern Washing-ton. However, i t seems to be extremely variable, especially in Grant and Lincoln Counties from which narrow and normal leaved specimens appear on the same sheets, and even single plants are not uniform, (Rogers et a l . 235, 236, 255). This variant has also been included with D. conjugens. Specimens examined. WASHINGTON: Columbia River at Clark's Ferry, Apr-i l 9, 1926, G.N. Jones 1598 (U.W.); Asotin Co.: northeast slope of Blue Mts., April 24, 1938, C.W. Sharsmith, 5550 (U.C); Chelan Co.: 10 miles north of Bntiat, April 18, 1931, J. Wm. Thompson, 5986 (U.W.); Douglas Co.: plains around Douglas, April 25, 1951, Elmer I. Apple gate, 6690 (U.C); Garfield Co.: Blue Mts., June 10, 1938, Don L. Peters, 568 (U.C); Blue Mts., June 13, 1936, L. Constance & H.F. Clements, 1751 (U.W.); Blue Mts. , 24 miles south of Pomeroy, May 26, 194 4, C.L. Hitchcock & C.V. Muhlick, 8518 (U.W.); Grant Co.: flats near Coulee City, April 21, 1955, R.C Roll- ins, 855 (U.C); flats near Coulee City, April 21, 1955, R.C. Rollins, 856 (U.C.); Coulee City, April 2L» 1955, E. Zaring, 73 (U.W.); Grand Coulee, April 3, 1940, H.T. Rogers et a l . , 256 (U.C, U.B.C); Grand Coulee, April 5, 1940, H.T. Rogers et al., 235 (U.C, U.W., U.B.C.); Kittitas Co.: Table Mt., 12 miles northeast of Liberty, June 12, 1938, C.L. Hitchcock et a l e ,  3624 (U.C,, U.W.); "Ellensburg (?)", April 23, 1899, C.7. Piper, 1018 (U.W.); north of Ellensburg, April 23, 1932, J. Wm. Thompson, 8225 (U.W.); 15 miles north of Ellensburg, April 4, 193k, J. Wm. Thompson, 5943 (U.W.); 63 Klickitat Co.: north of Goldendale, April 13, 1935, J. ttta. Thompson, 11325, (U.Wf; Falcon Valley, May 12, June 24, 1895, W.N. Suksdorf, 2202, s.n.. type or isotype of D. Hendersoni Gray var. leptophyllum Suks. (U.C.); mountains, west Klickitat Co., April, 1885, W.N. Suksdorf, 545 (U.C); Lincoln Co.S south side of Columbia River at Keller Ferry, April 4, 1940, H.T. Rogers et al . , 246 (U.C, U.W., U.B.C.); south side of Spokane River at its mouth, April 5, 1940, H.T. Rogers et al., 255 (U.C); south side of Spokane River at its mouth, April 5, 1940, H.T. Rogers et al., 256 (U.C.); Spokane Co.: Spokane, April 16, 1925, T.A. Bonser, s.n. (U.W.); open woods near Spokane, April 28, 1952, Sister Mary Milburge, 275 (U.W.); Walla Walla Co.: waste lands northeast of Walla Walla, April 9, 1956, WJR. Moore, 2 (U.C); Whitman Co.: south of Pullman, May 19, 1917, F.L. Pickett, 1098 (U.W.).; Pullman, May, 1895, L. Smith, s.n. (U.W.).; Yakima Co.:* Satus Pass, March 26, 1934, G.N. Jones, 4443 (U.C); near Wenas, April 15, 1940, J. Wm. Thompson, 14495 (U.C, U.W.); "Umptonum" Ridge, May 3, 1936, Harold W. Smith, 448 (U.W.); Klickitat meadows, June 19, 1899, J.B. Flett. 1210 (U. W.); OREGON: hillsides, eastern Oregon, May 25, June 8, 1898, Wm. C.  Cusick, 1845 (U.C); Harney Co.: Summit of Steens Mt., head of a tributary of Wildhorse Creek, May 22, 1929, Elmer I. Applegate, 5678 (U.C); head-j waters of Wildhorse Creek, summit of Steens Mt«, May 22, 1929, Elmer I.  Applegate, 5669 (U.C*).; west base of Steens Mt., May 20, 1929, Elmer I. Applegate, 5588 (U.C); south of Burns, May 2, 1927, L.F. Henderson, 9020 (U.C); Squaw Butte, May 1, 1941, G.J.C., s.n. (U.W.); Jackson Co.: between Jenny Creek and Keene Creek, Cascade Mts., April 14, 1925, Elmer I. Apple-r  gate, 4255 (U.C); Keene Creek Prairie, near summit of Cascade Mts., May 16 1985, Elmer I. Applegate, 4261 (U.C); Cascades, east of Ashland, July, 64 -1893, Mrs. R.M« Austin, 215, pro parte, (U . C . ) ; Klamath Co0s head of Denny Creek, Cascade Mts«, west of Klamath Lake, April 24, 1926, Elmer I, Apple gate, 4605 (U.C.); Fossil Lake region, between Ft. Klamath and the Summit, 1901, H.W. Furlong et. ale, s»n. (U.C.); Lake Co*: slopes of Little Honey Creek, eastern Oregon, May- 4, 1928, May Loveless, 86 (U.C.); Wasco Co*: near mouth of Deschutes River, Columbia Gorge, April 9, 1928, J*Wm»  Thompson 4086, (U.W.); IDAHO: Idaho Co*: east side of Snake River Canyon near mouth of Granite Creek, April 3, 1935, L. Constance et. a l a , 1010 (U. C ) ; 1$ miles up Granite Creek, Snake River Canyon, April 3, 1938, Jacque- line Packard, 567 (U.C); Owyhee Co*: 4 miles above convergence of Squaw Creek and Snake River, Snake River Canyon, May 13, 1937, Fred G.Meyert857 (U.W.); MONTANA: Fergus Co.: top of Grayhouse Peak, Big Snowy Mts., July 3, 1947, C.L. Hitchcock, 16072 (U.C, U.W.); Gallatin Co*: Bridger Canyon, April 21, 190O, Wyatt W. Jones, s.n. (U.C); Bozeman, April 29, 1901, Wyatt W. Jones, s.n. (U.C.); Bozeman, May 8, 1906, J.Wo Blankinship, 541a, (U.C.); Bridger Canyon, Bozeman, May 16, 1901, J.W. Blankinship, s*n.(U.W.); Bridger Canyon, Bozeman, May 27, 1899, J.W. Blankinship, s*n. (U.W*); Lewis and dark Co*: Helena, E.N. Brands gee, s.n. pro parte (U.C); S t i l l -water Co.: three and a half miles south of Dean post office, May 5, 1957, George Payer, 547 (U.W.). Glandular pubescent specimens: WASHINGTON: Spokane Co*: near Spsa-gle, April 26, May 17, June 50, 1916, Wilhelm Suksdorf, 8554 (U.CpU.W.); Spokane, May 1931, Sister Mary Milburge, s.n. (U.W.); near Colbert, April 28, 1932, Sister Mary Milburge, 274 (U.W.). 65 -8. Dodeoatheon columbianum n.sp. Herbage pubescent with long glandular hairs, inflorescence and upper part of scape glabrous; rootlets fleshy-fibrous from a small crown; leaf blades e l l i p t i c , oblanceolate, or spatulate, 2.0-12.5 cm. long, 0.8-3.5 cm. wide, apex rounded, rarely obtuse, base sharply contracted or narrowly cuneate, tapering to a winged petiole; petiole 0.8-7.5 cm. long; scape 3.0-20.0 cm. long; bracts 1.5-7.5 mm. long, acute to acuminate; flowers 1-6, pentamerous; calyx tube 1.5-3.0 mm. long, lobes 3.0-5.5 mm. long, narrowly acute to acuminate; corolla lobes rose or white, reflexed tube bearing an irregular purple line;Panthers 4.5-6.5 mm. long, purple,  usually becoming yellow toward their apices, filaments 0.5-1.0 mm. long, yellow, connective broad, transversely rugose, purple, becoming yellow at apex and bass; capsule cylindrical, about twice the length of the calyx, circumscissile, walls firm and opaque; seeds 1.5-2.0 mm. long, wingless. . Type locality: 1 mile due north of Natal, B.C., 4000 f t . , on a side h i l l , clay loam s o i l . Type: Crowsnest Pass region of B.C., May 11, 1950, A. Dzubin s.n. (U.B.C.). Habitat: grassland or open woods, 4000 - 7500 f t . Distribution: Rocky Mts. of Idaho, Montana, southeastern British Columbia, and southwestern Alberta; across southern Alberta to Saskatchewan. Map 5. Specimens examined: BRITISH COLUMBIA: mountains north of Crowsnest Pass, June 1, 1938, T.T. McCabe, 6503 (U.C); four miles east of Tffaaa,, May 18, 1938, T.T. McCabe, 6109 (U.C); Hewgate, May 16, 1943, W.B.. John-66 stone, B « n . (U.B.C.j; Newgate, close to Montana line, May 16, 1943, "Aw B. Johnstone, 15557, pro parte (EM.); Crowsnest Pass region, 1 mile due north of Natal, May 11, 1950, A. Dzubin, s.n. (U.B.C); Crowsnest Pass region, Natal, May 16, 1950, A. Dzubin, S.N. (U.B.C.); Cranbrook, April 11, 1915, C.B. Garrett, s.n. (P.M.); ALBERTA: Red Rock Canyon, Waterton Lakes Park, June 18, 1955, E.H. Moss, 5144 (U.B.C); Bobs Creek, Porcupine Hills, June 5, 1920, W.D. Cram, s.n. (N.M.); Jumping Pound Creek, June 11, 1897, Maooun, s.n. (N.M.); Vermillion Mt., Banff, July 6, 1891, Maooun, s.  n. (N.M.); Waterton Lake, Rocky Mts., July 50, 1895, Maooun, s.n., pro parte, (N.M.); Mt. Ingliamaldie, Banff District, July 18, 1916, D.Pelluet,  287 (N.M.); Banff, July, 1915 D. Pelluet, 91 (N.M.); SASKATCHEWAN; Cypress Hills Park, May 26, 1949, A.C Budd, 1299 (U.B.C.); Cypress Hills Park, May 27, 1949, A.C. Budd, 1510 (U.B.C.); IDAHO: Fremont Co.: 1 mile below Flat Rock Camp, south of west entrance to Yellowstone, 6 miles north of Pond Lodge, June 28, 1938, CJ.. Hitchcock et alo, 3850 (U.W.); Juniper Hills near St. Anthony, June 9, 1919, B.H. Q,uayle, 45 (U.C.); MONTANA: Flathead Co.: 1 mile south of Columbia Falls, April 12, 1941, H.T. Rogers 774 (U.C, U.W,}; Lewis and Clark Co.: south of Gibson Lake, about 25 miles northwest of Augusta, July 4, 1948, CL. Hitchcock, 18001 (U.C, U.W.); Meagher Co.5 Monument Peak, Little Belt Mts., Lewis and Clark National Forest, June 27, 1957, R. Bradley, 63 (U.W.); Missoula Co.: April 11, 1921, J.B. Kirkwood, 1242 (U.C); Missoula, Blackfoot Valley, near Ninemile Bridge, May 27, 1923, J.E. Kirkwood, 19595 (U.C); Miller Creek, 4 miles south of Missoula, April 20, 1938, F.A. Barkley, 2675 (U.C); Missoula, May, 1917, J.A. Hughes, 1241 (U.C.); Missoula, April 22, 1940, W. Dufour, 2 (U.C.) - 67 DISCUSSION OF D. CONJUGENS MD D. CGLUMBIANUM D. conjugens and D. columbianum stand more or less intermediate between the two main groups which are: those species having a tap root and very short filaments, and those species having a short crown and f i l a -ments at least 1 mm. long. The two species mentioned above possess the short crown, but their filament length corresponds to neither group, the filaments being evident, but usually only 0.5-0.8 mm* long. No live mat-erial of D. conjugens has been obtained and therefore nothing is known of of its chromosome count. D. columbianum is diploid, having 22 pairs of chromosomes from a count of first division anaphase. Unfortunately, no drawing has been made. With the exception of the variants mentioned under its descrip-tion, D. conjugens appears to be a uniform speoies extending from Montana westward through the Eastern Cordilleran System and across the Interior Plateaux to the Cascades Mountains. No specimens have been seen from west of the Cascades nor as far north as Canada* D. columbianum shows considerable variation in leaf shape, but otherwise appears uniform. The distinguishing characteristics of the two follow: D. conjugens D. columbianum 1. anthers 6.0-7.0 mm. long. anthers about 5.0 mm. long. 2. glabrous or nearly so. leaves thickly glandular pubesc-ent, inflorescence glabrous. 3. anthers dark purple throughout tips and bases of anthers yellow 4* glabrous or nearly so. leaves thickly glandular pubescent inflorescence glabrous. « 68 5* leaf apices acute* leaves almost spatulate* The ranges of the two overlap in Montana (Map 5), hut D . con- jugens extends west, p. columbianum north and east, and in the greater pert of the area of each, the other has not been found, P. columbianum is, with some hesitation, being treated as a new species« The east slope of the Rocky Mountains in Montana B e a m s to be an area of considerable variation in at least one species, P. pauciflorum, and an area where several species overlap* Rydberg (1900) has described, from this area, a number of so-called species, a l l apparently very much alike and a l l from the vicinity of Sand Coulee* Possibly one of these is the same as the group here called P. columbianum* Only study of the types can determine this point with certainty* If such were the case, the pres-ent name would, of course, be invalidated. However, since the material fits none of the Rydberg descriptions accurately, the new name is being applied pending further study. 9. Dodgdatheon Cusickii Greene Dodecatheon Cusickii Greene, Pitt* 3:73-74, 1890. Dodecatheon Meadia var. puberula Nutt., Journ. Acad* Nat. Sci. Phil* 7:48, 1834. Dodecatheon puberulentum Heller, Bull* Torr* Bot. Club 24:311, 1897. Dodecatheon puberulum (Nutt.) Piper, Contr. U.S. Nat. Herb. 11:445 1906. Dodecatheon Cusickii Greene var. album Suksd*, Werd. 1:30, 1927. Dodecatheon pauciflorm (Dur.) Greene var. Cusickii (Greene) Mason ex St. John, F l . S.E. Wash. Adj. Ida. 312, 1937. 69 Dodecatheon pauciflorum (Dur.) Greene var. Cusickii (Greene) Mason f. album (Suksd.) St. John, F l . S.E. Wash. Adj. Ida. 3 1 2 , 1 9 3 7 . Plants glandular pubescent throughout, especially in the inflor- escence; rootlets fleshy-fibrous from a small crown; leaf blades el l i p t i c , oblanoeolate, or linear, 1 . 5 - 5 . 5 cm. long, 0 0 3 » 1 # 9 cm. wide, margins entire, rarely dentate, apex rounded to acute, base narrowly cuneate, tapering to a variably- winged petiole; petioles 1 . 0 r 6 . 3 cm. long; scape 8.5-33.Q aaa. long; bracts 1 . 5 * 7 . 0 mm. long, broadly rounded to acuminate; flowers 2 - 1 4 , pentamerous; calyx tube 1 . 0 - 2 . 5 mm. long, lobes 2 . 0 - 5 . 5 mm. long,rounded to acuminate; corolla vivid rose-pink to white, with a fine red-orange scallop  on the reflexed tube; anthers 3 » 5 e 6 « 0 mm. long, yellow, noticeably contract- ed below the anthers; connective smooth, purple, often sharply contracted near the base to a narrowly acuminate apex; capsule narrowly ovoid, l-l£ times the length of the calyx, glandular pubescent, dehiscing by teeth, walls firm, opaque; seeds minute, 0 . 5 - 0 . 8 mm. long, wingless. Type locality: dry mountain ridges of eastern Oregon, 4000 f t . Habitat: blooming in moist grassy meadows of open woodland in areas becoming extremely dry in summer. Type: Cuslck 1527 Distribution: Interior Plateaux of south central and southeastern British Columbia; Washington, Idaho, and northern Oregon, less commonly extending into Montana. Map G, Chromosome count: from Princeton, B.C., meiotic, n - 23 or 24 (PI. 1 5 , 1 6 , 1 7 , 1 8 ) . Specimens examined: BRITISH COLUMBIA: Lake Botanie, Marble Mts. June 2 0 - 2 5 , 1 9 3 8 , J.W. and EJfl. Thompson, 72 (U.C, U.W.); south of Earn-Map 6 Dodeoatheon Cusickii PLATS 15 Dodeoatheon Cuaicfcii, Princeton, B.C. Camera lucida drawing, first division pro-phase of meiosis in a developing anther, showing 23 pairs of chromosomes. Ca.X 1800 Dodeoatheon Caslckll, Princeton, B.C. Photomiapograph, fi r s t division prophuee of meiosis In a developing anther, showing £9 pairs of chromosomes. Ca. X 10CO. PLATE 17 Dodeoatheon Cusick!l t Princeton, B.C. Camera lucida drawing, first division pro-phase of meiosis in a developing anther, showing 24 pairs of chromosomes. Ca.X 1600 PLATE 18 Dodecatheon Cusic&li, Princeton, B.C. Photomicrograph, same cell as Plate 17. Ca. X 1000 70 loops, May 29, 1938, J.W. and B.M. Thompson, 46& (U.W.); near International Boundary, Midway, April 11, 1905, W. Spreadborough, s.n. (N.M.); Kanaka Ban Eraser River, June 8, 1883, H i l l , a.n. (N.M.); Penticton, Okanagan, April 30, 1903, J.M. Macoun, s.n. (N.M.); Lytton, April 17, 1889, J.M«Macoun,s.n. (N.M.); Spence's Bridge, May 28, 1889, Macoun, s.n. (N.M.); Trail, May 20, June 18, 1902, J.M. Maooun, s.n. (N.M.); Meyer's Creek, west of Midway, May 10, 1905, J.M. Macoun, s.n. (N.M.); Sproat, June 20, 1890, Macoun s.n. (N.M.); Sproat, June 24, 1890, Macoun, s.n. (N.M.); EagLe Pass, west of Revel stoke, May 23, 1890, Macoun, s.n. (N.M.); west of Sophie Mt., July 11, 1902, J.M. Maooun, s.n. (N.M.)j Silver King Mine, Nelson, July 7, 1940, J.W Eastham, s.n. (U.B.C.); Penticton, May 3, 1939, J.W. Eastham, s.n. (U.B.C); Twin Lakes Valley, Penticton, May 10, 1940, J.W. Eastham, s.n. (U.B.C); autonit Anarchist Mt., Osoyoos, May 11, 1940, J.W. Basthsm, s.n. (U.B.C); Newgate, May 16, 1943, W.B. Johnstone, s.n. (U.B.C, Acc. No. 10782); Wardner, 49°N. 115°W., July 2, 1915 B.F. Murray, s.n. (U.B.C.); Vernon, April 27, 1913, W.H. Brit tain, s«n. (U.B.C.); Monte Creek, South Thompson, May 16, 1937, J. Bostook, 3>n. (U.B.C.); Kemloops, April 14, 1917, J.A. Wattle, s.n. (U.B.C); O^ilchena, 49°N. 120°W., June 22, A.H. Hutchin- son, s.n. (U.B.C); Salmon Arm, May 18, Miss Pits, s.n. (U.B.C}; Salmon Arm, May 17, 1917, Miss Pits, s.n. (U.B.C.); Tranquilla, May, 1937, V.C  Brink, s.n. (U.B.C); Botanie Valley, July, 1917, F. Perry, a.n. (U.B.C.); Okanagan, April 30, 1905, E l i Wilson, 441 (U.B.C.) Crawford Bay, May, 1940, H. Murray, s.n. (U.B.C); Erickson, Creston, May, 1940, H. Murray, s.n. (U.B.C); Erickson, Creston, May, 1940, E. Murray, s.n. (U.B.C); Botanie Valley, Lytton, 50°N. 121°W., May 30, 1949, T.M.C. Taylor, 2009 (U.B.C.); Merritt, 50°N. 120°W., June 2, 1949, T.M.C. Taylor, 2064 (U.B.C.); upper 71 Skoonkon Valley, June 13, 1914, Jo Davidson, Sen. (U.B.C.); Brisco, 50°N. o 116 W., E. Clark, s.n* (U.B.C.); between Pinantan and Pritchard, June 13 1935, T.T. McCabe, 2311B (U.C.); Anarchist Mt., Osoyoos, May 7, 1938, T.T.  McCabe, 5946 (TIC.); 3 miles north of Fort Steele, May 16, 1938, T.T. McCabe, 6832 (U.C.); 2 miles south of Merritt, May 3, 1938, T.T. McCabe, 5818 (U.C.) Fairview Rd., 7 miles west of Oliver, May 3, 1948, T.T. McCabe, 5807 (U.C); 3 miles below Lytton, junction of Eraser and Thompson rivers, April 19, 1934, T.T. McCabe, 825 (U.C.); 20^ miles east of Kamloops, May 14, 1935, T.T. McCabe, 2025 (U.C.); 6 miles northwest of and 500-1000 f t . above Pritchard, May 14, 1935, T.T. MoCabe, 2012 (U.C.); Alkali Lake Rd., near Junction with Chilcotin Rd., May 7, 1934, T.T. MoCabe, 919 (U.C); about 5 miles west of Lytton, April 17, 1935, T.T. McCabe, 1847 (U.C,); Fish Lake, south of Ksmloops, June 20, 1890, McEvoy, s.n. (K.M.); Vernon, April 20, 1898, Mrs. Dr. Morris, s.n. (U.B.C); Rossland, Kootenay, 1895, J. He si op, s.n. (U.B.C); Lytton, April 25, 1924, W.B. Anderson, s.n. (P.M.); Chase, May 23, 1922, W.B. Anderson, s.n. (P.M.); Adam's Lake, April 22, 1925, W.B.  Anderson s.n. (P.M.); Adam's Lake, April 28, 1924, W.B. Anderson, s.n. (P.M.) Kelowna, April 30, 1901, J.R. Anderson, s.n. (P.M.); Penticton, June, 1900, Miss E l l i s , s.n. (P.M.); near Princeton, May 28, 1950, G.A. Hardy,1 22136 (P.M.); Newgate close to Montana line, May 16, 1943, WAW.B. Johnstone, 15557, pro parte (P.M.); Lillooet, May 5, 1916, E.M. Anderson, s.n. (P.M.); Okanagan Landing, May 2, 1915, J .A. Munro, s.n. (P.M.); Anarchist Mt., May 1941, CO.. Carl, 13775 (P.M.); WASHINGTON: Adams Co.: below Palouse Falls, south west side of Palouse River, April 18, 1936, L. Constance and R.C.  Rollins, 1505 (U.C, U.W.); north bank of Palouse River, 3 miles east of Hooper, April 18, 1936, L. Constance and R.C. Rollins, 1498 (U.C , U.W.); 72 1-3/4 miles southeast of Maoall, April 18, 1946, R.G. Jeffrey, s.n. (U.W.); Asotin Co.: ^  mile south of Indian Tarn Creek, Blue Mts., June 1, 1937, L. Constance et al., 1869 (U.C); Chelan Co.: Cooper Mt. Rd., near Chelan, May 2, 1936, O.T. Edwards, 206 (U.C.); Tumwater Mt., Wenatchee Mts., May 23, 1931, J. Wm. Thompson, 6516 (U.W.); Tumwater Canyon near Leavenworth, April 24, 1932, C.B. Seeley, 149 (U.W.); Franklin Co.: Palouse Falls, April 8, 1923, H. St. John et al . , 5262 (U.C.); Garfield Co.: about 15 miles south of Pomeroy, Blue Mts., May 25, 1944, CL. Hitchcock and C.V.  Muhlick, 8289, (U.W.); Grant Co.: Coulee City, May 5, 1916, F.L. PicbBtt, 477 (U.W.); Lincoln Co.: south side of Spokane River at its mouth, April 18, 1940, H.T. Rogers, 277 (U.C, U.W.., U.B.C); Klickitat Co.: near Roose-velt, April 5, 1934, F.L. Pickett et al», 1436 (U.W.); Okanogan Co.: on toe reservation east of Omsk,.May 8, 1932, C.B. Fiker,.625 (U.W.); throughout county, Omak, 1931, CB. Fiker, Do 1 (U.W.); Sec. 8, T. 36 K. R.29 E., May 21, 1911, I.C. Otis, 479 (U.W.); Sec.8, T.36 N. R.29 E«, May 21, 1911, I.C Otis, 480 (U.W.); summit of Muckamuck Lookout, Tiffany Range, June 26, 1951, J. Wm. Thompson, 6989 (U.W.); Spokane Co.: Latah Creek, Spokane, April, 1922, Pearl Blue, s.n. (U.C.); near Spokane Bridge, May 4, July 11, 1916, Wilhelm Suksdorf, 8576 (UO., U.W.); Camp No. 2, near Rock Creek, May 28, 1895, J.H. Sandberg and J.B. Leiberg, 85 (U.C.); Spokane, April, 1951, Sister Mary Milburge, s.n. (U.W.); mouth of Hangman's Creek (Latah Greek), May 10, 1936, Sister Mary Milburge, 1084 (U.C); 10 miles south east of Spokane, April, 1931, Sister Mary Milburge, s»n. (U.W.); near Spo-kane, April, 1892, Aug. 11, 1892, Louis F. Henderson, s.n. (U.W.); Spokane, April 29, 1923, T.A. Bonser, s.n. (U.W.); near Mica, May 8, 1952, Sister  Mary Milburge, 509 (U.W.); Spokane, April 22, 1906, I.C. Otis, 417 (U.W.); - 73 near Spangle, May 10, 1916, Wilhelm Sukadorf, 8601, type of D. Casickii Greene var. album Sukadorf, (U.C); Stevens Co.. south, side of Columbia Hiver 2 miles below Gerame, April 22, 1940, H.T. Rogers, 513 (U.C., U.W., U.B.C.); Walla Walla Co.: north Hillside, Wallula Gap, April 4, 1934, F.L. Pickett et al., 1399 (U.W.); Wallula, April 6, 1923, H. St. John, 50 88 (U.W.); Whitman Co.: Almota, April 29, 1894, C.7. Piper, 1795 (U.W,); Granite Pt., south of Wawawai in Snake River Canyon, March 9* 1941, W.A.  Webber, 2018 (U.W.); between Wawawai and Granite Pt., March 15-14, 1915, F.L. Pickett, 206 (U.W.); Zamiak Butte, April 29, 1916, F.L. Pickett, 584 (U.W.); south of Pullman, May 19, 1917, F.L. Pickett, 1067 (U.W.); Pull-April 26, 1915, F.L. Pickett, 298 (U.W.); Rock Creek, west of Winona, April 10, 1924, H. St. John et al. , 5260 (U.C.); small creek above Almota, April 27, 1955, L. Constance et al., 1041 (U.C.); along the west side of Rock Lake, April 50, 1938, F.G. Meyer, 1439 (U.C.); Zamiak Butte, May 12, 1923, Aim strong and Thayer, s.n. (U.C.); above Wawawai, April 6, 1922, C.S.  Parker, 326 (U.C.); Wawawai, April 23, 1921, H. St. John, 5976 (U.C); Granite Pt., Truax, March 12, 1921, H. St. John and F.L. Pickett, 2998 (U.C); Yakima Co.: Rattlesnake Mt., May 8, 1901, J.S. Cotton, 357 (U.W.); OREGON: mountains, eastern Oregon, May 20, 1898, Tffia. Cusick, 1874 (U.C); Gilliam Co.: 5 miles west of Blalock, April 10, 1938, M.E. Peck, 19834 (U.C); IDAHO: Bonner Co.: north slope of Clark Fork River Canyon, 15 miles west of Sand Point, May 14, 1936, CL. Hitchcock. 2898 (U.W.)s Canyon Co.: Enmett, April 29, 1911, J. Francis Maobride, 793 (U.C); Zootenai Co.: Hauser, April 26, 1931, E.I. Apple gate, 6700 (U.C); Latah Co.: 2 miles south of Viola, May 2, 1937, Mj_ and L. Dillon, 781 (U.W.); Nez Perce Co.: south bank of Clearwater River, April 17, 1933, C.W. Shar-74 smith, 5528 (U.C.); near Lewiston, April 29, 1896, A.A. and E.G. Heller, 2985, type or isotype of D. puberulentum, (U.C); west of Kamaiah, Clear-water River, May 5, 1951, E.I. Applegate, 6756 (U.C.); Zamaiah, Clearwat-er River, Aug. 5, 1951, E.I. Applegate, 6754 (U.C.); between Culdesac and Winchester, May 1, 1951, E.I. Applegate, 6720 (U.C); MONTANA: Lewis and Clarke Co.: Helena, E.N. Brandegee, s.n. (U.C); Meagher Co.: top of Zing's H i l l , Little Belt Mts., Aug. 15, 1945, CL. Hitchcock and C.7. Muhlick,  12555 (U.W.); Missoula Co.: Missoula Prairie, May 17, 1925, J.E. Zirkwood, 1407 (U.C); west exposure of Mt. Sentinel, April 51, 1958, F.A. Berkley and James Salinas, 2670 (U.C, U.W.); Missoula Prairie, May 17, 1925, J.5.  Zirkwood, 1407 (U.C.). 10 Dodecatheon pauciflorum (Dur.) Greene. Dodecatheon pauciflorum (Dur.) Greene, Pitt. 2:72, 1890, pro parte. Plants of very variable size, from a few centimeters to 45 cm. ta l l ; glabrous throughout, except leaves rarely sparsely glandular pubes-cent; rootlets fleshy-fibrous from a short crown; leaf blades narrowly elliptic to oblanceolate, 0.5-11.0 cm. long, 0.2-5.5 cm. wide, margin en-tire, or occasionally irregularly repand, apex rounded to obtuse, base sharply contracted or narrowly cuneate and tapering to the petiole; pet-ioles 0.2-11.0 cm. long; scape 5.0-45.0 cm. long; bracts 2.0*8.0 mm. long, acute; flowers 1-15, pentamerous; calyx tube 1.5-2.5 mm. long, lobes 2.0-6.0 mm. long, rounded to acute; corolla rose-purple with a fine red-orange  scallop on the reflexed tube; anthers 3.0-7.0 mm. long, dark purple, f i l a -ments 1.0-5.0 mm. long, yellow, connectives purple, smooth, sharply con-stricted near their bases or tapering gradually to acuminate apices; cap-75 sules narrowly ovoid, once and a half to twice the length of the calyx, glabrous, dehiscing by teeth, walls firm, opaque; seeds 1,0-1,2 mm. long, wingless. Type locality: west of the Missouri River to the Rocky Mountains. Habitat: moist open meadows, saline marshes, and, in the eastern part of its range, around alkali sloughs; sea level to 10000 f t . Distribution: Rocky Mountains of British Columbia and Alberta, across Alberta and Saskatchewan, and into Manitoba; westward across Idaho and into western Washington, across southern Oregon through the Cascade Mount-ains; northward along the islands of the Washington and British Columbia coasts to the south coast of Alaska. Map 7. Chromosome counts: Mortlach, Sask., meiotic .... n - 22 (PI. 11 & 12). southeastern B.C., somatic 2n - approximately 44. (PI. 9 and 10). Montana diploid Lulu Island, near Vancouver . meiotic ., n - approximately 44. (PI. 13 and 14). Vancouver Island •••••• at least tetraploid. Eklutna Flats, Alaska ..»..»• tetraploid. Key to the varieties of Dodeoatheon pauciflorum. Anthers 3.5-5.0 mm. long, about twice as long as the filaments 10a. typicum. Anthers 5.5r7.0 mm. long, more than twice as long as the filaments .........»»....» • •• 10b. macro carpum. Map 7 Dodecatheon. p a u c i f l o r u m v a r * m a c r o carpum c i r c l e s v a r . t y p i c u m s q u a r e s PLATS 9 ft Dodeoatheon pauciflorum, Fairmont Springs, B.C. Camera lucida drawing, prophase of mitosis in a root tip pre treated with paradiohlorohenzene, showing approximately 44 chromosome a. Ca.X 3100 PLATS 10 Dodecathooa paueiflortra. Dot oh Creek, B.C. Camera lucida drawing, prophase of mitosis in a root tip pretreated with paradichloro-benzene, showing approximately 44 chromo-somes. Ca. X 2500. FLATS 11 t Dodecatheon pauciflorum, Mortlaeh, Saak. Cansr a lucida drawing, first division pro-phase of meiosis in a developing anther, shewing 22 pairs of chromosome a. Ca.X 3000 PLATE 12 Dodeoatheon pauciflorum, Uortlach, Saak Photomicrograph, fi r s t division prophase of meiosis in a developing anther, showing 22 pairs of chromosomes. Ca. X 1000 PLATS 15 eVi w Dodeoatheon pauoifLorum, Tar* maoro carpum, Lulu Island, near Vancouver, B* C* Camera lucida drawing, first division pro-phase of meiosis in a developing anther, showing approximately 44 pairs of chromo-somes. Ca. X 1800* FLATS 14 Dodeoatheon pauciflorum, var. maorocarpum, Lulu Island, near Vancouver, B. C. Photomicrograph, first division prophase of meiosis in a developing anther, showing approximately 44 pairs of chromosomes Ca. X 1000 - 76 10a. Dodeoatheon pauciflorum var* typlcum n.n. Dodeoatheon pauciflorum (Dur*) Greene var* typlcum n.n. Dodecatheon integrifolium Michx. var. vulgare Hook•, F l . Bor. Am. 2:118-119, 1840. Dodeoatheon Meadia L. Tar pauolflora Dur., PI* Pratt., 95, 1855 pro parte. Dodeoatheon pauciflorum (Dur.) Greene Tar. monanthum Greene, Pitt. 2:73, 1890. Dodeoatheon sallnum Nels., Bull. Torr. Bot* Club 26:131, 1899. Dodeoatheon unlflorum Rydb., Mam* N.Y. Bot. Gard. 1:307, 1900 Dodeoatheon philosoia Nels., Bull. Torr* Bot* Club 28:227, 1901. Dodeoatheon pauciflorum (Dur.) Greene subap. monanthua (Greene) Knuth, In Sngler, Daa Pflanz* 4:243, 1905* Dodeoatheon pauciflorum (Dur*) Greene aubsp* sallnum (Nels.) Knuth, In Engler, Das Pflanz. 4:243, 1905. Dodeoatheon vulgare (Hook.) Piper, Contr. U.S. Nat. Herb. 11:445, 1906. Dodecatheon pauciflorum (Dur.) Greene var. shoshonensls Nels., Bot. Gaz. 54:143, 1912. As In the species description except anthers 3.5-5.0 mm. long, about twice as long as the filaments. Type locality: as for the species. Habitat: moist open meadows or the edges of alkali sloughs, 2000-10000 f t . Distribution: As the species distribution east of the Cascade 77 -Mts. Caromosame count: see species description. Specimens examined: BRITISH COLUMBIA: southeast shore of Slocan Lake, May 12, 1933, T.T. McCabe, 6916 (U.C.); southeast corner of Columbia Lake, May 28, 1938, T.T. McCabe, 6395 (U.C); southeast corner of Columbia Lake, May 21, 1938, T.T. McCabe, 6172 (U.C); Little Sheep Creek Trail, Rossland, July 5, 1942, J.W. Eastham, 3 . n . (U.B.C); Corbin, 49°N. 114°W., A. Elett, 18 (U.B.C); "Crow Nest Pass", Rocky Mts., Maooun, s.n. (N.M.); ALBERTA: Lake Edith, Jasper Park, July 14, 1927, E.M. Kindle, s.n. (N.M.); Lake Louise Chalet, July 7, 1912, J. Smith, s.n. (U.B.C.); Banff, June 14, 1948, C.B.W. Rogers, 175 (U.B.C.); Banff, May 18, 1902, J.R. Anderson, s.n. (P.M.); Craigayle District, June 24, 1942, A.H. Brinkman, 5176 (U.B.C); . vicinity of Edmonton, June 28, 1917, W.C McCalla, 2592 (N.M.); Jasper, July 30, 1917, M.O. Malte, s.n. (N.M.); Jasper Park, June 9, 1930, H.M.  Laing, 546 (N.M.); Banff, vicinity of Basin, June 8, 1906, S. Brown, 14 (N.M.); Banff, June 13, July 6, 1899, Sanson, s.n. (N.M.); Seventeen Mile Hat, Banff, June 27, 1948, W.W. Mair, 98 (U.B.C.); Athabaska River, July 2, 1898, W. Spreadborough, s.n, (N.M.);. Athabaska River, June 26, 1898, W.  Spreadborough, s.n. (N.M.); Eossil Coulee, Milk River Ridge, June 22, 1883, Dawson, s.n. (N*M.) J Belly River, July 22, 1881, Dawson, s.n. (N.M.); Water-ton Lake, July 30, 1895, Macoun, s.n. pro parte (N.M.); Banff, June 27, 1891, Macoun, s.n. (N.M.); Bow River at Morley, Sept. 6, 1879, Macoun, s.n. (N.M.)jLake Louise, July 25, 1904, Macoun, s.n. (N.M.); Elbow River, June 25, 1897, Macoun, s.n. (N.M.); Calgary* June 5» 1897, Macoun, s.n. (N.M.); Pipestone Creek, Rocky Mts., June 29, 1904, Macoun, s.n. (N.M.); Wood t ft t "* Buffalo Park, MacKenzie Basin, about 59°57 30 N. 112°17 W.» June 15, 1928, 78 H»M. Raup, 5001 (N.M.); SASKATCHEWAN: Moose Mountain, June 4, 1884, J.M. Macoun, s.n. (N.M.); Wood Mountain Post, June 11, 1895, Macoun, s.n. (N.M.); Red Deer Lakes, July 20, 1879, Macoun, s.n. (N.M.); east of Prince Albert, July 6, 1896, Macoun, 3»n. (N.M.); Cypress Hills, June 25, 1894, Macoun, s.n. (N.M.); l£ miles east of McKague, June 22, 1940, A.J. Breit- ung, 579 (N.M.); Mortlach, June 12, 1950, G. Ledingham, 789 (U.B.C.); Indian Head, 1892, W. Spreadborough, s.n. - (N.M.); WASHINGTON: Asotin Co.: ^rnile south of Indian Tom Creek, Blue Mts., June 1, 1937, L. Constance et al., 1869 (U.W.); Garfield Co.: Blue Mts. near Pameroy, May 21, 1936, J.  Cromwell, 129 (U.W.); Lincoln Co.: east of Ores ton, June 23, 1933, J. Wm. Thompson, 9179 (U.W.); Whitman Co.: 2 miles west of Wilms, April 2, 1923, C.F. Lackey, s.n. (U.C.); Pullman, May 10, 1896, S.B. Piper, s.n. (UiW.); Pullman, May 3, 1923, E.A. Warren, 208 (U.W.); OEEGON: Eastern Oregon, June, 1899, Wm. C. Cusick, 2190 (U.C.); Oregon, Wm. C. Cusick, 1509 and 1528 (U.C); highest "Steins" Mts., July 5, 1898, Wm. C Cusick, 2007 (U.C); highest Powder River Mts., Aug., 1896, Wm. C Cusick, s.n. (U.C.); highest Blue Mts., July 20, Sept. 12, 1899, Wm. 0. Cusick, 2255 (U.C); "Steins Mts.", July 2, 1896, J.B. Leiberg, 2447 (U.C.); Baker Co.: East Pine Creek, near Cornucopia, Wallowa Mts., June 50, 1955, G.N. Jones, 7522 (U.C); Harney Co.: Blitzen Gorge, Steens Mts., July 16, 1935, J. Wm. Thompson, 12115 (U.W.); Steens Mts., June 14, 1946, B. Maguire and A.H. Holmgren, 26450 (U.C, U.W.); Josephine Co.: near Waldo, April, 1887, T. Howell, s.n. (U.W.); Lake Co.: Quartz Mt., T.38S. R16E., May 24, 1940, L.E. Detling,  4268 (U.C, U.W.); eastern slope of Crane Mt., July 7, 1932, E.I. Apple- gate, 7497 (U.C.); Klamath Co.: Swan Lake Valley, May 28, 1923, E.I.  Applegate, 5522 (U.C); region of Fossill Lake, between Lakeview and 79 Bryants, June, 1901, H.W. Furlong et a l . . s.n. (U.C.); Wallowa Co.: Wallowa Mt., Aug. 24, 1907, W.C. Cusick, 5222 (U.C, U.W.); Wheeler Co.: near Spray, Apr. 17, 1955, J. Wm. Thompson, 11558 (U.W.); IDAHO: Bannock Co.: Pocatello, May 4, 1956, Forrest Romero, s.n. (U.C.); Pocatello, March 29, 1936, R. Porter, s.n. (U.C); Blaine Co.: Smoky Mts., Aug. 15, 1916, J.F.  Macbride and E.B. Pay son, 5747, type or isotype of D. pauciflorum var. exquisitum (U.C); Clyde, July 11, 1916, J.F. Macbride and E.B. Payson, 5172 (U.C); Butte Co.: Pass Creek Gorge, about 10 miles north of Leslie, Lost River Mt., June 12, 1944, CL. Hitchcock and C.V. Muhlick, 8848. (U.W.); Custer Co.: "Dicky", T.9N. R.22 E., June 18, 1940, R.J. Davis, 2048 (U.C); head of Jim Creek, July 25, 1953, R.J.. Davis, 680 (U.C.); 2 miles east of Dickey, June 21, 1947, CL. Hitchcock, 15615 (U.C, U.W.); Ryan Peak, Boulder Mts., Sawtooth National Forest, Aug. 1, 1944, CL. Hitchcock and C.V. Muhlick, 10611 (U.C, U.W.); Lost River Mts., Leatherman Pass, head of the West Fork Pahsimeroi River, Aug. 16, 1944, CL. Hitchcock and C.V.  Muhlick, 11195 (U.W.); east end of MacKay Reservoir,. June 13, 1944, CL. Hitchcock and C.V. Muhlick, 8882 (U.W.); head of Rock Creek, northwest base of Mt. Borah, Lost River Mts., Aug; 12, 1944, CL. Hitchcock and C.V.  Muhlick, 10940 (U.W.)]} head of Morgan Creek, June 29, 1944, CL. Hitchcock and C.V. Muhlick, 9400 (U.W.); Gooding Co.: "Hogeraan" valley, vicinity of "thousand springs", May 10, 1959, A. Oronquist, 11115 (U.C); Hagerman, May 9, 1956, R. Porter, s.n. (U.C.); Hagerman Valley, May 9, 1936, Mas  Reese, s.n. (U.W.); Idaho Co.: 15 miles south of RiggLns on Little Salmon River, May 29, 1944, CL. Hitchcock and C.V. Muhlick, 8494 (U.W.); Sheep Creek 4 miles above the mouth, Snake River Canyon, May 16, 1936, J. Pack- ard, 75 (U.W.); foot of "He Devil" near 4th Sheep Creek Lake, T.23N. R.2W., June 13-30, 1937, J. Packard, 501 JU.C): Lemhi Co.: 10 miles east of Gilmore, June 25, 1947, CL. Hitchcock, 15768 {U.C, U.W.); 1 mile southr east of Blue Noes, head of Spring Creek, Northwest of Indianola Ranger Station, July 2, 1946, CL. Hitchcock and 0.7. Muhlick, 14554 (U.W.) J Eight Mile Creek, about 12 miles south west of Leadore, Lemhi Mts., June 24, 1944, CL. Hitchcock and C.V. Muhlick, 9254 (U.W.J; Owyhee Co.; 14 miles south-west of Mud Flat on Juniper Mt. Rd., June 7, 1946, B. Maguire and H. Holm-gren, 26538 (U.C); Silver City, June 17, 1911, J.F. Macbride. 911 (U.C); Twins Falls Co.: Shoshone Falls, June 24, 1912, A. Nelson and J.F. Mac- bride. 1729 (U.C); Valley Co.: 1 mile south of Landmark, June 27, 1946, CL. Hitchcock and C.V. Muhlick, 14067 (U.C, U.W.)} 5 miles west of Payette Lake, June 25, 1946, CL. Hitchcock and C.V. Muhlick, 15946 (U.W.); north end of Warm Lake, June 26, 1946, CL. Hitchcock and C.V. Muhlick, 14045 (U.W.); MONTANA: Old Hollowtop, near Pnny, July 7, 1897, P.A. Ryd- berg and E.A. Bessey, 4671 (U.C); about Divide Lake, Glacier National Park, July 4, 1954, B. Maguire, G. Piranian, 15797 (U.C); Logan Pass Glacier National Park, July 16, 1954, G.N. Jones, 5595 (U.C); head of Cottonwood Creek, Aug. 10, 1902, Blanklnship, s.n. (U.W.); Beaverhead Co.; above Torrey Lake, Pioneer Range, July 27, 1946, CL. Hitchcock and C.V.  Muhlick, 15089 (U.C, U.W.); above and to the west of Ore anno s Lake, head of Pintlar Creek, Anaconda Range, July 27, 1945, CL. Hitchcock and C.V. Muhlick, 12760 (U.C, U.W.)} ridge connecting Sheep and Black Lion Mts., Pioneer Range, July 50, 1945, CL. Hitchcock and C.V. Muhlick, 12976 (U.C. U.W.)} south of Lake Waukena, head of Rock Creek, Pioneer Mts., Aug. 1, 1945, CL. Hitchcock and C.V. Muhlick, 15095 (U .W. ) ; about 15 miles west of Wisdom, Gibbonsyille Rd., June 22, 1944, CL. Hitchcock and C.V. Muhlick, 81 9197 (U.W.); above AJax Lake, Bitter Boot Mts., July 23, 1945, C.L. Hitch*  cock and C.V. Muhlick, 12710 (UBW.); Deerlodge Co.. peak to south of Storm Lake, Anaconda Mts., July 21, 1946, C.L. Hitchcock and C.7. Muhlick, 14767 (U.C., U.W.); Storm Lake, west slope of Anaconda Mts., July 21, 1946, C.L.  Hitchcock and C.7. Muhlick, 14744 (U.W.); Ifergus Co.: floor of Half Moon Canyon at mouth, Big Snowy Mts., July 6, 1945, C.L. Hitchcock and C.7.  Muhlick, 12021, (U.C, U.W.); Flathead Co.: Lake McDonald, Aug. 10, 1932, B. Maguire, 984 (U.C); Gallatin Co.: east of Bozeman, May 26, 1901, J.W.  Blankinship s.n. (U.W.); Judith Basin Co.: Yogo Peak, Little Belt Mts., July 6, 1947, C.L. Hitchcock, 16150 (U.C, U.W.); Lewis and Clark Co.: Lincoln, Helena National Forest, July 1, 1948, CL. Hitchcock, 17907 (U.C, U.W.); east slope of China Wall, near north end of Lewis and Clark National Forest, Sun River Primitive Area, July 27, 1948, CL* Hitchcock, 18944 (U.W.); Helena, F.D. Kelsey, s.n. (U.C); Madison Co.: § mile north of Koch Peak, Taylor Mts., Aug. 2, 1946, CL. Hitchcock and C.7. Muhlick, 15208 (U.C, U.W.); along stream from Brandon Lake, Tobacco Root Mts., July 29, 1947, C.L. Hitchcock, 16985 (U.W.).; Meagher Co.: LinkPeak about 3 miles east of Williams Mt., Little Belt Mts., July 12, 1945, C.L. Hitch- cock and C.7. Muhlick, 12296 (U.W.); base of Baldy Mt., Big Belt Mts., x July 16, 1945, CL. Hitchcock end C.7. Muhlick, 12409 (U.C, U.W.); top of King's H i l l , Little Belt Mts., July 13, 1945, CL. Hitchcock and C7. Muh- lick, 12355 (U.C.).; summit of King's H i l l , Little Belt Mts., July 9, 1945, CL. Hitchcock and C.7. Muhlick, 12157 (U.W.); about 4 miles north of Castle City, Castle Mts., June 30, 1947, CL. Hitchcock, 15947 (U.C,U.W.); Missoula Co.: Missoula, June 18, 1924, J.B. Kirkwood, 1749 (U.C); § mile • west of Upper Holland Lake, Flathead Range, Flathead National Forest, July 15, 1948, CL* Hitchcock, 18389 (U.C, U.W.); Pattee Canyon near Missoula, 82 June 2| X93g| F.A. Berkley, 2452 (U.W.); Park Co.: 6 miles west of Four Mile Ranger Station, Boulder Canyon, July 12, 1947, CL. Hitchcock, 1633 6 (U.C, U.W.); 6 miles west of Four Mile Ranger Station, July 12, 1947, CL.  Hitchcock, 16536A (U.W.); Powell Co.: 6 miles south of Helmville, June 29, 1945, CL. Hitchcock and C7« Muhlick, 11707 (U.W.); Ravalli Co.: Sula, south of Darby, June 17, 1944, CL. Hitchcock and C.7. Muhlick, 9079 (U.W.); Stillwater Co.: north slope of Mt. Haystack, head of Boulder Creek, Absar* oka National Forest, Aug. 8, 1945, C.L. Hitchcock and C.7. Muhlick, 13455 (U.C«i U.W..J; Sweet grass Co.: 5 miles west of Rainbow Lake, Absaroka Plat-eau, July 16, 1947, CL. Hitchcock, 16427 (U.W.); near outlet of Granite Lake, head of Big Timber Creek, Crazy Mts., Aug. 5, 1945, C.L. Hitchcock and C.7. Muhlick, 15286 (U.W.)* 10b. Dodecatheon pauciflorum (Dur.) Greene var* macrocarpum (Gray) n. comb. Dodecatheon pauciflorum (Dur.) Greene var. macro carpum (Gray) n. comb. Dodecatheon Meadia L. var. macro carpum Gray, In Geol. Sur. Cal. Bot., ed. 2, v. 1:467, 1880, equivalent to ed. 1, 1876 (see D. frigidum). Dodecatheon macrocarpum (Gray) Knuth, In Engler, Das Pflanz. 4: 241*242, 1905. Dodecatheon macrocarpum (Gray) Knuth var. alaskanum Hult., Lunds Univ. Arssk.N.F., Sect. 2, 7 . 44:1289, 1948. As in the species description, except anthers 5.5-7.0 mm. long, more than twice as long as the filaments. Type locality: from Alaska southward. - 83 Habitat; moist open meadows or saline marshes, sea level •* 5000 f t . Distribution; along the coast of Oregon and ?/ashington, and the coast islands of Washington and British Columbia to Alaska. Chromosoms count; see species description. Specimens examine df ALASKA: Juneau, June 17, 1949, W.C. Frohne, 4965 (U.B.C.); Seward, June 9, 1949, W.C.Frohne, 4928 (U.B.C); Haines, June 15, 1949, W.C. Frohne, 4958 (U.B.C); Eklutna Flats, June 12, 1949, G,M. Frohne, 4945 (U.B.C); Eklutna Flats, June 20, 1949, G.M. Frohne, 4935, (U.B.C); Eklutna Flats, July 9, 1948, G-.M. Frohne, 49151 (U.B.C); shores of Tes Bay, June 50, 1895, Thos. Howell, 1648 (U.C.); Glacier Bay, Glacier Bay, June 28, 1907, Mrs. K. Stephens, 5 (U.C); Mole Harbor, Admir-alty Island, May-June, 1907, Mrs. K. Stephens, 94 (U.C.); Yes Bay, Aug. 25 1890, M.W. Gorman, s.n.'(U.C.); shores of Behm Canal, May 26, 1894, M.W.  Gorman, s.n. (U.C.); Deep Bay, Admiralty Is., June 22, 1939, W.J. Eyerdam, 5016 (U.C); Port Vita, Raspberry Strait, Raspberry Is., Kodiak group, May 31, 1945, W.J. Eyerdam, 5685 (U .W., N.M.); Afognak, Raspberry Strait, Rasp-berry Is., Kodiak, group, June 5, 1945, W.J. Eyerdam, 5686 (U .W., N.M.); Cordova, Aug., 1952, W.J. Eyerdam, s.n. (U .W.) ; Eklutna Flats, June 12, 1946, C Heller, s.n. tU.W.); Kodiak Is., May 31, 1897, J.M. Macoun, s.n. (N.M.); Middleton Isl., 59 50 N., June 15, 1892, -J.M. Macoun, s.n. (N.M.); Eagle River Flats, Anchorage, June 11, 1948, E. LePage, 25112 ( K . M . ) ; Se-ward, June U , 1941, J.P. Anderson, 4745 (N.M); Eklutna, June 26, 1941, J.P. Anderson, 6962 (N.M.); Shelter Is., Lynn Canal, May 11, 1941, J.P.  Anderson, 6430a and B (K.M.); Shelter Is., Lynn Canal, May 11, 1941, J.P. 84 Anderson, 6429 (N.M.); Montana Creek, near Juneau, May 23, 1941, J.p. And- erson, 6442 (N.M.); Juneau, June 7, 1940, J.P. Anderson, 6355 (N.M.); Jun-eau, May 18, 1940, J.P. Anderson, 6554 (N.M.); Palmer, June 10, 1944, J.P.  Anderson,,8425 (N.M.); YUKON: near Moosehide, May 20, 1914, Alice Bast- wood, 84 (U.C., N.M.) j BRITISH COLUMBIA: northern islands: Limestone Is-land, Queen Charlotte Is., June 9, 1915, CF. Newcombe, s.n. (U.B.C); Limestone Is., Queen Charlotte Is., May 27, 1925, CF. Neweombe, s»n. (P.M.); Porcher Is., Freeman Pass, Aug. 25, 1959, T^T. McCabe, 7545 (U.C); Larson Harbor, Banks Is., Aug. 18, 1959, T.T. MoCabe, 7529 (U.C); Calvert Is., Sorrow Island, Cape Calvert, June 1, 1957, T.T.McCabe, 4083 (U.C.); Calvert Is., Sorrow Islands, Cape Calvert, June 1, 1957 T.T. McCabe, 4082 (U.C); Calvert Is., Sorrow Islands, Cap Calvert, June 1, 1937, T.T. Mc- Cabe, 4081 (U.C); McKenny Is., northwest of Aristazabal Is., June 6, 1936, T.T. McCabe, 3426 (U.C); Borrowman Harbor, northwest of Aristazabal Is., June 5, 1936, T.T. McCabe, 3409 (U.C); Vancouver Island and southern coast; Victoria, June 1, 1872, Cowley, s.n. (N.M., pro parte); Mt. Copley, Cameron Lake, V.I., Aug. 13, 1949, D. Elvidge, s.n. (U.B.C); Woodward*s Landing, Lulu Is., May 13, 1925, T.R. Ashlee, s.n. (U.B.C); end of #5 Road, Lulu Is., May 18, 1924 T.R. Ashlee, s.n. (U.B.C.); end of #5 Road, Lulu Is., June 14, 1957 J.W. Eastham, s.n. (U.B.C.); end of #5 Road, May 14, 1941, J.W. Eastham, s.n. (U.B.C); Alberni, V.I. June 50, 1959, J.W. East- hem, s.n. (U.B.C); Victoria, April 9, 1958, J.W. Eastham, s.n. (U.B.C.); Yale, May 17, 1875, Macoun, s.n. (N.M., pro parte); Victoria, May 11, 1908 John Macoun, s.n. (N.M.); vicinity of ,,Ucleulet,,, May 25, 1909, John Macoun, s.n. (N.M.); Mt. Finlayson, May 6, 1908, John Macoun, s.n. (N.M.); Oak Bay, Victoria, June 5-May 14, 1908, John Macoun, s.n. (N.M.); Mt. Arrowsmith, - 85 7.1., July 17, 1887, John Macoun, s.n. (N.M.): Oak Bay, Victoria, May, 1913, John Macoun, s.n. (N.M.); Cedar H i l l , 7.1., May 26, 1887, John Macoun, s.n. (N.M.); Cedar H i l l , May 31, 1893, Macoun, s.n. (N.M.); Kennedy Lake, vicin-ity of "Ucleulet", July 23, 1909, John Macoun, s.n. (N.M.); Beacon H i l l , 7ictoria, May 25, .1913, John Macoun, s.n* (P.M.); Millstream, Nanaimo, July 1, 1914, John Macoun, s.n. (P.M.); Victoria, May 20,.1917, F. Perry, (U.B.C.); 7ictoria, April, 1895, A.J. Pineo, s.n. (U.B.C.)} Alberni, June 26, 1916, J.K. Henry, s.n. (U.B.C.); Victoria, May.l, 1897, J.K. Henry,  s.n. (U.B.C.); Victoria, March 10, 1912, J.K. Henry, s«n. (U.B.C.); Victor-ia, .April 6, 1915, Mrs. HiggLns, s.n. (U.B.C.); Victoria, May 1, 1917, Mrs.  J.T. Biggins, s.n. (U.B.C); Victoria, April 30, 1924, Mrs. J.T. Eiggins,  s.n. (U.B.C); end of #3 Road, Lulu Is., May 20, 1949, K.I. Beamish, 49102 (U.B.C.); end of #3 Road, Lulu Is., May 28, 1950, K.I. Beamish, 50105 (U.B.C); Uplands, Victoria, May 3, 1949, K.3A Beamish, 49101 (U.B.C.); Up-lands, 7ictoria, May 11, 1950, K.I. Beamish, 50101 (U.B.C.); Sayward, 7.1. June 3, 1950, 7. Krajina, s.n. (U.B.C.); Uplands, Victoria, May 3, 1924, C.F. Newcombe, s.n. (P.M.); Oak Bay, 7.1., April 22, 1924, G.A. Hardy, s.n. (P.M.); Chains Is., May 10, 1949, G.A. Hardy, s.n. (P.M.); Shawnigan Dist-trict, May 24, .1932, H. Toms, s.n. (P.M.); Mt. Toimie, 7.1., May 16, 1933, F. Keimode, s.n. (P.M.); Mt. Arrowsmith, 7.1., Aug. .1, 1917, F. Keimode,  a.n. (P.M.); KGksila, 7.1., June 13, 1897, J.R.Anderson, s.n. (P.M.); Comox, May 18, 1899, J.R. Anderson, s.n. (P.M.); Somass River, Alberni, June, 1916, W.R. Carter, 45 (PJ.M.); "Uchualisit", 7.1., 1915, W.B. Anderson,  s.n. (.M.); Goldstream, V.I., May'5,"1928, W.B."Anderson, s.n. (P.M.); Katz Landing, April 24, 1925, W.B. Anderson, s.n. (P.M.); WASHINGTON: American Lake, May 11, 1908, E. Todd, s.n. (U.W.); American Lake, May 11, 1908, 86 Bergsrson, s.n. (U.W.)} Clallam Co.; Starvation Flats, Mt. Angeles, July 17, 1951,J.Wm. Thompson, 7417 (U.C.)} Sequim, May 20, 1915, J.M. Orant, 3.n. (U.W*)} Island Co.: Goose Rock, Deception Pass State Park, Whidbey Island, April 23, 1936, H.W. Smith, 340 (U.C, U.W.)} Goose Rock, Whidbey Island, April 29, 1935, H.W. Smith, 66 (U.C.)} Goose Rock, "Whidby" Island, April 25, 1931, J. Wm. Thompson, 6096 (U.W.); "Whidby" Island, April 25, 1897, N.L. Gardner, s.n. (U.7J.); Pierce Co.: Muck Creek, near Roy, April 29, 1934, G.N. Jones, 4633 (U .W. ) ; near Roy, April 30, 1932, J. Wm. Thompson, 8245, (U.W.)} San Juan Co.: Brown's Island, near Friday Harbor, San Juan Island, May 5, 1956, B.D. Blanchard, 52 (U.C.)} Whatcom Co.: Lummi Rooks, April 28, 1959, W.C Muensoher, 9720 (U.C, U.W., U.B.C.)} Harts Pass, June 25, 1959, W.C Muensoher, 10062 (U.C.)} OREGON: Clackamas Co.; Oswego, Willamette River, May 2, 1886, L.F. Henderson, 641 (U.C.)} Jefferson Co.: Warm Springs Agency, June 25, 1944, E.V.A* Murphy, 78 (U.C.)} Multnomah Co.; bluffs of the Columbia River, near Multnomah Falls, May 8, 1936, S.L. Ander- son, s.n. (U.C). DISCUSSION OF D. CUSICKII, D. PAUCIEL0RUM and D. PAUCIFLORUM YAR. MACR0CARPUM D, Ousickii occurs mainly in the dry Interior Plateaux of Brit-ish Columbia and Washington. A few specimens have been seen from east of the Central Cordilleran System in British Columbia and even east of the Rockies in Montana (Map 6.). Throughout the major part of its range, this species is very con-stant in its characteristics, though, as has been previously suggested, there is a possibility of hybridization with D. conjugens where the two - 87 -ranges overlap in Washington, or with D. Hendersoni where the two ranges meet in the Columbia River Valley. Probably its nearest relative is D. pauciflorum, as the root type, capsule dehiscence, and filament length and colour of the two are alike. Likenesses in these characteristics, most stable in the genus, sug-gest that one might be a variety of the other. However, their generally distinct ranges (Maps 6 and 7), the striking glandular pubescence of p.  Cusiekii, especially in its inflorescence, and its very minute seeds, serve to distinguish them morphologically from one another. The main reason, however, for maintaining them as distinct spec-ies, at least t i l l further study can be made, is their apparent genetic constitution. Unfortunately, meiotic material is needed from many more' locations to establish definite conclusions, but the situation at present is as follows. P. pauciflorum from Fairmont Hot springs, southeastern British Columbia, is diploid, as shown by somatic counts from root tips, (PI. 9 and 10}• P. pauciflorum from Mortlach, Sask. has 22 pairs of chrom-osomes from meiotic counts, considered accurate (PI. 11 and 12.). As sug-gested below, these two groups also need further study, but at present are being considered the same. Here the number is variable as shown by meiotic counts in cells from within a single anther. Both 23 and 24 distinct ent-ities were counted (PI. 15-18). Each of these entities appears to be a pair of chromosomes, though such may not be the case, since, at first metaphase, one of them consistently lies off the plate (PI. 19, 20), behaving -in a manner unlike a normal pair of chromosomes. One possible explanation of this metaphase behaviour is the pre-sence of a univalent. A second possible explanation lies in the work done PLATE 19 Dodeoatheon Cusickii, Princeton, B.C. Camera lucida drawing, fi r s t division pro? phase of meiosis in a developing anther, showing in outline the group of chromosomes on the metaphase plate, one chromosome or pair lying off the plate. Ca. X 1700 PLATE 20 Dodeoatheon Cusickli, Princeton, B.C. Camera lucida drawing, first division early anaphase of meiosis in a developing anther, showing 22 pairs of chromosomes on the plate, one chromosome or pair lying on the edge of the plate* Ca. X 2500. on the existence of supernumerary or B-chromoscmes in various plants. Darlington and Upcott (1941), discussing B-chromosomes in Zea Mays, state that such iiromosames are found in some forty species of flowering plants. They are heterochromatic, therefore termed "inert'*, but though their be-haviour is somewhat irregular, they may pair with each other at meiosis, form chiasmata, and segregate. Ostergren (1947), in his paper on B-chromosomes in Anthoxanthum, cites cases from other workers in which the number of accessory chromosomes changes during the devlopment of the individual. He also states that in these cases the B-chramosomes usually l i e at the edge of the metaphase plate in mitosis. However, he discusses meiosis of only Anthoxanthum and here the number is regular and the behaviour normal* Whether such an explanation is applicable to D. Cusickii is a matter for study. Whatever the reason for the variation, the fact is evident that the chromosome number varies within an anther. Until more i3 known, then, of the genetic constitution of D. Cusickii and D. pauciflorum, the former is being left as i t now stands, an independent species. D. pauciflorum, like D. Jeffrey!, is a large, far-ranging, and apparently variable species. Populations in many parts of its range have been singled out and described as new species with a resulting confusion in synonymy. However, after examination of large numbers of specimens from the Prairies westward and north to Alaska, only one subdivision has been recognized, various attempts were made, including polygonal graphing (Davidson, 1947), to separate, for instance, the soroalled D. sallnum of the Prairies from D. pauciflorum of the Rockies in Montana. No constant differences were found. As stated above, "D. sallnum" from Mortlach, - 89 -Sask» has 22 pairs of chromosomes, P. pauciflorum of southern British Columbia has approximately 44 somatic chromosomes. No difference is indic-ated here, though, as has also been stated, much more cytologies! work is needed. Meanwhile, the two have been grouped together as D. pauciflorum var. typicum. With D. pauciflorum var. typicum also, there have been included a number of other variants whose differences from the species seem too slight to warrant taxonomic rank. Size has been used mainly as a basis for separation in two instances. True, small forms occur, but these are from high elevations, and a l l sizes of inter grades are also found. Slight pub-escence, too, has been made a criterion of species, but scattered specimens with varying degrees of pubescence on the leaves have been seen among Mont-ana materialo So small and unreliable a difference hardly seems signifi-cant. As indicated in the synonymy, then, a l l these variants are included with D. pauciflorum var. typicum. This variety extends across southern Oregon and through the Cas-cade Mountains. In the region of Klamath County, variation again appears, this time in a more stable characteristic, colour of filament, for here occur a few specimens whose filaments are purple. In the discussion of D. poeticum, i t was noted that this species occurs where the ranges of D. Cusickii and D. Hendersoni meet in the Columb-ia River Valley and that its pruple filament tube may have derived from the latter. A similar situation could occur in southern Oregon, for here the ranges of D. pauciflorum var. typcium and P.. Hendersoni meet. The fact may have no significance but cytological study would be of interest. In any case, as the variants here are very few and scattered, they have been in-- 90 eluded with D« pauciflorum var. typicum. From southern Oregon to northern Washington, along the coast, few specimens have been seen. However, from the Olympic Peninsula of Washington north along the islands to Alaska, distribution is fairly con-tinuous. This part of the species, with its northern range, is being des-ignated D. pauciflorum var. macro carpum (Map. 7) for the reasons given be-low. Cytological examination shows that plants on Lulu Island, near V a n c o u v e r , are tetraploid (PI. 13 a n d 14). Specimens from V i c t o r i a , B.C. are at least tetraploid, possibly hexaploid, and from Anchorage, Alaska, tetraploid. The pattern in this species seems to be the same as that of D. Jeffrey! and D. Henderson!; diploids in the south and east, polyploids along the north coast. Also, as in D. Jeffrey! and D. Henderson!, morpho-logical distinctions are very slight. Close comparison revealed no diff-erences between diploids and polyploids in major characteristics, except that the coast specimens have somewhat shorter filaments. Polygonal graphing (Davidson, 1947) of absolute quantitative characters and ratios of quantitative characters (PI. SI) shows a tendency of the coast forms to separate from the eastern in absolute anther length and ratio of filament to anther length. As a check, the same characters were graphed for popula-tions of Vancouver Island and Alaska, botljipolyploids, and here no separa^ tion is apparent (PI. 22)> On the basis, then, of this slight morphologic-al difference, in conjuction with their polyploidy and decidedly distinct range, the coast populations are being treated as a variety of the more eastern part of the species described by Dur and. Three variant populations are included here. On Lulu Island, Plate 21, 22 Polygonal graphs comparing quantitative characters from popu-lations of P. pauciflorum var. typicum and D« pauciflorum var. macro-carpum. The characteristics on one axis are in no way correlated with those on other axes as same represent ratios, others absolute measure-ments. Significance lies in the points on the axes, not in the areas^n-closed. Saoh point represents the average, for a particular characterist-ic, of the specimens on one herbarium sheet. Approximately 15 sheets (15 points) are used for each population on each graph. Plate 21. D. pauciflorum var. macro carpum, Vancouver Island, solid line D. pauciflorum var. typicum, Montana broken line Plate 22. P. pauciflorum var. macrocarpum, Vancouver Island, solid line P. pauciflorum var. macrocarpum, Alaska, broken line Plate 2 1 , 22 » a... leaf length,  H a t i 0 J leaf width B . Absolute length of scape r R*tin- l e n g t h o f s c a p 9 C . R a t i o . l e n g t h o f i Q a V e S length of calyx lobes D. Ratio: length of calyx tube T , T> length of anther B a t i o : length of filament F. Absolute length of anther PLATS 21 PLATE 22 91 near Vancouver, and at Alberni on Vancouver Island, Dodeoatheon grows in the brackish water of tidal flats, actually standing in water at high tide and never really on dry ground. In both instances the plants have longer scapes and petioles than the usual D. pauciflorum var. macrocarpum, but this may be a response to their environment, as otherwise they appear id-entical. Thosaon Lulu Island are known to be tetraploid (PI. 13 and 14); those from Alberni have not been studied cytologically. The third popula-tion, again probably an ecotype, is a group of dwarf plants at about 5000 ft . on Mt. Arrow smith, Vancouver Island. In stature they resemble the small plants of D. pauciflorum var. typicum at high altitudes in Montana, but since they occur within the range of var. macrocarpum, and since their genetic constitution is unknown, they, like the salt marsh populations, are included with the variety. CONCLUSION The study of the genus Dodeoatheon had as its object a reclassi-fication of the genus in Northwestern America with minor emphasis on its possible horticultural value. The classification, based on morphology, cytology, and distribution, is a division into large groups only, ten species and one variety. Lack of further subdivision is a deliberate effort to avoid adding to the previously existing confusion of synonymy, as i t is felt that the examination of seven to eight hundred herbarium specimens plus considerable cytological study has shown clearly the broad outlines of the genus, but much more cytogenetic work is necessary to est^ ablish the i n t e r - r e l a t i o n 3 h i p s of smeller populations. As for its horticultural possibilities, points both for and against may be mentioned. In its disfavour are the lack of a wide range 92 in morphological variation, especially colour, and the indication that int-erspecific hybridization may not be easily accomplished* On the other hand, the genus has undoubtedly many characteristics valuable in horti-cultural work* Some of these assets may be mentioned* It is suited to a variety of growing conditions: running water, bogs, extremely dry loca-tions, and alkaline soil; i t is apparently readily cross-fertilized or selfed; vegetative propagation is rapid. To the gardener, however, Dode- catheon* s greatest asset is the charm of its appearance: the t a l l scape and the umbel of gayly-coloured, nodding flowers with their re flexed petals and prominent stamens* Such a plant would be an addition to any garden. Besides producing a classification and a number of points of horticultural interest, the study has opened up a wide field for further work in regard to the evolutionary pattern within the genus* The facts as they stand are these* Dodecatheon ranges over most of North America and touches the Siberian Coast on Bering Strait. In the area with which this paper deals, the distribution is general except in northern B.C., Alberta, and Saskatchewan, and here, in part at least, its apparent absence may be due to an absence of collections. In this large area, the ten species re-cognized are generally quite distinct from one another even where their ranges overlap* Three possible exceptions to this situation have been noted, and in these instances hybridization has been suggested: 1. In the region of Spangle, Wash., where D. Cusickii and D.  conjugens overlap, a glandular pubescent conjugens appears. 2. In the Columbia River Valley, where D . Cusickii and D. Hend- erson! meet, D. poetlcum is found combining characters of the two. 3* In southern Oregon, where P.,pauciflorum var* typlcum and D. Henderson! meet, variants of the former occur showing the dark filaments of 93 the latter. Within the species, however, especially the two largest, D. pauc- iflorum and D. Jeffrey!, variation is more common. In both of these large species, within the area studied, the region of greatest variation is from the Cascades south of the American border to Montana. Canadian represent-atives north of this area, and a l l species west of the Cascades and Canad-ian Coast Range to Alaska, appe ar more uniform. A point of interest is the means by which species are distinguish-ed. Floral characteristics, usually the basis of distinction in a genus, here show l i t t l e stable difference. Instead, the most reliable characters, at least for primary separations, are roots and capsules. Somewhat the same situation holds in Delphinium, where roots and seeds are said by EWan (1945) to be most reliable. In seme genera, too, such as Lupinus and members of the family Umbelliferae, fruits provide the clearest distinc-tions. As in these instances, so in Dodecatheon, floral differences are useful for secondary separations. Physiological differences must serve, i f not to distinguish species, at least as isolating mechanisms for species of overlapping ranges often differ in their habitat or time of maturity. Within D. pauc- iflorum such ecological preferences are seen yet no sharp subdivisions could be found. Perhaps here is a species s t i l l evolving, in which isola-tion has not become complete. Cytologically, the genus appears to follow a distinct pattern to which each species conforms. Stated with the reservation that further i work may show a different situation, the pattern is as follows: in the Northwest, diploids occur in the area of greatest variation and extend north into Canada, from B.C. east of the Cascade Range to Saskatchewan; 94 polyploids follow the coast from somewhere south of the American border, along the American and Canadian coast islands to Alaska. The origin of such a pattern is worth considering. The area of variation is just south of the line of maximum glaciation. Have the poly-ploids moved into the glaciated regions since the last Ice Age? Hulten, (1948), suggests that D. frigidum, with its structly northern range, survived gLaciation in unglaciated Alaska and Yukon. Un-fortunately, its genetic constitution is unknown. Diploidy would best f i t the pattern suggested above. Also worth considering is the means of speciation within the gen-us. Polyploidy, though present, has as yet produced l i t t l e morphological effect, for in two of the three species, D. Hendersoni and D. Jeffrey!, where polyploidy has been found, the polyploids are morphologically in-distinguishable from the diploids unless in very minor characteristics. In the third, D. pauciflorum, alight evidence of differentiation exists, possibly a beginning. The similarity of diploid and polyploids suggests autoploidy, possibly from intraspecific crosses. However, even i f differentiation has not as yet occurred, poly-ploidy appears to have greatly increased the range of these three species and probably provided geographic isolation. This seems especially true of D. Jeffrey! and D. pauciflorum with their distribution to Alaska. Moreover, polyploidy may have provided genetic, as well as geographic isolation, but even with isolation established, accumulation of visible differences in an aitopolyploid, with its four (at least) chromosomes of each kind, must be slow. Polyploidy, then, whatever its future role in the evolution of the genus, cannot have been a major agent up to date. Some other mechanism has 95 -been at work in establishing the diploid species* Aside from polyploidy, as mentioned in the discussion of specia-tion, variation may have been produced by gene mutations or chromosomal changes, or a combination of the two. Only further study can solve the problem. This work, then, leaves much to be accomplished, before the gen-etic relationships within and between species can be understood. Yet these relationships form a basic part of further taxonomic work; they also fonn a basic part of horticultural Improvement; they indicate the means of speciation in the genus. Since purely cytological study is hampered by the small size of the chromosomes, further progress may be more easily made by combining cytology with breeding. The outcome of such work might be, not just a new ornamental or a more detailed classification, but also a greater insight into the course of evolution, at least in Dodeoatheon. Plate 23 Diagrams illustrating the terms used in specific descripti A. root type 1. short crown with extremely short fleshy rootlets, (D. Hendersoni only) 2. short crown with fleshy-fibrous rootlets 3. tap root with fie shy-fibrous rootlets B. leaf shape 1. orbicular 2. elliptical 3. ovate 4. lanceolate 5. oblonceolate 6. spatulate C. leaf apex 1. rounded 2. obtuse 3. acute 4* acuminate D. leaf base 1. subcordate 2. sharply contracted 3* broadly cuneate, tapering 4. narrowly cuneate, tapering E. leaf margin 1. entire 2. crenate with calluses in the sinuses 3. repand 4. dentate F. capsule shape 1. cylindrical 2. ovoid Gr. capsule dehiscence 1. circumscissile 2. dehiscing by teeth FLATS 23 A. ! C'a Aa.A. A4. i 96 Appendix I. Complete List of Material Contributed or Collected for Study of the Genus Dodecatheon. 1. PLANTS W. Arlidge •« D. columbianum ..... Seebe, Alt a. Dr. 7.C. Brink •••••• D. Henderson! Southern Vancouver Island. D. Cusickii ........ Cairn Mt., near CIInton,B.C. .»•»•••• Grasmere, B.C. D. columbianum ..... Grasmere, B.C. D. pauoifLorum Dutch Creek, B.C. • Fairmont Hot Springs, B.C. A. Budd • P. columbianum ..... vicinity of Cypress Hills,Alta. Mrs.H.C.Caverly ...<>. P. Henderson! ...... from her garden P. Charter P. pauciflorum var. macrocarpum ....... Mt.Tzouhalem, near Duncan, Vancouver Island. A. Dzubin ........... P. columbianum Natal, B.C. Mrs. Wm. Frohne ..... P. pauciflorum var. . macrocarpum ...... Eklutna Flats,near Anchorage, Alaska. . P. frlgidum ........ Thompson Pass, Alaska. Dr. C.L.Hitchoock ... P. Jeffrey! near Landmark, Ida. P. pauciflorum Montana. T. Midgloy .......... D. dentatum vicinity of Penticton, B.C. R. Pillsbury »• P. Jeffrey! Prince Rupert, B.C. W. Pringle •• P. Cusickii Grand Forks, B.C. 97 1. PLANTS cont'd*.• Dr. T.M.C. Taylor ... D.Cusickii •••••••• from his garden. D. Jeffrey! ....... Stevens Pass. Wash. Personal Collections: p. Cusickii • Richtar Pass.near Osoyooa, B.C. ....... Botanie Valley, Lytton, B.C. Aspen Grove, B.C. ....... Princeton, B.C. P. Jeffrey! •• Stevens Paas, B.C. ....... Forbidden Plateau, Vancouver Island. P. dentatum ....... vicinity of Penticton, B.C. P. pauciflorum var. macrocarpum •••••• Victoria, B.C. ....... Lulu Island,near Vancouver, B.C. • Mt. Arrow smith, Vancouver Island. 2. SEEDS (Specific names aa indicated by the contributor except those given by M. Jack and Dr. V.C. Brink). Dr. V.C. Brink D. Columbianum,.... Wasa, B.C. Curator, Kew Gardens. D. Media, D. Meadia var. integrifolium, D.aallnum, D. Lemoinei. M. Jack from a plant in his garden, possibly D. multiflorum. Dr. Janaki Ammal .... D. Clevelandi Seed bought from Lester Rowntree and E.K.Balls, Cal. P. Jeffrey! P. Hendersoni 98 2. SEEDS cont'd... D. patulum  D. al pinna 3. FIXED MATERIAL FOR CYTOLOGICAL STUDY W. Arldige ......... D. columbianum ••••• Saab*, Alta. Mrs. H.M. Charter .. D. Henderson! •••••• Duncan, Vancouver Island. D. Charter ......... D. Henderson! ...... Duncan, Vancouver Island. A. Dzubln .D. Columbianum Natal, B.C. 0. Fields D. Henderson! ...... Victoria, B.C. Dr. G. Ledinghem ... D. pauciflorum ..... Mortlach, Sask. 4. PRESSED PLANTS A. Budd D. columbianum ••••• Cypress Hills, Alta. D. Charter D. Hendersonl Duncan, Vancouver Island A. Dzubln D. columbianum ..... Natal, B.C. Dr. V. Krajina ••••• D. pauciflorum var macrocarpum ••••••• Sayward, Vancouver Island Dr. G. Ledlngham ... D. pauciflorum ..... Mortlach, Sask. Dr. J. Rattenbury •• D. Cleveland! ...... California. D. Henderson! ...... California. 5 . HERBARIA FROM WHICH PRESSED SPECIMENS WERE EXAMINED National Museum of Canada, Ottawa. Provincial Museum, Victoria, B.C. University of British Columbia, Vancouver, B.C. 99 5 . HERBARIA ERCM WHICH PRESSED SPECIMENS WERE EXAMINED cont'd.... University of California, Berkeley. University of Washington, Seattle. 100 BIBLIOGRAPHY 1 . A n d e r s o n , J . P . 1 9 4 9 , F l o r a o f A l a s k a a n d a d j a c e n t p a r t s o f C a n a d a , 7 1 1 . I o w a S t a t e C o l l . J o u r n . S c i . 2 3 : 1 3 7 - 1 8 7 . 2. B a b c o c k , E r n e s t B r o w n . 1 9 4 7 . The genus C r e p i s I . U n i v . C a l . P u b . B o t . 2 1 : 1 - 1 9 7 . 3 . B r a d l e y , M u r i e l V . 1 9 4 8 . A m e t h o d f o r m a k i n g a c e t o - c a r m i n e s q u a s h e s permanent w i t h o u t r e m o v a l o f t h e c o v e r s l i p . S t a i n T e c h . 2 3 : 4 1 - 4 4 . 4 . C l a u s e n . J . , K e c k , D . D . , a n d H i e s e y , W . M . 1 9 4 5 . E x p e r i m e n t a l s t u d i e s o n t h e n a t u r e o f s p e c i e s I I . C a m . I n s t . W a s h . P u b . 5 6 4 : 1 - 1 7 4 . 5 . D a r l i n g t o n , C D . 1 9 3 7 . R e c a n t a d v a n c e s i n c y t o l o g y . L o n d o n , J . & A . C h u r c h i l l . 6 . D a v i d s o n , J o h n IP. 1 9 4 7 . The p o l y g o n a l g r a p h f o r s i m u l t a n e o u s p o r t r a y -a l o f s e v e r a l v a r i a b l e s i n p o p u l a t i o n a n a l y s i s . Madrono 9 : 1 0 5 - 1 1 0 . 7 . D o b z h a n s k y , T h . 1 9 4 1 . G e n e t i c s find t h e o r i g i n o f s p e c i e s . New Y o r k , C o l . U . P r e s s . 8 . S w a n , J o s e p h . 1 9 4 5 . A s y n o p s i s o f t h e N o r t h . n m e r i c a n s p e c i e s o f D e l p h i n i u m . U n i v . C o l . S t u d . , s e r . D , 1 : 5 5 - 2 4 4 . 9 . F a s s e t t , Norman C 1 9 4 4 . D o d e c a t h e o n i n e a s t e r n N o r t h « m e r i c a . Jn. M i d i . N a t . 3 1 : 4 5 5 - 4 8 6 . 1 0 . F l o v i k , K . 1 9 4 0 . Chromosome numbers a n d p o l y p l o i d y w i t h i n the f l o r a o f S p i t z b e r g e n . H e r e d . 2 6 : 1 1 . G r a y , A s a . 1 8 7 6 . G E m o p e t a l a e . I n G e o l . S u r v . C a l . B o t . I . , C a m b r i d g e , U n i v . P r e s s , e d . 1 . ( O n l y e d . 2 , 1 8 8 0 , h a s been a v a i l a b l e , b u t t h e m a t e r i a l i s s t a t e d t o be t h e s a m e ) . 12«. G r a y , A s a . 1 8 7 8 . S y n o p t i c a l o f N o r t h j t o e r i c a : t h e G a m o p e t a l a e . e d . l ( O n l y e d . 2 , 1 8 8 6 , b e l o w , h a s been a v a i l a b l e , but t h e m a t e r i a l i s s t a t e d t o be t h e same.) 1 3 . G r a y , A s a . 1 8 8 6 a . S y n o p t i c a l f l o r a o f F o r t h . A m e r i c a : t h e G a m o p e t a l a e . W a s h i n g t o n , S m i t h o n i u n I n s t . , v . 1 , p t . 2 ond v . 2 , p t . 1 , e d . 2 1 4 . G r a y , A s a . 1 8 8 6 b . E s s a y t o w a r d a r e v i s i o n o f D o d e c a t h e o n . B o t . G a z . 1 1 : 2 3 1 - 2 3 4 . 1 5 . G r e e n e , E . L . 1 8 9 5 . Some s p e c i e s o f P o d s c a t h e o n . E r y t h . 3 : 3 7 - 4 0 , 7 1 , 7 2 . 1 6 . K a i l , F c r v e y M o n r o e . 1 9 1 2 . New and 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 I . U . C e l . P u b . B o t . 4 : 1 9 5 - 2 0 8 . - 101 17* Harland, S.C., 1934, The genetical conception of the species* Trop. Agr. llsSL-53. 18* Heiser, Charles B., Jr. 1949, Natural hybridization with particular referenoe to introgression* Bot. Rev, 15:645-687. 19. Hulten, Eric, 1948, Flora of Alaska and Yukon T i l l * Lunds Univ. Arssk.N.F., Avd. 2, Bd. 44:1203-1341. 20* Huxley, Julian. 1942. Towards the new aystemetics* In Huxley, J., ed*, The new systematics. Oxford, Clarendon Press. 21. Meyer, James R. 1945* Prefixing with paradichlorobenzene to f a c i l -itate chromosome study* Stain Tech* 20:121-125. 22* Ostergren, 0* 1947* Heterochromatic B<-chromosomes in Anthoxanthua. Hered. 35:261-296. 23. Pax and Knuth. 1905* In Engler, Das Pflanzenre ich 17* Englmann. 1900- unfinished. 24. Rydberg, P.A* 1900* Flora of Montana* Mem. N*Y. Bot. Gard.I. New York. 25* Silow, R.A.1944. The genetics of species development in the Old World cottons..Journ. Gen* 46:62^77. 26. Stebbins, O.L*, Jr. 1940. The significance of polyploidy in plant evolution, i t o . Nat* 74:54-66. 27. Stebbins, G. Ledyard, Jr. 1945. The cytological analysis of species hybrids II. Bot. Rev. 11:463-486. 28. Stebbins, G. Ledyard, Jr., 1947* Types of polyploids; their class-ification and significance. In Dams re c, M», ed. Advances in genetics I. New York, Academic Press. 29* Stephens, S.G. 1950. The internal mechanism of speciation in Gossypium. Bot. Rev. 16:115-149. 30. Upcott, Margaret. 1955. The cytology of triploid and tetraploid Lycopersicum esculentum. Journ. Gen. 31:1*19. . . a . . . . . . 


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