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BULBIFEROUS ACONITUM (RANUNCULACEAE) OF THE WESTERN UNITED STATES DONALD BRINK WOODS JENNIFER and E. A. Box 671327 P.O. AK Chugtak, 99367-1327, U.S.A. KINGSLEY STERN R. Department Biological of Sciences California State University Chico CA Chtco, 95929, U.S.A. urn Columbian.urn Nutt. IS a large and morpholo,gically variiible complex of popul; M ending from southern Canada northern to exico, ind i 1 ilbiferous pc.pulatjons have a restricted distr ibutior oc currmg in the Cascac 1, m )m northern southern Oregon, Kiamat hMour to the itair of northern Californi 1 IS asmall.disjianct group in the CahformaS:erra Nevad ithofLakeTahoe.The: a sot >us aconites ire probably the most distinctive group.wit hi n the A. columbianu < ? and they an treated herein as a subspecies. , RESUMEN Nutt. es un complejo grande variable morfologicamente I y c enden desde el sur de Canada hasta norte de Mexico, desc el y 'ork. Laspoblaciones bulbiferas tienen una distribucion restringid; lillera de la Cascada desde el norte al sur de Oregon, en las Montaiii; ilifornia, y un pequefio grupo disyunto en sur del Lago Tahoe al e Aconitum columbianum Nutt. polymorphic complex a of populations is occurring in the mountains of western North America from Colum- British Canada Chihuahua, Mexico and from bia, to California South Dakota. to Disjunct, outlying populations of A. columbianum occur also in the Driftless New area of Iowa and Wisconsin (Brink and Ohio and York 982), in (Brink 1 & Woods submitted for publication; litis 1965). Previous investigations among indicated extreme differences regional groups of populations, par- morphology and ticularly in floral other correlated This characters. variation was continuous, however, with no clearly defined breaks, populations if throughout the range of the species were considered in aggregate (Brink & deWet Bulbiferous populations 1975, 1980, 1981, 1982; Brink 1980). appeared be consistently somewhat regard since they to were different in this was The from other A. columbianum. purpose of this investigation distinct all bulbiferous aconites, determine the geographical distribution of these to taxonomy group. and review the of this to AND METHODS MATERIALS compared Aconitum populations of the United States were in field, Geographical and uniform nursery investigations. herbarium, laboratory, was determined by investiga- distribution of bulbiferous populations field CHSC, mapping from herbarium specimens (CAS, and by locations tion CLM, OSC, RM, UC, US, UTC, WILLU, WS). Acomtum plants ILL, JEPS, and from bulbiferous and non-bulbiferous populations were transplanted production grown growth chambers and greenhouses study bulbil to in under controlled conditions. we found no intergradation between bulbiferous In investigations field Hitchcock and non-bulbiferous Aconitum columbianum (however, see et al. Abrams The populations that we have seen were either 1964; 1944). producing and maturity with plants of sufficient size bulbiferous, all were completely non-bulbifer- conspicuous bulbils in the leaf axils, or they growth chambers and greenhouses, ous with no bulbil producing plants. In were produced Bulbils only plants from bulbiferous populations bulbils. Hickman Brink (1980) and illustrated in (199.3). an means of vegetative reproduction. In natural Bulbils are effective populations they to the ground late in the season where they sprout fall A may new produce one or vigorously, giving rise to plants. single leaf axil more Bulbil production usually greatest at about the middle of bulbils. is Most and bottom and diminishes toward the top of the plant. the stem, aerial On both and small percentage of bulbiferous plants bear bulbils flowers. a and completely supplants flower production, however, bulbifery plants, bulbils are produced to near the apex of the terminal inflorescence, in place A found Brink more in-depth description of bulbifery can be in of flowers. production should not be confused with the production of Bulbil (1975). one to several small daughter tubers at the first few nodes above the parent tuber, usually below ground. This occurs on a small percentage of the plants and non-bulbiferous populations. bulbiferous in all Bulbiferous Acomtum columbianum have a restricted distribution (Fig. 1), Nevada mountains of with one group of populations occurring in the Sierra Amador California south of Lake Tahoe in El Dorado, and Alpine counties; 350 away northern California in group beginning miles in and another ca. Klamath Mountains of Siskiyou and Del Norte counties. Bulbiferous the Oregon Cascade range populations extend from southern to northern in the Klamath, Douglas, Lane, Deschutes, Linn, Jefferson, Josephine, Jackson, in Hood Wasco, and River counties. Marion, Clackamas, Munz and Keck (1968) were uncertain whether the two disjunct groups of bulbiferous Acomtum in California (Fig. 1) should be treated as separate We morphological separating these can discern no consistent features taxa. two groups (Brink 1975). We Acomtum columbianum the have decided to treat bulbiferous at subspecific level, due in part to the nature of the variation within the species The complex extremely polymorphic (Brink whole. A. columbianum as a is & Wet The 1980, 1982; Brink de 1980). existence of geographical races led names and to a proliferation of published at the specific, subspecific varietal This nomenclatural proliferation was often the result of regional levels. which did not consider patterns of variation within the entire studies, Acomtum columbianum complex. Geographical races that appear quite dis- may from within regional context be indistinguishable other races tinct a Robust and diminutive hundreds even thousands of miles away. occurring or races that are regionally distinct are "connected" by intermediate races if the considered (Brink 1980, 1981, 1982). range of the species entire is We Acomtum polymorphic, have treated columbianum as one, large, complex. Only one group of populations consistently intergrading is which distinct from others. These are the bulbiferous aconites, occur in all We group and Oregon. have chosen to distinguish this as a California Rec- because geographically and morphologically distinct. subspecies it is would seem not to ognition of these bulbiferous aconites at the species level we have not accorded formal taxonomic be warranted, especially since recognition regional variants that are exceedingly diverse, but in charac- to that intergrade throughout the range of the complex (Brink 1982). ters Aconitum columbianum Nutt. Torrey and Gray, N. Amer. 1:34. in Fl. 1838. Tuber 6 x cm; "parent" tuber produces (rarely 2) "daughter" to ca. 1.5 1 Stem contiguous. tubers with a connective so short the tubers are essentially dm cm 2-30 Leaves 5-15 erect and stout to twining and reclining, long. broad, deeply 3-5 (7) cleft, lobes variously incised and toothed. Inflores- Flowers deep cence racemic, terminal, or terminal and axillary. typically Woods a , purple 18-50 to blue, occasionally white, yellowish, or blue-tinged; i mm mm high, helmet 11-34 high, pendent 6-16 sepals long. Aconitum columbianum columbianum la. subsp. Aconitum arizonkum Greene; A. bakeri Greene; A. cheirophyllum Greene; A. columbianum A A ochroleucumA. Nelson; columbianum suh?,p. pallidum Piper; divaricatum Rydberg; . . A. geranioides Greene; A. glaberrimum Rydberg; A. gracilentum Greene; A. helleri Greene; A. infectum Greene; A. insigne Greene; A. leibergii Greene; A. lutescens A. noveboracensevzv. quasiciliatum Fassett; A. ochroleucum Rydberg; A. obtusiflomm Greene; A. patens Rydberg; A. porrectum Rydberg; A. ramosum A. Nelson; A. robertianum Greene; A. sukaesium Greene; A. tenue Rydberg; A. tricorne Greene; A. uncinatum subsp. noveboracense (A. Gray) Hardin; A. vestitum Greene. meadows, Spring-fed bogs, seep areas, along streams, and in other wet areas in the mountains of western North America from British Columbia, Canada Chihuahua, Mexico 900-4,000 m. to at elevations of ca. Brit. Col., Wash., Ore., Calif, Nev., Mont., Idaho, Utah, Dak., Wyo., Ariz., Colo., S. and N. Mex. Disjunct, outlying, populations occur lower at elevations (as New low as 300 m), in Iowa, Wisconsin, Ohio and York. & Chromosome Number: n = [Wiens Halleck 1962, Cates 1968}, 2n = '^ & Kawano 16 [Longacre 1942, 1965, Crawford Gardner 1974, Brink 1975], & = 18 [Crawford Gardner 1974}. 2;z Aconitum columbianum viviparum lb. subsp. (Greene) Brink comb, nov. Basionym: A. viviparum Greene, Feddes Repert, Spec. Nov. Regni Veg. 7:2. OREGON: swamps 1909. Type: near Crater Lake, southern Oregon, Cmick 2912 WS!; (Lectotype, designated herein: isolectotype: UC!). NW Non Aconitum bulhtferumWo^fiW, Fl. Amer. 1:25. 1897. A. Wi-/>ra»2Reichenbach, & Ubers. Aeon. 55. 1819. A. howellii Nels. Macbr., Bot. Gaz. 56:473. 1913. A. NW & columbianum var. howellii (Nelson Macbride) C.L. Hitchcock, Vase. Pis. Pacif. OREGON: 2:231. 1964. marshes eastern slope Cascade Mountains near Mt. Hood, Howell. CALIFORNIA. A. hanseni Greene, Feddes Repert. Spec. Nov. Regni Veg. 1909. 7:3. Amador Hope Hope Co.: Valley, 1892, George Hansen. [Note: Valley actually in is Alpine County.] CALIFOR- A. platysepalum Greene, Feddes Repert. Spec. Nov. Regni Veg. 1909. 7:2. NIA: Mt. S side Shasta, northern California, Jul 1897, H.E. Brown. Spring-fed bogs, seep areas, meadows, along streams, and in other wet m areas at elevations of ca. 900-2,500 in the Cascade Range from norrhern to southern Oregon, in the Klamath Mountains of northern California, and Nevada with small disjunct group of populations in the California Sierra a south of Lake Tahoe. Acomtum These bulbiferous aconites were distinguished as hulbtferum first A homonym name Howell in 897 but this rejected as a later of bulbtferum 1 is , . Reichenbach, which part of A. vartegatum L., a European aconite. Three is names were published simultaneously in 1909 by E.L. Greene for specific One bulbiferous Acomtum. of these names (A vtviparum), was chosen to serve . new which basionym combination, as the for the subspecific refers to all among The was chosen from bulbiferous Aconitum columbianum. lectotype the syntypes of the basionym. and Bulbiferous populations in California have small flowers shallow nectaries (Brink 1975), and we suspected that this would be the case for all We bulbiferous Aconitum columbianum. found, however, that bulbiferous Mount Hood aconites in the region of Oregon, at the northern extreme of and deep the range of bulbiferous populations, have relatively large flowers Flower color in A. columbianum typically deep purple to blue; however, is color variants include completely white, yellowish, cream-colored, or blue- tinged at the edges of the sepals. Color variants occur occasionally within but white-flowered populations, blue-flowered populations, there are also and groups of populations. For example, a group of white-flowered, non- bulbiferous populations occurs in northeast Oregon and southeast Washing- Consequently, not surprising that there are also white-flowered ton. it is Amador bulbiferous populations, which occur in Alpine and counties of Representative Specimens, below). California (see nber: 2n = 18, 19, 20 [Brink 1975}. & Hope Aug Mayer (CHSC) Alpine 29 1974, Brink [flowers white, blue- Co.: Valley, s.n. Amador Dam, Aug McNeal 2730 (OSC) tinged at edges]. Co.: E of Silver Lake l4 1982, Dunn Aug Dorado {white-flowered]. Del Norte Co.: Creek, 6 1938, Keck 4793 (DS). El Co.: Lily Lake, 12-14 Aug 1978, Brink 1469 (CHSC); Echo Summit, 29 Aug 1974, Brink & & Mayer (CHSC). Siskiyou Co.: Red Rock Creek, 26 Jul 1974, Brink Brink s.n. s.n. OREGON. (CHSC); Kangaroo Lake and 24 1974, Brink (CHSC). vicinity, Jul s.n. Clackamas Co.: Clackamas Lake, 24 Jul 1927, Peck 15852 (DS). Deschutes Co.: Tumalo Creek, 12 Aug 1906, Whued 3162 (OSC). Douglas Co.: Diamond Lake, 17 Jul 1924, Applegate4l34 (OSC). Hood River Co.: Barlow Pass, Mount Hood, 3 Jul 1934, Thompson 1 11208 (US). Jackson Co.: WoodruffMeadows, 26jul 192^, Pendleton (OSC). Jefferson s.n. Camp Aug Co.: Sherman, 27 1962, Schoth 1929 (OSC). Josephine Co.: Lake Creek, Klamath Siskiyou Mountains, 18 Jul 1949, Whittaker SS217 (WS). Co.: Pole Bridge, Crater Lake Park, 31 Jul 1922, Applegate 3413 (UC). Lane Co.: Gold Lake, Sep 1962, 1 Monument Aug Marion Dennis 2433 (UTC). Linn Co.: Peak, 9 1947 Alters. (OSC). Co.: n. , Aug Dawn Wasco Meadow, 9 1978, 63 (OSC). Co.: Barlow Road, Sep 964, French Olallie 1 1 2604 (OSC). We acknowledge anonymous M, gratefully reviewers, Mayer, Bissell, L. J. J.M.J. deWet, R.A. Schlising, the herbaria from which loans were obtained OSC, UC, WILLU, We (JEPS, WS), and the herbaria that were visited. Museum appreciate receiving loans through the herbarium of the Cleveland of Natural History (CLM). Univ., Chico. 1980. Reproduction and variation in Aconitum (Ranuncuiaceae), wit! emphasis on California populations. Amer. Bot. 67:263-273. J. (Ranuncuiaceae) of the United States. Ph.D. diss. Univ. of Illinois, Urbana. 1982. Tuberous Acomtum (Ranuncuiaceae) of the continental United States morphological variation, taxonomy and disjunction. Bull. Torrey Bot. Club 109:1 3-23 and De Wet. J.M.J. 1980. Interpopulation variation in nectar production ir and Woods. Acomtum J.A. (Ranuncuiaceae). Flora of North America (submitted for publication), Intermountain Region. Utah Master's thesis. State Univ., Logan. Crawford, D.J. and R.C. Gardner 1974. Documented plant chromosome numben 1974:2. Sida 5:292-294. Hickman, Thejepson J.C. (Ed.) 1993. manual: Higher plants of California. University ol California Press, Berkeley. Hitchcock, M. OwNBEvandJ.W. C.L.A., A. Cronquist, Thompson. 1964. Vascular plants of the Pacific Northwest. Vol. Univ. of Washington 2. Press, Seattle. H.H. The Iltis, 1965. genus Gentianopsis (Gentianaceae): Transfers and phytogeographic comments. 2:129-154. Sida Kawano, S. 1965. Application of pectinase and cellulase in an orcein squash method. Bot. Mag. (Tokyo) 78:36-42. LoNGACRE, 1942. Somatic chromosomes oi Aconitum and A. D.J. noveboracense uncinatum. Bull. Torrey Bot. Club 69:235-239. A MuNZ, PA. and D.D. Keck. 1968. California flora and supplement. Univ. of California Press, Berkeley. WiENS, D. and D.K. Halleck. 1962. Chromosome numbers Rocky Mountain in plants. 1, Bot. Not. 115:455-464.