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Home , Boraginaceae, Mertensia, Mertensia longiflora, Mertensia oblongifolia, Pulmonaria

Great Basin Naturalist

Volume 58 Number 1 Article 4

1-30-1998

Identity of oblongifolia (Nutt.) G. Don () and its allies in western North America

Ahmed M. Warfa Brigham Young University

Follow this and additional works at: https://scholarsarchive.byu.edu/gbn

Recommended Citation Warfa, Ahmed M. (1998) "Identity of (Nutt.) G. Don (Boraginaceae) and its allies in western North America," Great Basin Naturalist: Vol. 58 : No. 1 , Article 4. Available at: https://scholarsarchive.byu.edu/gbn/vol58/iss1/4

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Great Basin Naturalist 58(1), © 1998, pp. 38-44

IDENTITY OF MERTENSIA OBLONGIFOLIA (NUTT) G. DON (BORAGINACEAE) AND ITS ALLIES IN WESTERN NORTH AMERICA

Ahmed M. Warfal

ABSTRACT.-The current status of Mertensia oblongifoUa (Nutt.) G. Don and its allied taxa is surveyed. On the bases of continuously coherent morphological characters and/or regionally correlated variations, more than 30 taxa, including , subspecies, varieties, and 1 forma, previously considered different from M. oblongifolia, are now placed under synonymy of this species. Those taxa currently known as M. fusiformis Greene, M. bakeri Greene, and M. bakeri var. osterhoutii Williams are among the new synonyms. Typification, , and morphological problems ofM. oblongifo­ lia are discussed.

Key words: Mertensia oblongifolia, typification, Wxorwmy, morphology, allied taxa.

Nuttall (1834) described and depicted Pul­ represents an important additional morpholog­ monaria oblongifolia from a collection ofplants ical feature in the . made by N.B. Wyeth in 1834 chiefly in the Machride (1916) also argued that M. oblon­ valleys of the Rocky Mountains, toward the gifolw had been misinterpreted. He examined sources of the Columbia River (corresponding fragments of a specimen in the Gray Herbar­ to present-day states ofIdabo and Wyoming). ium (GH) which were labeled, in Dr. Gray's As the Linnaean species of hand, "M. oblongifolia Nutt.! ex sp. Wyeth! (1753) in Nortb America were placed within misit Durand 1861." He noted that pedicels of Mertensia (Roth 1797), P oblongifolia Nutt. these fragments were very sparsely hispid; calyx was transferred by Don (1838) into Mertensia. divided nearly to the base, the lobes 5 mm Except for a few additions, Don maintained long, linear-Ianceolate; corolla-tube glabrous Nuttall's description of P oblongifolia for his within, 10 mm long, limb 5 mm long; fIlaments species and was followed by de Candolle as broad and as long as the anthers; style (1846), Gray (1875), and Coulter (1885). slightly exceeds. He concluded that the mor­ M. oblongifolia was later treated as Cerin­ phological characters of the fragments and thodes oblongifolium (Nutt.) Kuntze (1891). Nuttall's description agreed perfectly. Williams Kuntze's contemporary botanists, such as Nel­ (1937:124) also reported the above-mentioned son (1899, 1900), Rydberg (1899, 1900), and fragments in his monograph: "a fragment Piper (1906), and subsequent workers on the marked in Dr. Gray's hand ... is prohably Mertensia (Macbride 1916, Johnston from the type specimen, Wyeth (G)." The word 1932, Williams 1937, Higgins 1993) have rec­ probably indicates doubt as to the identity of ognized M. oblongifolia (Nutt.) G. Don as the the fragment, and actually Williams' doubt correct name. In fact, Cerinthodes oblongifol­ leads to lectotypification ofthe fragment. ium has remained inadequately known since However, the key problems in this study Kuntze's time and seems never to have been concern the typification, taxonomy, and mor­ mentioned again in the literature under Mer­ phology of the species, questions that I have tensia species in North America. examined in connection with a proposed revi­ De Candolle (1846:91) pointed out that M. sion of the genus Mertensia in North America oblongifolia was one ofthe least known species (Warfa in preparation). of the genus Mertensia, but added no further Pulmonaria oblongifolia was described by discussion. However, de Candolle's report that Nuttall (1834:43) as follows: "Glabriuscula, caule the leaves were more or less pubescent beneath simplici erecto, foWs lanceolato-oblongis

IMonle L. Beun Lile SdellCt: Museum, Brighiltn Young University, Provo, ur84602, USA (present address); 558 North Redwood Road #3-1, Salt Lake City, UT 84116, USA (permanent uddress),

38 1998] IDENTITY OF MERTENSIA OBWNGIFOLIA 39 obtusiuscuHs, superioribus acutis, floribus tubu­ apparently rejected by Rydberg (1922:732), loso-carnpanulatis paniculatis pedicellatis, caly­ who kept M. longiflora a separate species. cibus abbreviatis, laciniis linearihus acutis cili­ Rydberg's position was later supported by atis." Nuttall's description implies that he had Jepson (1925:842), Williams (1937:136), Davis seen a collection or a specimen with simple, (1952:592), and John (1956:348). Both Williams erect, and subglabrous stem, etc. In his foot­ and John not only recognized M. longiflora as note Nuttall reported: "Stem ... six to eight a species, but also recognized a number of inches; lower leaves commencing some dis­ synonyms under this species. However, the tance above the base of the stem ... and all status ofM. longiflora has remained at the spe­ more or less pubescent above; panicle formed cific level since then. ofaxillary approximating clusters offlowers ... ; M. foliosa Nelson (1899:243), erected from corolla bright blue; style somewhat exserted." a collection made by Evanston and again ten­ Nuttall thus explicitly stated that he studied a tatively identified and distributed as M. ablongi­ collection or at least a specimen with a com­ folia, was also placed in synonymy of M. ob­ plete habit "six to eight inches." His careful longifolio by Macbride (1916:18-19). Macbride examination of the position of the lower leaves placed M. nutans Howell, M. nevadensis A. above the base of the stem and other Nels., M. puhescens Piper, and M. nutans subsp. described features further confirms his posses­ suhealva Piper together with M. foliosa in syn­ sion of an entire specimen. Don (1838:372) also onymy of M. oblongifolia, making 3 new com­ mentioned a of 1/2 to 3/4 feet. Unlike binations: M. foliosa var. suheolva (Howell) both Nuttall and Don, Gray (1875:53), Mac­ Macbr., M. foliosa var. nevadensis (A. Nels.) bride (1916:17), and Williams (1937:123) appear Macbr., and M. foliosa var. puhescens (Piper) to have seen only the fragments of Nuttall's Macbr. Except for a few modifications, Mac­ specimen at the Gray Herbarium (GH). bride's synonyms under M. oblangifolia were 1 have seen Nuttall's plant collection at later supported by Williams (1937:123, 125, British Museum (BM) and the fragmentary 130). Contrary to Macbride, Rydberg (1922: specimen preserved at GH, the same scraps 732-733) treated M. faliasa and M. nutans as seen by Gray (1875), Macbride (1916), and different species from M. oblongifolia. Simi­ Williams (1937). The fragmentary specimen is larly, Tidestrom (1925:467) considered M. very poor, consisting mostly of dissected flow­ nevadensis, M. foliosa, and M. nutans subsp. ers and a single small leaf As correctly pointed subcalva entities of their own and recognized out by Macbride (1916), this fragmentary mater­ Pulmonario oblongifolia as the only ial is in accordance with Nuttall's description under M. oblongifolia. and the type specimen. Besides Macbride's observation on the rela­ On the same sheet of the type specimen at tionship between M. oblongifolia and M. foli­ BM are 2 other non-type specimens. Although osa, Nelson (1909) studied the affinities be­ these 2 latter specimens were collected much tween M. fusiformis Greene and M. eongesta later and originate from different localities, Greene on the one hand, and M. bakeri Greene, they agree with M. oblongifolia. However, as M. laterifolia Greene, and M. amoena A. Nels. duplicates of the type collection may possibly on the other. Based on these affinities, Nelson exist at the Herbarium of Kew Gardens (K) established 3 new combinations: M. papillosa and/or elsewhere, I choose to desigoate the fWJifarmis (Greene) A. Nels., M. bakeri omoeoo specimen deposited at BM as a lectotype and (A. Nels.) A. Nels., and M. bakeri laterifolia the fragmentary specimen preserved at GH as (Greene) A. Nels. Nelson then placed M. papil­ an isolectotype. lasa fusiformis under M. papillosa Greene, The synonymy ofM. oblongifolio has a long, while M. bakeri amoena and M. bakeri laterifo­ complicated bistory. lia were both placed under M. bakeri. He also Greene (1898:261) was based on a collection placed M. eongesta under M. papillosa, and M. made by Sandberg and Leiberg in 1893, tenta­ eaneseens Rydb. under M. bakeri. Nelsons com­ tively identified and distributed as M. oblongi­ binations and synonymy arrangements were folia. It was placed in synonymy ofM. oblongi­ apparently rejected by both Rydberg (1922: folia by Piper (1906:479), who was followed 734, 1932) and Tidestram (1925:467), who by Macbride (1916:18). This synonymy was treated M. bakeri, M. fWJiformis, M. amoena, 40 GREAT BASIN NATURALIST [Volume 58 and M. la.te!ifolia as species. While Rydberg in this study. As a result of this review, I pres­ placed M. congesta under M. fusiformis, M. eutly treat the species M. oblnngifolia, M. bak­ secundorum Cockerell under M. laterifolia, eri, M. fusiformis, M. foliosa, and M. amoena, as and made nomenclatural transfer of M. well as most of their current synonyms, as a canescens into M. cana Rydb., Tidcstrom placed single morphologically variable but allied group M. paniculata Val: nivalis S. Wats. under M. (see Taxonomic Remarks and Variations). There­ bakeri. As did both Rydberg and Tidestrom, fore, M. oblnngifolia is the only species recog­ Williams (1937:100, 118) considered M. bakeri nized iu this study, while M. fusiformis, M. and M. fu.siforrnis separate species each with a bakeri, and M. bakeri var. osterhoutii Williams number of synonyms. Contraly to Rydberg, are among its new synonyms. Williams placed M. secundorum under M. Although I have not yct examined the M. lanceolata (Pursh) A. DC. and M. laterifolia Inngif/Dra type specimeu (Sandberg & Leiberg under M. bakeri. s. u.) at the herbarium of Notre Dame (ND), Johnston (1932:84-85), aware of the strict its current synonyms, such as M. puk;he/la Piper ecological relationship between M. foliosa and (1906), M. pulchella subsp. glauca Piper (1906), its environments, studied this relationship M. horneri Piper (1906), M. Inngiflnra var. horn­ carefully and affirmed that in response to the eri Macbride (1916), and M. Inngiflora var. pul­ environment, this species exhibited 3 phases chella Macbride (1916), have been examined of morphological variation that correspond to and found to be closely allied to M. ohlongifo­ (1) M. foliosa, (2) M. foliosa var. subcalva, and lia. However, as I have not consulted the type (3) M. foliosa var. amoena (A. Nels.) Johnston, material of M. longiflnra, these taxa are uot respectively. Furthermore, he provided a more included iu this study. Both M. longiflnra aud complete set of synonyms under each of these its synonyms will be placed either in syn­ taxa and suggested that M. foliosa VUl: sub­ onymy to M. ohlongifolia or as infraspecific calva was hetter named M. foliosa var. sub­ taxa to it. calva f. rnacl'J1idei, and M. cusickii Piper and M. praecox Smiley, currently placed under M. eplicata Macbride as M. foliosa var. anwena M. ohlongifolia, is now considered different £ cusickii (Piper) Johnston. M. ob/nngifolia was from this species but rather close to M. arizon­ not mentioned in Johnston's paper. ica Greene. Also, M. stenolnba Greene (1901) In his monumental work, A Monograph aud M. symphytoides Greeue (1901), both cur­ of the Genus Mertensia in North America, rently synonyms to M. ohlongifolia., were not Williams (1937) published the followiug uew treated in this study because I was unable to combinatious uuder M. ohlnngifolia: M. oblnn­ examine the type specimens of these taxa, gifolia var. nevarknsis (A. Nels.) Williams, aud which are probably at the herbarium of ND as M. oblnngifolia var. amoena (A. Nels.) Williams. indicated by Williams (1937:126, 130) and/or He recognized 26 synonyms under M. ohlon­ elsewhere. However, M. praecox, M. sterwloha, gifolia and its varieties (Williams 1937:123, aud M. syrnphytoulcs will be treated together 125, 130), as did Davis (1952:592). Higgins with the remaining taxa of the genus Merten­ (1993:88) later found Williams' varieties of M. sia in North America. oblongifolia ideutical to the species aud placed MATEI\IALS AND METHODS these infraspecific taxa into synonymy. Despite the extensive literature available This paper is based on a study ofherbarium on the genus Mertensia in North America, the type material obtained on loan from BM, BRY, identity of M. ob/nngifolia and its relatiouship CAS, F, GlI, ORE, RM, US, and WILLU with M. bake";, M. fusiformis, etc., have received herbaria (abbreviations according to Holm­ little attention. Lack of information exchange gren et al. 1990), as well as all literature avail­ and/or discordant opinions among early con­ able on the subject. In addition, I consulted a tributors may have overshadowed the signifi­ large set ofM. oblnngifolia collections, deposited cance of this relationship among the taxa in at BRY and representing tlle states and coun­ question. ties in which the species occurs. The purpose of this paper is to review all Only well-developed flowers, nutlets, and literature available on the above-mentioned vegetative parts were used for measurements. taxa and examine all type specimens of all taxa Floral parts (when small) were measured 1998] IDENTITY OF MERTENSIA OBLONGIFOLIA 41

under a Bausch & Lomb stereomicroscope stan, Contr. Arnold Arb. No. 3:85. 1932, syn. nov.; Williams, after softening in ethanol alcohol; a ruler Ann. Mo. Bot. Card. 24:130. 1937. M. oblongifolia val'. amoena (A. Nels.) L.a. Williams, Ann. Mo. Bot. Card. scaled in mm was used for measuring larger 24:130. 1937, syn. nov. Type locality: Montana, Madison plant parts. County, Monida, Clen Creek, Yellowstone Park, In this study I have generally followed tax­ 16.V1.1899, Nelson & Nelson 5413, (RM! holotype, BRY!, onomic concepts commonly used in taxonomic CHi, US! isotypes). revisions based mainly on herbarium material. M. intermedia Rydberg, Mem. N.Y. Bot. Card. 1:335, I consider morphologically coherent units to 1900, syn. nov.; Williams, Ann. Mo. Bot. Card. 24:125. 1937. Type locality: Montana, Bridger Mountains, 17-18.VI.1897, be species; if considerable intraspecific varia­ Rydberg & Bessey 4873 (NY holotype, not seen, CH! iso­ tion is evident, I generally discuss it under type). Taxonomic Remarks and Variations. All syn­ M. congesta Creene, Pl. Baker. 3:17. 1901 (article not onyms are listed in chronological order under seen), syn. nov.; Williams, Ann. Mo. Bot. Card. 24:100. the species. 1937. Type locality: Colorado, Poverty Ridge, above Cimarron, 13.Y1.1901, Baker 129 (ND holotype, not seen, CHI, RM!, US! isotypes). M. oblongifolia (Nutt.) G. Don, Gen. Hist. 4:372. M. lateriflam Greene, PI. Baker. 3:18. 1901 (article not 1838; de Candolle, Prodr. 10:92. 1846; S. Watson, U.S. seen), syn. nov.; Williams, Ann. Mo. Bot. Card. 24:118. CeDI. ExpI. 40th Par. (Bot. King's Exped.) 5:238. 1871; 1937. M. bakeri lat-eriflora (Creene) Nelson, Coult. & Nels., Gray, Proc. Am. Acad. 10:53. 1875; Coulter, Man. Bot. Man. Ry. Mt. Bot. 432. 1909, syn. nov.; Williams 1937:118. Rocky Mt. 262. 1885; Rydberg, ~em N.Y. Bot. Card. Type locality: Colorado, Carson, western Colorado, 1:336. 1900; Macbride, ContI'. Gray Herb. N. S. No. 48:17. 21.VII.1901, Baker 334 (CH! hololotype, RM!, US! iso­ 1916; Tidestrom, Contr. U.S. Nat. Herb. 25:467. 1925; types), Williams, Ann. Mo. Bot. Card. 24:123. 1937; Davis, Fl. M. nutans Howell, Fl. N. W. Am. 491. 1901 (article not Idaho 592. 1952; Higgins, Utah Fl. 88. 1993. Pulmonaria seen), syn. nov.; Williams, Ann. Mo. Bot. Card. 24:125. oblongifolia Nuttall, Jour. Acad. Nat. Sci. Phila. 7:43. 1834; 1937. M. foliosa val'. subcalva (Piper) Macbride, Contr. C. Don, Gen. Hist. 4:372. 1838; de Candolle, Prodr. 10: 2. Cray Herb. K S. No. 48:18. 1916. Type locality: Oregon, 1846. CerintllOdes oblongifolium (Nutt.) Kuntze, Rev. Cen. on high hills, near Coldendale, IV1878 (20.rVl882), How­ PI. Pt. 2:436. 1891. Type locality: IdahoJ\Vyoming, North­ ell s. n. (ORE! holotype). ern Andes, Towards Sources ofColumbia River, 'Wyeth s. n. M. coronata A. Nelson, Bull. Torr. Bot. Club 29:403. (BM! lectotype, CH! isolectotype, designated here). 1902, syn. nov.; Williams, Ann, Mo. Bot. Card. 24:125. 1937. M. paniculata (Ait.) G. Don var. nivalis S. Watson, U.S. Type locality: Wyoming, Sweetwater County, 9.V1.1900, CeoI. Expl. 40th Par. (Bot. King's Exped.) 5:239. 1871, Nelson 7071 (RMl holotype, CHI, ORE! isotype). syn. nov. M. nivalis (Wats.) Rydberg, Mem. N.Y. Bot. Card. M, cusickii Piper, Bull. Torr. Bot. Club 29:643. 1902, 1:336. 1900, syn. nov. Type locality: Utah, Bear River syn. nov.; Williams, Ann. Mo. Bot. Card. 24:130. 1937. M. Canyon, VII1.1869, Watson 844 (CHI holotype). foliosa val'. amoena f. cusickii (Piper) Johnston, Contr. M. bakeri Creene, Pittonia 4:90. 1899, syn. nov.; Arnold Arb. No. 3:85. 1932, s)'n. nov.; Williams, Ann, Mo. Williams, Ann. Mo. Bot. Card. 24:118. 1937. Ty-pe locality: Bot. Card. 24:130. 1937. Type locality: Oregon, Stein's Colorado, southern Colorado, summit of Mt. Hayden, Mts., eastern Oregon, 18.VI.1901, Cusick 2582 (article! 14.VII.1898, Baker, Earle & Tracy 576 (ND holotype, not specimen) (US! holotype, CHI, ORE!, RM! isotypes). seen, CHI, RM!, US! isotypes). M. nevadensis A. Nelson, Proc. BioI. Soc. Wash. 17:96. M. foliosa A. Nelson, Bull. Torr. Bot. Club 26:243. 1899, 1904, syn. nov.; Williams, Ann< Mo. Bot. Card. 24:125. 1937. syn. nov; Williams, Ann. Mo. Bot. Card. 24:125. 1937. Type M.foliosa val'. nevadensis (A. Nels.) (A. Nelson) Macbride, locality: Wyoming, southwest \Vyoming, on the sagebrush Contr. Cray Herb. N. S. No. 48:19. 1916, syn. nov.; slopes in the foothills, 28.V1897, Evanston 2951 (RM! Williams, Ann. Mo. Bot. Card. 24:125. 1937. M. oblongifo­ holotype, CH! isotype). Zia val'. nevadensis (A. Nels.) L.a. Williams, Ann. Mo. Bot. M. /usiformis Greene, Pittonia 4:89. 1899, syn. nov.; Card. 24:125. 1937, syn. nov.; Davis 1952:592; Higgins Williams, Ann. Mo. Bot. Card. 24:100. 1937. M. papillosa 1993:88. Type locality: Nevada, Hunter Creek Canyon, 5 fusiformis (Greene) A. Nelson, Coult. and A. Nelson, Man. miles west of Reno, 16.V1903, Kennedy & True 711 (RMl Ry. Mt. Bot. 421. 1909, syn. nov.; Williams, Ann. Mo. Bot. holotype). Card, 24:100. 1937. Type locality: Colorado, Bob Creek, M. myosotifolia Heller, Colo. Agr. Exp. Sta. Bull. (Fl. West La Plata Mountains, 28.V1.1898, Baker, Earle & Colo.) 100:292. 1906 (article not seen), syn. nov; Williams, Tracy 206 (ND holotype, not seen, Fl, CHI, RM!, US! iso­ Ann. Mo. Bot. Card. 24:118. 1937. M. lanceolata val: types). myosatifolia (Heller) Macbride, Contr. Cray Herb. N. S. M. tubiflora Rydberg, Bull. Torr. Bot. Club 26:544. No 48:15. 1916, syn. nov. Type locality: Colorado, Eagle 1899, svn. nov.; Williams, Ann. Mo. Bot. Card. 24:125. County, Red Cliff, 26.Y1.1900, Osterhout 2164 (MO lecto­ 1937. Type locality: Wyoming, Big Horn Mountains, head­ type, selected by Williams, not seen, CHi isolectotype). waters of the Tongue River, VII.1898, Tweedy 119 (NY M. nutans subsp. subcalva Piper, Contr. U.S. Nat. holotype, not seen, CHI, US! isotypes). Herb. (FL \iVash.) 11:479. 1906; Williams, Ann. Mo. Bot. M. amoena A. Nelson, Bot. Caz. 30:195. 1900, syn. Card. 24:123. 1937. M. foZiosa var. subcalva (Piper) nov.; Williams, Ann. Mo. Bot. Card. 24:130. 1937. M. bak­ Macbride, ContI'. Cray Herb. N. S. No. 48:18. 1916; eri amoena (A. I\els.) A. Nelson, Coult. & A. Nelson, Man, Williams, Ann. Mo. Bot. Card. 24:123. 1937. M. foliosa Ry. Mt. Bot. 422. 1909, syn. nov.; Williams, Ann. Mo. Bot. val', subcalva f. rnacbridei (Macbr.) Johnston, Contr. Arnold Card. 24:30. 1937. M. foliosa var. amoena (A. Nels.) John- Arb. 3:84. 1932, syn. nov.; Williams, Ann. Mo. Bot. Card. 42 GREAT BASIN ATIJRALIST [Volume 58

24:123. 1937. lYpe locality: Washington, Yakima Region, 0.5-2.5(4.5) em, attenuated or tapering, rarely Rattlesnake Mts., 29.1Y.1901, Cotton 328 (US! ho]otype, rounded at the base, acuminate to obtuse, CHI, RM! isotypes). M. pubescem Piper, Contr. U.S. Nat. Herb. (Fl. Wash.) rarely rounded at the apex, entire, scabrous or 11:479. 1906, non de Candolle 1846, syn. nov.; Williams, sparsely to densely ciliate at the margins, gla­ Ann. Mo. Bot. CurcI. 24:130. 1937. M. foliosa var. pubes­ brous to minutely scabrous on both sides, or cen~ (Piper) Macbride, Contr. Gray Herb. N. S. No. 48:19. sparsely to densely pubescent above, glabrous 1916; Williams, Ann. Mo. Bot. Card. 24:130. 1937. Type to scabrous beneath, or densely pubescent on locality: Washington, Douglas County, WaterviJIe. 23.rv.1900, Whited 1214 (US! holotype. ORE! h"Otype). both sides; midrib prominent; winged, M. refrG{;tQ, Nelson, Bot. Gaz. 56:69. 1913, syn. nov.; 6--12 em long, glabrous or pubescent all over. WilIillJJls. Ann. Mo. Bot. Card. 24:118. 1937. Type locality: congested, becoming panicled Colorado, Wagon Wheel Cap. 28.VH. 1912, Griffin 139 with age, with few branches to rather crowded, (RM! holotype. CH! Lsotype). formed of axillary approximating clusters of M. eplicata Macbride,. Contr. Gray Herb. N. S. No. 48:16. 1916, syn. nov. Type locality: Idaho, Boise County, flowers; peduncles up to 6 em long; pedicels Dry Buck, 10.V1911, Macbride 856 (RMl holotype). very slender and often drooping, 1-10 mm long, M. nelsonii Macbride, Contr. Gray Herb. N. S. No. glabrous or pubescent; calyx divided nearly to 48:19. 1916; Williams, Ann. Mo. Bot. Card. 24:123. 1937. the base, 3-8 mm long, enlarging in fruit, gla­ Type locality: Nevada, Elko County, Jarbidge, 9.YII.1912, brous or pubescent, lobes 5, 2-5 mm long, Nelson & Macbride 1995 (RM! holotype, CHI isotype). M. bakeri vnr. subglabra Mil.cbride & Payson, Contr. narrowly linear to lanceolate-triangular, acumi­ Cmy Herb. N. S. No. 49:66. 1917; Williams, Ann. Mo. Bot. nate to acute, sparsely to densely ciliate or Card. 24:l23. 1937. Type locality: Idaho, Custer County, hispid at the margins. Plant hermaphrodite; Josephus Lakes, 3.VIIl.1916, Macbride & Payson 3544 flowers bright blue, occasionally subtended by (CH! holotype, CAS!, RM! isotypes). lanceolate foliar ; corolla tubular-cam­ M. oblongifolta var. nimbata Macbride. ContT. Gray Herb. N. S. No. 53;18. 1918 (article not seen), syn. nov.; panulate, up to 15 mm long, tube 5-12 X 3 rom, Williams, Ann. Mo. Bot. Card. 24:130. 1937. Type locality: lobes 4-5 mm long, obtuse; attached Montana, Bozeman, 18.Y.1893, Cottschalch s. n. (CHI at the throat of corolla, free part of ftlaments holotype). 2-4 mm long, usually dilated, crests or append­ M. cooperae Peck, Torreya 32:151. 1932 (article not ~een), syn. nov; Williams. Ann. Mo. Bot. Card. 24:130. ages in the throat between the bases ofthe ftl­ 1931. Type locality; Oregon, Harney County. 6 miles west aments conspicuous, with a lO-toothed ring at of R~ey, Vl.I922, Cooper 11127 (WILLU! holotype, CHI the hase of the tube; anthers 1.2-2 mm long, isotype). oblong and straight; style 10 rom long, usually M. Weri var. osterhotdii Williams, Ann. Mo. Bot. enclosed or somewhat exserted; nutlets 3 rom Card. 24:120. 1937, syn. nov. Type locality: Colorado, Grand County, Sulphur Springs, 8.VI.1906, Osterhout long, alveolar and white spotted, strongly muri­ 3225 (RM! holotype. CHI ootype). cate, rugose. DISTRlBUTlON.-Mertensia oblongifolia is Perennial 10-50 em tall, with fairly woody, widespread throughout the Mountain and thick, short, erect or vertical rootstocks, usu­ Pacific states ofNorth America. ally branched at the summit; roots numerous HABITAT.-M. oblongifolia is known in slender, fibrous, intermingled with few large clumps and moist open slopes. It is also found woody ones, and the l-..everal crowns closely on plains, hillsides, and/or mountains with covered or clothed with dead brown leaf bases pine woods. It has an altitudinal range from and dead petioles; stems 1 or more from each 7800 to 13,000 feet (2377-3962 m). elongated crown, straight and simple, ascend­ TAXONOMIC REMARKS AND VARlATIONS.­ ing to erect, slightly to fairly conspicuously Mertensia oblongifolia is one of the most mor­ striate or angled, smooth or rough, glabrous or phologically variable species ofthe genus. The densely pubescent with fine, relatively long, variation is probably correlated with geologi­ spreading or closely appressed or crisped­ cal and/or ecological responses. The subconi­ retroflexed hairs. Leaves alternate, green, thick, cal to conical, or shortly fascicled to cushion­ occasionally ample, radical or lower leaves shaped, or rarely tapering rootstock (caudex) commencing some distance above the base of of most type specimens of synonyms examined the stem, few, scattered, petiolate, the upper­ supports such variation. Basal and upper leaves, most numerous or crowded at the summit, often monomorphic in shape, size, and pubes­ sessile to subsessile, with lamina linear-Iance­ cence for most synonyms ofM. oblongifolia, or olate to lanceolate-oblong or spatulate to nar­ rarely dimorphic in some type specimens such rowly oblong-ovate, rarely elliptic, :>-12 x as M. cusick;; (Cusick 2582) and M. eplicata 1998] IDENTITY OF MERTENSIA OBWNGIFOUA 43

(Macbride 856), both at RM, further confirm LITERATURE CITED this variation. Regarding the indumentum, M. oblongifolia varies from entirely glahrous to DE CANDOLLE, A.P. 1846. Borragineae. In: A.P. de Can~ dolle, Prodromus Systematis Naturalis Regni Veg­ completely pubescent. etabilis 10. Paris. VERNACULAR NAMEs.-Bluebell, bluebells, COCKERELL, T.D.A. 1907. A new Mertensia from Colorado. spindle bluebell, western bluebell. Muhlenbergia 3:68. ___.. 1918. Notes on the flora of Boulder County, Colo­ ACKNOWLEDGMENTS rado. Torreya 8(73):177-183. COULTER, J.M. 1885. Manual of the of the Rocky Mountains. Ivison, Blakeman, Taylor, and Company, This study has been performed entirely at New York and Chicago. the Herbarium ofthe Monte L. Beao Life Sci­ DAVIS, RJ. 1952. Flora of Idaho. Brigham Young Univer­ ence Museum of Brigham Young University sity Press, Provo, UT (BRY). I am deeply grateful to Dr. Staoley L. DON, G. 1838. A general history of the dichlamydeous 4. Longman and Co., London. Welsh, Curator of tbe Herbarium, for invalu­ GRAY, A. 1875. Contributions to the botany of North able advice, instructive suggestions, and con­ America. Notes on Boraginaceae. Proceedings ofthe structive criticism of the manuscript. I also American Academy ofArts and Sciences, New Series thank Dr. Duane Atwood, Assistaot Curator, lOAS-52. ___;. 1886. Synoptical flora of North America 2(1). 2nd aod staff of the Herbarium for arranging all edition. Ivison, Blakeman, Taylor, and Company, New loans to BRY York. I express tbanks to Elder Alexander Morri­ Greene, E.L. 1898. New or noteworthy species XXI. Pit­ son and Dr. Ron Gerrison, both of the Corpo­ tonia 3:257-263. __,-.' 1899. West American Asperifoliae IV. Pittonia ration of the President, The Church of Jesus 4B6-97. Christ of Latter-day Saints, for constant inter­ .,.,-__.. 1901. Plantae Bakerianae 3. Washington. est and encouragement and for first approach­ HIGGINS, L.G. 1993. Boraginaceae. Pages 66-92 in S.L. ing BYU concerning my research activities. Welsh et al, editors, A Utah £lora. 2nd edition revised. Similar thanks go to the Deao of the College Brigham Young University Print Services, Provo, UT. HOLMGREN, PK., N.H. HOLMGREN, AND L.C. BARNETT. of Biology and Agriculture, Chairman of the 1990. Index herbm'iorum, part 1, The herbaria of the Department of Botaoy and Range Science, world. 8th edition. Regnum Vegetable 120. and Director of the Life Science Museum, all JEPSON, WL. 1925. A manual of the £lowering plants of of BYU, for hosting my research and placing California. University of California Press, Berkeley and Los Angeles. all necessary facilities at my disposal. JOHN, H.S. 1956. Flora of southeastern Washington and of I am much obliged to directors and cura­ adjacent Idaho. Revised edition. Students Book Cor­ tors of the herbaria mentioned under Materi­ poration, Pullman, WA. als aod Methods for loan of specimens. JOHNSTON, I.M. 1932. Notes on various ofthe west­ ern United States. Contributions from the Arnold All literature or references needed for the Arboretum ofHarvard University 3:83-98. work have been obtained through painstaking KUNTZE, O. 1891. Revisio Generum Plantarum 2. Leipzig. work by Dr. Nathan M. Smith, Science Librar­ LINNAEUS, C. 1753. 1. Edition 1. iao of the Life Science Museum, as well as the Holmiae. staff ofthe Harold B. Lee Library ofBYU. MACBRIDE, J.E 1916. The true Mertensias of western North America I. Contributions from the Gray Herb­ Also, maoy tbanks are due my friends aod arium of Harvard University, New Series 48:1-20. colleagues at the Monte L. Bean Life Science MACBRIDE, J.E, AND E.B. PAYSON. 1917. New or otherwise Museum for support and encouragement, par­ interesting plants from Idaho IV Contributions from ticularly, Dr. Douglas Cox, Assistant Director the Gray Herbarium of Harvard University, New Series 49:60-72. of the museum, and Ms. Terry Simmons, mu­ NELSON, A. 1899. New plants from Wyoming VII. Bul­ seum secretary, for introducing me to comput­ letin of the Torrey Botanical Club 26:236-250. ers and related facilities. _~,,' 1900. Contributions from the Rocky Mountain Comments and suggestions on the final draft Herbarium I. Botanical Gazette 30(3):189-203. _~,," 1902. New plants from Wyoming XIV Bulletin of of the manuscript by Dr. Larry C. Higgins of the Torrey Botanical Club 29:400-406. St. George, Utah, Ms. JoAnne Abel of the . 1904. New plants from Nevada. Proceedings of Great Basin Naturalist, and Dr. Mats Thulin of --c'''he Biological Society ofWashington 17:91-98. Uppsala University are appreciated. ___.. 1909, In: J.M. Coulter and A. Nelson, editors, New Finaocial support has been given by the manual of botany of the central Rocky Mountains (vascular plants). Cincinnati and New York. Corporation of the President, The Church of _-co' 1913. Contributions from the Rocky Mountain Jesus Christ ofLatter-day Saints. Herbarium XIII. Botanical Gazette 56(1):63-71. 44 GREAT BASIN NATURALIST [Volume 58

NELSON, A., AND T.D.A. COCKERELL. 1903. Three new . 1904. Studies on the Rocky Mountain Flora XIII. plants fi·oIU New Mexico. Proceedings ofthe Biolog­ -~B;;-u'l1etin ofthe Torrey Botanical Club 31:631-655. ical Society ofWashington 16:45-46. __~,. 1922. Flora of the Rocky Mountains and adjacent NU'lI'ALL, T. 1834. A catalogue of a collection of plants plains. Hafner Publishing Co., New York made chiefly in the valleys of the Rocky Mountains ---c;" 1932. Flora of the prairies and plants of central or Northern Andes, towards the source of the Colum­ North America. New York Botanical Garden, New bia River. Boragineae. Journal of the Academy of York Natural Science of Philadelphia 7:43-45. TIDESTROM, I. 1925. Flora of Utah and Nevada. Contribu­ OSTEHIIOUT, G.E. 1917. A new Mertensia. Torroya 17: tions from the United States National Hcrbm'ium 175-176. 25,7-635 PIPRR, C.v 1902. New and noteworthy northwestem plmlts WATSON, S.,.IIT AL. 1871. Botany. In: C. King, editm; United VII. Bulletin ofthe Toncy Botanical Club 29:642-646. States geological exploration of the fortieth parallel. ___' 1906. Flora of the State ufWashington. Boragina­ Government Printing Of6ce, Washingtoll, DC. COlle. Contributions from the United States National WILLIAMS, L.O. 1937. A monograph of the genus Merten­ Herbarium 11:412-486. 8m in North America. Annals ofthe Missouri Botani­ ROTH, A.W. 1797. Catalecta Botanica 1. Leipzig. cal Garden 24:17-159. RYDllEHG, PA. 1899. New species from the western United States. Bulletin of the Torrey Botanical Club Received 7 May 1997 26,541-546, Accepted 11 August 1997 . 1900. Catalogue of the flora of Montana and the --~~~e'llowstone National Park Boraginaceae. Memoirs ofthe New York Botanical Garden 1:326-337.