Phlox Longifolia Nuttall (Polemoniaceae) Complex of North America

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Theses and Dissertations

1977-07-18

Phlox longifolia Nuttall (polemoniaceae) complex of North America

Frederick J. Peabody Brigham Young University - Provo

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BYU ScholarsArchive Citation Peabody, Frederick J., "Phlox longifolia Nuttall (polemoniaceae) complex of North America" (1977). Theses and Dissertations. 7955. https://scholarsarchive.byu.edu/etd/7955

This Thesis is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. tYK ),oi PHLOX LONGIFOLIA NUTTALL (POLEMONIACEAE) COMPLEX OF NORTH AMERICA ifJZ

A Thesis Presented to the Department of Botany and Range Science Brigham Young University

In Partial Fulfillment of the Requirements for the Degree Master of Science .

by Frederick J. Peabody August 1977 This manuscript of a journal article, by Frederick J. Peabody accepted in its present form by the Department of Botany and Range Science of Brigham Young University as satisfying the thesis requirement for the degree of Master of Science.

ii The Great Basin Naturalist PuBLISHED AT PRovo, UTAH, BY BRIGHAM YOUNG UNIVERSITY

ISSN 0017-3614

VoLUME39 March 31, 1979 No. I

PHLOX LONGIFOLIA NUTT. (POLEMONIACEAE) COMPLEX OF NORTH AMERICA•

Frederick J. Peabody'

ABSTRACT.- Over 1,000 herbarium specimens including 24 type specimens were examined in an attempt to achieve a clearer understanding of the Phlox longifolia complex. Four variables were measured for each specimen and the data were statistically analyzed by discriminant analysis. Using a previously published system of classification for the complex, approximately 73 percent of the measured variation among the specimens was accounted for by that system. The clustering patterns produced in this first analysis indicated that a more conservative approach would be advisable. The formulation and subsequent application of a modified system produced a grouping of specimens that accounted for 95 percent of the measured variation for the four morphological characters considered. From four species and seven subspecies previously recognized, one species with five varieties is proposed. The typification of Phlox longifolia Nutt. and other related taxa is resolved.

The classification of plants belonging to tistical Package for the Social Sciences avail- the Phlox longifolia Nutt. complex has been able at Brigham Young University Computer fraught with difficulties from the very begin- Center. Over 1,000 specimens from nine ning of the nomenclatural history of the western United States herbaria were exam- group. Infraspecific taxa have been attri- ined and included in the sample. Twenty-four buted to one species and then another, new type specimens from an additional 13 her- species have been proposed to deal with the baria were also examined. Classical methods great range of diversity within the group, and of taxonomic research were employed in re- confusion has arisen as to the level or rank of viewing type material and adjusting the no- recognition best suited for any one entity. menclature in order to bring the treatment of The present study has two basic objectives: this complex into agreement with the first, to correlate recent collections and cur- national Code of Botanical Nomenclature rent descriptions with the type material; and (Stafleu et al. 1972). second, to review the classification of the The variables tested were the following: complex in light of a broad sample. vestiture of the inflorescence herbage, corolla The method proposed for the solution of tube length, leaf length, and leaf width. Over the problem is twofold: A test of the most re- the past 160 years of taxonomic history these cently proposed classification system four morphological features have proved to (Wherry 1955), and then a similar test of a be adequate for separation into infraspecific modified system proposed by me. Statistical taxa. They are also easily measured from analyses were conducted with the use of the dried and mounted herbarium specimens. IBM 360 computer using the SPSS V602 pro- Following the descriptions of each taxon in gram for discriminant analysis from the Sta- the taxonomy section is a citation of repre-

1A thesis submitted in partial completion for the degree Master of Science. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602; present address: Department of Botany and Plant Pathology. Iowa State University, Ames, Iowa 50010.

1 2 GREAT BASIN NATURALIST Vol. 39, No. 1 sentative material examined, followed by a tures. Subsequent workers influencing this number indicating the total number of speci- complex inch,ide: E. Nelson (1899), Brand mens examined and placed within that taxon. (1907), Jones (1895, 1908), A. Nelson (1909, The standard abbreviations of herbaria are 1912, 1924, 1931), Wherry (1938, 1939, 1940, those of Holmgren and Keuken (1974). Type 1941, 1942, 1943, 1944, 1955, 1956), Peck specimens were examined and photographed (1941), Jepson (1943), and Mason (1951). and the photographs deposited in the Herba- Over the past 163 years, 16 species and 36 in- rium of Brigham Young University (BRY); fraspecific taxa have been named within this they are indicated by an asterisk(") following complex. Of the 52 names proposed in this the herbarium symbol designation in the list group only 5 are recognized as valid in the of synonyms. The standard abbreviations of present study. serial publications are those of the Torrey Botanical Club (1969). The standard abbreviations of separate works are those of Law- TAXONOMY OF PHLOX LoNGIFOLIA COMPLEX rence (1968). Statistical information is avail- able from the author upon request. Taprooted perennials, 0.3-5 dm tall, from a lignous base; leaves cauline, opposite, ACKNOWLEDGMENTS sometimes alternate above, 5-70 (100) mm long, 1-7 mm wide, linear to oblanceolate; The author wishes to acknowledge the upper cauline leaves glabrous, pubescent or many herbaria and their staffs for the loan of glandular-pubescent; lower cauline leaves material for examination. Appreciation is also glabrescent, basally connate or distinct; pedi- extended to Dr. Stanley L. Welsh for valu- cels (5) 10-40 (50) mm long, glabrous, pub- able assistance in research and preparation. escent, or glandular-pubescent; calyx of 5 The constant support of Robyne, my wife, is aristate to cuspidate, basally connate sepals deeply appreciated. with prominent herbaceous midribs, glabrous except on adaxial surface, pubescent or NoMENCLATURAL HISTORY glandular-pubescent, distinct chartaceous intercostal membranes either plicate or flat; Thomas Nuttall (1834) described Phlox corolla tube (8) 10-28 mm long, usually longifolia from material gathered by Wyeth glabrous but sometimes sparsely pilose within 1833 from the "valleys of the Rocky out, the limb 5-15 mm long, incisorate, erose, Mountains generally". Heller (1897) elevated or entire; stamens unequally inserted on the Torrey's (1859) P. speciosa var. ? stansburyi corolla tube, included or scarsely excerted; to species rank in a cryptic note which af- style 3-cleft, the ovary 3-loculed, the seeds 1 fords little information as to its salient fea- (2-4) per locule.

KEY TO VARIETIES

I. Inflorescence herbage glabrous except calyx within, or pubescent on upper .leaf margins, pedicels and calyx without; leaves linear ...... P. longifolia var. longifolia Inflorescence herbage glandular-pubescent, rarely simply pubescent (var. longipes and var. viridis); leaves linear to oblanceolate ...... 2 2(1). Corolla tube 22-28 mm long; southwestern New Mexico westward across Arizona to southern and western Utah, Nevada and eastern California ...... P. longifolia var. stansburyi

Corolla tube 12-19 mm long; variously distributed throughout western North America ...... 3 March 1979 PEABODY: NORTH AMERICAN PHLOX 3

3(2). Leaves (even upper) distinctly oblanceolate, 3-5 mm wide; inflorescence herbage glandular-pubescent; internodes well spaced; stems stout and often trailing; southwestern South Dakota westward to eastern California ..... P. longifolia var. brevifolia

Leaves linear to linear-lanceolate, 1-3 mm wide; inflorescence herbage rarely nonglandular-pubescent; internodes well spaced or congested; stems slender and usually ascending; variously distributed throughout western North America ...... 4 4(3). Leaves 45-70 mm long, 2.5-4 mm wide; internodes well spaced; northern New Mexico and Arizona through Neva,da, Utah, and western Colorado, narrowly extending into southern Idaho and Oregon ...... P. longifolia var. longipes

Leaves 16-35 mm long, 1-2 mm wide; internodes usually congested; Wash- ington, Oregon, Idaho, extending narrowly into extreme northern Utah and Nevada ...... P. longifolia var. viridis

Phlox longifolia Nutt. var. longifolia P. longifolia Nutt. ssp. humilis (Dougl. ex Hook.) Wherry, Proc. Acad. Nat. Sci. Philadelphia 90: Phlox longifolia Nutt., J. Acad. Nat. Sci. Philadelphia 7: 135. 1938. 41. 1834. Holotype: Valleys of the Rocky Mts. P. longifolia Nutt. ssp. calva Wherry, Proc. Acad. Nat. 0 0 generally, 1833, Wyeth s.n. (BM ). Isotype (K ). Sci. Philadelphia 90: 136. 1938. Holotype: Idaho; P. humilis Dougl. ex Hook., Fl. Boreali-Amer. 2: 72. Butte Co., 13 miles by road southwest of Darling- 1838. Holotype: Oregon; barren sandy plains of ton (43°411h'; 113°341h'), 21 June 1931. E. T. 0 0 the Columbia, 1826, Douglas s.n. (K ). Wherry s.n. (PH ). P. speciosa Pursh var. B Dougl. ex Hook., Fl. Boreali- P. kmgifolia Nutt. ssp. typica Wherry, pro. typ. Notul. Amer. 2: 72. 1838. Holotype: Oregon; on the Nat. Acad. Nat. Sci. Philadelphia 87: 5. 1941. summit of the Blue Mountains and subalpine P. longifolia Nutt. ssp. cortezana (A. Nelson) Wherry, range of the Rockies near perpetual snow, no Notul. Nat. Acad. Nat. Sci. Philadelphia 87: 5. 0 date, Douglas s.n. (K ). 1941. P. sabini Hook. pro syn., Fl. Boreali-Amer. 2: 72. 1838. P. grahamii Wherry, Brittonia 5: 60. 1943. Holotype: P. speciosa Pursh var. linearifolia Hook., Hooker's J. Bot. Utah; Uinta Co., talus slopes west side of Green Kew Gard. Misc. 3: 289. 1851. Holotype: Valley River, south of mouth of Sand Wash, 4500 ft., 0 of the Kooskooskie River and adjoining plains, no elevation, 27 May 1923, Graham 7884 (CM ). 0 date, Douglas s.n. (K ). P. longifolia Nutt. ssp. a-longifolia Wherry, pro. typ. P. linearifolia (Hook.) A. Gray, Proc. Amer. Acad. Arts 8: Morris Arb. Bull. 3: 90. 1956. 255.1870. Plants short to tall, 0.5-5.0 dm; internodes P. longifolia Nutt. f. humilis (Doug). ex Hook.) Voss, Vil- more or less congested or well spaced; leaves morin Blumengartn 1: 681. 1894. P. longifolia Nutt. ssp. marginata Brand, Das opposite, upper leaves glabrous or pubescent, 0 Pflanzenreich 4" : 65. 1907. Holotype: Oregon; nonglandular, (10) 20-50 (60) mm long, steep grassy slopes near Snake River, where it is 1.0-2.5 mm wide, linear to linear-lanceolate; 0 common, 23 May 1901, Cusick 2517, (G ). calyx glabrous or pubescent with intercostal P. longifolia Nutt. ssp. marginata Brand var. humilis (Dougl. ex Hook.) Brand, Das Pflanzenreich 4'"': membranes either plicate or flat; corolla tube 66. 1907. 12-16 mm long (Fig. 1). P. longifolia Nutt. ssp. linearifolia (Hook.) Brand, Das 0 Representative material.- Huntley 889 Pflanzenreich 4" : 66. 1907. P. patula A. Nelson, Univ. Wyoming Puhl. Sci., Bot. 93: (WTU); Parker 578 (OSC); Davis 334 (IDS); 47. 1924. Holotype: Colorado; Platte Canyon, 19 Vickery 550 (ARIZ); Cronquist 6237 0 May 1894,A. Nelson 1589, (RM ). (COW); Tlwmpson 11316 (MONTU); Brown P. marginata (Brand) A. Nelson, Amer. J. Bot. 18: 434. 3780 (UNM). 645. 1931. Distnoution.- Central and eastern Wash- P. cortezana A. Nelson, Amer. J. Bot. 18: 434. 1931. Holotype: Colorado; Montezuma Co., roadside ington, eastern and central Oregon, southern between Cortez and Mesa Verde National Park, Idaho, southwestern Montana, southwestern 0 11May1925, A. Nelson 10436 (RM ). . Wyoming, Nevada. Utah, western Colorado, 4 GREAT BASIN NATURALIST Vol. 39, No. 1

Fig. 1. Phlox longifolia Nuttal var. longifolia. IsoTYPE: Valleys of the Rocky Mts. generally. Wyeth s.n., no date 0 (BMl }. Measure bar equals 1 cm. March 1979 PEABODY: NORTH AMERICAN PHLOX 5 extreme northwestern New Mexico, northern tion of the intercostal membranes of the ca- Arizona, and extreme Califor- lyx, naming this species P. linearifolia (Hook.) nia. This nonglandular fotm is the most abun- A. Gray. dant variety in northern and mesic localities With the passage of years since the work (Fig. 2). of Gray (1870) and the activity of post-Gray- Notes- Nuttall (1834:) described Phlox an monographers, the taxonomy of this group longifolia based on a specimen with long became extremely obscured. Nuttall's type leaves as compared to the caespitose repre- was disregarded, Gray's Phlox linearifolia fell sentatives of the genus, sic. P. caespitosa, P. out of use and bi- or trinomials were applied hoodii etc. Douglas (1838) described P. hu- to erroneous taxa. The longer-leaved and tall- milis based on material shnilar to that of Nut- er P. linearifolia became commonly known as tall's P. longifolia, selectling this epithet be- P. longifolia, leaving the type of P. longifolia cause the leaves were smaller than those of P. to be erroneously renamed at various in- speciosa already described by Pursh (1814) fraspecific levels. Among these renamings and Lindley (1830). Gray (1870) rectified the are: P. longifolia ssp. marginata var. humilis situation by placing Do111glas's P. humilis in (Dougl. ex Hook.) Brand (1907), P. longifolia synonymy with Nuttall's P. longifolia. At the ssp. humilis (Dougl. ex. Hook.) Wherry same time Gray proposed a new species (1938), and P. longifolia f. humilis (Dougl. ex which had longer leaves and distinct replica- Hook.) Voss (1894). This unfortunate condi-

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Fig. 2. Distribution of Phlox kmgifolia Nutt. var. longifolia over western North America. 6 GREAT BASIN NATURALIST Vol. 39, No. 1 tion has continued to the present day, caus- P. stansburyi (Torr.) Heller ssp. superba (Brand) Wherry, ing taxonomic confusion within the complex. Notul. Nat. Acad. Sci. Philadelphia 113: 4. 1942. Wherry (1943) proposed Phlox grahamii Plants moderately tall, 1.5-4.0 dm, inter- Wherry as an intermediate form between nodes well-spaced, 3-6 cm long; leaves oppo- two genera (Phlox and Microsteris). His rea- site, upper leaves somewhat glandular-pub- sons were that the plant in question (Graham escent, 22-39 mm long, 2.1-3.8 mm wide, 7884) combines morphological characters of linear to linear-lanceolate; calyx glandular- each genus. It is perennial and rather tall pubescent with the intercostal membranes with well-spaced internodes and a distinct usually flat; corolla tube 22-28 mm iong corolla tube like Phlox. The leaves, however, (Fig. 3). are relatively small, few, and some are lobed Representative material-Arnott 37, near the base; the corolla tube is strongly (WTU), Maguire 25123 (OSC), Ferris 8054 flaring at the base and the inflorescence is (MONTU), Hershey 2826 (UNM), Wooton sparse as in the genus Microsteris. s.n. (COLO), Wooton s.n. (ARIZ). 38. Upon examination of this specimen, which Distribution- Western New Mexico, Ari- unfortunately is the only specimen of this zona, western Utah, Nevada, and extreme species ever collected or reported, I have eastern California (Fig. 4). found that it is copiously infested with a rust Notes- The taxon stansburyi proves to fit (Puccinia plumbaria Peck) according to iden- into P. longifolia quite well because it has tification made with Arthur (1962). The base many characters in common with other vari- of the corolla tube is filled with hyphae, eties. The long corolla tube usually cited as causing it to flare, and one of the few upper the distinguishing feature merely represents a leaves bears a number of aecia imbedded in point in a continuum of corolla tube length its epidermis that caused it to lobe abnor- within this complex. Other features are essen- mally as it elongated in growth. The general tially the same as in P. longifolia (sensu lato). health of the plant was obviously very poor As defined here var. stansburyi occurs not and all of the features noted by Wherry as re- only in southwestern New Mexico and adja- sembling the genus Microsteris are, in my cent Arizona (Wherry 1956) but also in opinion, the result of teratology. Phlox northern Arizona, Utah, and Nevada. As it grahamii appears to be a diseased P. long- passes northward it appears to assume some ifolia var. longifolia and is therefore reduced of the characteristics of other varieties, most to synonymy. noticeably reduction in leaf size and length of the corolla tube.

Phlox longifolia Nutt. var. stansburyi (Torr.) A. Gray Phlox longifolia Nutt. var. brevifolia (A. Gray) A. Gray Phlox longifolia Nutt. var. stansburyi (Torr.) A. Gray, Proc. Am. Acad. Arts 8:255. 1870. Phlox longifolia Nutt. var. brevifolia (A. Gray) A. Gray, · P. speciosa Pursh var. ? stansburyi Torr., Rep. U.S. Mex. Synop. Fl. N. Amer. Vol. 2 Pt. 1: 133. 1878. Bound. Surv. 2: 145. 1859. Holotype: New Mexi- P. longifolia Nutt. var. stansburyi (Torr.) A. Gray f. co; Dona Ana Co., gravelly hills near the Organ brevifolia A. Gray, Proc. Amer. Acad. Arts 8: 255. Mountains, 30 April 1852, Bigelow s.n. (NY). 1870. Lectotype: Nevada; Ormsby Co., near Car- P. stansburyi (Torr.) Heller, Bull. Torrey Bot. Club 24: son City, 1865. Anderson s.n. (GH°). 478. 1897. P. longifolia Nutt. var. stansburyi (Torr.) A. Gray subvar. P. longituba Heller, Muhlenbergia 2: 228. 1906. Isotype brevifolia (A. Gray) Watson, C. King, Report of (?): California; Inyo Co., Sierra foothills west of the Geographical Expl. 40th Parallel 5: 261. 1871. Bishop in coarse granite sand, 23 May 1906, Hel- P. stansburyi (Torr.) Heller var. brevifolia (A. Gray) E. ler 8320 (BM"). Nelson, Wyoming Agric. Exp. Sta. Annual Rep. P. (Torr.) Heller ssp. eu-stansburyi Brand, 9: 27.1899. Das Pflanzenreich 4'"': 66. 1907. P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand var. P. superba Brand, Das Fflanzenreich 4"": 67. 1907. Holo- brevifolia (A. Gray) Brand, Das Pflanzenreich 0 type: Nevada; Nye Co., Tonopah, May 1905, 4" : 66. 1907. Brown s.n. (UC"). P. grayi Wooton & Standley, pro. syn. Contr. U.S. Nat. P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand f. Herb. 16: 161. 1913. longituba (Heller) Wherry, Notul. Nat. Acad. P. longifolia Nutt. ssp. brevifolia (A. Gray) H. Mason, Nat. Sci. Philadelphia 113: 4. 1942. Abrams Ill. Fl. Pac. States 3: 409. 1951. March 1979 PEABODY: NORTH AMERICAN PHLOX 7

Fig. 3. Phlox lnngifolia Nutt. var. stansburyi (Torr.) Gray. ToPOTYPE: New Mexico, Dona Ana Co., in the Organ Mountains. E. 0. Wooton s.n. 28 May 1905 (WTU•). Measure bar equals 1 cm. 8 GREAT BASIN NATURALIST Vol. 39, No. 1

Plants shorts, often trailing, 1-3 dm tall, one rank to another betrays its complexity. internodes well spaced, 1-4 (5) cm long; Of all varietal names applied over the past leaves opposite, upper leaves glandular-pub- years, Gray's var. brevifolia antedates all and escent, 16-30 mm long, 3-5 (6) mm wide, is chosen as the legitimate name. Because distinctly oblanceolate; upper sometimes lin- Gray indicated no type in the publication of ear-lanceolate; calyx glandular-pubescent, in- this variety, a lectotype (Wherry 1955) has tercostal membranes usually flat; corolla tube been designated (the reader is referred to the 13-15 mm long (Fig. 5). list of synonyms). It is estimated that the Representative material.- Cronquist 8894 range of this taxon extends further eastward (WTU), Lenz 21826 (OSC), Forewood s.n. than any other within this complex, crossing (K), Welsh 9689 (BRY), Christensen s.n. the Contin.ental Divide through Wyoming (BR Y), Blauer 7 (BR Y), Palmer 308 (GH), and into the Black Hills of southwestern Anderson s.n. (GH). 67. South Dakota (fide Forewood 1888). Distribution.- Extreme southwest South Dakota, central western Colorado, northern Arizona, Utah, Nevada, and extreme eastern Phlox longifolia Nutt. var. longipes California (Fig. 6). (M. E. Jones) M. E. Peck Notes.- Variety brevifolia has been named Phlox kmgifolia Nutt. var. longipes (M. E. Jones) M. E. under innumerable combinations over its tax- Peck, Mann. Higher PLS. Oregon 571. 1941. onomic history. Gray's (1870; 1878) and Wat- P. linearifolia (Hook.) A. Gray var. longipes M. E. Jones, son's (1871) transference of this taxon from Contr. W. Bot. 12: 53. 1908. Holotype: Idaho;

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Fig. 4. Distnbution of Phlox longifolia var. stansburyi (Torr.) Gray over western North America. March 1979 PEABODY: NORTH AMERICAN PHLOX 9

Washington Co., Weiser, 28 April 1900, Jones P. longifolia Nutt. ssp. ccnnpacta (Brand) Wherry, Proc. 0 s.n. (POM ). Acad. Nat. Sci. Philadelphia 93: 135. 1938. P. longifolia Nutt. var. puberula E. Nelson, Wyoming P. longifolia Nutt. ssp. longipes (M. E. Jones) Wherry, Agric. Exp. Sta. Annual Rep. 9: 26. 1899. Holo- Proc. Acad. Nat. Sci. Philadelphia 93: 135. 1938. type: Wyoming; Uinta Co., Evanston, 5 June P. viridis E. Nelson ssp. longipes (M. E. Jones) Wherry, 0 1898, A. Nelson 4544 (RM ). Morris Arb. Bull. 3: 88. 1955. P. viscid.a E. Nelson, Wyoming Agric. Exp. Sta. Annual P. viridis E. Nelson ssp. ccnnpacta (Brand) Wherry, Bail- Rep. 9: 25. 1899. Holotype: Oregon; Columbia eya 4: 98. 1956. Co., Blue Mountains, 15 July 1896, Piper 2397 Plants moderately tall 1.5-4.0 dm, inter- 0 (US ). nodes somewhat congested or well-spaced P. stansburyi (Torr.) Heller ssp. ccnnpacta Brand var. vis- 0 1-5 cm long; leaves opposite, upper leaves cida (E. Nelson) Brand, Das Pflanzenreich 4" : 67.1907. glandular-pubescent, 3-10 (15) mm long, P. stansburyi (Torr.) Heller ssp. ccnnpacta Brand var. 1.7-3.6 mm wide, linear to linear-lanceolate; puberula (E. Nelson) Brand, Das Pflanzenreich 0 calyx glandular-pubescent with the inter- 4" : 67. 1907. P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand var. costal membranes usually flat; corolla tube brevifolia (A. Gray) Brand subvar. microcalyx 14-18 mm long (Fig. 7). 0 Brand, Das Pflanzenreich 4" : 67. 1907. Holo- Representative material.- Whites 1037 type: Arizona; Yavapai Co., Prescott Mountain 0 (WTU), Peck 7790 (OSC), Davis 3082 (IDS), District, 1876, Palmer 391 (G ). Cottam 5062 (ARIZ), Clokey 7630 (MONTU), P. puberula (E. Nelson) A. Nelson, Manual Bot. Rocky Mts.: 397. 1909. Clark s.n. (UNM), Hitchcock 20432 (COLO). P. longifolia Nutt. var. filifolia A. Nelson. Bot. Baz. 122. (Crawfordsville) 54: 143. 1912. Holotype: Idaho; Distribution.- Eastern Oregon, southern Blaine Co., Ketchum midst sagebrush, stream Idaho, Utah, western and southern Colorado, bottoms, alt. 5887, 19 July 1911, A. Nelson 1192 0 Nevada, northern Arizona, northwestern (RM ).

Fig. 5. Phlox longifolia Nutt. var. brevifolia Gray. COLLECTION: South Dakota, Black Hills. W. H. Forwood s.n. 0 1887 (K ). Measure bar equals 1 cm. 10 GREAT BASIN NATURALIST Vol. 39, No. 1

New Mexico, and extreme eastern California P. stansburyi (Torr.) Heller ssp. compacta Brand var. (Fig. 8). puberula (E. Nelson) Brand subvar. viridis (E. 250 Notes.- Even though this taxon is found in Nelson) Brand, Das Pflanzenreich 4 : fn. 1907. P. longifolia Nutt. ssp. viridis (E. Nelson) Wherry, Notul. northern as well as southern areas it appears Nat. Acad. Nat. Sci. Philadelphia 87: 5. 1941. to be more abundant in southern Utah, Ari- P. viridis E. Nelson ssp. a-viridis Wherry, pro. typ. Mor- zona, New Mexico, and southwestern Colo- ris Arb. Bull. 3: 88. 1955. rado. The long, narrow leaves and glandular pubescence are distinctive. It appears that Plants short 1-3 dm, stem much branched var. longipes may be a transitional form be- from the base, internodes 1-2 cm long; leaves tween the pubescent var. longifolia and the opposite, upper leaves glandular-pubescent, glandular-pubescent var. stansburyi. The 16-30 mm long, 1.0-1.5 mm wide, linear; ca- overlapping pattern in distribution and mor- lyx glandular-pubescent, intercostal mem- phology would point to this possibility. branes usually flat; corolla tube 13-15 mm long (Fig. 9). Phlox longifolia Nutt. var. viridis Representative material.- Hitchcock (E. Nelson) Peabody stat. nov. 17409 (WTU), Peck 25922 (OSC), Davis 99- 36 (IDS), Stevens 150 (BRY), Stevens 161 Phlox viridis E. Nelson, Wyoming Agric. Exp. Sta. An- (BRY), Platt 157 (BRY). 142. nual Rep. 9: 2.5. 1899. Holotype: Washington; Distribution.- Central Washington, cen- Kittitas Co., Ellensburg, 20 May 1897, Piper 2689 (WS"). tral and southeastern Oregon, southern

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Fig. 6. Distribution of Phlox longifolia Nutt. var. brevifolia Gray over western North America. March 1979 PEABODY: NORTH AMERICAN PHLOX 11

Idaho, extreme northern Utah and Nevada LITERATURE CITED (Fig. 10). ABRAMS, L. 1954. Illustrated flora of the Pacific States. Notes.- Easily distinguished by its fine, Vol. 4, Stanford University Press, Stanford, Cali- narrow leaves, multiple branches, and usually fornia. ARTHUR, J. D. 1962. Manual of the rusts of the United congested internodes, var. viridis is primarily States and Canada. Hafner Publishing Co., New of northern distribution, extending southward York. 438 pp. along the western slope of the Rocky Moun- BENTHAM, G. 1845. Polemoniaceae. In: A. de Candolle, tains into northern Utah and Nevada. Prodromus 9: 302-322. BRAND, A. 1907. Polemoniaceae. In: Das Pflanzenreich 0 4" : 1-203.

Fig. 7. Phlox longifolia Nutt. var. longipes (M. E. Jones) Peck. HoLOTYPE: Idaho, Washington Co., Weiser. M. E. 0 Jones s.n. 28 April 1900 (POM ). Measure bar equals 1 cm. 12 GREAT BASIN NATURALIST Vol. 39, No. 1

CHAUDHRI, M. N., I. H. VEGTER, AND c. M. DEWAL. __ 1851. Catalogue of Mr. Geyer's plants collected 1972. Index herbariorum, Part II-Collectors in the upper Missouri. Hooker's J. Bot. Kew (E-H). Regnum Vegetable, Vol. 9. Kemink en Gard. Misc. 3: 273-305. Zoon N. V. Utrecht, Netherlands. 296-473. JEPSON, W. L. 1925. Manual of the flowering plants of CRONQUIST, A. 1964. In: C. L. Hitchcock, et al. Vascular California. Associated Student Stores, Berkeley, plants of the Pacific Northwest. Univ. Wash. California. 1238 pp. Puhl. Biol. 17(4): 1-510. --· 1943. A flora of California, Vol. 3. University of FISHER, R. A. 1936. The use of multiple measurements California Press, Berkeley, California. 284 pp. in taxonomic problems. Ann. Human Genetics 7: JONES, M. E. 1895. Contributions to western botany. 179-188. Contr. W. Bot. 8: 1-43. GRAY, A. 1870. Revision of the North American Polemo- --· 1908. Contributions to western botany, No. 12. niaceae. Proc. Amer. Acad. Art. 8: 247-282. Contr. W. Bot. 12: 1-100. --· 1878. Polemoniaceae. Synoptical flora of North KuNTZE, 0. 1891. Rev. Gen. Bot. Pars. 2. H. Sturtz, America, Vol. II, Pt. 1. John Wilson and Sons, Wursburg. Cambridge, Massachusetts. LANJOuw, J., AND F. A. STAFLEU. 1954. Index herbario- llELLER, A. A. 1897. Notes on plants of New Mexico. rum, Part II, Collectors (A-D). Regnum Vegeta- Bull. Torrey Bot. Club 24: 477-480. bile, Vol. 2. Kemink en Zoon N. V., Utrecht, --· 1906. Polemoniaceae. Muhlenbergia 2: 228-235. Netherlands. 1-174. HOLMGREN, P. K., AND W. KEuKEN. 1974. Index her- --· 1957. Index herbariorum, Part II, Collectors bariorum, Pt. I, The herbaria of the world. Reg- (E-H). Regnum vegatabile, Vol. 9. Kemink en num vegatabile, Vol. 92. Dosthoek, Scheltema Zoon N.V., Utrecht, Netherlands. 175-295. and Holkema, Utrecht, Netherlands. 397 pp. LAWRENCE, G. M. H., ed. 1968. Botanico-Periodicum- Hoon:R, W. J. 1829-1834. Flora Boreali-Americana. Huntianum. S-H Service Agency, Inc., New York, Vol. I. G. Bohn, London. 298. pp. 1063 pp.

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Fig. 8. Distribution of Phlox longifolia Nutt. var. longipes (M. E. Jones)Peck over western North America. March 1979 PEABODY: NORTH AMERICAN PHLOX 13

LINDLEY, J. 1830. "Phlox speciosa." In: Edward's Bot. __ 1931. Phlox longifolia and Phlox stansburyi and Reg. 16: pl. 1351. their immediate relatives. Amer. J. Bot. 18: MASON, H. L. 1925. Abrams illustrated flora of the Pacif- 441-442. ic States. Stanford University Press, Stanford, NELSON, E. 1899. Revision of the western North Ameri- California. can Phloxes. Wyoming Agric. exp. Sta. Annual NELSON, A. 1909. "Phlox puberula." In' Nelson, A. and J. Rep., No. 9: 1-36. M. Coulter, New manual of botany of the central NIE, N. E., ed. 1975. Statistical package for the social Rocky Mountains (vascular plants). American sciences. 2d ed. McGraw-Hill Book Co., New Book Co., New York. 646 pp. York. -- 1912. "Phlox longifolia var. filifolia." In: Bot. NUTIALL, T. 1834. Plants of the Rocky Mountains. J. Gaz. (Crawfordsville) 54: 143. Acad. Nat. Sci. Philadelphia 7: 5-60. --· 1924. Taxonomic studies by Aven Nelson I. PEcK, M. E. 1941. A manual of the higher plants of Ore- Phloxes, new and old. Univ. Wyoming Puhl. Sci. gon. Binfords and Mort, Portland, Oregon. 866 Bot. 1(93): 47-68. PP· --· 1931. New species from mountains and deserts. --· 1941. Validation of new combinations. Madrono Amer. J. Bot. 18: 431-441. 6: 135-136.

Fig. 9. Phlox longifolia Nutt. var. viridis (E. Nelson) Peabody. HoLOTYPE: Washington, Kittitas Co., Ellensburg. C. 0 V. Piper 2689 20 May 1897 (WS ). Measure bar equals 1 cm. 14 GREAT BASIN NATURALIST Vol. 39, No. I l'uRsH, F. 1814. Flora Americae-Septentionalis. ln two --· 1939. Four southwestern subspecies of Phlox. J. volumes. White, Cochrane and Co., London. Wash. Acad. Sci. 29: 517-519. 91-AFLEU, F. A. 1967. Taxonomic literature. Regnum veg- --· 1940. Geographic relations in the genus Phl,ox. etabile, Vol. 52. Inter-documentation Co. AG, Bartonia 20: 12-14. Utrecht, Netherlands, 566 pp. · -- 1941. Phloxes of Idaho. Notul. Nat. Acad. Nat. --· 1972. International code of botanical nomencla- Sci. Philadelphia 87: 5-13. ture. Regnum vegetabile, Vol. 82. A. Oosthoek's --· 1940. A provisional key to the Polemoniaceae. Uitgeversmaatschappij N. V., Utrecht, Nether- Bartonia 20: 14-17. lands. 426 pp. --· 1942. Phloxes of Nevada. Notul. Nat. Acad. Nat. ToRREY BoTANICAL CLUB. 1969. Index to American bot- Sci. Philadelphia 113: 4-11. anical literature. ln four volumes. G. K. Hall and -- 1943. Microsteris, Phlox, and an intermediate. Co., Boston, Massachusetts. Brittonia 5: 60-63. TORREY, J. 1859. Botany of the boundary. In: Emory, Re- --· 1944. New Phloxes from the Rocky Mountains port on the United States and Mexican boundary and neighboring regions. Notul. Nat. Acad. Nat. survey, Vol. 2. C. Wendell, Printers, Washington, Sci. Philadelphia 146: 1-11. D.C. --· 1955. The genus Phlox. Morris Arb. Bull. 3: Voss, J. 1894. Phl,ox l,ongifolia f. humilis. In: Vilmorin 1-174. Blumengarten 1: 681. --· 1956. Validation of new combinations in Phl,ox. WATSON, S. 1871. The Botany. In: C. King, Report of Baileya 4: 97-98. the geographical exploration of the fortieth paral- WOOTON, E. 0., AND P. C. STANDLEY. 1913. New Plants lel. Vol. 5. U.S. Government Printing Office, from New Mexico-Polemoniaceae. Contr. U.S. Washington, D.C. Nat. Herb. 16: 160-162. W11ERRY, E. T. 1938. Phloxes of Oregon. Proc. Acad. Nat. Sci. Philadelphia 90: 133-140. •

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Fig. 10. Distribution of Phl,ox l,ongifolia Nutt. var. viridis (E. Nelson) Peabody over western North America. PHLOX LONGIFOLIA NUTTALL (POLEMONIACEAE)

COMPLEX OF NORTH AMERICA

Frederick J. Peabody

Department of Botany and Range Science

M.S. Degree, August 1977

ABSTRACT

Over 1,000 herbarium specimens including 24 type specimens were examined in an attempt to achieve a clearer understanding of the Phlox longifolia complex. Four variables were measured for each specimen and the data were statistically analyzed by discriminant analysis. Using a previously published system of classification for the complex approximately 73% of the measured variation among the specimens was accounted for by that system. The clustering patterns produced in this first analysis indicated that a more conservative approach would be advisable. The formulation and subsequent application of a modified system produced a grouping of specimens that accounted for 95% of the variation for the four morphological characters considered.

four species and seven subspecies previously recognized, one species with five varieties is proposed. The typification of Phlox longifolia Nuttall and other related taxa is resolved.

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