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NOMENCLATURAL INNOVATIONS IN PHLOX (POLEMONIACEAE), WITH UPDATED CIRCUMSCRIPTION OF P. CAESPITOSA, P. DOUGLASII, P. MISSOULENSIS, AND P. RICHARDSONII PDF

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Preview NOMENCLATURAL INNOVATIONS IN PHLOX (POLEMONIACEAE), WITH UPDATED CIRCUMSCRIPTION OF P. CAESPITOSA, P. DOUGLASII, P. MISSOULENSIS, AND P. RICHARDSONII

NOMENCLATURAL INNOVATIONS PHLOX (POLEMONIACEAE), IN WITH UPDATED CIRCUMSCRIPTION OF AND RICHARDSONII DOUGLASII, MISSOULENSIS, CAESPITOSA, P. P. P. P. James Locklear H. 7431 Briarhurst Circle USA. Nebraska Lincoln, 68506, Phlox condensata subsp. covillei (E.E. Nelson) Locklear, comb, et stat. nov.; P] Locklear, comb, et stat. nov.; Phlox villosissima subsp. latisepala (Wherry) L< Phloxes are components of a remarkably diverse array of plant communities and ecological systems across common North America, and are often among the most herbaceous species in the associations in which they Examples Olympic Mountains, occur. include Phlox in subalpine grasslands of the muscoides in diffusa P. Wyoming Rocky Mountain and Basin rock outcrop communities, pulvinata in alpine subulata fell-fields, P. P. and United in barrens glades in the eastern States. A Wherry The most recent monograph on the genus Phlox (Polemoniaceae) that of (1955). forthcoming is number manuscript on nomenclatural changes proposed here Phlox (Locklear, in prep.) requires for a of taxa, to make these names available for the work. Four of these are changes in rank from variety to subspecies who consistency with concepts of rank in the forthcoming work and those of Wherry, used "subspecies" for in preference to "variety." The concept of subspecies used here to delimit population systems that exhibit is and geographic variation distinctive ecological associations. new proposed combination updated the of condensata subsp. requires Justification for P. covillei cir- new cumscriptions of caespitosa and douglasii, which are provided here. Justification for the proposed P. P. updated combination of richardsonii subsp. borealis requires circumscription of richardsonii, also provided. P. P. an updated circumscription provided. Additionally, of missoulensis is P. NOMENCLATURAL Journal of the Botanical Research Institute of Texas 3(2) comb, ina Coville subsp. lutescens (S.L. Welsh) Locklear, et stat. nov. basiqnym: Phlox aus- & - May Rimrock blackbrush, ash, squawbush, comm., 11 1983, S.L. Welsh, B. Welsh M. Chattzrley 21972 (holotype: BRY; elev., ft paper on austromontana complex, Welsh (1985) recognized four varieties—the first-described In his the P. and most widespread form austromontana) plus three intergrading expressions (vars. jonesii, lutescens, (var. and much more occurring in the canyon country of southern Utah and ad- limited distribution prostrata) of jacent Arizona. These taxa are distinguished at the rank of subspecies because of their discrete distributions been and unique The combination austromontana subsp. prostrata has published ecological associations. P. (Wherry previously 1939). comb, Phlox condensata Gray) E.E. Nelson subsp. covillei (E.E. Nelson) Locklear, et stat. nov. Basionym: (A. Wherry Mountains, 08 Jul 1891. E Covilk with F Funston 1801 (holotype: destroyed by fire: CAS; neotype, designated by 1942: US!; V. B, GH! K, NY, internet image! PH, digital image!). isotypes: from The taxonomic history of condensata complex. Gray (1870) described caespitosa var. condensata P. P. is Rocky Mountains Southern type material collected by Charles Christopher Parry in alpine habitat in the NY, PH, Nelson (1899a) of Colorado (Hololtype: GH!; Isotypes: ISC internet image! digital image!). Elias Wherry elevated to species rank as condensata. (1941, 1955) reduced to P. caespitosa subsp. condensata, P. it it As but Cronquist (1959) elevated back to species rank. detailed in the circumscription of P. caespitosa that it rank condensata treated here the of species. follows, at is P. White Mountains Nelson described from type material collected in the of eastern Elias (1899) covillei P. West occur and subsequent exploration of the Intermountain revealed this entity to in alpine/ California, subalpine situations in a number of other high elevation mountain ranges of the Great Basin as well as in the Wherry taxon but Cronquist Sierra Nevada. (1942, 1955) affirmed recognition of this at the species level, Wherry reduced synonymy under condensata. (1969) disagreed, defending recognition of (1959) to P. it P. from on morphological and complete geographic separation condensata the basis of differences P. covillei am Rocky Mountains, but Cronquist (1984) was not convinced, noting, unable to distinguish the of the "I Wherry Colorado from more western While morphological differences cited by are not plants materials." because profound (see key below), the taxon covillei warrants recognition as a subspecies of its entirely pine separate range, relationship paralleled in the similar bi-centric distribution pattern of the bristlecone a and Intermoun- complex, with the Southern Rocky Mountain expression recognized as Pinus aristata the with pine tain expression as Pinus longaeva (Bailey 1970). Phlox condensata is strongly associated bristlecone New and Mexico and subalpine woodland, subsp. condensata with Pinus aristata in the Rockies of Colorado mountain and Nevada. subsp. with Pinus longaeva in Great Basin ranges in California covillei NELSON TO THE OF PHLOX CONDENSATA GRAY) KEY SUBSPECIES (a. E.E. New cence herbage glandular-pubescent; Southern Rocky Mountains Colorado and Mexico in made by confused and condensata the >ting ecological literature in relation to is difficult floristic P. lature surrounding this species and subspecies. Literature pertaining to subsp. condensata (generally its Cox 1933 Baker Boyce 2005; Cockerell 1903; idensata) includes these: 1983; et (as P. caespitosa); al. & & Langenheim n Hartman Rottman 1988 Kiener 1962 Mitchell 1979; 1985, pulvinata); 1967; (as P. 647 Nomenclatural innovations Phlox Locklear, in & & Weber Wittmann Wooton Shepherd 1975 2001a, 2001b; Standley (as caespitosa)] (as caespitosa)] P. P. 1915 caespitosal Literature pertaining to subsp. covillei (generally as P. covillei) includes these: Billings (as P. & & & Thompson Sydoriak 2001 condensata)] Chabot Billings 1972; Clokey 1951; Goodrich 1957; Botti (as P. 2004 Krantz 1994 Kimball condensata); 1981 Kartesz 1988; condensate)] (as condensate); et (as P. (as al. P. P. & & Mooney Marchand Linsdale 1952; Lloyd Mitchell 1973; Major Taylor 1977; 1973; et al. 1962; et al. & & & Munz Wilken Munz Keck Nelson Kennedy 1906 Patterson 1993 1974; 1959; (as P. dejecta)] (as P. Peinado 2005; Pemble 1970; Sharsmith 1940 (as caespitosa var. condensate); Spira 1991 condensata)] et P. al. & & Wenk Wherry Mooney Dawson 2007 Wright 1965. 1942; condensata); condensata); (as (as P. P. comb, Phlox pilosa subsp. longipilosa (Waterf.) Locklear, et stat. nov. Basionym: Phlox longpihsa Waterf., Rhodora L. and from Waterfall (1971) recognized this entity as related to but distinct pilosa. Taylor Taylor (1981) P. endemic Wichita in the reduced to a variety of pilosa. Phlox pilosa subsp. longipilosa is to granitic soils P. it main Mountains region southwestern Oklahoma, remote from the distribution of pilosa to the east. of P. It shown complex distinguished from the broader pilosa as in the following key. is P. many sometimes Stems per plant numerous, with 0-1 8 leaf-nodes; uppermost leaves and bracts alternate, as 1 mm 2-4 as 7 to 8 being so; inflorescence stems and calyx densely pilose with long jointed hairs mostly long; somewhat awn Phlox pilosa subsp. twisted calyx-lobes apically bearing relatively long, longipilosa Locklear (Waterf.) Phlox sensu pilosa lato on from by John Richardson while serving Hooker described type material collected (1838) richardsonii P. BM, Captain John Franklin's Second (1825-1827) Overland Expedition to the Arctic (holotype: K; isotypes: Cape on GH! NY, Richardson the type in 1826 near Bathurst the image! internet image!). collected digital northwest Canada. Arctic seacoast of up by Linnaeus The has long been with that of described (1753) identity of richardsonii tied sibirica, P. P. Hooker most southern Although (1838) from original material collected in "Asia boreali," likely in Siberia. by Lay and considered the two species he identified specimens of richardsonii collected in 1826 distinct, P. no from Kotzebue Sound on the Alaskan coast as noting the plants differed "in respect" Collie sibirica, at P. The from North American continued treatment phloxes the Arctic as material collected in Asia. of P. sibirica well into the twentieth century, as exemplified by the works of such notable students of the Arctic flora as and Hulten Porsild (1939a, 1939b) (1948, 1968). Wherry North American from recognizing (1955) separated the phloxes of the Arctic sibirica, P. bo- P. and Major morphological differences subsp. subsp. richardsonii richardsonii. richardsonii alashensis, P. realis, P. Wherry by Wherry are reflected in the key below. Not noted by are significant ecological differences, cited being with mid- graminoid-dominated steppe communities, primarily in the associated elevation, sibirica P. (Ermakov Grubov Knyaszev cold, dry interior plateaus of Middle Asia et al. 2002, 2006; 2001; et al. 2007; and Krasheninnikov Mirkin 1985; Peshkova 1972, 2001; Sochava 1945; Yurtsev 1963, 1967, 1972), 1937; et al. with dwarf-shrubland occurring almost above the Arctic Circle in association alpine entirely richardsonii P. and herb communities. alpine cm and branched, forming 8-1 from Plant an upstanding suffrutescent perennial, sparingly diffusely tufts 5 tall cm the erect-ascending flowering shoots 5-1 long with 5 leaf-nodes below the inflorescence; taproot, ca. a h of Texas Institute 3(2) mm; m 20-40 ova pedicel length North complex forms western x richardsonii recognized here as a of intergrading largely limited to the is and and occupying Arctic, differing morphologically in stature size of floral parts distinct floristic :rican shown under )ns of the Arctic. As in the key below, three subspecies are recognized here the first-published new combination subsp. e of richardsonii, necessitating the of P. richardsonii borealis. P. PHLOX HOOK. OF RICHARDSONII SUBSPECIES E mm Longest (4-)8-25 pedicels Ion 1. borealis (Wherry) Locklear coastal areas of the Bering Strait p. mm -6 Longest pedicels long; core 1 I . The distribution of the richardsonii complex ranges from the Bering Sea region of Alaska east to northwest P. The from Canada, mostly north of the Arctic Circle, with a few highly limited occurrences reported Asia. known Canada from few northwest first-described expression (subsp. richardsonii) relatively stations in is Canada, from Cape and northern Alaska. The core of range along the Arctic seacoast of Bathurst in its is mainland promontory on Mackenzie Northwest the District of of the Territories, east to Clifton Point, a Nunavut occurs on Banks Island of the Canadian Arctic Archipelago. Floristic the coast of Territory. also It Aiken and includes ecological literature pertaining to subsp. richardsonii (generally as richardsonii) these: P. & MaCoun Holm Kuc Freedman 1990 Johansen 1999; 1922; 1924; 1974, 1996; et al. et al. (as P. sibirica); Holm 1921. and mountains Phlox richardsonii subsp. alaskensis occurs in the higher interior foothills of Alaska, Yukon, and Northwest occurs most abundantly in the Brooks Range of northern Alaska, the Territories. It but also in the Alaska Range of central Alaska and in the Mentasta, Nutzotin, and Wrangell mountains in Yukon Mountains and Richardson Mountains south-central Alaska. also occurs in the Ogilvie of the Ter- It 2006a ritory. Floristic and ecological literature pertaining to subsp. alaskensis includes these: Carlson et al & P Cody 2002 Cook Roland 2002 Cooper 1989 (as alaskensis); et al. (as P. alaskensis); (as P. richardsonii); & Drew Shanks 1965 1952 Juday 1989 Jordal (as alaskensis) (as sibirica); (as P. sibirica); (as P. sibirica) P. P. ; ; 1972 Spetzman 1959 Walker 1989, 1991, 1994 Porsild (as P. alaskensis); (as P. sibirica); et al. (as P. sibirica). Seward Peninsula northwest Phlox subsp. distributed along the coast of the of richardsonii borealis is on Bering has been reported the Asiatic side of the Alaska, in the vicinity of the Strait. It also (as P. alaskensis) & Bering in the southeastern portion of the Chukotka Peninsula of the Russian Far East (Balandin Strait far On American Razzhivin 1980; Razzhivi 1986, 1994). side of the occurrences of richardsonii subsp. Strait, P. known Norton Sound and Kotzebue Sound, and from Cape Krusenstern, from are coastal areas of borealis Cape Lisburne, Cape Prince of Wales, Cape Sabine, and Cape Thompson. Floristic and ecological literature 2006b Hansen 1951 Carlson pertaining to subsp. borealis includes these: et al. (as P. alaskana); (as P. sibirica); Muir Hutchison 1934 Johnson 1966 Kelso 1989 1917 (as (as P. sibirica); et al. (P. sibirica); (as P. sibirica); P. 1994 1963 1939b Razzhivin 1986, Shetler sibirica); Porsild 1939a, (as P. sibirica); (as P. alaskensis); (as P. subsp. sibirica borealis). show genus The geographic distribution of these subspecies parallels with patterns of speciation in the Douglasia (Primulaceae) (Kelso 1992; Kelso 1994). also roughly corresponds to the extent of three et al. It — Nomendatural innovations Phlox 649 Locklear, in by floristic subdivisions of the Arctic described Yurtsev (1994), each reflecting regional differences in envi- ronment and climate. Subspecies richardsonii associated with the Central Canada subprovince, which is is markedly continental. Subspecies alaskensis associated with the North Alaska subprovince, which encom- is more passes the continental parts of the Brooks Range and northern the Richardson Mountains, foothills, its and which Alaska with the Arctic slope of cold, true Arctic climate, a region experiences colder winter its summer temperatures than coastal areas but also experiences more heat and dryness. Subspecies borealis associated with the Beringian Alaska subprovince, which encompasses the Alaskan coast of the Bering is notably the Seward Peninsula and coastal areas of Kotzebue Sound, Norton Sound, and the Chukchi Strait, warm Sea. This subprovince has an oceanic climate due to the influence of a sea current. While the sum- mer and more climate along the Bering Strait generally colder, windier, overcast than continental areas is of Alaska, the overall climate is less harsh than in the interior mountains due to the proximity of the sea. on on Chukotka Species borealis also occurs the Asiatic side of the Bering Strait the coast of the Peninsula A number in the parallel Beringian Chukotka subprovince. of vascular plants are considered "Beringian" characteristic of or endemic to the Bering Sea region on both the American and Asiatic sides of the Strait more (Kelso 1987, Yurtsev 1994), with the floras of these coastal areas being diverse than that toward the (Kelso interior 1989). Phlox subulata setacea comb, subsp. Locklear, nov. L. (L.) et stat. Basionym: Phlox setacea Sp. 1:153. 1753; Phlox l., PI. name on Linnaeus established the setacea in Species Plantarum (1753) based a figure (table 98, figure 3) P. published in Plukenet's Phytographia (1691). The epithet setacea was subsequently misapplied by Curtis (1798) in the Botanical Magazine for what was an illustration and description of (Loddiges 1823; nivalis P. Sweet 1827) and the two names became entangled until Wherry (1929, 1937) sorted out the confusion and Wherry and reinstated (1929) interpreted setacea as a glabrate expression of subulata P. nivalis. P. P. later name synonymy (1955) placed the Linnaean in under subulata subsp. subulata. Reveal (1982) located et P. al. specimen Herbarium Museum, London which a pertinent in the Sloane of the Natural History (BM-SL), they recognized typotype Plukenet Noting specimen matched as the of the figure. that the clearly the attributes of Wherry what (1929) treated as subulata Reveal (1982) demonstrated that the Linnaean var. australis, et P. al. name epithet setacea has priority over and resurrected subulata a proposed by Brand australis, P. var. setacea, This taxon recognized here the rank of subspecies. (1907). is at Phlox yillosissima (A. Gray) Small subsp. latisepala (Wherry) Locklear, comb. nov. Basionym: PhloxpiiosaL. subsp. Gray (1870) described drummondii from type material collected by Charles Wright in 1851 var. villosissima P. on Nueces the "Pebbly bars of the [River]" in southern Texas (holotype: GH!). Small (1903) elevated to it species rank, but considered villosissima an annual like drummondii. Whitehouse (1935) relocated still P. P. by Nueces where the likely of Wright's collection searching crossings of the River near Uvalde, Texas site she found "numerous plants that closely matched the type" growing in the chalky gravel beds of the river and determined than an that the entity a perennial rather annual, providing further merit for recognition is Wherry recognized but as a species. (1955) the distinctiveness of this taxon, treated as pilosa subsp. it P. riparia. Levin from by (1968) reinstated villosissima as a species distinct pilosa, reproductively isolated ecol- P. P. & Smith and karyology (Levin 1966; Levin 1967) experimental crosses (Levin 1966). Phlox a villosissima is tetraploid (2n = 28) and appears to be an allotetraploid. These data, coupled with those from studies of seed & & protein chemistry (Levin Schaal 1970) and plant pigment (flavonoid) chemistry (Levy Levin 1971, be between 1974) indicate villosissima could the stabilized derivative of past hybridization pilosa subsp. P. P. 650 Journal of the Botanical Research Institute of Texas 3(2) Evidence and drummondii subsp. pilosa mcallisteri. froi P. more complicated this hypothesis but also indicate relati 2002). Amos by Arthur Nelson described aspera from type material collected Heller in central (1899) Elias P. Texas. Brand (1907) discerned a relationship with villosissima, treating the taxon as P. drummondii subsp. P. Wherry drummondii subvar. aspera. (1955, 1966) also recognized a close relationship be- var. villosissima proposing subsp. tween the two, but, as with P. villosissima, treated P. aspera as a variant of P. pilosa, P. pilosa homonym was and Levin (Wherry unavailable subspecific noted Nelson's aspera a as a epithet). latisepala and occupied same noting the entity (1968) reinstated aspera, citing the justifications as for villosissima P. P. morphology and Both Levin an between and in terms of ecology. intermediate position villosissima pilosa P. P. on Edwards and Wherry considered the combined range of this pair of taxa to be centered the Plateau of common and most western portion the plateau Texas, with villosissima/?. pilosa subsp. riparia in the of P. P. more and occurring the north. subsp. to east aspera/?. pilosa latisepala and with two with each other Ferguson (1998) cited possible local integration of these entities P. pilosa noted by Wherry reason maintaining them the subspecies level pending further study. [1966]) as for at (also While combined both under In the most recent treatment of Phlox in Texas, Turner (1998) P. villosissima. may show morphological and these taxa intergrade where their ranges overlap, they distinctive differences The complex key ecological associations toward the outer portions of the overall range of the (see below). and recognized here subspecies of taxon formerly treated as aspera pilosa subsp. latisepala is as a P. vil- P. P. new combination. necessitating the losissima, SMALL GRAY) (A. ton notably of species range, ri> subsp. latisepala (Wherry) Locklear range portion of species OF PHLOX NUTT. CIRCUMSCRIPTION CAESPITOSA Wyeth on by 1833 described from type material collected Nathaniel Jarvis in Nuttall (1834) caespitosa P. Oregon Expedition (1832-1833). Nuttall described the type locality the return (eastbound) leg of his first 20 th The on dry adding, "Flowering about the of April." date "April "Flat-Head the sides of as river, hills," 22 nd " written on the holotype specimen label (BM) and on isotypes (GH, PH) of caespitosa. According P. is Wyeth journal, 22 April 1833 was the day he departed "Flathead Post," also called "Flathead House," to 's trading post historians place not on the Flathead River but on the Clark Fork River near the present-day a A number more town Thompson northwestern Montana (Sanders County) (McKelvey 1955). of of Falls in Thompson made been recent collections of caespitosa have in the vicinity of Falls. P. open-growing form Wherry complex (1941, 1955) interpreted caespitosa as a consisting of a taller, P. more occurring middle elevations in the Rocky Mountains, plus three shorter, compact (subsp. caespitosa) at under and occurring higher elevations alpine/ expressions (subspp. condensata, platyphylla, pulvinata) at subalpine conditions. Cronquist (1959) revised this complex, raising the high elevation forms to the rank and of species as condensata and pulvinata (incorporating subsp. platyphylla) interpreting P. caespitosa P. P. woodland and savanna. open-growing lower primarily in Pinus ponderosa species of elevations, as a taller, and Wherry with considering condensata 1965a, 1965b, 1969) took exception these revisions, (1962, P. pulvinata mere high altitude expressions or "ecads" of caespitosa. However, the ecological divergence of P. P. and from pronounced, and, following Cronquist, condensata these alpine/subalpine entities caespitosa is P. P. pulvinata are recognized here as separate species. P. — Nomendatural innovations Phlox Locklear, in cm and branched, forming open 5-15 the erect-ascending Plant a subshrub, sparingly diffusely tufts tall, 1. cm montane woodland, flowering branches 2.5-7.7 long, with internodes typically apparent; associated with Phlox savanna and grassland < more compact growth forming mats or cushions Plant a caespitose perennial herb, habit pulvinate, or less 1, cm cm upper 2-7.5 the flowering shoots 1-3(-5) long; associated with alpine or subalpine ecological tall, mm 5-10 packed sely together; (eaves appressed, erect, Rocky Mountain Northern As circumscribed here, caespitosa a species of the System, chiefly in the P. is Rocky Mountains southeastern Columbia, northern Idaho, and western Montana. occurs most in British It on Mountains and intermoun- abundantly in Montana, particularly the eastern slope of the Bitterroot in the m 800-1500 tain valleys of the Rocky Mountain Trench. occurs at lower to middle elevations (mostly but It } up 2040 m) piedmont and mountains, and with morainal in the in association features of river valleys. to common woodland and Phlox strongly associated with Pinus ponderosa savanna, but also in caespitosa is is where and montane, and valley grasslands in the region Pseudoroegneria spicata, Festuca idahoensis, foothill, graminoids. Festuca campestris are the chief made and by con- the Interpreting ecological literature in relation to caespitosa difficult floristic P. is name fused nomenclature surrounding this plant, particularly in connection with the douglasii (discussed P. name below), but with added problems coming from the literature of alpine ecology where the caespitosa P. has been misapplied condensata and pulvinata. Literature pertaining to caespitosa includes these: to P. P. P. & & On Shaw Habeck Lackschewitz 1991; Lesica 2002; Root 1972 (as pulvinata); 1979. P. CIRCUMSCRIPTION OF PHLOX D Hooker (1838) described douglasii in his Flora Boreali-Americana. As one of the first of the small-leaved P. name cushion-forming phloxes described from western North America, the subsequently served douglasii P. many number which were as catch-all for later-discovered, superficially similar species, a of treated as va- some nomendatural including or subspecies of point in their histories, andicola, rieties douglasii at P. P. P. and Further confusion was caused by the emergence of the term "Douglasii Hybrids" austromontana, P. diffusa. which was wide cushion phlox in horticultural literature in the early 1900s, applied to a variety of cultivars much Wherry on of probable hybrid origin. brought needed clarity to this situation through his papers the and 1955 monograph which he recognized western phloxes (1938, 1941) his in douglasii as a distinct P. on Columbia Northwest. species with a distribution centered the Plateau in the Pacific synonymy under under Cronquist (1959) reduced douglasii to caespitosa, justifying this treatment his P. P. discussion of pulvinata which he separated from caespitosa. Cronquist described the type of caespitosa P. P. P. known name was as "a compact plant of the taxon usually as P. douglasii." Since the P. caespitosa published by Nuttall (1834) four years before Hooker (1838) published douglasii, Cronquist reasoned that "the latter P. Wherry and (1962, 1965a) considered this "a serious misunderstanding," calling douglasii caespitosa P. P. among Wherry two "about as distinct species as can exist the Microphloxes." Indeed, (1955) placed these species in independent subsections of the genus Douglasianae and Caespitosae. Despite Wherry's protests, most subsequent and ecological workers in the Pacific Northwest have followed Cronquist. Based on floristic BM, examination of the type material of both douglasii (holotype: K, digital image!; isotypes: digital image! P. BM, GH! GH! NY, internet image! PH, digital image!) and caespitosa (holotype: digital image!; isotypes: P. Washington, NY, internet image! PH, digital image!), plus study of these entities in the field douglasii in (P. 652 Journal of the Botanical Research Institute of Texas 3(2) As by Oregon, and California; caespitosa in Montana), they are recognized here as separate species. noted P. Wherry and dark needle-shaped (1962, 1965a, 1969), the leaves of douglasii are green, slender (acicular P. or acerose), and covered by long gland-tipped hairs, while the leaves of caespitosa are pale green, relatively P. broad and and bear coarse glandless along the margins. The distribution (linear-oblong), thickish, cilia flat on Columbia while centered in the Northern of douglasii centered the Plateau, that of caespitosa is P. is P. Rocky Mountains. made and by the con- Interpreting floristic ecological literature in relation to P. douglasii is difficult was fused nomenclature surrounding this species. Prior to Cronquist (1959), this taxon treated as P. Wherry or the recognized by (1938, 1955) as a reduced expression of the former latter douglasii rigida, P. under the name subsp. Following Cronquist's revisions, most workers abandoned these douglasii rigida. P. names Applegate Culver in favor of caespitosa. Literature pertaining to douglasii includes these: 1938; P. P. & 1964 Daubenmire Daubenmire 1968 Driscoll 1964a, 1964b; Eckert (as (as caespitosa); P. diffusa); P. & Munz Keck Peck 1957 (as Hall 1967; Howell 1903; 1959 (as douglasii subsp. rigida); 1941; P. diffusa); P. & The Piper 1906 (as rigida); Piper Beattie 1914 (as P. rigida); St. John 1937 (as P. douglasii var. rigida). P. following key separates from and, additionally, from a similar-appearing douglasii caespitosa P. P. P. diffusa, species that occurs in range. and Northern and Dng) with coarse glandless thickish, cilia; flat, w mostly lacking or fine; ia OF PHLOX CIRCUMSCRIPTION Wherry from by Merton Reed 1939 on Waterworks (1944) described missoulensis the type collected in P. J. Montana PH, Cronquist reduced Hill near Missoula, (holotype: digital image!). (1959) this entity to kelseyi P. Wherry which relationship and reduction with (1962, 1965b) strongly disagreed, calling var. missoulensis, a on missoulensis "a striking endemic...wholly unrelated" to kelseyi. In her thesis the P. kelseyi complex, P. P. Campbell (proposing but not (1992) followed Cronquist in treating missoulensis as a variant of kelseyi P. P. publishing subsp. but cited "extremely different habitats" plus "complete geographical kelseyi missoulensis), P. isolation coupled with differences in 10 morphological characters" in support of recognition of missoulensis and as a distinct entity within the complex. Phlox missoulensis kelseyi are recognized here as sepa- kelseyi P. P. endemic Northern Rocky Mountains rate species (see key below). Phlox missoulensis to the in west-central is where Anaconda, Montana, chiefly along or just west of the Continental Divide occurs in the Garnet, it and and and on mountains with Lewis Clark, Sapphire ranges small in the vicinity of Missoula, outlying occurrences east of the Continental Divide in the Little Belt Mountains. and Leaves dark green, linear to linear-lanceolate, thinnish but with subacerose ciliate surficially pilose with tip, gland-tipped exposed mountain slopes and habitat dry to stony and shallow fine ridges; xeric; soils hairs; Phlox missoulensis and and (becoming wrinkly Leaves pale green, te cuspidate, thickish succulent, fleshy line; on drying), the margin slightly thickened, basally c glabrous to moderately pubescent; mountains and intermountain basins and habitat seasonally moist to wet; fine- valleys soils alluvial, flats; Phlox kelseyi to highly-alkaline aline ACKNOWLEDGEMENTS The author wishes to acknowledge the curators and staff of the following herbaria for the loan of specimens . 653 Nomenclatural innovations Phlox Locklear, in MONTU, CAN, OKLA, RSA-POM, ALA, BM, and from PH, and/or providing images label data specimens: K, WS. NEB, was Thanks Robert Kaul, curator of especially helpful in securing these loans. also to the curators of GH, RM, US work from BRIT, TEX/LL, for access to their collections. This has benefited greatly conversations Andrew with Carolyn Ferguson. thanks anonymous reviewers as well as Hipp, Barney Lipscomb, Finally, to Guy Nesom and comments on Tony Reznicek, and valuable review helpful the manuscript. for M CL M.C G.W. Aiken, S.G., J. Dallwitz, L.L. Consaul, McJannet, L J. Gillespie, R.L. Boles, Argus, J.M. Gillett, PJ. Scott, R. Elven, LeBlanc, A.K. Brysting, and H. Solstad. 999 and onwards. Flora of the Canadian Arctic Archipelago: descriptions, 1 and information Version: 29th April 2003. http://www.mun.ca/biology/delta/arcticf/'. illustrations, retrieval. Monument, Applegate, 1938. Plants of the lava beds National California. Amer. Midi. Naturalist 19 (2): E.I. 334-368. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Ann. Missouri Bot. Gard. Bailey, D. 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Vegetation-soil relationships Eckert, in in R.E., Jr. Oregon Oregon. PhDThesis, State University. Ermakov, M. Cherosov, and Gogoleva. 2002. Classification of ultracontinental boreal forests central Yakutia. N., P. 9-440. Geobot. 37:41 Folia and and and M. Valachovic 2006. Vegetation of the rock outcrops screes the forest-steppe Ermakov, Chytry, in N., B. and Western Sayan southern Phytocoenologia 36:509-545. steppe belts of the Altai Mts., Siberia. L Ferguson, CJ. 1998. Molecular systematics of eastern Phlox (Polemoniaceae). PhDThesis, University of Texas at Austin. DNA A and 2002. chloroplast phylogeny of eastern Phlox (Polemoniaceae): implications Ferguson, CJ. Jansen. R.K. 324-1 of congruence and incongruence with the phylogeny. Amer. Bot. 89:1 335. ITS J. North American Phlox (Polemoniaceae) and 1999. Relationships of eastern Ferguson, Kramer, R.K. Jansen. C.J., F. on sequence 6-631 based ITS data. Syst. Bot. 24:61 V W. tundra vegeta- Freedman, Zobens,TC. 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