INTRODUCING THE SPOTTED SAXIFRAGES: SAXIFRAGA SECT. BRONCHIALES, SECT. NOV. (SAXIFRAGACEAE) ERIC G. DECHAINE

RHODORA, Vol. 116, No. 965, pp. 25–40, 2014 E Copyright 2014 by the New England Botanical Club DOI: 10:3119/13-09; first published online April 9, 201...
Author: Scarlett Marsh
1 downloads 1 Views 255KB Size
RHODORA, Vol. 116, No. 965, pp. 25–40, 2014 E Copyright 2014 by the New England Botanical Club DOI: 10:3119/13-09; first published online April 9, 2014.

INTRODUCING THE SPOTTED SAXIFRAGES: SAXIFRAGA SECT. BRONCHIALES, SECT. NOV. (SAXIFRAGACEAE) ERIC G. DECHAINE Western Washington University, Biology Department, 516 High St., Bellingham, WA 98225 e-mail: [email protected] ABSTRACT. Recent molecular phylogenetic analyses have revealed that Saxifraga sect. Trachyphyllum as currently circumscribed is polyphyletic, with the designated type, S. aspera, falling outside of the more speciose clade. Several other lines of morphological, geographic, and ecological evidence also distinguish the two distantly related groups from one another. In combination, these data necessitate the circumscription of a new section, S. sect. Bronchiales, which is described herein. The section is hypothesized to have arisen in the southern Rocky Mountains of North America, followed by northwestward expansion across Beringia and diversification among refugia.

Key Words: arctic-alpine, Asia, Beringia, endemism, Haida Gwaii, Japan, Pacific Northwest, Rocky Mountains, Saxifraga sect. Bronchiales, S. sect. Trachyphyllum, Saxifragaceae, Siberia

Plants of the genus Saxifraga L. (Saxifragaceae Juss.) are found primarily in the arctic, montane, and temperate environments of the northern hemisphere, but also in South America and northern Africa. The approximately 400 plus species of saxifrages are defined by a more or less consistent floral structure of five sepals, five petals, ten stamens in two whorls (outer opposite sepals and inner opposite petals), two carpels that are united at least in part, and an absence of a free hypanthium, though vegetative characters vary considerably (Brouillet and Elvander 2009; Gornall 1987; McGregor 2008; Webb and Gornall 1989). The taxonomic diversity within the genus has been subdivided into fifteen sections based primarily on variation in morphological characters (Gornall 1987; Webb and Gornall 1989), but further redefined by molecular phylogenetic analyses (Conti et al. 1999; Soltis et al. 1996; Vargas 2000; Zhang et al. 2008). The taxonomy of Saxifraga sect. Trachyphyllum has been rewritten several times, with modern taxonomists agreeing that the section is in need of thorough revision (Brouillet and Elvander 2009; Elven et al. 2007; McGregor 2008; Webb and Gornall 1989; Zhmylev 1988). Gaudin (1828) described sect. Trachyphyllum as being low, alpine herbs with lanceolate, spinulose-ciliate leaves and 25

Rhodora rhod-116-965-02.3d 17/4/14 01:34:43

25

Cust # 13-09

26

Rhodora

[Vol. 116

white flowers, and assigned S. aspera L. as the type for the section that also included S. bryoides L. Koch (1837) recircumscribed sect. Trachyphyllum to include S. aspera, S. bryoides, S. tenella Wulfen, and S. aizoides L., and provided a more detailed description. A few attempts were made to reclassify the members of sect. Trachyphyllum (among other Saxifraga) and place them in the genera Ciliaria Haworth or Leptasea Haworth, but those revisions were not followed (Don 1822; Haworth 1821; Small 1905; Weber 1982). The work of Engler and Irmscher (1919) broadened the description of sect. Trachyphyllum to also include taxa from Siberia and North America: S. bronchialis L. [including several infraspecific taxa that had been described at the rank of species (Don 1822; Small 1905)] and S. tricuspidata Rottb. (with three varieties). This account was followed by a number of regional assessments. For instance, Losina-Losinskaja (1939) and Siplivinsky (1971) added several Siberian taxa to the section. North American additions were provided by Calder and Savile (1959), who described the Haida Gwaii endemic S. taylorii Calder & Savile. Khokhryakov (1979) and then Zhmylev and Khokhryakov (1985) subdivided the section into subsections and series based on morphological characters. At that time, Zhmylev and Khokhryakov (1985) had suggested the possibility of dividing sect. Trachyphyllum into two sections—one section for subsect. Fibrosophyllae Zhmylev & A.P. Khokhr. (that included S. aspera, the type for sect. Trachyphyllum, and S. bryoides) and another section containing the remainder of the species. Later, Gornall (1987), followed by Webb and Gornall (1989) and McGregor (2008), provided a synthesis that included all known members of sect. Trachyphyllum to date. Multiple lines of evidence support the division of sect. Trachyphyllum and the circumscription of a new section. First, subsect. Fibrosophyllae can be distinguished morphologically from the other taxa in sect. Trachyphyllum by leaves with fibrous margins (hence the name of the subsection) and forward-pointing cilia, conspicuous buds in the leaf axils at flowering, and a large yellow blotch at the base of their otherwise white petals. Second, both members of subsect. Fibrosophyllae are restricted geographically to the mountains of Europe (Alps, Appenines, Carpathians, and Pyrenees), whereas all the other species in sect. Trachyphyllum have a trans-Beringian distribution, ranging from the Rocky Mountains of North America across Beringia and westward to the mountains of Russia, Japan, and China (McGregor 2008; Webb and Gornall

Rhodora rhod-116-965-02.3d 17/4/14 01:34:43

26

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

27

1989). Third, subsect. Fibrosophyllae are distinct ecologically in their dependence upon silicaceous substrates. In addition to the aforementioned morphological, geographic, and ecological differences, recent molecular phylogenies clearly show that species traditionally associated with sect. Trachyphyllum are distributed across two distantly related clades. DeChaine et al. (2013) used chloroplast trnL-F and nuclear ITS (internal transcribed spacer, including the entire ITS1-5.8s-ITS2 region) genetic sequence data in both maximum likelihood and Bayesian analyses to reconstruct phylogenies for Saxifraga, focusing on 19 members (including all accepted species) of sect. Trachyphyllum. All genetic markers and phylogenetic analyses agreed that the section was polyphyletic. One of the inferred clades was restricted to subsect. Fibrosophyllae and included the type species for sect. Trachyphyllum, S. aspera, whereas the remaining taxa were nested within a separate, strongly supported monophyletic clade (referred to therein as ‘‘Clade T2’’) positioned basally within the genus (DeChaine et al. 2013). The latter clade is the focus of this paper. All members of the new section of Saxifraga described herein were previously included in sect. Trachyphyllum (Gaud.) Koch, the Rough-Leaved Saxifrages, because they usually exhibit stiff hairs along the margins and tips of their evergreen leaves that give them a rough appearance and feel. Now, morphology, geography, ecology, and phylogeny (DeChaine et al. 2013) all warrant the description and circumscription of sect. Bronchiales DeChaine, sect. nov. TAXONOMIC TREATMENT

Saxifraga L. sect. Bronchiales DeChaine, sect. nov. TYPE: Saxifraga bronchialis L. Members of sect. Bronchiales, the Spotted Saxifrages, are characterized (see Figure 1) as: Plants evergreen perennial herbs, loosely to densely mat-forming. Flowering stems leafy, terminal. Leaves in basal rosette and cauline, alternate; blades linear to lanceolate, usually stiff, entire or 3-toothed, apiculate or strongly mucronate apically, glabrous, sometimes glandular with multiseriate hairs; lime-secreting hydathodes absent. Inflorescence panicle or sometimes reduced to a solitary flower. Flowers regular; sepals 5, erect to ascending; petals 5, 6 oblong, white to pale

Rhodora rhod-116-965-02.3d 17/4/14 01:34:43

27

Cust # 13-09

28

Rhodora

[Vol. 116

Figure 1. Illustration of typical characters of Saxifraga sect. Bronchiales. A. Flower of S. austromontana (DeChaine 12-1250, WWB). B. Flower of S. derbekii (Zhmylev s.n., ALA). C. Whole plant of S. funstonii (DeChaine 11-37, WWB). D. Leafy rosette of S. trachyphyllum (DeChaine 12-176, WWB) showing 3-lobed leaves. E. Leafy rosette of S. bronchialis (Kharkevich 25, UBC). F. Leafy rosette of S. omolojensis (Roland 92-120, ALA). G. Leafy rosette of S. funstonii (DeChaine 11-921, WWB).

Rhodora rhod-116-965-02.3d 17/4/14 01:34:43

28

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

29

yellow, usually with yellow, orange, red, or pink-purple spots; stamens 10; ovary 6 superior; styles 2; stigmas 2. Capsule ovoid to conical. Seeds oblong. SIMILAR TAXA.

Members of sect. Bronchiales are readily distinguished from sect. Trachyphyllum because the latter have leaves with fibrous margins and forward-pointing cilia, conspicuous buds in the leaf axils at the time of flowering, and a large yellow blotch at the base of their white petals. KEY DISTINGUISHING SECT. BRONCHIALES AND SECT. TRACHYPHYLLUM

1a. Leaves usually with cartilaginous margins; buds not conspicuous at flowering; and petals with spotting or no coloring, but not a yellow blotch at base . . . . . . . . . . sect. Bronchiales 1b. Leaves with fibrous margins and forward directed cilia; conspicuous buds in the leaf axils at the time of flowering; and a large yellow blotch at the base of their white petals . . . . . . . . . . . . . . . . . . . sect. Trachyphyllum DISTRIBUTION AND HABITAT.

Saxifraga sect. Bronchiales is a transBeringian section, occurring in the mountains of western North America (though not in the Sierra Nevada of California), from New Mexico to Alaska including Oregon and Washington and eastward from Alaska across boreal and arctic North America to Newfoundland and Labrador (Canada), and in Greenland. In Eurasia, populations are found across boreal and arctic Russia, from the Kuril Islands, Sakhalin, Kamchatka, and Chukotka, north to Wrangel Island and west to the Ural Mountains, in Heilongjiang and Nei Mongol China, and south to the alpine environs of Japan. Like other members of Saxifraga, plants primarily inhabit arctic, alpine, and sub-alpine environments, but also occur at lower elevations on islands around the North Pacific, such as Haida Gwaii (Canada). Plants generally grow on rocky cliffs, talus and scree slopes, and gravelly flats. NOMENCLATURAL HISTORY

Many of the species of sect. Bronchiales are readily distinguishable, but questions regarding some taxonomic ranks and the

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

29

Cust # 13-09

30

Rhodora

[Vol. 116

number of subspecies, varieties, and forms beg further scrutiny. Based on the genetic lineages inferred by DeChaine et al. (2013) and previous taxonomic assessments, the following species are included within sect. Bronchiales. 1. Saxifraga austromontana Wiegand, Bull. Torrey Bot. Club 27: 389 (1900). TYPE: Santa Fe Canon, 9 miles east of Santa Fe, New Mexico, 8000 ft, 16 Jun 1897, A.A. & E.G. Heller 3775 (SYNTYPE: DAO not seen). Syn.: Ciliaria austromontana (Wiegand) W.A. Weber, Phytologia 51: 371 (1982); Leptasea austromontana (Wiegand) Small, N. Amer. Fl. 22: 153 (1905); Saxifraga bronchialis L. subsp. austromontana (Wiegand) Piper, Contr. U.S. Natl. Herb. 11: 313 (1906); S. bronchialis var. austromontana (Wiegand) M. Peck, Man. Pl. Oregon 365. 1941. REPRESENTATIVE SPECIMENS EXAMINED: CANADA. Alberta: Jasper National Park, Signal Mountain, elev. 7400 ft, 53uN, 118uW, 15 Aug 1963, Fluvian 169 (UBC); near Jasper, 05 Jul 1939, Moss 4810 (UBC); Banff National Park, 11 Jul 1952, Jenkins 1333 (UBC); west of Waterton Lakes National Park, Starvation Peak area, elev. 2073–2438 m, 49u029N, 114u169W, 30 Jul 1958, Taylor (UBC); Banff National Park, Sentinel Pass, elev. 8550 ft, 51u209N, 116u109W, 01 Aug 1962, Vrugtman 620138 (UBC). British Columbia: Trail along Ewart Creek, 18 Jul 1976, Hainault 7907 (WTU); Lillooet area, Noaxe Lake, elev. 6000 ft, 51u049N, 122u459W, 01 Aug 1961, Beamish 610715 (UBC); near Penticton, Green Mtn., 49u039N, 124u209W, 03 Jul 1937, Stonor s.n. (UBC); Garibaldi Park, Black Tusk Meadows, 49u589N, 123u 019W, 01 Aug 1951, Jackson s.n. (UBC); northwestern British Columbia, Atlin Provincial Park, Teresa Island, Birch Mountain, south cirque, 59u269N, 133u499W, 31 Jul 2001, Latimer 59 (UBC); Rocher Deboule Mtns., 01 Jul 1952, Harrison 15 (UBC); Salvus, 2 miles northeast along highway between Prince Rupert and Terrace, 17 Jul 1954, Calder 13112 (UBC). UNITED STATES. Colorado: Ouray County, south of Ouray, junction of Engineer Mountain Jeep Road with Hwy 550, elev. 8200 ft, 24 Jun 1962, Taylor 4713 (UBC); Hinsdale County, Southern Gunnison Basin: San Juan Mountains: Alpine Gulch Trail from ca 1 trail mi S of trailhead to intersection of Alpine Gulch and Grassy Mountain trails, ca 3–4 air mi SW of Lake City, 37.9934uN, 107.3669uW, 29 Aug 1999, Arnett 7859 (RM); Gunnison County, Southern Gunnison Basin: San Juan Mountains: ca 2.3 air mi S of Powderhorn, ca 7.4 air mi ESE of Gateview, ca 10.5 air mi NNW of Cathedral, 38.2369uN, 107.1026uW, 20 Aug 1990, Arnett 7192 (RM); Hinsdale County, Southern Gunnison Basin: San Juan Mountains: along the Grizzly Gulch Trail from ca 0.5–0.6 mi from trailhead, ca 1.1–1.7 air mi E-ENE of Whitecross Mountain, 37.9356uN, 107.4584uW, 03 Aug 1999, Arnett 6232 (RM); Jackson County, Northcentral Colorado: Medicine Bow Mountains: ca 1/4 mi N of Montgomery Pass and to the W, ca 6 air mi ENE of Gould, ca 23 air mi SE of Walden, 40.5465uN,

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

30

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

31

105.9039uW, 05 Jul 2000, Nelson 50216 (RM); Bridal Veil Falls near Telluride, 24 Jun 12, DeChaine 12-1250 (WWB). Idaho: Fremont County, Targhee National Forest: Centennial Mountains: summit of Sawtell Peak, ca 10 mi NNW of Island Park, 44.5688uN, 111.4479uW, 10 Jul 1990, Evert 19558 (RM); Custer County, Sawtooth National Recreational Area (White Clouds), North of the lower Boulder Chain Lakes, elev. 10200 ft, 16 Jul 1980, Taylor 7129 (WWB). Montana: Beaverhead County, Above Torrey Lake, Pioneer Range, 27 Jul 1946, Hitchcock 15058 (WTU); Ravalli County, Bitterroot Mtns, Palisade Peak Ranger Station, elev. 8400 ft, 10 Aug 1946, Hitchcock 15351 (WTU). Washington: Whatcom County, At Twin Lakes, north of Shuksan, 31 Jul 1962, Mattson s.n. (WWB); Whatcom County, Top of Skyline Ridge, elev. 6200 ft, 13 Aug 1971, Nerbovig s.n. (WWB); Whatcom County, Tomyhoi Peak, elev. 6700 ft, 11 Aug 1970, Taylor 2508 (WWB); Olympic Mtns., Seven Lakes Basin, 24 Aug 1935, Jones 8229 (WWB); Stevens County: Calispell Peak, T34N R42E S15; 16, 21, 11 Jul 1979, Althauser s.n. (WWB). New Mexico: Sandoval County, Jemez Mountains: Bandelier National Monument: upper Alamo watershed, ridge of Rabbit Mountain, S of NM Hwy 4 and W of Forest Road 289, 16 Aug 2002, Jacobs 5164 (RM).

2. Saxifraga bronchialis L., Sp. Pl. 1: 400. 1753. TYPE: (LECTOTYPE: LINN not seen, Herb. Linn. No. 575.37, Webb 1987). Syn.: Ciliaria bronchialis (L.) Haw., Saxifrag. Enum. 43 (1821); Leptasea bronchialis (L.) Komarov., Fl. Kamtschatka ii. 217 (1929); Saxifraga stelleriana Merk ex Ser. in A. P. de Candolle & A. L. P. P. de Candolle, Prodr. 4: 46 (1830); S. bronchialis subsp. stellerana Malyschev, Fl. Sibir. 7: 178 (1994); S. algisii T.V. Egorova & Sipliv., Novosti Sist. Vyssh. Rast. vi. 241 (1970); S. kolymensis A. P. Khokhr., Novosti Sist. Vyssh. Rast. 15: 158 (1979); S. stelleriana Merk ex Ser. var. pubifolia Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 93: 96 (1988); S. balandinii Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 104: 40 (1999). Infraspecific taxa: Saxifraga bronchialis subsp. anadyrensis (Losinsk.) Kozhevn., Bot. Zhurn. (Moscow & Leningrad) 67: 1368 (1982); S. anadyrensis Losinsk., Fl. URSS ix. 180, 487 (1939); S. bronchialis subsp. compacta (Adams) Malyschev, Fl. Sibir. (Berberidac.-Grossulariac.) 7: 177 (1994); S. compacta Adams, Nouv. Mem. Soc. Imp. Naturalistes Moscou 3: 239 (1834). REPRESENTATIVE SPECIMENS EXAMINED: RUSSIAN FEDERATION. Amur Oblast: Spurs of Yam-Alin’ mountain range, upper Selemdzha River, elev. 1700 m, 03 Aug 1983, Yakubov s.n. (UBC). Kamchatka Krai: Koryaksky Natsionalny Okrug, Olyutorsky Region, the upper Maly Kubaveyem river, 06 Jul 1976, Barkalov s.n.

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

31

Cust # 13-09

32

Rhodora

[Vol. 116

(UBC); Olyutorsky Region, head of Malosh River, Kyuvubyem, 06 Jul 1976, Varkalev 255 (UBC); Penzhinsky region, near Manily village, on rocks of right bank of Penzhin river, 01 Jul 1975, Kharkevich & Buch s.n. (ALA); Northeast of Achayvayan Olyutorsky Province, 61.56uN, 172.08uE, 15 Jul 1976, Horskov s.n. (UBC). Koryaksky: District of Manily, Penzhinskiy Guba, right shore of Penzhinskaya River, 62.33uN, 165.48uE, 01 Jul 1975, Kharkevich 25 (UBC); Karaginsky Island, Utuvayan River NW of Mt Visokaya, 02 Aug 2005, Eriksen & Topel 1640 (ALA); 01 Aug 1975, Kharkevich s.n. (ALA). Sakha Republic (Yakutia): Tiksi S, valley and small mountain E of town, 71u38.1-59N, 128u51.0-89E, 01 Aug 2004, Solstad & Elven 04/1109 (ALA); Lena r. west bank: plateau mountain 3–5 km w of Chekurovka village, 71u03.59N, 127u28.09E, 16 Jul 2004, Solstad & Elven 04/ 0436 (ALA).

3. Saxifraga caulescens Sipliv., Novosti Sist. Vyssh. Rast. 8: 151 (1971). TYPE: Trans-Baikal, pine forest, surroundings of Czita, 15 May 1908, P.S. Michno (LE not seen). REPRESENTATIVE SPECIMENS EXAMINED: RUSSIAN FEDERATION. Kamchatka: Tolbachik Volcano, track to camp site on south side, 55u439N, 160u119E, 13 Jul 2003, Brummitt & A. Brummitt 20811 (E).

4. Saxifraga cherlerioides D. Don, Trans. Linn. Soc. London 13: 382 (1822). TYPE: Camtschatka, Merk (ISOTYPE: LE not seen). Syn.: Leptasea cherlerioides (D. Don) Small, N. Amer. Fl. 22: 153 (1905); Saxifraga bronchialis subsp. cherlerioides (D. Don) Hulte´n, Ark. Bot., n.s. 7: 68 (1968); S. bronchialis var. cherlerioides (D. Don) Engl., Monogr. Saxifraga 216 (1872); S. pseudoburseriana Fisch. ex Torr. & A. Gray, Fl. N. Amer. 1: 565 (1840). Infraspecific taxa: S. cherlerioides subsp. ascoldica (Sipliv.) Vorosch., Florist. issl. v razn. raonakh SSSR (A. K. Skvortsov) 174 (1985); S. ascoldica Sipliv., Novosti Sist. Vyssh. Rast. 8: 156 (1971). REPRESENTATIVE SPECIMENS EXAMINED: CANADA. Yukon Territory: Ogilvie and Wernecke Mountains, 15 miles NNW of Mount Gibben, elev. 5300 ft, 64u559N, 139u199W, 12 Jul 1984, Cody & Ginns s.n. (UBC). RUSSIAN FEDERATION. Sakhalin Oblast: Taliki River, elev. 78 ft., 54u16.499N, 142u49.1139E, 14 Aug 2001, Stiles 2001-231 (WTU); Northern tip of Sakhalin Island, From unnamed river north of Muni River, to Cape Bakianiy, elev. 0 ft, 54u23.6249N, 142u39.7349E, 07 Aug 2001, Joneson 3148 (WTU); Sakhalin Island, on the Schmidta Peninsula at northern end of island, slopes of Peak 604, 4 kilometers southwest of Tri Brata Mountain, elev. 1968 ft, 54u3940.680N, 143u43926.640E, 12 Aug 2003, Legler 1222 (WTU). UNITED STATES. Alaska: Alinchak Peninsula, east coast of Alaska Peninsula, 57u479N, 155u169W, 02 Aug 1989, Lindstrom s.n. (UBC).

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

32

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

33

5. Saxifraga derbekii Sipliv., Novosti Sist. Vyssh. Rast. 8: 157 (1971). TYPE: North Sea of Ochotensis, on the beach of Eirineici Bay, 9 Jul 1912, Th. A. Derbek 2 (LE not seen). Syn.: Saxifraga spinulosa Royle subsp. derbekii (Sipliv.) Vorosch., in A. K. Skvortsov (ed.), Florist. issl. v razn. raonakh 175 (1985). Infraspecific taxa: S. derbekii Sipliv. subsp. xerophylla A.P. Khokhr., Novosti Sist. Vyssh. Rast. 15: 161 (1979); S. cherlerioides D. Don var. xerophylla (A.P. Khokhr.) Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 93: 97 (1988); S. khokhrjakovii Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 104: 40 (1999). REPRESENTATIVE SPECIMENS EXAMINED: RUSSIAN FEDERATION. Magadan Oblast: 4 km west of Ola Settlement, 12 Jul 1972, N.S. Probatova s.n. (UBC); 28 Aug 1983, P. Zhmylev s.n. (ALA); 28 Jul 1983, P. Zhmylev s.n. (ALA).

6. Saxifraga funstonii (Small) Fedde, Just’s Bot. Jahresber. 33: 613 (1905). TYPE: Disenchantment Bay, vicinity of Yakutat Bay, Alaska, Funston 91 (NY not seen). Syn.: Leptasea funstonii Small, N. Amer. Fl. 22: 154 (1905); Saxifraga bronchialis subsp. funstonii (Small) Hulte´n, Kongl. Svenska Vetensk. Acad. Handl., ser. 3, 8: 12 (1929); S. bronchialis var. funstonii (Small) Rosend., Bot. Jahrb. Syst. 37: 73 (1905); S. bronchialis var. purpureomaculata Hulte´n, Fl. Alaska Yukon 5: 907–908 (1945); S. firma Litv. ex Losinsk., Fl. URSS 9: 184, 488 (1939); S. bronchialis var. purpureomaculata Hulte´n, Acta Univ. Lund. 2, 41, no. 1: 907 (1945); S. codyana Zhmylev, Bjull. Moskovsk. Obac. Isp. Prir., Otd. Biol. 97(1): 95–96 (1992); S. bronchialis subsp. codyana (Zhmylev) Cody, Canad. Field-Naturalist 108: 94 (1994). REPRESENTATIVE SPECIMENS EXAMINED: CANADA. British Columbia: Mount Edziza, west of Eddontenajon on Stewart-Watson Lake Road, elev. 6000– 7000 ft, 557uN, 130uW, 14 Jul 1973, Beamish 730459 (UBC); Stikine Regional District, Atlin Provincial Park, Teresa Island, Birch Mountain, saddle behind north knoll, elev. 4900 ft, 59u259N, 133u459W, 06 Jul 1974, Buttrick 474 (UBC). Northwest Territories: Ridge west of Canoe Lake, elev. 1750 ft, 66u129N, 135u559W, 27 Jul 1966, Lambert (UBC). Yukon Territory: British Mountains, Tributary of Firth River, 69u079N, 140u159W, 07 Jul 1980, Cody 27166 (ALA, UBC); Carcross, Montana Mountain, elev. 5400 ft, 60u039N, 134u429W, 30 Jun 1968, Beamish 681488 (UBC).

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

33

Cust # 13-09

34

Rhodora

[Vol. 116

RUSSIAN FEDERATION. Chukotka Republic: Wrangel I., Gusinaia R., western part of Wrangel I., middle course of the river, Leningradsky Creek (left tributary), 08 Jul 1970, Zhukova & Petrovsky s.n. (ALA); Wrangel I., middle current of Neozhidannaya R., Krupny Cr, 16 Jul 1970, Zhukova & Petrovsky s.n. (ALA). Kamchatka Krai: Ust-Bolsheretski region, Pauzhetka, Kambalnyi ridge, 26 Jul 1969, T.I. Nechaeva s.n. (UBC). Sakhalin Oblast: East/central Sakhalin; Longri River basin, elev. 54–250 ft., 54u16.8189N, 142u48.6649E, 15 Aug 2001, Stiles 2001-297 (WTU); Northeast part of Sakhalin Island. Sea of Okhotsk side. Environs of Taliki River, elev. 383 ft., 54u16.4989N, 142u48.6399E, 14 Aug 2001, Joneson 3389 (WTU); Sakhalin Island, central part; 45 kilometers east-southeast of Tymovskoye and 15 kilometers southeast of Mt. Lopatina; in headwaters of Krebtovyy River 1.5 kilometers northeast of Chamginsky Pass, elev. 2624 ft., 50u45923.820N, 143u179210E, 28 Jul 2003, Legler 964 (WTU). UNITED STATES. Alaska: Philip Smith Mountains, Yukon River-Prudhoe Bay Haul Road, 68u099N, 149u269W, 04 Aug 1982, Murray 8538 (ALA); Solomon Quad, Seward Peninsula Mi 50 Kougarok Rd, 06 Jul 1993, Murray, Yurtsev & Kelso 11307 (ALA); Northeast Alaska, Okpilak Valley, 2 kilometers west of West Okpilak Lake, elev. 1902 ft., 69uN, 144uW, 05 Jul 1985, Grulke 1090 (WTU); North Slope County, Okpilak Valley, 1 km west of West Okpilak Lake, elev. 1902 ft., 69uN, 144uW, 16 Aug 1984, Grulke (WTU); Skagway-Hoonah-Angoon County: Point Hope, Northwest Alaska, 68u219 166u459, 12 Jul 1977, Taylor 6066 (WWB); North Slope Borough, De Long Mountains, 07 Jul 2011, DeChaine 11-37 (WWB); North Slope Borough, Pt. Lay, 07 Jul 2011, DeChaine 11-921 (WWB).

7. Saxifraga kruhsiana Fisch. ex Ser., in A. P. de Candolle & A. L. P. P. de Candolle, Prodr. 4: 46 (1830). TYPE: near Ijiga, Kruhs (LE not seen). Syn.: Saxifraga cherlerioides D. Don var. microglobularia (A.P. Khokhr.) Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 93: 97 (1988); S. microglobularia A.P. Khokhr., Biology Rast. and Valium. D. North East 15: Fig. 2 (1981). REPRESENTATIVE SPECIMENS EXAMINED: RUSSIAN FEDERATION. Kamchatka Krai: 20 Aug 1974, Kharkevich s.n. (ALA). Magadan Oblast: 130 km N of Magadan, upper part of Ola River (left bank) on the Olskoje Plateau, 60u419N, 15u189E, 1554 m, 23 Jul 1988, Elias & Murray 11244 (ALA); 31 Aug 1973, C. Kharkevich s.n. (ALA).

8. Saxifraga nishidae Miyabe & Kudoˆ, Trans. Sapporo Nat. Hist. Soc. vi. 170 (1917). TYPE: Rocky alpine, Hokkaido, Prov. Ishikari, Mt. Yubari, at Gamaiwa, 7 Aug 1916, S. Nishida (SAP not seen). Syn.: Saxifraga arinae Zhmylev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 93: 97 (1988). REPRESENTATIVE SPECIMEN EXAMINED: Gamaiwa, 06 Aug 1989, Brooks 89-38 (E).

JAPAN.

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

Hokkaido:

34

Mt.

Yubari,

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

35

9. Saxifraga omolojensis A.P. Khokhr., Novosti Sist. Vyssh. Rast. 15: 160 (1979). TYPE: Magadan region, 140 km southeast of Omolon (fl. Avlandja), steep, rocky cliffs, 5 Aug 1976, A.P. Khokhryakov (MHA not seen; ISOTYPE: LE not seen). REPRESENTATIVE SPECIMENS EXAMINED: RUSSIAN FEDERATION. Magadan Oblast: Upper Kolyma R. Valley, ‘Aborigen’ field station vicinity, 61u309N, 149u309E, 480–550 m, 22 Jul 1992, Roland 92-120 (ALA); Upper Kolyma R. Valley, ‘Aborigen’ field station vicinity, 61u309N, 149u309E, 480–550 m, 06 Jul 1992, Roland 92-65 (ALA).

10. Saxifraga rebunshirensis (Engl. & Irmsch.) Sipliv., Novosti Sist. Vyssh. Rast. 8: 155 (1971). TYPE: on rocks, Kamchatka, D. Merk (LE not seen). Syn.: Saxifraga bronchialis f. rebunshirensis Engl. & Irmsch. in A. Engler, Pflanzenr. IV. Fam. 117, II: 461 (1919); S. bronchialis var. rebunshirensis (Engl. & Irmsch.) Hara, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot., 6: 65 (1952); S. cherlerioides var. rebunshirensis (Engl. & Irmsch.) Hara, J. Jap. Bot. 13: 174 (1937); S. bronchialis f. laxa (Hara) T. Shimizu, New Alp. Fl. Japan in Color 2: 356 (1983); S. bronchialis f. togakushiensis (Hara) Toyok. & Nosaka, J. Asahikawa Univ. 3: 158 (1975); S. cherlerioides f. laxa Hara, J. Jap. Bot. 13: 175 (1937); S. cherlerioides f. togakushiensis (Hara) Ohwi, Bull. Natl. Sci. Mus. Tokyo 33: 74 (1953); S. cherlerioides var. togakushiensis Hara, J. Jap. Bot. 13: 175, f. 44-G (1937); S. yuparensis Nosaka, J. Fac. Sci. Hokkaido Univ., Bot. 9: 149 (1974); S. bronchialis var. yuparensis (Nosaka) T. Shimizu, New Alp. Fl. Japan in Color 2: 356 (1983). REPRESENTATIVE SPECIMEN EXAMINED: RUSSIAN FEDERATION. Sakhalin Oblast: Moneron Island of the Sea of Japan, Vicinities of Malki settlement, 09 Jul 1974, Stepanova s.n. (UBC).

11. Saxifraga spinulosa Adams, Me´m. Soc. Imp. Naturalistes Moscou 5: 96 (1817). TYPE: In alpine region of Baikal, between villages of Kultuk and Karazay, August s.n. (MW not seen). Syn.: Saxifraga bronchialis subsp. spinulosa (Adams) Hulte´n, Kongl. Svenska Vetensk. Acad. Handl. III, 8: 14 (1929); S. submonantha A.P. Khokhr. & Kuvaev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 99(1): 124 (1994). REPRESENTATIVE SPECIMENS EXAMINED: RUSSIAN FEDERATION. Kamchatka Krai: Koryaksky Natsioinalny okrug, Olyutorsky region, upper Maly Kubaveyem river, 06 Jul 1976, Barkalov & Kozhevnikov s.n. (ALA). Khabarovsk Krai: Ayano-Maisky district west, 5 Aug 1979, Kharkevich s.n. (ALA); Ayano-Maisky

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

35

Cust # 13-09

36

Rhodora

[Vol. 116

district west, 13 Jun 1979, Kharkevich s.n. (ALA). Magadan Oblast: Kontakt Field Station at Kolymskaja Water Balance station near Kulu, 425 km NN of Magadan, logged floodplain on Itrikan River, 10 km SW of station, 24 Jul 1988, Elias 11254 (ALA); Tuva Tes-Khem district, 07 Aug 1979, Daniel s.n. (ALA).

12. Saxifraga taylorii Calder & Savile, Brittonia 11: 248 (1959). TYPE: British Columbia, Queen Charlotte Is., below Mt de la Touche, near head of Fairfax Inlet, Tasu Sound, west coast of Moresby Island, 16–17. viii. 1957. J.A. Calder and D.B.O. Taylor 23511 (HOLOTYPE: DAO not seen; ISOTYPE: K not seen). REPRESENTATIVE SPECIMENS EXAMINED: CANADA. British Columbia: Queen Charlotte Islands, 9 km west northwest of Queen Charlotte City, Mount Raymond, elev. 914 m, 53u169N, 132u149W, 09 Jul 1997, Douglas 13250 (UBC); Queen Charlotte Islands, Moresby Island, 3 km west of Apex Mountain, elev. 914 m, 52u429N, 131u539W, 19 Aug 1997, Lomer 97446 (UBC); Queen Charlotte Islands, Moresby Island, ridge above blue Heron Bay and Fairfax inlet, on Gwaii Hanaas National Park boundary, elev. 600 m, 52u419150N, 131u589170W, 19 Aug 1998, Lomer 98-241 (UBC); Queen Charlotte Islands, 20 miles south of Moresby Camp, Moresby Island, elev. 2000 ft., 52u559N, 132u39W, 29 Jul 1957, Calder 23085 (WTU).

13. Saxifraga tricuspidata Rottb., Skr. Kiøbenhavnske Selsk. Laerd. Elsk. 10: 446 (1770). TYPE: Greenland 1739. Egede. (HOLOTYPE: C not seen). Infraspecific taxa: Saxifraga tricuspidata Rottb. f. albiflora Engl. & Irmsch., in A. Engler, Pflanzenr. IV. Fam. 117, II: 464 (1919); S. tricuspidata f. ligulata Savile & Calder, Canad. Field-Naturalist 66: 105 (1952); S. tricuspidata f. lutescens Engl. & Irmsch., in A. Engler, Pflanzenr. IV. Fam. 117, II: 464 (1919); S. tricuspidata f. micrantha (Sternb.) Calder & Savile, Brittonia 11: 243 (1959); S. tricuspidata var. micrantha Sternb., Revis. Saxifrag. 2: 62 (1831); S. tricuspidata f. subintegrifolia Abrom., Biblioth. Bot. 8, Heft 42: 36 (1899); S. tricuspidata var. subintegerifolia (Abrom.) Polunin, J. Bot. 76: 100 (1938); S. tricuspidata f. woodruffii Calder ex Cody, Canad. Field-Naturalist 64: 92 (1950). REPRESENTATIVE SPECIMENS EXAMINED: CANADA. Alberta: Mt. Coliseum, 2 miles north of Nordegg, elev. 2042 m, 4 Jul 1949, Cawley s.n. (UBC); Signal Mt., Jasper National Park, elev. 1524 m, 52uN, 118uW, 15 Aug 1953, Florian 144 (UBC). British Columbia: Found near Highway 7, vicinity of border, Chilkat Pass, 28 Jul 1974, R.J. Taylor 5183 (WWB); Toad Mt. ridge, mile 422 Alaska Highway, 49uN, 117uW, 21 Jun 1980, Gilmore 2 (UBC).

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

36

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

37

Manitoba: Some miles out of flim flon, 16 Jul 1955, Krivda 1515 (UBC). Northwest Territories: Banks Island, east of 1st camp, elev. 30 m, 30 Jul 1965, Krajina s.n. (UBC); Inuvik airfield, elev. 61 m, 68u169N, 133u329W, 01 Jul 1965, Krajina 65071936 (UBC); Frobisher Bay, 63u459N, 67u159W, 01 Jul 1964, D.E. Swales s.n. (UBC); Jacobsen McGill base camp, ca. 3.5 miles southwest base, Axel Heiberg Island, 79u239N, 90u429W, 20 Jul 1961, Parmelee 1172 (UBC); Ellesmere Island vicinity of Lake Hazen, 81u459N– 82u109N, 68u309W–72u459W, 01 Aug 1959, Powell 678 (UBC). Nunavut: Baffin Island, Auyuittuq National Park, 14 Jun 12, DeChaine 12-176 (WWB). Yukon Territory: West Dawson, 15 Jun 1949, Calder 3066 (WTU). GREENLAND. West Greenland, north Disko, Kugangup ivnarta, elev. 20 m, 70u179N, 53u559W, 24 Jul 1983, Byrge 742 (UBC). UNITED STATES. Alaska: Cape Krusenstern, Northwest Alaska, 67u99 163u449, 07 Jul 1977, Taylor 6008 (WWB); Central Brooks Range, Nign River headwaters, elev. 2500 ft, 16 Jul 1979, Gustafson s.n. (WTU); Misheguk Mountain Quad, DeLong Mtns, Inaccessible Ridge Kagvik Cr. headwaters, 04 Jul 2002, Parker & Beattie 11852 (ALA); Beachey Point Quad, W side Kuparuk R., S side Service City, 07 Aug 1982, Walker 82-3 (ALA); within 3 mile radius of Camp Denali, 18 Jun 1959, Bucknell & Hunter s.n. (ALA).

14. Saxifraga vespertina (Small) Fedde, Just’s Bot. Jahresber. 33: 613 (1906). TYPE: Washington, Grays Harbor Co., Baldy Peak, Conard 302 (HOLOTYPE: NY not seen). Syn.: Ciliaria vespertina (Small) W.A. Weber, Phytologia 51: 371 (1982); Leptasea vespertina Small, N. Amer. Fl. 22: 153 (1905); Saxifraga bronchialis L. var. vespertina Rosend., Bot. Jahrb. Syst. 37: 73 (1905); S. bronchialis L. subsp. vespertina (Small) Piper, Contr. U.S. Natl. Herb. 11: 313 (1906). REPRESENTATIVE SPECIMENS EXAMINED: UNITED STATES. Oregon: Saddle Mountain, 22 Mar 1978, Schofield s.n. (UBC); Clatsop County, Saddle Mountain, 05 Jun 1979, Bohm 1319 (UBC); Columbia Gorge, Horsetail Falls, 20 Jun 2012, DeChaine 12-725 (WWB).

Biogeographic hypotheses. The distribution of taxa and the molecular phylogeny of the section provide the foundation for a general hypothesis on the biogeographic history of sect. Bronchiales. The phylogenetic tree inferred for sect. Bronchiales [referred to as Clade T2 in DeChaine et al. (2013)] suggests that the section originated in the alpine environs of the southern Rocky Mountains of North America, and a subsequent north-westward expansion promoted genetic divergence across Beringia and into Asia. There is strong support for the North American arctic-alpine species, Saxifraga austromontana and S. tricuspidata, being positioned basally within sect. Bronchiales. Indeed, the basal-most lineages of

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

37

Cust # 13-09

38

Rhodora

[Vol. 116

the section, individuals of S. austromontana, reside in the southern Rocky Mountains of New Mexico and Colorado, implying that this region set the stage for the origin and early evolution of the group. As populations of sect. Bronchiales expanded northwards and westwards across Beringia and into Asia, additional lineages arose. Multiple authors have noted the likely importance of isolation in refugia in promoting divergence among the members of this section (Calder and Savile 1959; Losina-Losinskaja 1939; Siplivinsky 1971), and their hypotheses were further supported by the biogeographic analyses of DeChaine et al. (2013). Indeed, geography underlies evolutionary relationships in this group: the North American arcticalpine (S. austromontana and S. tricuspidata), coastal Cascadia of northwestern North America (S. taylorii and S. vespertina), Beringia (S. codyana and S. funstonii), the North Pacific (S. cherlerioides), the southeastern coast and islands of Asia (S. caulescens, S. nishidae, and S. rebunshirensis), the Okhotsk region (S. derbekii and S. omolojensis), and more central Asia (S. bronchialis, S. kruhsiana, and S. spinulosa). Several refugia around the Pacific Rim, including Beringia (Hulte´n 1937), those south of the ice in western North America [the southern Rocky Mountain Refugium, Klamath region, Haida Gwaii (Shafer et al. 2010; Soltis et al. 1997; Swenson and Howard 2005)] and eastern Asia [the mountains of Japan and neighboring islands (Ikeda et al. 2006; Ito 1981; Koidzumi 1919)], and west of Beringia in Central Asia (Hong 1993; Zhu and Rowe 1987) likely promoted divergence following the expansion of the section out of North America. This biogeographic framework sets the stage for further analyses into the role of dispersal and refugial isolation as a driver of genetic divergence and the rise of endemics in sect. Bronchiales. ACKNOWLEDGMENTS. Funding for this study was provided by the National Science Foundation (ARC-0714232 and DEB-1256832). I am grateful for valuable discussions with Charles Davis (HUH) regarding the taxonomic treatment presented in this manuscript. In addition, I would like to thank Kanchi Ghandi (HUH) and two anonymous reviewers for their constructive comments on this manuscript. LITERATURE CITED

BROUILLET, L. AND P. E. ELVANDER. 2009. Saxifraga, pp. 132–142. In: Flora of North America Editorial Committee, eds.. Vol. 8. Flora of North America North of Mexico. Oxford University Press, New York, NY, and Oxford, UK.

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

38

Cust # 13-09

2014]

DeChaine—Saxifraga sect. Bronchiales, sect. nov.

39

CALDER, J. A. AND D. B. O. SAVILE. 1959. Studies in Saxifragaceae. II. Saxifraga sect. Trachyphyllum in North America. Brittonia 11: 228–249. CONTI, E., D. E. SOLTIS, T. M. HARDIG, AND J. SCHNEIDER. 1999. Phylogenetic relationships of the Silver Saxifrages (Saxifraga, Sect. Ligulatae Haworth): Implications for the evolution of substrate specificity, life histories, and biogeography. Molec. Phylogen. Evol. 13: 536–555. DECHAINE, E. G., S. A. ANDERSON, J. M. MCNEW, AND B. M. WENDLING. 2013. On the evolutionary and biogeographic history of Saxifraga sect. Trachyphyllum (Gaud.) Koch. (Saxifragaceae Juss.). PLoS ONE 8: e69814. DOI:10.1371/journal.pone.0069814. DON, D. 1822. Monograph of the genus Saxifraga. Trans. Linn. Soc. London 13: 341–452. ELVEN, R., D. F. MURRAY, V. YU, AND B. A. YURTSEV, eds. 2007. Panarctic flora: Annotated checklist of the panarctic flora (PAF) vascular plants. Published on-line by the Panarctic Flora Project, R. Eleven, editor-in-chief, Univ. Oslo, Norway, Website (nhm2.uio.no/paf/). Accessed Mar 2013. ENGLER, H. G. A. AND E. IRMSCHER. 1919. Saxifragaceae: Saxifraga, pp. 449– 709. In: A. Engler Das Pflanzenreich IV. Fam. 117, II. W. Engelmann, Leipzig, Germany. GAUDIN, J. 1828. Saxifraga taxon Trachyphyllum. Flora Helvetica. Vol. 3: 85, 108– 110. Turici, Sumptibus Orellii, Fuesslini, et Sociorum, publishers, 1828–33. GORNALL, R. J. 1987. An outline of a revised classification of Saxifraga L. Bot. J. Linn. Soc. 95: 273–292. HAWORTH, A. H. 1821. Saxifrage¨arum Enumeratio. R. and A. Taylor, Shoe Lane, London, UK. HONG, Q. 1993. Floristic interrelations of the arctic and alpine tundras in eastern Asia and western North America. Acta Phytotax. Sinica. 31: 1–16. HULTE´N, E. 1937. Outline of the History of Arctic and Boreal Biota during the Quaternary Period. Lehre J. Cramer, New York, NY. IKEDA, H., K. SENNI, N. FUJII, AND H. SETOGUCHI. 2006. Refugia of Potentilla matsumurae (Rosaceae) located at high mountains in the Japanese archipelago. Molec. Ecol. 15: 3731–3740. ITO, K. 1981. Alpine Plants in Hokkaido. Seibundo Shinkosha, Tokyo, Japan. KHOKHRYAKOV, A. 1979. Kamnelomki sektsii Trachyphyllum Gaud. na severovostoke Azii. Novosti Sist. Vyssh. Rast. 15: 157–164. KOCH, W. D. J. 1837. Synopsis Florae Germanicae et Helveticae. F. Williams, Frankfurt, Germany. KOIDZUMI, G. 1919. Genetic and floristic phytogeography of the alpine flora of Japan. Bot. Mag. (Tokyo) 33: 193–222. LOSINA-LOSINSKAJA, A. 1939. Saxifraga, pp. 138–199. In: V. L. Komarov, ed., Flora of the U.S.S.R., Vol. 9. Botanical Institute of the Academy of Sciences of the U.S.S.R., Moscow and Leningrad, Russia. MCGREGOR, M. 2008. Saxifrages: The Definitive Guide to 2000 Species, Hybrids, and Cultivars. Timber Press, Inc., Portland, OR. SHAFER, A. B. A., C. I. CULLINGHAM, S. D. COˆTE´, AND D. W. COLTMAN. 2010. Of glaciers and refugia: A decade of study sheds new light on the phylogeography of northwestern North America. Molec. Ecol. 19: 4589–4621. SIPLIVINSKY, V. N. 1971. Aziatskie kamnelomki sektsii Trachyphyllum Gaud. Novosti Sist. Vyssh. Rast. 8: 147–158.

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

39

Cust # 13-09

40

Rhodora

[Vol. 116

SMALL, J. K. 1905. Saxifragaceae, pp. 81–158. In: N. L. Britton, ed., North American Flora, Vol. 22. New York Botanical Garden, New York, NY. SOLTIS, D. E., M. A. GITZENDANNER, D. D. STRENGE, AND P. S. SOLTIS. 1997. Chloroplast phylogeography of plants from the Pacific Northwest of North America. Pl. Syst. Evol. 206: 353–373. ———, R. K. KUZOFF, E. CONTI, R. GORNALL, AND K. FERGUSON. 1996. MatK and rbcL gene sequence data indicate that Saxifraga (Saxifragaceae) is polyphyletic. Amer. J. Bot. 83: 371–382. SWENSON, N. G. AND D. J. HOWARD. 2005. Clustering of contact zones, hybrid zones, and phylogeographic breaks in North America. Amer. Naturalist 166: 581–591. VARGAS, P. 2000. A phylogenetic study of Saxifraga sect. Saxifraga (Saxifragaceae) based on nrDNA ITS sequences. Pl. Syst. Evol. 223: 59–70. WEBB, D. A. 1987. Typification of the Linnean species of Saxifraga. Bot. J. Linn. Soc. 95: 259–272. ——— AND R. J. GORNALL. 1989. Saxifrages of Europe with notes on African, American, and some Asiatic species. Christopher Helm Publishers Ltd, London, UK. WEBER, W. A. 1982. New names and combination, principally in the Rocky Mountains flora. II. Phytologia 51: 369–376. ZHANG, D.-J., S.-Y. CHEN, Q.-B. GAO, Y.-Z. DUAN, AND S.-L. CHEN. 2008. Circumscription and phylogeny of Saxifraga sect. Ciliatae: Evidence from nrDNA ITS sequences. J. Syst. Evol. 46: 667–675. ZHMYLEV, P. YU. 1988. Notes of rockfoils (Saxifraga L.) of the section Trachyphyllum Gaud. Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 93: 91–99. ZHMYLEV, P. AND A. KHOKHRYAKOV. 1985. K sistematike sektsii Trachyphyllum Gaud. roda Saxifraga L. (Saxifragaceae). Novosti Sist. Vyssh. Rast. 22: 123–125. ZHU, T.-C. AND J. S. ROWE. 1987. A comparison of alpine tundra floras: N. E. China and N. W. North America. Linzer Biol. Beitr. 19: 29–41.

Rhodora rhod-116-965-02.3d 17/4/14 01:34:49

40

Cust # 13-09

Suggest Documents