Sedum oregonense ( Watson 1882: 373 ) Peck (1941: 361)

Zika, Peter F., Wilson, Barbara L., Brainerd, Richard E., Otting, Nick, Darington, Steven, Knaus, Brian J. & Nelson, Julie Kierstead, 2018, A review of Sedum section Gormania (Crassulaceae) in western North America, Phytotaxa 368 (1), pp. 448-450 : 448-450

11. Sedum oregonense ( Watson 1882: 373) Peck (1941: 361) . Figs. 2D , 5E–F , 6E–G , 9B , 13C , 38

Cotyledon oregonensis Watson (1882: 373) . Gormania watsonii Britton in Britton & Rose (1903: 29). Echeveria watsonii (Britton) Nelson & Macbride (1913: 476). Sedum watsonii (Britton) Tidestrom in Dayton (1927: 119, as “watsoni”).

Type: — UNITED STATES. Oregon: [likely Hood River Co., on the E side of Barlow Pass in the White River drainage], northern Oregon, Cascade Mountains near Mount Hood , White River Hill , Barlow Pass , July 1880, J. Howell & T. J. Howell s.n. (lectotype, designated by Clausen [1942: 35], GH; isolectotypes, DS, G ex herb. de Candolle internet image, LE internet image, ORE, US internet image, YU 69005 internet image) .

The protolog said the type was collected in June 1880, but the type specimens were all labeled July 1880. The detailed locality data came from the duplicate at ORE, which was labeled in Thomas J. Howell’s hand as an isotype. Several of the isotypes listed his brother Joseph Howell as a co-collector .

Additional specimens examined: CALIFORNIA. Del Norte County: Sanger Peak road, 1646 m, 1 July 1972, Howe s.n. ( SD) ; E of Sanger Peak , 1585 m, 27 June 1976, Denton 3987 ( OSC, WTU) ; Bear Basin Butte , 1605 m, 5 July 2012, Otting CWG-122 ( JEPS, WTU) ; SE of Chimney Rock , 1400 m, 19 July 1950, Tracy 19202 ( UC) ; WSW of Sawtooth Mountain , 1300 m, 9 June 2013, Zika 26205 ( OSC, WTU) ; near Peak Eight , 1425 m, 24 June 2012, Zika 25928 ( HSC, JEPS, RSA) ; NW of Peak Eight , 1270 m, 10 June 2014, Zika 26621 & Brainerd ( OSC, WTU) . Del Norte / Siskiyou county border: Poker Flat, 26 June 1937, Leach 4713 ( ORE) . Siskiyou County: Salmon Mountains , above Taylor Lake, 2165 m, 21 August 1939, Carter 1467 ( DS) ; Salmon-Trinity Alps Primitive Area , Caribou Lake, 2040 m, 27 July 1955, Wiggins 13521 ( DS, WS) ; Devils Punchbowl , 1525 m, 31 August 1963, Wheeler 8322 ( RSA) ; Red Buttes Wilderness , SW of Buck Peak, 1675 m, 11 August 2011, Brainerd 1916 ( OSC) ; Bolan Lake Road , 1455 m, 1 July 2012, Roché CR-1 ( OSC) ; Salmon Mountains , English Peak, 1768 m, 4 August 1969, Thorne 63383 & Oettinger ( RSA) ; Salmon Mountains , Etna Pass, 1860 m, 11 July 2013, Brainerd & Otting CWG-242 ( CAS, CHSC, DAV) ; same site, 19 June 2014, B. L. Wilson 18106 ( WTU) ; Salmon Mountains , Russian Wilderness, S of Little Duck Lake, 2235 m, 8 July 2013, Brainerd & Otting CWG-247 ( MO, RENO, WTU) ; same site, 2210 m, 8 July 2013, Brainerd & Otting CWG-248 ( BH, MICH, OSC) ; E of Little Duck Lake , 2040 m, 8 July 2013, Brainerd & Otting CWG-250 ( F, GH, UBC) ; Siskiyou Mountains , Sterling Mountain, Soda Creek, 1770 m, 3 July 1934, Wheeler 2847 ( RSA) ; Copper Mountain , 1720 m, 27 June1976, Denton 3990 ( WTU) ; Siskiyou Mountains , WNW of Copper Butte, 1610 m, 19 July 2011, Zika 25647 A ( CAS, F, JEPS, RSA, UCR, WTU) ; Cook and Green Pass, 1735 m, 11 June 1991, Zika 11197 ( OSC) ; S of Cook and Green Pass, 1225 m, 13 July 2013, Brainerd & Otting CWG-251 ( OSC) ; E of Cook and Green Pass, 1400 m, 13 July 2013, Brainerd & Otting CWG-252 ( WTU) ; road to Big Meadow trailhead, 1915 m, 12 July 2013, Brainerd 2190 & Otting ( OSC) ; Red Rock Mountain , 2185 m, 2 September 1974, Mastrogiuseppe & Mastroguiseppe 172 ( OSC) ; Siskiyou Wilderness , above Raspberry Lake, 1745 m, 28 August 2012, Roché & Korfhage CR-2 ( SOC) ; Siskiyou Wilderness , Preston Peak ridge, 2020 m, 28 August 2012, Roché & Korfhage CR-3 ( DAV) . Trinity County: Trinity Alps , Salmon-Trinity Primitive Area , Sapphire Lake, 2286 m, 10 August 1954, Kruckeberg 3737 ( WS, WTU) ; N of Dedrick , 1340 m, 24 June 1976, Denton 3961 ( WTU) ; trail to Canyon Creek Lakes, 1340 m, 24 June 1976, Denton 3967 ( HSC, OSC, WTU) ; same trail, 1606 m, 24 June 1976, Denton 3968 ( WTU) ; Trinity Alps , N of Grizzly Lake, larvae of Callophrys mossii feeding on flowers, 1980 m, 26 July 1993, Emmel 1279 ( RSA) . OREGON. Clackamas County: Mazama Lodge , 21 July 1927, Leach 1281 ( ORE) ; High Rock , 1475 m, 11 July 2012, Zika 25964 ( F, GH, HSC, KANU, MO) . Curry County: en route to Babyfoot Lake Botanical Area , 1315 m, 29 June 1976, Denton 4000 ( WTU) . Deschutes County: 21 miles SW of Bend , 2 September 1976, Crosby 1036 ( OSC) ; Deschutes National Forest , Sisters Road, 7 August 1931, Henderson 13992 ( ORE) ; Soda Creek , 15 July 1947, Ireland 2659 ( ORE) . Deschutes or Lane County: W of North Sister, August 1957, Van Vechter 151 ( OSC) . Douglas County: Cascade Range , Willamette National Forest , Rigdon Point proposed Research Natural Area, 1 August 1991, Brainerd & Newhouse s.n. ( OSC) ; Huckleberry Mountain , 22 August 1978, Christy 2388 ( ORE) ; Bald Ridge , 22 July 1975, Crosby 263 ( OSC) ; Gooseberry Burn , Twin Lakes Trail, 10 July 1941, Detling 4966 ( ORE) ; Umpqua National Forest , S of Black Rock, 29 September 1993, Fosback & Fosback 713 ( OSC) ; Calapooya Range , E of Loletta Lakes, 1630 m, 12 August 2010, Halse 8054 ( RSA); Umpqua National Forest, 1920, Ingram 1410 ( OSC) ; Abbott Butte , 6 July 1899, Leiberg 4261 ( ORE) ; Black Rock Lookout, 23 July 1944, Overlander s.n. ( OSC) ; Rogue-Umpqua Divide Wilderness , above Rocky Ridge Trail, 1835 m, 9 August 2012, Roché & Korfhage RRR ( SOC) . Douglas or Jackson County: Crater Lake Road, 12 August 1896, Leiberg 2937 ( ORE) ; near Abbott Butte , 21 June 1972, Mitchell 67 ( OSC) . Hood River County: Mount Hood region , head of Bluegrass Ridge, 31 July 1927, Thompson 3323 ( DS, WILLU) ; same site, Thompson 3322 ( WTU) ; same site, 6 August 1927, English Jr. 864 ( BH) ; Mount Hood National Forest , near Bottle Prairie, 21 July 1933, Jones 4193 ( POM, WTU [2 sheets]) . Jackson County: Siskiyou Mountains , Pilot Rock, 18 June 1928, Applegate 5520 ( DS) ; Ragsdale Butte , 1370 m, 29 June 1939, Hitchcock 5015 & Martin ( BH, GH, RSA, WS, WTU [2 sheets]) ; Mount Ashland , 19 July 1913, Peck s.n. ( WILLU) ; same site, 28 July 1935, Thompson 12365 ( CAS, WILLU, WTU) ; same site, 5 July 1957, Dennis 578 ( OSC) ; same site, 10 June 1976, Denton 3842 ( WTU) ; same site, 4 July 1976, Denton 4030 ( WTU) ; W of Mount Ashland , 2010 m, 22 August 1976, Denton 4049 ( WTU) ; Observation Gap , 2135 m, 29 June 1977, Denton 4101 ( WTU) ; Big Red Mountain , 2040 m, 15 July 1950, Whittaker SS-391 ( WS) ; Rogue River National Forest , Grayback Mountain, 1745 m, 3 July 1976, Denton 4011 ( OSC, WTU) ; Deadmans Point, 1980 m, 28 July 1977, Denton 4109 ( WTU) ; W of Deadmans [ Point ], 1900 m, 4 July 1976, Denton 4025 ( HSC, OSC, WTU) ; Dutchman Peak , 2165 m, 28 July 1977, Denton 4108 ( WTU) ; E of Dutchman Peak , 1995 m, 22 August 1976, Denton 4048 ( OSC, WTU) ; same site, 28 July 1977, Denton 4104 ( WTU) ; N of Prospect , 21 June 1925, Pendleton s.n. ( OSC) ; Cascade Mountains , July 1927, Sherwood s.n. ( WILLU) ; WNW of Dutchman Peak , 1820 m, 30 July 2011, Zika 25666 & Wilson ( CAS, DAV, HSC, KNFY, NY, OSC, SOC, WTU) . Jefferson County: Brush Creek and Cabot Creek Canyons, 5 August 1959, Swedberg 168 ( OSC) . Jefferson or Linn County: Mount Jefferson , Echo Lake, 19 July 1907, Gorman 2783 ( OSC) . Jefferson or Marion County: divide west of Whitewater Creek, 20 August 1920, Gorman 5097 ( ORE) . Josephine County: Siskiyou National Forest , near Babyfoot Lake Botanical Area, 1345 m, 29 June 1976, Denton 3994 ( OSC, WTU) ; WSW of King Mountain , 950 m, 20 July 2011, Zika 25657 ( NY, UC, WS, WTU) ; Klamath Mountains , King Mountain, 1500 m, 13 July 2013, Brainerd & Otting CWG-243 ( HSC, KANU, WTU) ; E of Tannen Lake , 29 July 1976, Crosby 826 ( OSC) ; near Bolan Lake , 20 July 1945, Peck 23874 ( WILLU) ; Bolan Mountain , 30 July 1976, Crosby 837 ( OSC) ; same site, 26 June 1976, Denton 3977 ( OSC, WTU) ; same site, 26 June 1976, Denton 3979 ( OSC, WTU) ; Bolan Lake , 1640 m, 26 June 1973, Denton 2754 ( WTU) ; Mount Elijah , 24 June 1949, Detling 6494 ( ORE) ; same site, 10 July 1949, Whittaker SS-144 ( WS) ; Lake Mountain Trail , 20 July 1950, Detling 6722 ( ORE) ; Grayback Mountain , 13 July 1930, Henderson 12830 et al. ( ORE) ; near Wilderville , 26 May 1923, Savage s.n. ( ORE) . Klamath County: Crater Lake [National Park], 22 August 1896, Gorman s.n. ( ORE) ; same site, 25 August 1902, Cusick 2974 ( ORE) ; same site, 30 August 1916, Peck 4664 ( OSC) ; same site, 30 August 1916, Heller 12608 ( BH, OSC) ; Crater Lake [National Park], Cathedral Cliff, 19 August 1896, Gorman s.n. ( ORE, WILLU) ; Crater Lake [National Park], Vitae Falls, 1830 m, 21 July 1935, Thompson 12248 ( CAS, POM, WILLU, WTU) ; same site, 2010 m, 30 July 1943, Clausen 4996 & Trapido ( BH [3 sheets]) ; same site, 2073 m, Hutchison 948 ( SD) ; [Crater Lake National Park], Garfield Peak , 11 August 1919, Sweetser s.n. ( ORE) ; same site, 19 July 1935, Applegate 9847 ( WILLU) ; same site, 2285 m, 30 July 1940, Clausen 5006 & Trapido ( BH [2 sheets]) ; same site, 5 July 1976, Denton 4031 ( WTU) ; Lakeview Mountain , 20 August 1926, Anderson 318 ( OSC) ; Union Creek , 21 June 1928, Sipe s.n. ( OSC) ; Cowhorn , 27 July 1948, Menefee 10 ( OSC) . Lane County: Linton Creek , 19 July 1926, Anderson 316 ( OSC) ; South Sister Peak , 26 June 1934, Andrews s.n. ( ORE) ; Bohemia Mountain , 24 June 1924, Constance s.n. ( ORE) ; same site, 17 September 1946, Baker 3290 ( OSC) ; Fairview Mountain , 28 September 1947, Baker 5139 ( OSC) ; O’Leary Mountain , 27 July 1938, Detling 3417 ( ORE) ; same site, 12 July 1965, Clausen 651050 ( BH) ; same site, 16 July 1965, Clausen 651060 ( BH, GH) , Clausen 651068 ( BH) ; E of Deer Butte , 1192 m, 24 July 1965, Clausen 651078 ( BH) , Clausen 651080 ( BH) ; same site, 12 August 1965, Clausen 651172 ( OSC) ; Moolak Mountain , 1 August 1941, Detling 5084 ( ORE) ; Tenas Lake , 23 July 1942, Detling 5394 ( ORE) ; Rebel Peak , 6 July 1951, Detling 7008 ( ORE) ; Bohemia Camp , 1August 1936, Gilkey & Powell s.n. ( OSC) ; Horsepasture Trail , 19 July 1936, Henderson s.n. ( ORE) ; Willamette National Forest , Olallie Trail, 27 July 1938, Ireland 1252 ( ORE) ; NW of Benson Lake , 11 September 1948, Ireland 2858 ( ORE) ; Fairview Mountain , 27 May 1934, King s.n. ( ORE) ; Horsepasture Mountain , 1 July 1914, Peck 4666 ( WILLU) ; N of Mink Lake , 1580 m, 22 July 1965, Clausen 651073 ( BH) , Clausen 651076 ( BH) ; SE of French Mountain , 1390 m, 19 June 2016, Zika 27948 ( GH, OSC) . Linn County: Willamette National Forest , N of junction Highway 126 and Belknap Springs Road, 6 July 1976, Denton 4032 ( OSC, WTU) ; E of junction Highways 126 and 20, 1145 m, 6 July 1976, Denton 4034 ( WTU) ; Carpenter Mountain , 23 July 1941, Detling 5061 ( ORE) ; Clear Lake , 29 June 1943, Gilkey & Smith s.n. ( OSC) ; Cascade Range , Willamette National Forest, Bruno Meadows, 17 July 1998, Halse 5432 ( BH, OSC, RSA) ; road to Clear Lake , 24 July 1953, Hitchcock 20077 ( OSC) ; Deer Butte Trail, 1 July 1942, Hopson s.n. ( OSC) ; Iron Mountain , 28 August 1960, Mason 532 ( OSC) ; same site, 1465 m, 2 August 1975, Denton 3704 ( WTU) ; same site, 6 July 1976, Denton 4035 ( WTU) ; S of Cone Peak , 1440 m, 6 July 2012, Zika 25959 ( RSA, UBC, WTU) ; Mount Jefferson , 14 August 1919, Peck 9163 ( WILLU) ; Nash Crater , 1949, Roach s.n. ( OSC) ; W of junction of Routes 20 and 22, 10 May 1957, Steward 7346 ( OSC) ; Route 126, E of Route 20, 1100 m, 10 July 2012, Zika 25962 ( BH, CAS, DAV, JEPS) . Marion County: Mount Jefferson , Jefferson Park, August 1933, Leach 4436 ( ORE) ; same site, 7 July 1927, Evinger s.n. ( OSC) ; Santiam National Forest , head of Whitewater Creek, 10 September 1916, Flory ( OSC) ; Mount Jefferson , Park Butte, 8 August 1933, Leach 4437 ( ORE) ; S of Breitenbush , 10 August 1935, Peck 18779 ( WILLU) . Wasco County: Mount Hood National Forest , Bulo Point, 1435 m, 7 July 1976, Denton 4041 ( WTU) .

Distribution and ecology: — Sedum oregonense grew in the Cascades and Siskiyous ecoregions of Oregon ( Meyers et al. 2015), and the northern Klamath Ranges ecoregion of California (Jepson eFlora 2017). It was recorded from Mount Hood, Oregon, to northern Trinity County, California, a north-south distance of 500 km and the largest distribution of any member of Sedum section Gormania ( Fig. 37 ). A common and wide-ranging species of ledges, stony ridgelines, and subalpine rocky meadows, it was capable of colonizing recent lava flows and roadcuts. Sedum oregonense was known from elevations of (760–) 1067–2470 m in California. In Oregon the elevational range was 880–2300 m, where it occurred on the crest as well as the west slopes of the Cascade Range, within the Cascades ecoregion ( Meyers et al. 2015). Bedrock substrates were varied, and according to herbarium labels included volcanics, granite, diorite, and schist. Unlike most taxa in section Gormania , Sedum oregonense did not seem limited by dispersal or recruitment difficulties over most of its range. For example, over time it successfully recolonized the Crater Lake area of the southern Oregon Cascades ( Zika 2003) after the massive eruption of Mount Mazama 7700 years ago, estimated to have buried 13,000 square kilometers under 15 cm or more of ash and pumice ( Williams 1942). Once established, S. oregonense often spreads vegetatively, and many populations were vigorous and large. In this way, ecologically, it was more like widespread western species such as S. spathulifolium and S. oreganum .

Notes: — Sedum oregonense ( Fig. 38 ) was a variable taxon, in its cytology and morphology. Its characteristic habit was with numerous branching stolons, bearing loose rosettes with obovate, rounded to retuse leaves. The stem leaves were elliptic to oblong or obovate, and truncate at the base, or suborbicular and slightly clasping; some populations exhibited both forms of stem leaves. Inflorescence branches and the foliage were usually slightly glaucous. The inflorescences varied from flat-topped to capitate or cylindrical. The sepals were relatively short. The ascending acute petals were relatively broad distally, and usually white, but sometimes cream or pale yellow in the southern portion of the range. The corollas usually lacked pink or red color. The fresh anthers were yellow, generally becoming white or brown with age.

Over most of the northern portion of its range Sedum oregonense was the only representative of Sedum section Gormania . The numerous loose rosettes, with easily observed internodes between the leaves, distinguished it from similar species in northern California, S. flavidum and S. marmorense .

Heteroploidy, different chromosome ploidy levels within a single taxon, was known within some Sedum species outside section Gormania ( Uhl 1962) , such as S. lanceolatum , S. stenopetalum Pursh (1814: 324) , and S. wrightii A. Gray (1852: 76) . Nonetheless, Denton (1982) divided most white-flowered Sedum by chromosome number into two taxa, tetraploid S. obtusatum subsp. retusum , sometimes with suborbicular stem leaves, and hexaploid S. oregonense , often with oblong stem leaves. Northwards in the Oregon Cascades, polyploids she considered S. oregonense had either suborbicular or oblong stem leaves ( Denton 1982; this study). Denton studied populations along a 20 km transect between Mount Ashland and Dutchman Peak, in the Siskiyou Mountains of Jackson Co., Oregon. Her vouchers at WTU ( Table 3) documented chromosome counts of 2n = 30, 60, and 90 ( Denton 1979b; Denton & Kerwin 1980). We revisited Denton’s transect over four field seasons, and were unable to find consistent morphological characteristics to distinguish between the three chromosome races. Therefore, we treated all those populations as part of S. oregonense and considered the species a heteroploid. Denton and Kerwin (1980: 905) noted that the hexaploid and tetraploid populations were “morphologically most similar” though they placed them in different species. Even if it was considered appropriate to distinguish tetraploid and hexaploid S. oregonense taxonomically, the name S. obtusatum subsp. retusum was misapplied (see the nomenclatural notes for S. sanhedrinum ).

In her monograph, Denton (1982) cited duplicates from Ragsdale Butte, Jackson Co. (Hitchcock & Martin 5015 WTU) as both Sedum laxum subsp. laxum and as S. oregonense . The specimens had the same habit and habitat, and did not differ in technical details of the foliage, stem leaves, or inflorescence, so we interpreted all replicates of that gathering as S. oregonense .

Figures

FIGURE 2. In full sun, loose open rosettes, with easily visible internodes, were characteristic of sterile shoots for three species. A–B. Sedum citrinum, Del Norte Co., California (Zika 26185). C. Sedum kiersteadiae, diffuse habit with numerous stolons typical of this species and the next, Siskiyou Co., California (Zika 26289). D. Sedum oregonense, Clackamas Co., Oregon (Zika 25964).

FIGURE 5. Stem leaves longer than wide, with a truncated base, were typical of most species of Sedum section Gormania. The stem leaves were commonly broadly obovate to elliptic or oblong, and were flattened or nearly cylindrical.A. Sedum citrinum, Del Norte Co., California (Zika 26185). B. Sedum laxum subsp. laxum, Del Norte Co. (Zika 25927). C. Sedum oblanceolatum × oregonense, Siskiyou Co., California (Zika 25647B). D. Sedum obtusatum, Nevada Co., California (Zika 26271). E–F. Sedum oregonense, Josephine Co., Oregon (Zika 25657). G. Sedum sanhedrinum, Lake Co., California (Zika 26639 & Brainerd).

FIGURE 6. Suborbicular stem leaves were typically found in Sedum laxum subsp. heckneri, S. eastwoodiae, in many populations of S. flavidum, and in scattered populations of S. oregonense. The leaf base was often slightly clasping on suborbicular stem leaves. Stem leaf shape varied across populations, growing seasons, and on individual stems. A–C. Sedum flavidum. A–B. Trinity Co., California (Zika 25922 & J.K. Nelson). C. Humboldt Co., California (Wilson, Otting, & Darington CWG-118). D. Sedum laxum subsp. heckneri, Jackson Co., Oregon (Zika 11199). E–G. Sedum oregonense. E–F. Clackamas Co., Oregon (Zika 25964). G. Del Norte Co., California (Zika 25928).

FIGURE 9. The sepals were short compared to the length of fresh petals in most species. In all species the sepals elongated after anthesis, as the petals wilted. A–C. Relatively short sepals in early anthesis. A. Sedum citrinum, Del Norte Co., California (Zika 26193). B. Sedum oregonense, Jackson Co., Oregon (Zika 25666 & Wilson). C. Sepals elongating post-anthesis, Sedum oblanceolatum, Siskiyou Co., California (Zika 25650A). D–F. Relatively long sepals in early anthesis. D–E. Sedum paradisum subsp. paradisum. D. Shasta Co., California (Lindstrand III & Van Susteren NSR-8). E. Trinity Co., California (Zika 25920 & J.K. Nelson). F. Sedum paradisum subsp. subroseum, Sierra Co., California (Zika 26279).

FIGURE 13. Distal half of the petals broad and ascending. A. Sedum flavidum, Trinity Co., California (J.K. Nelson JKN-12-2). B. Sedum obtusatum, Nevada Co., California (Zika 26271). C. Sedum oregonense, Del Norte Co., California (Otting CWG-122). D. Sedum paradisum subsp. subroseum, Plumas Co., California (Wilson, Janeway & Zika CWG-14).

FIGURE 37. Distribution of Sedum obtusatum, S. oregonense, S. paradisum subsp. paradisum, S. paradisum subsp. subroseum, based on verified herbarium specimens.

FIGURE 38. Sedum oregonense. A. Habit, with numerous branched sterile shoots, Josephine Co., Oregon (Zika 25657). B–D. Stem leaves, varied from much longer than wide to ± as long as wide, truncated or slightly clasping at base. Colors varied from red to pink, orange, purplish, green or gray. B. Zika 25657. C. Linn Co., Oregon (Zika 25962). D. Clackamas Co., Oregon (Zika 25964). E. Loose rosettes, with easily visible internodes between leaves of the sterile shoots, slightly glaucous, Del Norte Co., California (Otting CWG-122). F. Nodding or arching young inflorescences, pre-anthesis, Lane Co., Oregon (Zika 27948). G–I, K. Inflorescences, varied from flat-topped to capitate or cylindrical. J–K. Flowers, petals typically white, rarely pale yellow, petals ascending, anthers yellow, Josephine Co. G. Del Norte Co. (Zika 25928). H–I, K. Josephine Co. H, K. Zika 25657. I–J. Brainerd & Otting CWG-243.