Caltha biflora Candolle (1818: 310)

Wefferling, Keir M. & Hoot, Sara B., 2017, Species circumscription of the Caltha leptosepala polyploid complex (Ranunculaceae) based on molecular and morphological data, Phytotaxa 316 (3), pp. 201-223 : 212-213

Caltha biflora Candolle (1818: 310) . C. leptosepala var. biflora (Candolle) Lawson (1884: 69) . C. leptosepala subsp.

biflora (Candolle) Smit (1969: 26) .

Type:— CANADA. British Columbia: Northwest coast of North America, inland behind Banks Island (between Haida Gwaii and mainland), 1792, Menzies s.n. (holotype BM!). Fig. 1 .

Caltha leptosepala var. howellii Huth (1892: 68) . C. howellii (Huth) Greene (1899: 79) . C. rotundifolia (Huth) Greene var. howellii (Huth) Davis (1900: 15) . C. biflora subsp. howellii (Huth) Abrams (1944: 175) . C. leptosepala subsp. howellii (Huth) Smit (1973: 143) .

Type:— U.S.A. California: near Colby, Butte County, 1896, R. M. Austin s.n. (neotype, designated here, NDG!). Fig. 3 .

Caltha malvacea Greene (1899: 75) .

Type:— U.S.A. Washington: Cascade Mountains, 1838–1842, Wilkes Expedition 484 (Lectotype US!).

Morphology and cytology:—Leaf blades 0.7–1 × as long as wide, reniform to orbicular (rarely emarginate), often diplophyllous (especially in the southern part of the range), margins crenate (in the north) to subentire (in the south) ( Figs. 1 , 3 ); flowers (1–)2(–3) per stem; filaments filiform, 0.1–0.2(–0.3) mm wide (Giblin et al. in press), narrower and often longer than anthers ( Figs. 1 , 11A ); pollen pantoporate to pantocolpate ( Fig. 7 ); follicles with stylar beak 0.1–0.2 mm in length (Giblin et al. in press); hexaploid (2 n = 6 x = 48, Wefferling et al. 2015; holoploid genome size ca. 10–18 pg/2 C, Wefferling et al. 2017).

Taxonomic notes:—Regarding the original type specimen of Caltha leptosepala var. howellii , collected by J. Howell in 1882 in the Cascade Mountains and deposited in “HGB” (HGB was Howell’s abbreviation for “Herbarium generale Berolinense” at Berlin, The Botanic Garden and Botanical Museum Berlin-Dahlem, current acronym B; B. Hellenthal, Museum of Biodiversity and Greene-Nieuwland Herbarium, pers. comm.): “…If the specimen has ever been part of our herbarium [B] then it was probably lost in WW2” (R. Vogt, Botanischer Garten und Botanisches Museum Berlin-Dahlem, pers. comm.). Another specimen collected by Howell (NDG17323) would be an ideal neotype, but lacks clear morphology of the carpels, among other important features. Therefore, NDG17325 ( Fig. 3 ), collected by R.M. Austin near Colby, Butte Co. in 1896 is selected as neotype as (1) it is mentioned by Greene (1899), (2) the leaves bear resemblance to those drawn in Huth’s (1892) treatment (his figure 8), and (3) it is a complete specimen showing stamens, nearly mature and immature carpels, etc. The diplophyllous nature of the leaves ( Huth 1892) is not so clear in this specimen, but one upturned auricle can be seen in the leftmost leaf.

Geographic distribution:—Lowland to subalpine in and west of Coast Ranges and Cascade-Sierra axis; from southeast Alaska south through coastal British Columbia, Washington, Oregon, California, and western Nevada ( Wefferling et al. 2017, Giblin et al. in press).

Additional specimens examined (*pollen sampled for SEM, n = 12; †ploidy level estimated via FCM or chromosome count [ Wefferling et al. 2017], n = 27):— CANADA: British Columbia: N. end of first firebreak on Croman Rd. Woss Area Vancouver Island, 50.175566, -126.363954, elevation 390 m, Stevenson S. 518*, 6/5/1975 ( V). British Columbia: 14.5 miles on road from Shawnigan Lake to Port Renfrew , 48.616667, -123.899996, elevation 580 m, Calder , J. A. and K. T. MacKay 29381, 21-May-1961 (US). British Columbia : Mountain at head of McClintock Bay , Masset Inlet , Graham Island , 53.640189, -132.574431, elevation 310 m, Calder , J. A., D. B. O. Savile , and R. L. Taylor 21584*, 18-Jun-1957 ( V). British Columbia , Vancouver Island , Brooks Quadrant , Harris Peak , Brooks Peninsula , 50.225, -127.725, elevation 825 m, Ogilvie , R. T., W. J. Schofield and R. J. Hebda 848912, 9-Aug-1984 ( V). British Columbia : Three Arm creek , San Juan River Valley , 48.587475, -124.124074, elevation 900 m, Hebda , R. and G. Allen 91-17, 9-Jul-1991 ( V). British Columbia : ca. 1/ 4 mile beyond end of Feona Rd. ( Canadian Forest Products’ setting #M-50), ca. 20 miles S of Woss , northern Vancouver Island , 49.930832, -126.563567, elevation 823 m, Bavis , P. 770608-001, 8-Jun-1977 ( V). British Columbia , North Coast , Princess Royal Island , NW end of island, ridge E of Home Bay , 53.274999, -129.05722, elevation 639 m, Marr , K. L. and C. Copley KM6340 , 25-Jun-2005 ( V). U.S.A.: Alaska: Thorne Bay , Lyman Anchorage Cove, 55.6, -132.58, elevation 91 m, Koval, V. L. 5*, 19-May-1991 ( ALA). Alaska: Deer Mtn., 3.5 miles southeast of Ketchikan , 55.38, -131.57, elevation 730 m, Jaques 1209, 2-Jul-1972 ( OSC). Alaska: Ketchikan, Harriet Hunt Lake road, 55.478649, -131.607508, elevation 230 m, Williams, M. 3094, 5-Jul-1972 ( WTU). California: Kimshew Point quad. High Cascade Range, below the head of Keyser Creek and just below the dirt road, 0.5 km E of Bald Mountain Lookout , 39.9525, -121.4775, elevation 1695 m, Janeway, L. P. 7564*, 26-May- 2002 ( CSU). California: Lassen National Forest ( NF), 3.3 miles on rd. 110, off Hwy. A21 , 40.502731, -121.129142, elevation 1920 m, Meyer, K. M. and A. Townesmith 145 †, 21-Jul-2007 ( DAV). California: McKay Camp Meadows, 40.91, -123.04, elevation 1524 m, Mesler 909, 12-Jul-2011 ( HSU). California: Dunsmuir Quad. Bog around Cedar Lake , ca. 11 mi. SW of Mt. Shasta City, 41.2109, -122.5045, elevation 1737 m, Taylor, M. S. 2778, 4-Jun-1980 ( MO). California: Kaiser Wilderness, Sierra NF, George Lake drainage, 37.295611, -119.169917, elevation 2737 m, Wefferling, K. M. and L. Woo 21*, 27-Jun-2012 ( UWM). California: Kaiser Wilderness, Sierra NF, 20 m from Kaiser Pass Rd. , 37.300556, -119.104806, elevation 2743 m, Wefferling, K. M. 25 †, 28-Jun-2012 ( UWM). California: Shasta-Trinity NF, meadow above and NW of Upper Gumboot Lake, 41.209722, -122.516306, elevation 1904 m, Wefferling, K. M. and L. Woo 30* †, 3-Jul-2012 ( UWM). California: Shasta-Trinity NF, E of Pacific Crest Trail , above Fawn Meadow, 41.239972, -122.508556, elevation 1997 m, Wefferling, K. M. and L. Woo 33 †, 5-Jul-2012 ( UWM). Nevada: Lake Tahoe Basin Mgmt Unit ( USFS), Incline Lakes area , ca. 0.5 mi. SW of Incline Lake, 39.289, -119.934, elevation 2500 m, Christie, K. 1808, 15-Jun-2010 ( ASC). Nevada: Hobart Creek Reservoir, Lake Tahoe State Park , 6 miles E of Carson City , 39.1928, -119.8697, elevation 2347 m, Johnson , J. M. 097*, 1-Jun-1997 ( NYBG). Oregon: Rogue River NF, Oregon hwy 230 to FS road 6510 to 6515, two miles south of summit of Hershberger Mountain , 43.0197, - 122.4519, elevation 1676 m, Baldwin, C. 428, 28-Jun-1994 ( ID). Oregon: Mt. Hood , at Government Camp on Hwy 26, 45.3, -121.76, elevation 1158 m, Chambers, K. L. 1687, 31-May-1961 ( NYBG). Oregon: Cascade Range, Willamette National Forest , Bruno Meadows area , about 5 air miles SE of Idanha, 44.6473, -122.0011, elevation 1244 m, Halse, R. R. 5869 †, 13-Jul-2000 ( OSC). Oregon: Cascade Range, Willamette National Forest , Bruno Meadows, along F. S. Road 2234, about 5 air miles SE of Idanha, 44.6474, -122.007, elevation 1244 m, Halse, R. R. 4746, 9-Jul-1994 (RSA). Oregon: Fanno Bog, 44.851718, -123.596, elevation 850 m, Wilson, B. L. 6986, 11-Jun-1994 ( SRP). Oregon: Klamath NF, Mt Ashland , Pacific Crest Trail , 42.075694, -122.726861, elevation 2000 m, Wefferling, K. M. and L. Woo 38 †, 6- Jul-2012 ( UWM). Oregon: Umpqua NF, Abbott Butte, just outside Rogue-Umpqua Divide Wilderness, 42.941167, - 122.547417, elevation 1790 m, Wefferling, K. M. and L. Woo 39* †, 7-Jul-2012 ( UWM). Oregon: Umpqua NF, side of Silent Creek near Diamond Lake inlet, 43.126361, -122.161111, elevation 1586 m, Wefferling, K. M. 43 †, 8-Jul-2012 ( UWM). Oregon: Mt. Hood NF, near trailhead for Elk Meadows and Sahalie Falls , 45.322361, -121.634111, elevation 1359 m, Wefferling, K. M. and L. Woo 44 †, 16-Jul-2012 ( UWM). Oregon: Mt. Hood Wilderness , Mt. Hood NF, on trail to Paradise Park along PCT, 45.357445, -121.752278, elevation 1686 m, Wefferling, K. M. 61 †, 8-Aug-2012 ( UWM). Oregon: Umpqua NF, Umpqua side of Rogue-Umpqua Divide, S end Donegan Prairie, 42.920087, -122.590767, elevation 1605 m, Wefferling, K. M. and L. Woo 42 †, 7-Jul-2012 ( UWM). Oregon: Mt. Hood NF, along creek below Mt. Hood Meadows Drive , 45.326963, -121.660281, elevation 1585 m, Wefferling, K. M. 47s †, 16-Jul-2012 ( UWM). Oregon: Mt. Hood Wilderness , NW end of Elk Meadows, 45.34525, -121.620528, elevation 1570 m, Wefferling, K. M. 68* †, 1-Jul-2013 ( UWM). Oregon: Mt. Hood Wilderness , NW end of Elk Meadows, 45.34525, -121.620528, elevation 1570 m, Wefferling, K. M. 69* †, 1-Jul-2013 ( UWM). Washington: Mink Lake , Olympic NP, 47.949332, -123.868271, elevation 940 m, Harthill, M. P. 1069*, 11-Aug-1972 ( RSA). Washington: Mink Lake , Olympic NP, 47.9479, -123.8677, elevation 950 m, Harthill, M. P. 988, 11-Aug-1972 (RSA). Washington: Gifford Pinchot NF, Indian Heaven Wilderness, outside of Carson , 46.00317, -121.78873, elevation 1424 m, Meyer A. s. n., 24-Sep-2011 ( UWM). Washington: Alpine Lakes Wilderness Mt. Baker-Snoqualmie NF, near outlet of Rainy Lake , 47.514215, -121.537258, elevation 1116 m, Wefferling, L. 1, 10-Jun-2012 ( UWM). Washington: Mt. Baker-Snoqualmie NF, Heather Lake , 48.071806, -121.783917, elevation 740 m, Wefferling, K. M. 48 †, 21-Jul-2012 ( UWM). Washington: Alpine Lakes Wilderness, Mt. Baker-Snoqualmie NF, between Mason and Kulla Kulla Lakes, 47.426806, -121.54625, elevation 1341 m, Wefferling, K. M. 50* †, 22-Jul-2012 ( UWM). Washington: Alpine Lakes Wilderness, Mt. Baker-Snoqualmie NF, outlet of Nimue Lake , 47.539556, -121.397167, elevation 1138 m, Wefferling, K. M. and L. Wefferling 52, 23-Jul-2012 ( UWM). Washington: Alpine Lakes Wilderness, Mt. Baker-Snoqualmie NF, below Denny Mountain , 47.436972, -121.456333, elevation 1114 m, Wefferling, K. M. and B. Wefferling 54 †, 30-Jul-2012 ( UWM). Washington: Gifford-Pinchot NF, below road W of Takhlakh Lake , 46.274691, -121.606949, elevation 1265 m, Wefferling, K. M. 55 †, 1-Aug-2012 ( UWM). Washington: Gifford-Pinchot NF, Takhlakh Meadow, 46.270444, -121.58875, elevation 1382 m, Wefferling, K. M. 57 †, 1-Aug-2012 ( UWM). Washington: Gifford-Pinchot NF, Takhlakh Meadow, 46.269707, -121.588677, elevation 1400 m, Wefferling, K. M. 58 †, 1-Aug-2012 ( UWM). Washington: Gifford-Pinchot NF, Takhlakh Meadow, 46.269707, - 121.588677, elevation 1400 m, Wefferling, K. M. 59s †, 1-Aug-2012 ( UWM). Washington: Sauk Mountain , 48.523121, -121.598597, elevation 1550 m, Shrum, J. SM5 †, 22-Jul-2015 ( UWM). Washington: Olympic National Park , Boulder Creek , lower site, 47.976209, -123.697533, elevation 677 m, Hunter, G. BL4 †, 15-Jul-2015 ( UWM). Washington: Olympic National Park , Little River , 48.04704, -123.504876, elevation 330 m, Hunter, G. LR1 †, 15-Jul-2015 ( UWM). Washington: Gifford-Pinchot NF, Takhlakh Meadow, 46.268447, -121.586248, elevation 1400 m, Wefferling, K. M. CR1 * †, 1-Jan-2014 ( UWM). Washington: Gifford-Pinchot NF, Babyshoe Pass, 46.268139, -121.604472, elevation 1320 m, Wefferling, K. M. and L. Wefferling 72s, 15-Jul-2014 ( UWM). Washington: Gifford-Pinchot NF, Takhlakh Meadow, 46.268447, -121.586248, elevation 1400 m, Wefferling, K. M. and L. Wefferling 94s †, 15-Jul-2014 ( UWM). Washington: Gifford-Pinchot NF, Babyshoe Pass, 46.268139, -121.604472, elevation 1320 m, Wefferling, K. M. and L. Wefferling 74s, 15-Jul-2014 ( UWM). Washington: Olympic Peninsula, Rd. #303 between Rugged Ridge and Pine Mt., 47.948646, -123.961192, elevation 884 m, Buckingham, N. 02023, 29-Jun-1979 ( WTU). Washington: Cascade Mountains of Western Washington , Mt. Gardner , collected at the intersection of roads 155, 155.5, and 155.6, 47.3667, -121.5514, elevation 1100 m, Stout, T. SAR4 †, 23-Jun-2001 ( WTU) .

Figures

FIGURE 1. Holotype of Caltha biflora, collected by A. Menzies, 1792. “Northwest coast of America, inland behind Banks Island.” (between Haida Gwaii and mainland British Columbia, Canada). Housed at The Natural History Museum, London, England (BM565604). Inset photo shows a single narrow, filiform filament; magnified 5×.

FIGURE 3. Neotype of Caltha leptosepala var. howellii, collected by R.M.Austin, July 1896. “Colby, Butte County, Northern California” (U.S.A.). California.Housed at the Greene-Nieuwland Herbarium, Notre Dame, U.S.A. (NDG17325). Inset photo shows narrow filaments; magnified 5×.

FIGURE 7. Pollen of Caltha biflora, determined by morphology and molecular data (Wefferling & Hoot, unpublished data). ††ploidy level determined by chromosome counts (Wefferling et al. 2015); †ploidy level estimated by flow cytometry (Wefferling et al. 2017). A. U.S.A.: Alaska, Koval, V.L. 5 (ALA). B. U.S.A.: Nevada, Johnson, J.M. 097 (NYBG). C. U.S.A.: California, Janeway, L.P. 7564 (CSU). D. U.S.A.: California, Wefferling, K.M. and L. Woo 21 (UWM). E. U.S.A.: California, Wefferling, K.M. and L. Woo 30† (UWM). F. U.S.A.: Washington, Wefferling, K.M. 50† (UWM). G. Canada: British Columbia, Calder, J.A., D.B.O. Savile, and R.L. Taylor 21584 (V). H. U.S.A.: Oregon, Wefferling, K.M. 69† (UWM). I. U.S.A.: Washington, Wefferling, K.M. CR1†† (UWM). Size bars = 6 μm.

FIGURE 11. Close-up photographs of Caltha flowers.A.C.biflora. U.S.A.:Washington,Alpine Lakes Wilderness.Photo by L.Wefferling. B. C. chionophila. U.S.A.: Wyoming, Medicine Bow National Forest. C. C. leptosepala. U.S.A.: Alaska, Chugach National Forest.