Sedum paradisum ( Denton 1978: 236 ) Denton ex B. L. Wilson subsp. subroseum B. L. Wilson & Zika, 2018

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

12b. Sedum paradisum ( Denton 1978: 236) Denton ex B. L. Wilson subsp. subroseum B. L. Wilson & Zika , subsp. nov. Figs. 3C–D , 9F , 13D , 15D , 40A–F , 41 .

Species nostra Sedum paradisum subsp. paradisum aemulans, differt axe inflorescentiae juvenili nutante, a Sedum obtusatum subsp. obtusatum floribus albis postea saepe pallide erubescentibus recedens.

Type: — UNITED STATES. California: Plumas County, dry serpentine ridgetop 1 air mile NE of Pilot Peak , 2005 m, 6 August 2011, P. F . Zika 25721 & L. P . Janeway , B. L . Wilson (holotype, WTU; isotypes, CAS, CHSC, OSC, UC) .

Rhizomes and stolons to 18 cm long, 1.7–5.1 mm diameter; sterile leafy shoots often numerous. Rosettes 13–55 mm diameter. Rosette leaves usually dense, in shady situations sometimes loosely arranged with visible internodes, slightly glaucous, at least when young, green, grey-green, orange to red, or purple, strongly flattened dorsiventrally, obovate to oblanceolate, cuneate, 10–31 × 6–16 mm, 2–3.5 mm thick, apices obtuse or notched, less commonly truncate or acute. Stem leaves ascending, slightly glaucous, at least when young, and colored like the rosette leaves, 9–20 × 3–11 mm, 1.5–3.5 mm thick, flattened, truncate at base, narrowly oblong, oblong-oblanceolate, or obovate to oblanceolate, apices acute or obtuse. Fertile stems pink to reddish or orange (green), 6–21 cm tall, nodding or bent in bud, usually erect in flower and fruit. Inflorescences 2.5–13 × (1–) 2–7 cm, narrowly cylindrical panicle-like cymes or flat-topped and corymbiform, subglobose if depauperate, with 4–10 branches, proximal branches ascending or spreading, 15–60 mm long, solitary at inflorescence nodes. Inflorescence bracts resembling stem leaves, but smaller, 2.5–18 × 1–8.5 mm, bases truncate, tips acute or blunt. Flowers (3–)10–49 per inflorescence, fresh flower diameter (5–) 7–12 mm, flowers 5-merous, erect, calyx green, pink or brown, 3.6–6.4 × 3.7–6.1 mm, sepals fused basally 0.8–2.2 mm, free sepal tips 2.3–4.9 mm long, apex narrowly acute, acute or obtuse. Fresh petals fused at base 1.4–3.3 mm; petals 6.5–10.1 mm long, 2.7–3.9 mm wide at mid-length, creamy-white, white, or greenish-white, base often dull light pink, midvein pink to red (especially in bud), apices or bases often pink or red, especially with age. Fresh petal blade v-shaped or trough-shaped in cross section, broad, distal half ovate or widely deltoid, at half its length slightly spreading circa 30° from floral axis, apex sometimes apiculate with subterminal mucro 0–0.2 mm. Stamens 10, when fresh shorter to longer than petals, filaments greenish-white or white, aging red, fresh anthers narrowly elliptic or oblong to elliptic-oblong, 1.2– 1.8 × 0.6–0.85 mm, yellow, aging white or less often gray, brown, orange, or blackish. Nectaries shallowly crescent-shaped, sunken in middle, white to dark yellow, 0.8–1.6 × 0.3–0.6 mm. Ovaries 4.0– 6.3 mm, erect, fused 0.5–1.4 mm, maturing into 5 dark brown erect follicles, 4.2–6.7 mm, with erect to slightly curved style remnant forming a narrow beak 0.6–1.3 mm, follicles fused 1.1–2.0 mm at base, containing 19–24 seeds. Seeds medium brown, oblanceolate, shiny, striate, 1.1–1.35 × 0.4–0.5 mm, including stipe 0.05–0.25 mm.

Paratypes:— UNITED STATES. CALIFORNIA. Butte County: below Pilot Peak, on Quincy-Laporte Road , 1930 m, 6 August 2011, Wilson 16817 & Janeway, Zika ( OSC, WTU) . Plumas County: near La Porte , 915 m, 29 June 1968, Ahart s.n. ( CAS) ; N side of Pilot Peak , 20 September 2011, Ahart 17693 ( WTU) ; Plumas-Eureka State Park , 1592 m, 28 May 1961, Uhl 962 ( BH) ; Eureka Peak , 1695 m, 31 July 1964, Clausen 641125 ( BH) ; W of Eureka State Park , 1660 m, 13 June 1976, Denton 3878 ( OSC, WTU) ; SSW of Graeagle , 1830 m, 14 June 1976, Denton 3884 ( WTU) ; Bucks Lake , summer 1975, Griggs 310 ( CAS) ; Lakes Basin area , 1980 m, 11 August 1950, Weatherby 1510 ( RSA) ; Little Jamison Creek , 1600 m, 27 July 1964, Clausen 641107 ( BH) , Clausen 641108 ( BH) , Clausen 641110 ( BH) , Clausen 641111 ( BH) , Clausen 641114 ( BH) ; Jamison Creek , 1700 m, 6 August 2011, Wilson, Janeway & Zika CWG-14 ( DAV, JEPS, RENO, RSA, WTU) ; Quincy-Laporte Road , 1930 m, 6 August 2011, Wilson, Zika & Janeway CWG-15 ( HSC, US) ; WNW of Mills Peak , 1885 m, 28 June 2013, Zika 26283 ( DAV, HSC, JEPS, UCR) . Sierra County: NE of Gold Lake , 1990 m, 29 July 1964, Clausen 641120 ( BH, GH) ; Sierra Buttes , 29 July 1964, Clausen 641127 ( BH, GH) , 641115 ( BH, GH), 641116 ( BH); same site, 14 June 1976, Denton 3899 ( OSC, WTU) ; E of La Porte , 1700 m, 28 July 1982, Ahart 3700 ( CAS) ; Packer Lake , 1940 m, 28 June 2013, Zika 26279 ( CAS, CHSC, RSA, SD) .

Distribution and ecology: —Restricted to the Northern High Sierra Nevada ecoregion (Jepson eFlora 2017), California, north of the Middle Yuba River, at elevations of 915–2005 m ( Fig. 37 ). Habitats included rocky slopes, ridgelines, and dry cliffs, in full sun to partial shade, on a variety of bedrock types including peridotite, andesite, and granite. Common associates included: Abies magnifica A. Murray bis, Aconogonon phytolaccifolium (Meisn. ex Small) Rydb. , Calochortus leichtlinii Hook. f. , Cryptogramma acrostichoides R. Br. , Eriogonum umbellatum var. nevadense Gand. , E. ursinum S. Watson var. ursinum , Myriopteris gracillima , Pellaea bridgesii Hook. , Penstemon deustus , P. newberryi A. Gray , Pinus contorta Douglas ex Loudon , P. jeffreyi Balf. , P. lambertiana Douglas , Poa secunda J. Presl , Silene bernardina S. Watson , and Spiraea splendens É.N. Baumann ex K. Koch.

Etymology: — Sedum paradisum subsp. subroseum , or Plumas stonecrop, was named for its flowers, which usually turned pinkish and with age gave the entire inflorescence a rosy appearance.

Notes: — Sedum paradisum subsp. subroseum ( Figs. 40A–F , 41 ) differed from S. paradisum subsp. paradisum in its bent young inflorescence axis, slightly shorter sepals, more strongly red-tinted foliage, and its more eastern distribution in the Sierra Nevada range.

Sedum paradisum subsp. subroseum was separable from S. obtusatum in the Sierra Nevada by flower color; petals of S. paradisum subsp. subroseum were white while those of S. obtusatum were pale yellow to deep yellow. The two were allopatric. The southernmost known station of S. paradisum subsp. subroseum was at Sierra Butte, north of the Middle Yuba River, and the northernmost known station of S. obtusatum was near Jackson Meadows Reservoir, 11 km distant and south of the Middle Yuba River.

Figures

FIGURE 3. Species typically with dense rosettes in full sun occasionally formed loose rosettes in sheltered microsites, often associated with shade, talus, or protected crevices. A–B. Sedum patens vegetative shoots, Del Norte Co., California. A. Typical dense form (Zika 26987). B. Loose form (Zika 26609 & Brainerd). C–D. Sedum paradisum subsp. subroseum, Sierra Co., California (Zika 26279). C. Typical dense form. D. Loose form.

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 15. The color of the petals, anthers, filaments, and carpels often changed after anthesis.A. Sedum albomarginatum, after anthesis petals turned orange-brown; Butte Co., California (Zika 25867). B. Sedum flavidum, buds deep pink especially along the keel, faded to white at anthesis, then aged to a uniform deep pink; Humboldt Co., California (Wilson, Otting, & Darington CWG-118). C. Sedum laxum subsp. laxum, after anthesis green carpels turned dark red, red anthers turned white; Del Norte Co., California (Zika 25936). D. Sedum paradisum subsp. subroseum, after anthesis green carpels turned dark red, yellow anthers turned white; Plumas Co., California (Wilson, Zika & Janeway CWG-15). E. Sedum rubiginosum, after anthesis red-based yellow petals turned pallid; Tehama Co., California (Zika 26234).

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

FIGURE 40. A–F. Sedum paradisum subsp. subroseum. A. Habit, showing stolons and dense rosettes, and consistently elongated proximal and distal stem leaves (Zika 26283). B. Flower, side view, showing ascending petals and elongated sepals. C. Flower, top view, showing broad upper petal and acute tips. B–C, E–F. Zika 26279. D. Petal tip enlargement, showing subterminal mucro projecting on the right side, and the hooded petal tip on the left side (Zika 26283). E. Rosette leaf. F. Stem leaf, with truncated base; in the field the stem leaves were often a dull gray, but varied to green or pink. G–M. Sedum patens. G. Habit, showing loose rosettes, stem leaves longer than wide, and widely branched inflorescence. H. Flower, side view, showing spreading petals. I. Flower, top view, showing narrow gradually tapered upper half of petals, and acuminate tips. G–I. Zika 26609 & Brainerd. J. Enlargement of petal tip, showing subterminal mucro projecting on right, hooded petal tip on left (Zika 26608 & Brainerd). K. Dehisced follicles with attenuated apex. L. Rosette leaf. M. Stem leaf, with truncated base. K–M. Zika 25935.

FIGURE 41. Sedum paradisum subsp. subroseum. A–D. Habitat. A. Plumas Co., California (Zika 25721 Janeway & Wilson). B. Sierra Co., California (Zika 26279). C–D. Plumas Co. (Zika 26283). E. Young inflorescences, nodding prior to anthesis on right, straightening as coming into flower on left; stem leaves elongated and overlapping proximally and distally; Plumas Co. (photo: Tim Hanson). F. Erect flowering shoots in full sun, Plumas Co. (Zika 25721 Janeway & Wilson). G–H. Lax growth form in partial shade on mossy ledges, sheltered vegetative shoots with loose rosettes; Sierra Co. (Zika 26279). I–J. Typical dense rosettes in full sun, showing variation in color, Plumas Co. (Zika 25721 Janeway & Wilson). K. Rosette leaves, bottom row; stem leaves, upper row; Sierra Co. (Zika 26279). L. Inflorescence, Plumas Co. (Wilson, Janeway & Zika CWG-14). M–P. Flowers, showing fresh anthers yellow to almost white, after dehiscence white or brownish, ascending petals white or pinkish, aged pink to dark pink, distal half of petal broad, acute. M. Green metallic bee visiting flowers, Plumas Co. (Zika 26283). N–O. Sierra Co. (Zika 26279). P. Two flowers and a bud, fresh anthers almost white, subapical mucro on petal tips, and as flower matured, sepals elongated, lower half of petals with deeper color; Plumas Co. (Wilson, Zika & Janeway CWG-15).