Sedum sanhedrinum Berger (1930: 451)

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

15. Sedum sanhedrinum Berger (1930: 451) . Figs. 1F , 4E , 5G , 14E–F , 44D , 45

Gormania retusa Rose in Britton & Rose (1903: 31). Cotyledon retusa (Rose) Fedde in Schumann & Fedde (1904: 828). Sedum laxum (Britton in Britton & Rose 1903: 29) Berger (1930: 451) subsp. retusum (Rose) Clausen (1942: 39) . Sedum obtusatum Gray (1868: 342) subsp. retusum (Rose) Clausen (1975: 375) .

Type: — UNITED STATES. California: Lake Co., Sanhedrin Mountains , 5000 feet [1525 m], 6 August 1902, A. A . Heller s.n. (lectotype, designated by Clausen [1942: 39], US; isolectotype, NY) .

Not Cotyledon retusa ( Lindley 1847: 306) Baker in Saunders (1869: sub. t. 64, no. 22).

Not Sedum retusum Hemsley (1880: 51) .

We were not certain where Amos Arthur Heller (1867–1944) collected the lectotype of Gormania retusa ( Sedum sanhedrinum , Fig. 44 ), which was labeled “Lake Co., Sanhedrin Mountains, alt. 5000 feet [1524 m], 6 August 1902, A. A. Heller s.n. ” ( Fig. 44A ). His bound collecting notebook for 1902, archived at WTU, did not have an entry for the type of Gormania retusa . His collecting trip in the vicinity of Sanhedrin Mountain was between 14 July and 11 August 1902 (Heller 5855 to Heller 6048). His next collections were dated 18–20 August in Sonoma County (Heller 6049–6056). The Gormania retusa lectotype at US ( Figs. 44A, D ), and the isolectotype at NY ( Figs. 44B–C ), were dated 6 August, both in a handwriting that does not match the hand in Heller’s notebook ( Fig. 44E ). There were three entries in Heller’s book for 6 August (Heller 6036–6038), all from “Thistle Glade,” on the west side of Snow Mountain ( Fig. 44F ). The exact location was uncertain, but likely along the lower drainage of Thistle Glade Creek, which drains the western flank of Snow Mountain, entirely within Lake Co., from 1850–5600 feet [564–1707 m] elevation. Because Heller’s notebook (Heller 6038, Fig. 44E ) clarified these collections [ Apocynum Linnaeus (1753: 213) , Helenium Linnaeus (1753: 886) , and Solidago Linnaeus (1753: 878) ] were from the “west base of Snow Mountain,” this would be the wrong habitat and too low an elevation for Sedum sanhedrinum , which is known from 4600 feet [1402 m] and higher.A more appropriate habitat for Sedum sanhedrinum would be the “northerly slope of Mount Sanhedrin opposite Mt. Hull” where he collected Aquilegia Linnaeus (1753: 533 ; Heller 6041, Fig. 44G ) on 7 August, and labeled it from Lake Co. His collecting book ( Fig. 44E ) lists Heller 6041 on 9 August, not 7 August, adding some uncertainty. Most of the northern slope of Mount Sanhedrin, N of the summit at Big Signal Peak, and at elevations above 1400 m, is in present day Mendocino Co., but was part of Lake Co. in Heller’s day. Heller also gathered specimens in the Sanhedrin summit region earlier in his trip, on 20, 22, 24, and 28 July, but any of those dates created a larger discrepancy with the label data on the type of G. retusa .

In summary, we believe there were minor inconsistencies on the lectotype label for Gormania retusa at US. The handwriting on the US and NY types did not belong to Heller, and so it is understandable if some of the details were not a perfect match for Heller’s notebook. It appears Heller was at low elevations on 6 August ( Figs. 44E–F ), but at suitable high elevations on either 7 or 9 August ( Fig. 44E ), perhaps the correct collection date for the type. All Heller’s printed labels credited his collections to Lake Co. ( Consortium of California Herbaria 2017), though some were in modern day Mendocino Co. The type locality may be the north slope of Mount Sanhedrin ( Fig. 44G ), at ca. 5000 feet [1524 m], and this was probably in modern Mendocino Co., not Lake Co. Sedum sanhedrinum was extant but uncommon on Mount Sanhedrin, along the ridge east of Big Signal Peak (Steven Darington, obs.; Hutchison 913). Finally, Heller’s numbered gatherings in his collecting book indicated sets of 15–30, which Heller was offering for sale (at 5 cents each). The lack of a collection number on the type suggests that the stonecrop was uncommon in 1902, and so Heller did not gather it as a large set of replicates for sale.

Additional specimens examined: — UNITED STATES. CALIFORNIA. Glenn County: Bear Rock , 1510 m, 26 June 2013, Wilson & Otting CWG-230 ( OSC, WTU) . Lake County: Snow Mountain , 1 July 1929, Baker 3541 ( UC) ; Mendocino National Forest, Forest Service Road M1 , 1585 m, 14 July 1977, Smith 9330 et al. ( RSA, WTU) ; W slope of Goat Mountain , 1490 m, Dean 5042 ( UCR) ; Hull Mountain , 1955 m, 26 June 2013, Wilson & Otting CWG-231 ( OSC, WTU) ; same site, 12 June 2014, Zika 26639 & Brainerd ( OSC) . Mendocino County: summit Mount Sanhedrin , 1885 m, 21 July 1962, Hutchison 913 ( BH, JEPS) ; ENE of Bald Mountain , larvae of Parnassius smintheus near [subsp.] sternitzkyi feeding on plants, 1860 m, 22 July 2001, Emmel 1731 ( UCR) ; Monkey Rock , 1920 m, 26 June 2013, Wilson & Otting CWG-232 ( HSC, JEPS) ; same site, 11 June 2014, Zika 26637 & Brainerd ( CAS, CHSC, SBBG) ; NE of Eel River Work Station , 1485 m, 27 June 2013, Wilson & Otting CWG-233 ( DAV) ; same site, 11 June 2014, Zika 26629 & Brainerd ( OSC, WTU) ; W of Little Baldy , 1415 m, 11 June 2014, Zika 26633 & Brainerd ( OSC, WTU) ; Hull Mountain , 1890 m, 12 June 2014, Zika 26638 & Brainerd ( WTU) . Tehama County: South Yolla Bolly ( Mount Linn ), 2285 m, 16 July 1955, Barbe 175 ( UC [2 sheets]) ; South Yolla Bolly Peak , 14 July 1963, Clausen 63206 ( BH) ; South Yolla Bolly Mountains, Valentine Ridge , 18 June 1963, Clausen 63212 ( BH) ; same site, 18 June 1963, Clausen 63212 ( BH) ; same site, 30 June 1963, Clausen 63209 ( BH) ; same site, 3 August 1963, Clausen 63208 ( BH) ; same site, 14 August 1963, Clausen 63400 ( BH) ; Tomhead Lookout , 2040 m, 2 August 2011, J. K. Nelson JKN-2 ( JEPS, RSA) ; same site, 13 July 2012, Zika 25970 ( WTU) ; same site, 27 June 2013, Zika 26244 ( WTU) ; Government Flat , 1740 m, 27 June 2013, Wilson & Otting CWG-234 ( CAS, NY) .

Distribution and ecology: — Sedum sanhedrinum was restricted to Glenn, Lake, Mendocino, and Tehama counties, California ( Fig. 25 ). The populations ranged from the Yolla Bolly Mountains in the north to Goat Mountain in the south, all in the High North Coast Ranges ecoregion (Jepson eFlora 2017), at elevations of 1415–2286 m. Substrates included serpentine and schist. Habitats were rocky slopes, ridgelines, talus, boulder fields, and rocky creek banks, usually in dry to very dry sites, in full sun or, less commonly, in light shade.

Morphological notes: — Sedum sanhedrinum ( Fig. 45 ) was often a stout plant with dense rosettes of acute to truncate leaves. The petals were erect or spreading about 10° from the floral axis, and pale to bright yellow, often with an apricot tinge near the base. It was apparently closely related to S. paradisum subsp. paradisum . Both had overlapping stem leaves, elongate towards and into the inflorescence. Both had relatively long sepals, dense rosettes, and similar variation in flower color. However, they were allopatric, with S. paradisum subsp. paradisum ( Fig. 37 ) found northwest of S. sanhedrinum ( Fig. 25 ). Living plants also differed in the angle of petals, which were ascending in S. paradisum and essentially erect in S. sanhedrinum . Early in the growing season, before the flowers opened, S. sanhedrinum flowering stems nodded, while they were erect in S. paradisum subsp. paradisum .

Nomenclatural notes: —Understanding Sedum sanhedrinum was difficult for several reasons. The name was published overseas, and was unknown to most California botanists. The type specimen was discolored and its stem leaves fell off ( Fig. 44D ), complicating its interpretation. Location information for the type specimen was vague. The name S. obtusatum subsp. retusum , founded on S. sanhedrinum , was employed broadly ( Clausen 1975, Denton 1982), based on a description ( Denton 1982) that did not fit plants collected at the type locality. Thus there were many specimens and literature reports of S. obtusatum subsp. retusum that we referred to more northern entities, especially S. oregonense and S. kiersteadiae .

The name Sedum obtusatum subsp. retusum was applied to plants with white or pale yellow petals, growing in the North Coast Range and Klamath ecoregions, north to the Siskiyou Mountains just north of the Oregon border ( Denton 1979b, 1982, 1993; Denton & Kerwin 1980; Boyd & Denton 2012). Those plants were reported to have loose rosettes and to form extensive stoloniferous mats ( Denton 1982). That appeared to be a misinterpretation. In our view, most specimens reported as S. obtusatum subsp. retusum ( Denton 1982) were S. oregonense , which had loose rosettes, a spreading stoloniferous habit, and white flowers. Denton (1979b, 1982; Denton & Kerwin 1980) restricted her S. oregonense species concept to hexaploid plants. Based on chromosome counts, Denton interpreted tetraploid colonies as S. obtusatum subsp. retusum , but we considered them to be S. oregonense ( Table 3). We revisited some of Denton’s northern tetraploid populations and found no morphological markers to separate them from diploid or hexaploid S. oregonense .

Figures

FIGURE 1. Dense rosettes on vegetative shoots, without visible internodes between the leaves, were found in most species. A. Sedum flavidum, Trinity Co., California (Zika 26646 & Brainerd). B. Sedum laxum subsp. laxum, Del Norte Co., California (Zika 25927). C. Sedum oblanceolatum, Jackson Co., Oregon (Zika 25636 & Lang). D–E. Sedum rubiginosum, Tehama Co., California. D. Wild plant (Zika 26238). E. Cultivated plant (Wilson & Coberly CWG-10). F. Sedum sanhedrinum, Mendocino Co., California (Zika 26629 & Brainerd).

FIGURE 4. Water stored in the stem leaves appeared to be used in the reproductive effort. Stem leaves withered as flowering progressed. Proximal stem leaves shriveled before those near the inflorescence. A. Sedum laxum subsp. heckneri, Siskiyou Co., California; proximal stem leaves withered, distal still plump (Zika 25640). B. Sedum patens, Del Norte Co., California; proximal stem leaves shriveling during anthesis (Zika 26608 & Brainerd). C–D. Sedum rubiginosum, Tehama Co., California. C. Turgid stem leaves at the initiation of flowering (Wilson & Coberly CWG-10). D. Shriveled stem leaves at late anthesis (Zika 26234). E. Sedum sanhedrinum, Tehama Co., proximal stem leaves withering first (Zika 25970).

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 14. Cultivated individuals often had petals more widely spreading than in the wild. A–B. Sedum flavidum corollas, Humboldt Co., California (Brainerd & Otting CWG-103). A. Ascending petals, wild plants. B. Spreading petals, cultivated plants. C–D. Sedum laxum subsp. laxum corollas, Del Norte Co., California. C. Erect petals, wild plants (Brainerd & Otting CWG-107). D. Ascending petals, cultivated plants (Brainerd & Otting CWG-104). E–F. Sedum sanhedrinum corollas, Tehama Co., California. E. Erect petals, wild plants (Zika 26244). F. Ascending petals, cultivated plants, Tehama Co. (J.K. Nelson JKN-2).

FIGURE 25. Distribution of Sedum flavidum, S. kiersteadiae, S. rubiginosum, and S. sanhedrinum, based on verified herbarium specimens.

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

FIGURE 44. Gormania retusa type locality. A. Handwritten label of lectotype, Sanhedrin Mts., Lake Co., alt. 5000 feet, 6 August 1902, A.A. Heller s.n. (US). The label handwriting does not match Heller’s handwriting in his collection notes. B. Handwritten isolectotype label at NY, in the handwriting of Nathaniel Britton (Amy Weiss pers. comm.). C. Label from inside the fragment packet, isolectotype (NY). D. Lectotype specimen (US), shadows showing where two inflorescences were once present. E. Heller’s 1902 collecting book at WTU, with no listing for Gormania (or Sedum). 4 August, Hullville on Eel River (Heller 6024–6035). 6 August, West base of Snow Mountain (Heller 6036–6038). 9 August, Northerly slope of Sanhedrin (Heller 6039–6042). F. Printed label of Heller 6038 (NY 2339077 ex herb. BKL), 6 August 1902, from low elevation at “Thistle Glade, west side of Snow Mountain”. G. Printed label of Heller 6041 (NY 2743052 ex herb. BKL), 7 August 1902, from “northerly slope of Mount Sanhedrin opposite Mt. Hull.” There was no entry for 7 August 1902 in Heller’s collection book, where he listed 6041 on 9 August.A, D. Images reproduced with permission, courtesy of United States National Herbarium, Smithsonian Institution. B–C, F–G. Images reproduced with permission, they belong to the C. V. Starr Virtual Herbarium of The New York Botanical Garden (http://sweetgum.nybg.org/science/vh/).

FIGURE 45. Sedum sanhedrinum. A. Habit, showing large rosette leaves and ascending stem leaves, Mendocino Co., California (Zika 26629 & Brainerd). B–D.Variation in stem leaves, spreading to ascending, broadly to narrowly obovate, bases truncated, slightly glaucous, green to purple or pink. B. Zika 26629 & Brainerd. C–D. Mendocino Co. (Zika 26633 & Brainerd). E. Dense and small rosette leaves, Tehama Co., California (Zika 26244). F. Capitate inflorescence, with dense flowers, Lake Co., California (Wilson & Otting CWG-231); inflorescences were also cylindrical (e.g., Fig. 45D, type) or flat-topped. G–K. Variation in flowers, corolla color pale yellow, orangeyellow, or bright yellow, faded to pink, orange-brown, pale brown, or yellow-brown, white or dull red; anthers yellow or, rarely, dull orange-red, faded to pink, reddish, white, or dull yellow; sepals elongated after anthesis. G. Tehama Co. (Wilson & Otting CWG-234). H. Zika 26629 & Brainerd. I. Zika 26633 & Brainerd. J–K. Petals erect, tips acute to blunt, Tehama Co. (Zika 25970).