SEDUM JURGENSENII (HEMSL.) MORAN (CRASSULACEAE)

REID MORAN

Herbarium,

Sonoma State University,

Rohnert Park, CA 94928

Abstract. Sedum jurgensenii grows in east central Mexico at 1100 to 2900 m and is common in cultivation. It has usually been placed in Villadia sect. ALTAMIRANOA as V. elongata (Rose) R. T. Clausen or later as V. jurgensenii (Hemsl.) H. Jacobsen. It also includes A. necaxana Fröd. and ap­parently S. brandtianum Poelln. In ALTAMIRANOA it is unique in its hispidulous herbage and (fide C. H. Uhl) its chromosome number of n = 23 and is remarkable for its elongate stems and its clasping arrow-head leaves. S. jurgensenii subsp. attenuatum subsp. nov. differs in its strongly ascending and imbricated, narrower and more sharply acute leaves. The type (Moran & Kimnach 7776, SD) is from a limestone canyon at San Hipolito, Puebla.

Sedum jurgensenii is a tall, weak and re­laxed, fuzzy plant with white flowers, common in the mountains of Hidalgo, Mexico, and neighboring states and often grown in Mexico and abroad. It has been treated in Villadia and, having the flowers in cymes, in section ALTAMIRANOA (e.g., Calderón, 1974, 1979; Clausen, 1959), where it was long known as V. elongata (Rose) R. T. Clausen and more recently as V. jurgensenii (Hemsl.) H. Jacobsen. However, Altamiranoa now appears to be a polyphyletic group, whose species seem best dispersed in Sedum (Moran, 1996). H. 't Hart (1995) defined Sedum as a paraphyletic group of some 350 species, in two subgenera. In his classification Sedum jurgensenii would fall close to S. bourgaei Hemsl. and S. goldmanii (Rose) Moran (= Villadia batesii (Hemsl.) Baehni & J. F. Macbr.), both of which he showed (Fig. 1) in the Acre-clade of subgenus Sedum. No SEM photo of the seeds is available.

Since other plants of ALTAMIRANOA are all glabrous, this one is distinctive in its hispidulous herbage: stems and upper leaves may sometimes look nearly glabrous, but leaves of young shoots in particular are unmistakably bristly. It is remarkable also for its elongate and often sprawling stems (Fig. 3) and its somewhat clasping fat arrowhead-like leaves. Also, it has a unique chromosome number: in 21 collections from more or less throughout the range, aside from one probable trisomic, Dr. Charles H. Uhl has found n = 23, as in no other species of Altamiranoa or Villadia.

Fig. 1. Flowering branch of Sedum jurgensenii ssp. jurgensenii, San Diego, 29 July 1962, X l.8. From Pachuca, Hidalgo, Mexico (M&K 7782); topotype of Altamiranoa elongata Rose.

Sedum jurgensenii (Hemsl.) Moran, Haseltonia 4: 46.1996.

Weak hispidulous perennial, branching near base and above, with roots not tuberous-thickened. Peren­nial stem base woody, rarely to 8 mm thick, dark brown or finally grayish, the lower third of plant at flowering time bearing bristly new shoots 0.5-3 cm long, with ± 25-50 imbricated leaves in 3 and 5 spi­rals. Flowering stems erect to trailing and rooting, l-5( —7) dm long, 0.5-1 mm thick above. ± 2 mm thick below, green to pink or dark red, glabrous to sparsely or closely puberulent with non-glandular trichomes mostly 0.1-0.2 mm long, closely leafy, the leaves in 3 and 5 spirals, near base of stem often as­cending but mostly spreading, on hanging stems reflexed even to 45°, the internodes 0.5—1 mm long, the leaf sites ca. 2 mm wide and 1.5 mm high, the attach­ment oval, ± 0.3—0.5 mm wide. Leaves triangular-ovate to -lanceolate (or linear when dried), acute, cordate-clasping or somewhat sagittate, 4—8( —15) mm long, 2-3 mm wide, 1.3—2 mm thick, rounded dorsally. convex ventrally, with acute to obtuse mar­gins, variously hispidulous, the lower densely so, with trichomes mostly 0.1—0.3 mm long, the upper often more sparsely so and with shorter trichomes. Cyme 2-5 or ultimately even to 8 cm wide, of 2-5 or more mostly close-set cincinni, each l-3(—8) cm long, with 2—12( —20) flowers, scarcely circinate, the buds and flowers all ± erect, sessile or subsessile. Sepals erect and ± appressed to corolla, equal in length, ovate-lanceolate, acute, free to and around base but scarcely spurred, 2.5-5 mm long. 1—1.5 mm wide, 0.5-1 mm thick, rounded dorsally. ± flattened ventrally, with acute margins, hirsutulous mostly on margins and keel with trichomes to 0.1 mm long, so sometimes ± ciliate. Corolla white, in bud 4-6 mm long, with segments imbricate, in anthesis campanulate to open, 5-8 mm wide, the tube 1—1.5 mm long, ± 3 mm wide, the segments elliptic, broadly acute, 3-4.5 mm long, 1.6-2 mm wide, obtusely keeled, the keel often with a few short trichomes above, ending in a short blunt apiculus. Stamens erect, the filaments white, subterete, ± 0.3 mm thick, 2.5—4 mm long above corolla base, adnate 1—1.8 mm. the epipetalous slightly shorter and adnate slightly higher; anthers red. ovoid to oblong, 0.6—0.8 mm long. Nectar glands deep yellow, truncate, 0.35-0.5 mm high, 0.6-1 mm wide above, slightly narrowed below, 0.1—0.15 mm thick above where thickest. Gynoecium greenish white, 3.5-4.5 mm high, 2 mm thick, the pistils erect, lanceolate, connate ± 0.5 mm, the styles 1.5-2 mm long, soon separating; ovules 15-25, 0.4-0.45 X 0.2 mm. Follicles brown, erect, with beaks out-turned. Chromosomes: n = 23. Flowers Sept.-Dec.

This widespread plant varies somewhat throughout its range. Leaves are mostly ca. 5-6(-7) mm long in wild plants but may be up to 10 mm long when those same plants grow in the greenhouse. The only collection seen from Guanajuato (Lau 079) has the longest leaves, in the cultivated specimen up to 15 mm; and one from Querétaro (Boutin 3437), apparently field collected, has leaves to 12 mm. It is not clear whether long leaves are characteristic for this part of the range. Several collections from Oaxaca, Puebla, and Veracruz have the leaves more strongly ascending than usual and more sharply acute: these are separated below as a new sub­species, subsp. attenuatum.

Fig. 2. Inflorescence of Sedum jurgensenii ssp. jurgensenii. 15 July 1961, X 1. Same plant as Fig. 1.

Sedum jurgensenii subsp. jurgensenii

(Figs. 1-4)

Cotyledon jurgensenii Hemsl., Diagn. Pl. Nov. 1: 9. 1878.

Altamiranoa (?) jurgensenii (Hemsl.) Rose in Britton & Rose, N. Amer. Fl. 22: 50. 1905.

A. elongata var. jurgensenii (Hemsl.) Fröd. [illegitimate name], Acta Horti Gotob. 10(App.): l43. 1936.

Villadia jurgensenii (Hemsl.) H. Jacobsen, Nat. Cact. Succ. J. 13: 76. 1958.

TYPE: MEXICO, south Mexico without locality, 1843-44, C. Jurgensen 616 in part (holotype K!).

Altamiranoa elongata Rose in Britton & Rose, Bull. New York Bot. Gard. 3(9):31. 1903. Cotyledon elongata (Rose) Fedde, Bot. Jahresber. Just 31(1): 825. 1904.

Villadia elongata (Rose) R. T. Clausen, Bull. Torrey Bot. Club 197. 1940.

TYPE: MEXICO, Hidalgo, mountains above Pachuca, 2850 m, 1 June 1899, J. N. Rose & W. Hough 4461 (holotype US 346433!; isotype NY!).

?Sedum brandtianum Poelln., Repert. Sp. Nov. 33: 241. 1933.

TYPE: None cited, the plant cultivated, of unknown origin.

Altamiranoa necaxana Fröd., Acta Horti Gotob. 10(App.): 144, figs. 1158-1165, pl. 97. 1936.

V. necaxana (Fröd.) H. Jacobsen, Nat. Cact. Succ. J. 13: 76. 1958.

TYPE: MEXICO, Puebla, shaded cliffs at 1100 m, barranca below Salto Chico, Necaxa, 12 February 1932, H. Fröderström & K. Hulté,n 850 (holotype at S not seen; isotypes GH!, MICH!, NY!).

ILLUSTRATIONS: Cited above.

Fig. 3. Downhill plant of S. jurgensenii ssp. jurgensenii, San Diego, 12 Dec. 1970, X 0.25. From Cerro Santuario, Hidalgo

Mature leaves mostly spreading ± perpendic­ular to stem, or on pendent stems reflexed and so ascending in reverse (Fig. 4), triangular-ovate to-lanceolate, acute. Sepals ovate-lanceo­late, acute. Petals acute.

Fig. 4. Cyme of plant shown in Fig. 2, X l.8. Note reflexed (and hence "inversely ascending") leaves of this downhill branch.

Distribution: Cliffs and rocky slopes. Hidalgo [and Tamaulipas?] to Guanajuato, Puebla, and Veracruz, at 1100-2900 m. Selected collections. GUANAJUATO. Redamaderas, A. Lau 079, HBG41310 (Uhl 2582*) (BH). HIDALGO. Clay on moist limestone mountainside 35 mi N of Zimapan, F. A. Barkley 17M304 (TEX); moist broadleaved evergreen forest, 1470 m, 7.2 km SW of Tlanchinol, B. Bartholomew. L. Landrum, H. W. Li & T. S. Ying 3425 (CAS, C,II, NY); between Jacala and Zimapán, 1980 m, D. E. Breedlove & F. Almeda 59408 (CAS); sandstone cliffs, 2320 m, Barranca el Salto 4 km N of Encarnación, R. T. Clausen 78-16 (CU); dry rocky slope between Real del Monte and Pachuca, 2745 m, R. T. Clausen & J. L. Edwards 7433 (CU, GH, UC); rocky roadcut 2 mi N of Durango, D. B. Dunn 17388 (CAS, NY); 7 km W of Tenango de Doria, J. Gimate L. 752 (CAS, MICH); Mpio. Zimapán, rocky walls and floor of Barranca de San Vicente, 2000 m, H. E. Moore Jr. 1300 (BH, GH); slopes and summit of red sandstone peak, Cerro de las Canteras, S of Actopán, 2700 m, H. E. Moore Jr. 1501 (BH, GH, US); dry rocky pine-oak woods, Puerto de la Zorra, N of Jacala, 1500 m, H. E. Moore Jr. 1677 (BH, GH); slopes and ravines by Río Pantolán, between Zacualtipán and Olotla, 1600-2000 m, H. E. Moore Jr. 5335(BH); rocks in wet woods S of Zacualtipán, 1950 m, R. Moran 10064* (ARIZ, BH, CAS, DH, CU. GH, RSA, SU, TEX, UC, US); Encarnación, R. Moran 14784 (H. Fittkau) (BH); Pachuca, 2400 m, R. Moran & M. Kimnach 7782* (CAS, CU, SD) (Figs. 1. 2); on rocks in forest and on cliffs above, Cerro Santuario, 2600 m, R. Moran & C. H. Uhl 13398* (CAS, SD, US) (Figs. 3, 4); Pachuca, C. R. Orcutt (US); Sierra de Pachuca, 9500 ft, C. G. Pringle 8233 (CU, GH, NY, UC, US); 9777(CU, GH, NY, US); Sierra de Pachuca, J. N. Rose & ?R. Hay 5608 (NY); mts. above Pachuca, J. N. Rose & J. H. Painter 6756 (GH, NY, UC, US); matorral xerofilo 4 km NE of Pachuca, 2600 m, J. Rzedowski 29200 (ARIZ, CAS, MICH, US); Pachuca, C. H. Uhl 1471* (BH, CU); Cantil de Tambor, 22 mi N of Jacala, 5400 ft, C. H. Uhl 1866* (BH); Barranca de Marmoles, 16 mi N of Zimapán, C. H. Uhl 2345* (BH). PUEBLA. Planted at roadside shrine 1.5 mi N of Acatepec, 2100 in. R. Moran & M. Kimnach 7728a (SD); shaded rocks in canyon, 8 mi F. of Huauchinango, C. H. Uhl 1555* (BH); volcanic cliffs 2 mi S of Zacatlán, 7000 ft, C. H. Uhl 2350*; rocks in arroyo at 2075 m, 4.2 km E of Aquixtla, C. H. Uhl 2356* (BH). QUERÉTARO. Limestone around sinkhole, Mex. Hwy. 120 near SLP boundary, 5800 ft, F. C. Boutin 3437 (Uhl 2317*) (BH, HNT, SD); abundante sobre peñas, 1630 m, El Calvario, 1 km W of El Aguacate, H. Rubio 269 (MICH); broken lime­stone, summit 16 mi ENE of Landa de Matamoros, 5600 ft, C. H. Uhl I860* (BH); limestone 0.5 mi W of San Joaquin & 23 mi NE of Vizarrón, 7750 ft, C. H. Uhl 2130* (BH). SAN LUIS POTOSÍ. Near Xilitla, 5800 ft. F. Boutin 3437 (C. H. Uhl 2317*) (BH); occasional on rocks at 1900 m, Pico el Agujón, Sierra Equiteria, R. Moran & C. H. Uhl 13380 (SD?); igneus rock and soil, mesophytic forest, 1.5 mi SW of Final de Amoles, J. M. Smith, B. L. Turner, & M. A. Whalen 824 (TEX). TAMAULIPAS. Acacia woods by stream, 54 mi S of Ciudad Victoria, J. S. Wilson 12316 (TEX) [elevation ap­parently very low for this plant, seemingly less than 300 m; locality therefore questionable]. VERACRUZ. Limestone cliffs 5 km WSW of Maltrata, 1800 m, R. T. Clausen TMV-0-M1 (BH); canyon of Gemapa River, 5 km N of Coscomatepec, R. T. Clausen TMV-0-G1 (CU).

The subsp. jurgensenii is occasionally grown in Mexico, sometimes beyond its natural range; and it has been grown in the United States and abroad since its introduction by Dr. J. N. Rose nearly a century ago. In specimens from cultiva­tion the leaves are often 10 mm long, though not wider than usual.

Hemsley (1880) reported that this species and Cotyledon mexicana [= Sedum goldmanii] were both distributed as Jürgensen 616, which sug­gests to me that the two may have been collect­ed together. Quite possibly then the type came from the same region as the type of A. elongata. According to Standley (1922: 181), little is known of Jürgensen, who collected for Henri Galeotti in Mexico and especially in Oaxaca, after Galeotti had returned to Europe in 1840.

With the type of Altamiranoa elongata, Rose also cited C. G. Pringle 8233, from the same area. Rose transferred Cotyledon jurgensenii to Altamiranoa with a query, saying it was known only from the type specimen; but the holotype leaves little doubt that they are the same.

According to Fröderström (1936), Dr. K. von Poellnitz had sent him living plants of Sedum brandtianum, grown by him and by Haage in Erfurt, and distinguished by Prof. Huber as a new species. Dr. R. L. Praeger had already writ­ten Fröderström of his belief that it was identi­cal with A. elongata. From the original descrip­tion, this seems quite probable. Von Poellnitz (1933) mentioned sending living plants to the botanical gardens at Dahlem, Kiel, and Kew; and it is possible that some herbarium speci­mens may be found.

Fröderström (1936) wrote that Altamiranoa necaxana Fröd. was "closely allied to A. elon­gata and perhaps only a glabrous form of this from a less extreme region." In fact, he labeled three isotypes (GH, MICH, NY) as "var." necax­ana of A. elongata. Although he described the plant as glabrous, these isotypes are somewhat puberulent, and especially the leaves of new shoots are characteristically bristly. The plant seems not to differ otherwise. In a near topotype (Uhl 1555) Dr. Uhl found n = 23, as usual.

Fig. 5. Flowering branch of S. jurgensenii ssp. attenuatum, San Diego, 2 Sept. 1961, X 1.8. Part of type collection, from San Hipolito, Puebla (M&K 7776).

Sedum jurgensenii subsp. attenuatum R. Moran, subsp. nova

(Figs. 5, 6).

TYPE: MEXICO, Puebla, limestone canyon on NE edge of San Hipolito (near 18°55'N, 97°53'W), 2200 m, 26 November 1959, Reid Moran and Myron Kimnach 7776* (holotype SD 56940; isotype BH) (Figs. 5, 6).

A subsp. jurgensenii foliis adscendentibus imbricatisque, angustioribus, subteretibus subacuminatis, sepalis subacuminatis, petalisque anguste acutis differt.

All leaves strongly ascending and imbricated, nar­rowly triangular-ovate or the upper triangular-lanceo­late, subterete, sharply acute or subacuminate. Sepals slender-acute or subacuminate. Petals narrow-acute.

Fig. 6. Flowers of S. jurgensenii ssp. attenuatum, San Diego, 2 Sept. 1961, X 2. Same plant as Fig. 5.

Distribution: Puebla to Oaxaca and Veracruz at 1800-2500 m. Collections: OAXACA. Abundance, encinar, 2200 m, Cerro Gato, E of Guadalupe Membrillos. Dist. Huajuapan, P. Tenorio L. 18206 (CAS). PUEBLA. Sierra Negra E of Coxcatlán, F. Otero 127; lava flow by Mex. Hwy. 144 near Esperanza and 6 mi S of Ciudad Serdán, 8400 ft, C. H. Uhl 1897* (BH); rocks and loose gravelly soil on steep slope of crater with lake, Atenco, 2500 m, C. H. Uhl 2364* (BH). VERACRUZ. SE exposure on limestone cliffs 5 [or 0.5] km WSW of Maltrata, 1800 m, R. T. Clausen TMV-0-M1 (BH).

This plant is generally similar to subsp. jurgensenii, and on the basis of three counts, ac­cording to Dr. Uhl, it shares the unique chromo­some number of n = 23. However, the leaves of mature stems are strongly ascending and imbri­cated like those of the new shoots and are nar­rower and more acuminate (Fig. 5). The subsp. jurgensenii also grows in Puebla and Veracruz.

Acknowledgements

The curators at ARIZ, BH/CU, CAS/DH, GH, HNT, K, MICH, NCC. NY, POM/RSA, SD, TEX, UC, and US, have lent specimens and have helped in various other ways. Charles H. Uhl has kindly permitted the use of his valuable cytological data; and he, Henk 't Hart, Charles Quibell, and Myron Kimnach have made use­ful suggestions for this paper. I sincerely thank them all for their help.

References

Calderón de Rzedowski, G. 1974. Las Crasuláceas del Valle de México. Cact. Suc. Mex. 19: 49, 51-63, figs. 30-35.

---------. 1979. Crassulaceae. In J. Rzedowski & G. Calderón de Rzedowski, Flora Fanerogamica del Valle de México 1:238-254, fig. 42.

Clausen, R. T. 1959. Sedum of the Trans-Mexican Volcanic Belt. Cornell Univ. Press. Ithaca, New York.

Fröderström, H. 1936. The genus Sedum L., Part 4 [American species]. Acta Horti Gothob. 10(app.): 1-181, figs. 1-1360, pl. 1-115.

Hart, Henk 't. 1995. Infrafamilial and generic classifi­cation of the Crassulaceae. Chapter 10. Pp 159-172 in H. 't Hart and U. Eggli, eds., Evolution and systematics of the Crassulaceae. 1-192, figs. Backhuys Publishers, Leiden.

Hemsley, W. B. 1880. Botany: Crassulaceae. In F. D. Godman and O. Salvin, eds., Biologia Centrali-Americana... 1: 387-399, T19-21.

Moran, R. 1996. Altamiranoa into Sedum (Crassu­laceae). Haseltonia 4: 46.

Poellnitz, K. von. 1933. Sedum brandtianum spec. nov. Repert. Spec. Nov. 30: 241-242.

Standley, P. C. 1922. Trees and shrubs of Mexico (Fagaceae-Fabaceae). Contr. U.S. Natl. Herb.23 (2): i-xxxvii, 171—515.

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*) Chromosome count by C. H. Uhl from this collection.

© Haseltonia, 1997