Nolina palmeri S. Watson. var. brandegeei Trelease Proceedings

Hernández-Sandoval, Luis & Rebman, Jon P., 2018, The Genus Nolina (Asparagaceae) of the Baja California Peninsula, and the Recognition of a New Species Combination, Systematic Botany (Basel, Switzerland) 43 (3), pp. 717-733 : 721-727

Nolina palmeri S. Watson. var. brandegeei Trelease Proceedings of the American Philosophical Society 50:209. 1911.

This taxon was previously recognized as an infraspecies

( Trelease 1911) and as a synonym of N. palmeri , a species considered by Hochstätter (2010) and Thiede (2012) within the section Microcarpae with acaulescent plants, and narrow leaves with coiled tips, but our study found enough morphological differences to elevate in taxonomic rank to the species level.

Nolina brandegeei is quite different from N. palmeri because it is an arborescent plant with linear leaves that have no narrowing or constriction above the base, leaf margins with erect and forked denticles, leaf tips shredded, fruits 0.6 cm long, 0.9 to 1 cm wide, fruit peduncles 6–12 mm long and articulated near the base, and flowers present in summer with fruits mature in fall, seeds exposed at maturity. This taxon resembles N. beldingii , but that species has leaves with a constriction above the base, leaf margin with curved and nonforking denticles ( Fig. 5 ), leaf apices entire, fruits 1 cm in diameter or more, flowers produced in spring with fruits maturing in summer and seeds not exposed in mature fruits.

Arborescent plant, 2 to 6 m high, trunks 20–30 cm diam, usually branched with apical rosettes, old specimens much branched; branches separate from each other at 30° to 40°; bark gray, fissured, forming small rectangles. Rosettes massive ca. 1.5 m in diam, with erect leaves, reflexed at maturity, persistent and covering the branches, but not the trunk. Leaves linear, not narrowing or constricted above the base, 95 to 120 cm long, 1.0– 1.5 cm wide at the middle, flat in cross section, green to glaucous green; leaf bases triangular, 4.5–7 cm long, 5.0 at the base, and 1.3–1.7 at the top; margins denticulate; teeth 0.2 mm, erect to curved, forked or branched, sometimes fused. Inflorescence 1 to 3.5 m long, 30 cm wide, ovoid; scape 15 to 70 cm long, 1.5–2.0 cm diam; bracts linear to long triangulate, 47 cm long decreasing apically in size to 20 cm; inflorescence branches perpendicular to ascending, up curved below the middle, 43 cm long, decreasing apically in size to 10 cm, subtended by bracts 17 to 6 cm long; branchlets 14 to 3 cm long, the final branchlets 5 to 8 cm long. Staminate flowers campanulate, 2.5 mm diam, 2 to 3 per node on pedicels 2 to 3.5 mm articulated below the middle, surrounded by wide lanceolate membranaceous bracteoles, 1.5 to 3 mm long, persistent, whitish to yellowish, margins laciniate; tepals oblong to oblong-lanceolate, 2.5 mm long, 1 mm wide, reflexed at maturity, apex with a glandular pilose area 0.1 mm long; filaments 1.5 mm long, anthers 1 to 1.2 mm long. Pistillate flowers campanulate, 2.5 mm diam, 1 to 2 per node, on pedicels 3 to 3.5 mm long, articulate below the middle, surrounded by lanceolate membranaceous bracteoles 1.5 to 3 mm long, persistent, yellowish, margins laciniate; tepals lanceolate, 2 mm long, 1.5 mm wide, thickened at the middle of the base, style very short, stigmas 3-branched; nectaries septal, at the base of the ovary. Fruits depressed, 1(–2) per node, 6 mm long, 10 mm wide, on peduncles 6 to 12 mm long, articulate near the base, tepals persistent, the internal adpressed covering the nectaries, the external reflex. Seed 1(2 or 3) per fruit, ovoid to ellipsoid, ca. 3.5 mm long, 2.5 to 3 mm wide, exposed at maturity, maroon. Figure 6 ; Table 1.

Ecology and Distribution —This species occurs on northfacing slopes, on steep hills and in canyons with alkaline igneous rocks in the sierras de La Giganta, San Francisco, Guadalupe, Las Palmas, and San Bruno in the state of Baja California Sur ( Fig. 7 ). The plants grow mostly in Quercus forests and desert scrub with Brahea brandegeei (Purpus) H. E. Moore , Fouquieria columnaris (Kellogg) Curran , Ebenopsis confinis (Standl.) Barneby & J. W. Grimes , Prosopis sp. , and Croton sp. , from 700 to 1900 m altitude on volcanic rocks. The plants flower from June to September, and set fruits from October to December.

To assess the identity of the proposed taxon, a set of problems was found within the herbarium collections concerning the probability that this taxon could be either part of N. palmeri or of N. bigelovii . According to Brandegee (1889), he collected one specimen of an unknown Nolina at San Julio on April 19–20 of the same year. He stated that the plant has “leaves only of an arborescent branching species, 15 feet high with light green narrow leaves 3–4 feet long.” These details, and the fact that he also cited a young specimen of N. bigelovii from Ubi (Yubay), show evidence that he knew the species, and that it was a different taxonomical entity ( Table 1). Recently, Hochstätter (2010) presented an article on Nolina , considering N. palmeri var. brandegeei as a synonym for N. palmeri . But, the paper appears to be mainly a bibliographic revision and not a detailed taxonomic treatment. However, though N. brandegeei has similarities with N. palmeri , including the exposed seeds at maturity, it differs greatly in its arborescent habit, larger and flat leaves, leaf margin denticles forked, shredded and not coiled leaf tips, and larger fruits ( Fig. 4 ). In addition, the type specimen of Nolina brandegeei (Trelease) L. Hern. comb nov. deposited at MO should be relabeled as collected at San Julio, Baja California Sur, on April 19–20, 1889.

Representative Specimens Examined — Mexico. — BAJA CALIFORNIA SUR: San Julio, 11 Apr. 1889, Brandegee s/n ( UC) ; Ubi, Agua Bonita , 8 May. 1889, Brandegee s/n ( UC) ; Sierra La Giganta, above Los Encinos , 1 Mar. 1939, H. S. Gentry 4292 ( GH, MO) ; Ridge northwest of main peak, Cerro de La Giganta , 23 Nov. 1947, A. Carter 2038 ( LL, MEXU, US) ; North slope of Cerro San Juan, 4 Feb.1964, R. Moran 11584 ( SD) ; Summit of Volcán las Tres V´ırgenes, 12 Feb. 1964, R. Moran 11691 ( SD, US) ; North slope, summit of Cerro de la Higuera, Sierra San Francisco , 21 Feb. 1964, R. Moran 11723 ( SD) ; Summit, Cerro Natividad, Sierra San Francisco , 24 Feb. 1964, R. Moran 11751 ( SD) ; At pass ca. 3 miles west of ex-mission Guadalupe on trail to San Pedro, 11 Mar. 1964, R. Moran 11790 ( CAS, SD) ; North slope at summit of Cerro Azufre , 20 Oct. 1971, R. Moran 18736 ( SD, US) ; North slope of Cerro Barranco, Sierra de Guadalupe , 23 Oct.1971, R. Moran 18822 ( SD) ; Sierra de La Giganta, Cerro Mechudo , the southernmost peak of the Sierra, 2 Nov. 1971, R. Moran 18892 ( BM, MEXU, SD) ; N slopes of NE side of Volcán Tres V´ırgenes, 12 Apr. 1973, J. Henrickson 8993 ( SD) ; Sierra de Las Palmas , W de San Bruno (31 miles), 20 jun. 1973, H. S. Gentry 23319 ( AZ, DES, CAS, MEXU, US) ; Rancho La Laguna, Sierra San Francisco , 23 Nov. 1976, R. Moran 23824 ( SD) ; Sierra San Francisco, Mesa San Jorge, ca. 7–7.5 km W of San Francisco & 16 km MW of Santa Marta, 9 jun. 1984, J. Dice 512, 515, 516 ( CAS, MEXU, SD) ; Sierra de San Francisco, Mesa de San Jorge, ca. 4.21 km (by road) WSW of the Village of San Francisco de la Sierra , 23 Jun. 1986, C. Dice , 675 ( GH, SD) ; Sierra de San Francisco, west of the town of San Francisco de La Sierra , along road to Rancho Sorpresa, 24 Apr. 1994, W. Hogdson 8185 A ( DES, SD) ; Sierra de San Francisco, west of the town of San Francisco de La Sierra , on the road to Rancho Sorpresa, 24 Apr. 1994, J. Rebman 2627 ( BCMEX, SD) ; Sierra San Francisco, Cerro Bola , 3 Sept. 1995, W. Hogdson 9582, 9589a ( DES; SD) ; San Francisco de la Sierra, 23 Oct 1997, J. Rebman s/n ( HCIB, SD) ; West of Mulegé ; cumbre de San Pedro ; between the Ex-mision Guadalupe and San Juan de las Pilas , 29 Oct.1997, J. León de la Luz s/n ( BCMEX) ; Sierra de Guadalupe: West of Mulegé : cumbre de San Pedro between ex-mission Guadalupe and San Juan de las Pilas, 29 Oct. 1997, J. Rebman 4716 ( HCIB, SD) ; Sierra de San Francisco , 1.5 km al NW de Santa Ana, Nov. 2004, O. Baltasar s/n, ( QMEX) ; Sierra de la Giganta, Campamento de la Sabanilla del Mechudo , 11 Nov. 2007, J. L. León de la Luz 10539 ( HCIB, SD) ; Hernández 2007 ( QMEX). NOLINA INTERRATA H. S. Gentry Madro ~ no 8:181. 1946 . TYPE:

USA: California, San Diego Co. Slope west of Dehesa

School, 5 Aug. 1945, H. S. Gentry 7330 (Holotype SD,

isotypes: AZ, CAL, MICH).

Plant rhizomatous, subterranean stems growing in a horseshoe shape by branching always to the same side; bark reticulate forming pentagonal pyramids 0.5 cm deep, 1 cm wide. Leaf rosettes subsessile, 40 to 90 cm diam, 15 to 25 leaves per rosette. Leaves linear, 48 to 109 cm long, 0.7–1 cm wide, helicoidal, glaucous green, scabrid; leaf margins with two sizes of denticles, the larger ones separated ca. 0.5 mm; leaf bases triangular to long deltate, 6–7 cm long, 3–3.5 cm wide in the inferior part, 1.2–1.5 cm wide in the superior part, yellow straw to dark brown with age, persistent, recurved; leaf apex entire. Inflorescences paniculate, largely ovoid to largely ellipsoid, 0.8 to 2 m long, 40 to 42 cm wide; scapes 35 to 45 long, 0.8 to 1.3 cm diam at the base; scape bracts linear, 15 to 30 cm long, separated among them by 10 to 15 cm; inflorescence branches compound, 37 cm long, decreasing apically in size to 9 cm, pedunculated, with 2 branchlets at the base, all subtended by papery linearlanceolate bracts, 14 cm long, decreasing apically in size to 1 cm; branchlets 3 to 8 cm long, the apical ones 8 to 15 cm long, all subtended by amplexicaule papery bracts, 7 to 10 mm long. Staminate flowers 2 to 3(–4) per node, 3 to 3.5 mm diam, on pedicels 4 to 5 mm long, wider in the apex, articulate above the middle, surrounded by membranaceous bracteoles, 2 to 3 mm long, margins dentate; tepals lanceolate, 2 to 3 mm long, 1.5 mm wide, reflexed from the middle at maturity, white to creamy, the midvein yellow green; filament 1 to 1.2 mm long, anthers 0.8 to 1 mm long; ovary undeveloped, on a small stipe. Pistillate flowers 2 per node, 2 mm in diam, on pedicels 1.8 to 2 mm long, articulate around the middle, surrounded by membranaceous bracteoles 2 to 4 mm long, margins dentate; tepals lanceolate, 2 to 2.5 mm long, 1 to 1.5 mm wide, white to creamy with purple dots or blotches around the midvein; staminoids 1 mm long; ovary 1.5 to 2 mm diam, stigmas sessile, 3-lobulate. Fruits slightly depressed, 0.8 to 1 cm long, 1 to 1.2 cm wide, on peduncles 6 to 7 mm long, articulate below the middle; fruit lobes papery, brown with purple reddish spots at the middle, with weak margins; apical notch 2.5 to 3 mm deep; tepals persistent, the external ones reflexed. Seeds ovoid, asymmetrical, 4 to 4.5 mm long, 2 to 2.5 cm wide, grayish green to brown at maturity, microreticulate, punctate, and sometimes with papillate trichomes, hilum suprabasal with a conic caruncle. Figure 8 .

Ecology and Distribution —This species grows in sandy to deep soils on slopes with gabbro outcrops, 340 to 460 m in elevation, in chaparral vegetation with Adenostoma fasciculatum Hook. & Arn., Arctostaphylos sp. , Rhus ovata S. Watson and Ceanothus sp. The individuals of N. interrata can withstand periodic wildfires.

It is an endemic and endangered species of San Diego County, California and the northwestern part of Baja California, Mexico ( Fig. 2 ). The plants flower from April to August, and set fruits from August to October.

Representative Specimens Examined — Mexico. — BAJA CALIFORNIA: Rancho de la Cruz, 6 Km ENE of San Antonio de Las Minas , 12 Aug. 1981 R. Moran 29794 ( CAS, MO, TEX) ; Rancho de la Cruz, 6 Km ENE of San Antonio de Las Minas , 12 Aug. 1981 R. Moran 29795 ( MEXU, MO, SD, UC) ; 1 km NW of ranch house, Rancho de la Cruz, 6 Sept. 1981, R. Moran 29836

(SD); Upper Canyon Arce , 2 km SW of Rancho de la Cruz, 6 Sept. 1981, R. Moran 29841 ( SD) ; 3 km SSW of Rancho La Cruz , 6 Sept. 1981, R. Moran 29845 ( SD) ; Upper portion of Ca ~ nón Arce, ca. 2.1 km of Rancho de la Cruz, ca. 10 km N of Ensenada, J. Dice 621 ( CAS) ; ca. 1 km of Rancho La Cruz , 13 km N of Ensenada, J. Dice 700 ( SD) .

Figures

FIG. 5. Leaf margins. A. Nolina beldingii. B. N. bigelovii, with a separate leaf fiber. C. N. brandegeei. D. N. interrata. E. N. palmeri.

FIG.6. Nolina brandegeei. A. Habitat. B. Plant habit. C. Solitary plant. D. Leaf apices. E. Plant habit with staminate inflorescence. F. Male flower showing the apical trichomes in the tepal (from voucher J. Dice 515). G. Fruit (from voucher W. Hogdson 9582). Photos: Jon Rebman. Illustrations: Luis Hernández- Sandoval.

FIG. 7. Distribution of Nolina brandegeei in the Baja California Peninsula.

FIG. 4. Distribution of Nolina bigelovii in the Baja California Peninsula.

FIG.8. Nolina interrata. A.Habit. B.Plants with inflorescences.C.Inflorescence bracts. D. Inflorescence architecture.E.Staminate inflorescence branch.F. Staminate flowers. Photos A, E, F: Jon Rebman; B–D: Luis Hernández-Sandoval.

FIG. 2. Distribution of Nolina interrata and N. beldingii on the Baja California Peninsula.