Salvia subpatens Epling (1939: 97)

González-Gallegos, Jesús Guadalupe & Gama-Villanueva, Olga Jazmín, 2013, Resurrection of Salvia species (Lamiaceae) recently synonymized in Flora Mesoamericana, Phytotaxa 151 (1), pp. 1-24 : 17-22

Salvia subpatens Epling (1939: 97) ( Figs. 1 and 2A–B ). Type: MEXICO. Estado de México. Temascaltepec :

near Tejupilco, 1340 m, 27 July 1933, G. B. Hinton 4376 (holotype UC!, isotypes ASU, BM, GH, K!, US!). Fig .

13 A and B.

Salvia viscidifolia Epling (1940: 517) . Type: MEXICO. Guerrero. Coyuca de Catalán: Río Frío Diamantes , 2100 m, 26 September 1937, H.B. Hinton et al. 10725 (holotype UC!, isotype K!) .

Perennial herb, erect, 40–80 cm tall, arising from tuberous fusiform roots; stems pilose. Leaves mostly sessile, or with petioles up to 2 mm long, pilose; leaf blade lanceolate, oblong-lanceolate to ovate, 4.3–15 × 1.4–6.3 cm, acute at the apex, rounded to truncate at the base, serrate margin, pilose or with appressed hairs, and covered profusely with sessile glandular dots on both surfaces. Inflorescences in racemes (5–) 11–22 cm long, with 3–13 verticillasters each, these 2–6-flowered, lowermost nodes 1.3–2 cm apart, floral axis pilose to hirsute and covered wih glandular-capitate hairs. Floral bract lanceolate to linear-lanceolate, 3.2–11.4 × (0.4–) 1–1.7 mm, persistent, attenuated at the apex, truncate at the base, margin entire, covered with some glandular-capitate hairs. Flowers with pedicels (2.6–)4–5(–7.1) mm long (up to 9 mm long in fruit), pilose and with some glandular-capitate hairs, glandular punctate. Calyx (4.4–) 7.2–11.1 mm long, 3.1–6.5 mm wide at the throat (up to 13.3 × 8.8 mm in fruit), green and irregularly purple tinged, pilose and covered with glandular-capitate hairs, internally with antrorse hairs toward the apex, upper lip 5-veined and shortly trimucronate, lobes of the lips acute, border of the lips ciliated. Corolla deep blue with white nectar guides on the lower lip, glabrous except for the upper lip and beneath the lower one which are sparsely pilose; tube 8.5–11.8 mm long, (3.6–) 4.5–6.4 mm wide at the throat, slightly constrained toward the base, ventricose, internally epapillate; upper lip 5.6–8.5 mm long, lower one (7.3–)11.4–15.3 × 5.5–12 mm, deflexed. Stamens 2, exserted 9–12 mm from upper corolla lip; filament 2–3(–6) mm long; connective 16.3–19 mm long, thin and not ornate; theca 2.7–3(– 4) mm long; a pair of staminodes above and behind filament attachment to the corolla. Gynobasic horn absent; style 2.1–3.3 cm long, glabrous, lower stigmatic branch acute. Mericarp ovoid, 2.2–3 × 1.5–2.5 mm, light brown and irregularly dark brown marbled, smooth and glabrous.

Distribution, habitat and phenology: Salvia subpatens grows in the Mexican states of Estado de México, Guerrero, Jalisco and Michoacán (fig. 14). It inhabits pine-oak, oak, pine and montane cloud forests, from (1600–) 2100–2900 m elevation. It shares habitat with Abies religiosa , Arbutus occidentalis McVaugh & Rosatti , A. xalapensis , Castilleja mcvaughii N.H.Holmgren , Comarostaphylis glausescens , Pinus durangensis Martínez , P. hartwegii Lindl. , Quercus crassifolia Bonpl. , Q. laurina Bonpl. and Symplocos citrea . It flowers and fructifies from late July to the beginning of January.

Discussion: Salvia patens (fig. 13 C and D) and S. subpatens belongs to section Blakea (1939: 94) ; the species contain therein are morphologically very similar, they are grouped mainly because they share tuberous fusiform roots, persistent floral bracts, trimucronate upper calyx lips, showy deep blue corollas with the upper lip generally arquate, glabrous styles and lower stigmatic branch very short to almost absent. Klitgaard (2012) regards the type specimens of S. subpatens as possibly representing distal fragments of individuals of S. patens , since the leaves are reduced and sessile as the distal leaves in the latter species. Hence, she reduced S. patens to the synonymy of S. subpatens . However, the examination of more than just type specimens and field exploration reveals that such proposal is not justified. Salvia subpatens differs in several other characters that can not be embraced by the morphological variation of S. patens . The stems lack of glandular-capitate hairs, though they can rarely present them in the last internodes before the inflorescence (vs. covered with glandularcapitate hairs throughout the stem), sessile leaves or with petioles up to 2 mm long along the stem (vs. leaves with petioles 2–10 cm long that are progressively reduced toward the inflorescences till being absent), lanceolate, oblong-lanceolate to ovate leaf blades (vs. ovate-deltoid to deltate), generally narrower leaves (1.4–6.3 cm vs. 5–14 cm), shorter calyces [(1.4–) 7.2–11.1 mm vs. 11.8–16(–20) mm], smaller corolla tubes [8.5–11.8 × (3.6–) 4.5–6.4 mm vs. 16–24 × 7–10 mm], shorter upper (5.6–8.5 mm vs. 23–37 mm) and lower [(7.3–) 11.4–15.3 mm vs. 16–24 mm] corolla lips, exserted stamens from upper corolla lip (vs inserted), shorter connective [16.3–19 mm vs. (3.4–) 4–4.3 mm], and shorter style (2.1–3.3 cm vs. 5–6 cm).

Salvia subpatens does not grow in the area covered by the Flora Mesoamericana project (fig. 14).

Representative specimens examined ( Salvia subpatens ): MEXICO. Jalisco Cuautitlán: mountains E of Manantlán , about 15 miles SSE of Autlán by way of Chante, 8300 ft, 27 July 1949, R. L . Wilbur & C. R . Wilbur 1873 ( MICH!); top of sharp crest of the Sierra de Manantlán Oreintal just E of Cerro Las Capillas, along road from Cerro La Cumbre to Los Jardines , 19 km due SSE of El Chante , 19º33’15”N 104º9’0”W, 2800 m, 7 January 1980, P . D. Sorensen & P . Metekaitis 7911 ( ENCB!, MEXU!, UC!, WIS!); top of sharp crest of the Sierra de Manantlán Oreintal just E of Cerro Las Capillas, along road from Cerro La Cumbre to Los Jardines , 19 km due SSE of El Chante , 19º33’15”N 104º9’0”W, 2800 m, 10 October 1980, H. H . Iltis & R . Guzmán-M. 3214 ( ENCB!, IBUG!, UC!, WIS!); Cerro de Las Capillas, junto a la cabaña de los guarda incendios, Sierra de Manantlán , 19º33’15.6”N 104º8’53.6”W, 2880 m, 21 January 2011, J. G . González-G. & F.J. Santana Michel 850 ( IBUG!, ZEA!). Tecalitlán: km 41 de la brecha que conduce a Jilotlán de Dolores, a partir de la carr. 110, campamento maderero Plan de Lego , Altitlán , 1940 m, 18 July 1986, A . Rodríguez-C. 413 (IBUG!). Michoacán. Coalcomán: Sierra de Torricillas. 2200 m, 13 October 1938, G. B . Hinton et al. 12369 ( MICH!, UC!); Sierra de Torricillas (sic), 2150 m, 24 July 1939, G. B . Hinton et al. 13989 ( MICH!) .

Representative specimens examined ( Salvia patens ): MEXICO. AguascaIientes. Calvillo: Barranca Las Cazuelas, Sierra Fria, 2350 m, 2 September 1981, C . Cuéllar-R. 23 ( HUAA!). Guanajuato. León: frente al Área Natural Protegida Sierra Los Lobos, km 18 de la carretera León-San Felipe , 21º20’N 101º40’W, 2500 m, 6 October 2008, M . Harker et al. 3858 ( IBUG!). Hidalgo. Omitlán de Juárez: poblado de Guerrero, km 17 carretera Méx. 105 entre Pachuca y Huejutla, 20º9’27.3”N 98º39’27.6”W, 2548 m, 16 September 2011, A . Rodríguez-C. et al. 6455 ( IBUG!). Jalisco. Lagos de Moreno : cerro de El Espía, 3 km al S de la presa Juan Vaquero y 6.3 km al SE de Ciénega de Mata , en una cañada que desemboca a la presa, 21º42’38.4”N 101º45’19.1”W, 2450 m, 20 October 2011, J. G . González-G. & F. Pérez 1128 ( IBUG!). Michoacán. Senguío : aproximadamente 1 km al SW de la Cortina de la Presa Chincua, en camino Chincua-Seguío, m, 29 August 1989, I . García-R. 2885 ( GUADA!, IBUG!). Ucareo: Las Peñas del Bellotal , 9 October 1987, H . Díaz-B. 4214 ( IBUG!). Querétaro. Amealco : 2 km a l s de Laguna de Servín, 2650 m, 25 August 2001, E . Carranza & S . Zamudio 6214 ( IBUG!). Colón : Vertiente S del Cerro Zamorano, cerca de Trigos, 2800 m, 2 September 1987, J . Rzedowski 44500 ( IBUG!). San Luis Potosí. Villa de Zaragoza: alrededores de Sierra de Álvarez, entrando por San Francisco , 22º1’48.4”N 100º36’46.8”W, 2352 m, 12 September 2011, A . Rodríguez-C. et al. 6364 (IBUG!).

Salvia unicostata Fernald (1900: 501) Type: MEXICO. San Luis Potosí. Without specific locality, 6000–8000 ft, 1878, C. C. Parry & E. Palmer 760 (holotype GH!, isotypes BM, E, F!, K!, MO!, NY!, P!, PH, US!) .

Perennial herb, erect, 15–50 cm tall; stems glabrous, interpetiolar space pilose. Leaves sessile; leaf blade linear, 2.4–8.2 × 0.1–0.5 cm, acute to attenuated at the apex, slightly attenuated at the base, margin entire and revolute, verrucose above and sparsely hispidulous beneath on the midvein, the rest glabrous, midvein thicken and white, the other are not visible; wih axillary fascicles of several immature leaves. Inflorescences in racemes 7–15(–33) cm long, with 3–9 verticillasters, these 2–6-flowered, lowermost nodes 2.4–4.5 cm apart, floral axis covered with glandular capitate hairs. Floral bract ovate, ovate-lanceolate to rhomboid, (4–)5.7– 6.6(–10.2) × 2.8–3.2 mm, caudate at the apex, attenuated and then truncate at the base, margin entire and ciliated, covered with glandular capitate hairs outside. Calyx 5.2–6.1 mm long, 3.1–4 mm wide at the throat (up to 6.2–7 × 5–8.7 mm in fruit), green to bluish green, covered with glandular capitate hairs and black sessile glandular dots, internally verrucose, upper lip 5–7-veined and trimucronate, lobes acute and spinulose. Corolla deep blue with white nectar guides on the lower lip, upper lip and ventral portion of lower one pilose, the rest glabrous to glabrescent; tube 5–6.2 mm long, 3.1–3.8 mm wide at the throat, slightly ventricose, not constrained at the base, internally epapillate; upper lip 3.6–4.1 mm long, lower lip (8–)9.6–10.5 × (7.4–) 8–9.8 mm, deflexed. Stamens 2, included; filament 1.4–1.6 mm long; connective 4.5–5 mm long, geniculate near midportion; theca 1.4–1.6 mm long; two staminodes present above and behind filament attachment to corolla tube. Gynobasic horn 0.7–0.9 mm long; style 8.5–8.9 mm long, evidently thicker toward the apex and densely pilose on dorsal side, lower stigmatic branch acute. Mericarp ovoid, 1.5–1.7 × 1–1.2 mm, amber-brown, smooth and glabrous.

Distribution, habitat and phenology: Salvia unicostata is restricted to the Mexican Plateau region, reported from the states of Jalisco, Nuevo León, San Luis Potosí and Tamaulipas; but probably also present in Aguascalientes. It dwells in dwarf oak, pine, oak-pine, and cedar forests, from (1795–)2300–2500(–3400) m elevation. It shares habitat with Juniperus deppeana Steud. , Nasella mucronata (Kunth) R.W.Pohl , Pinus cembroides Gordon , P. montezumae Lamb. , P. teocote Schltdl. & Cham. , Quercus chihuahuensis Trel. , Q. eduardi Trel. , Q. greggii (A.DC.) Tre. , Q. grisea Liebm. , Q. laeta Liebm. , Q. mexicana Bonpl. , Q. opaca Tel. , Q. potosina Trel. , Q. sideroxyla Bonpl , Sisyrinchum tenuifolium Humb. & Bonpl. ex Willd. , Stevia lucida Lag. , Yucca filifera Chabaud ; it is sympatric with other Salvia species as S. axillaris Moc. & Sessé , S. macellaria Epling , S. microphylla Kunth , and S. prunelloides Kunth. It flowers and fructifies from the end of May to October.

Discussion: Salvia unicostata belongs to section Uliginosae ( Epling 1935: 52) Epling (1939: 54), this is noteworthy because the calyx is covered with dark amber sessile glandular dots and the upper calyx lip is trimucronate. Klitgaard (2012) reduced S. unicostata to the synonymy of S. reptans , a species that belongs to section Farinaceae . Nonetheless, additionally to the distinctive characters of Uliginosae exhibited by S. unicostata , this can be distinguished by means of its clearly opposite leaves (vs. pseudo-whorled in S. reptans ), thicken and whitish midvein (vs. slender and green), revolute leaf margin (vs. not revolute), floral bracts (outer surface), floral axis, pedicels and calyces covered with glandular-capitate hairs (vs. hispidulous), calyx lobes stiff at the apex like a tiny spine (vs. acute and slender), shorter corolla tube [5–6.2 mm vs. 7.9– 8.5(–10) mm], shorter upper corolla lip (3.6–4.1 mm vs. 5.2–6.2 mm), geniculate connective (vs. ornate at midportion with a retrorse acute tooth), shorter connective (4.5–5 mm vs. 6.6–7.7 mm), shorter filament (1.4– 1.6 mm vs. 2.5–2.8 mm), and shorter style (8.5–9 mm vs. 12.2–13 mm).

Salvia unicostata grows in semiarid habitats in the Mexican Plateau and surrounding portions of Sierra Madre Oriental; hence it is not present in the area covered by the Flora Mesoamericana project. Furthermore, the kind of habitat occupied by S. unicostata differs from that of S. reptans , which grows in wet soils.

Representative specimens examined: MEXICO. Jalisco. Lagos de Moreno : cerro de El Espía, 3 km al S de la presa Juan Vaquero , 21º42’45”N 101º45’17.9”W, 2300 m, 20 October 2011, J. G . González-G. & F. Pérez 1121 ( IBUG!) . Nuevo León. Arramberi : La Escondida-San Francisco, 1795 m, 23 June 1993, G. B . Hinton 23079 ( Herbario Hinton !); San Josecito, below, 2395 m, 31 June 1999, G. B . Hinton et al. 27393 (Herbario Hinton!) . Nuevo León. Zaragoza-Puerto Pino , above, 2790 m, 26 June 1978, G. B . Hinton et al. 17408 (Herbario Hinton!) . Zacatecas. Pinos: Ejido La Purísima, aproximadamente 4 km al SW del pobaldo Cerrido de Dolores , 2540 m, 4 sep 2013, L. F . Colin-Nolasco & J. F . Guerrero-Rodríguez 868 ( IBUG!) .

Figures

FIGURE 1. Salvia brachyodonta, A) inflorescence, B) leaves (pictures taken by J.G. González-Gallegos).

FIGURE 2. Distribution of Salvia brachyodonta represented by an individual track (green line) and collection points (circles). The northern boundary of Flora Mesoamericana is highlighted with a red line.

FIGURE 13. Salvia subpatens, A) habit, B) corolla. Salvia patens, C) corolla, D) habit (picture A taken by L.M. González-Villarreal, B by J. Padilla-Lepe, C and D by A. Castro-Castro).