Salvia ramamoorthyana Espejo Serna (1993: 92) emend. J.G.González

González-Gallegos, Jesús Guadalupe, 2015, Salvia ramamoorthyana and S. omissa (Lamiaceae), two names for two old and largely confused species from Mexico, Phytotaxa 236 (3), pp. 215-225 : 217-218

Salvia ramamoorthyana Espejo Serna (1993: 92) emend. J.G.González

— Salvia nepetoides auct . Fernald (1900: 529), Epling (1939: 44–45), Espejo Serna & Ramamoorthy (1993: 88–89) et González-Gallegos & Gama-Villanueva (2013: 13–15), non Kunth (1817: 299).

Inter species sectionis Sigmoideae laminis foliorum rotundatis, truncatis vel cordatis ad basim, calycibus capitato-glandulosis, pedicellis capitato-glandulosis et axibus floralibus capitato-glandulosis, calycum labiis superioribus 5 vel 7-nervatis ab eis recedens.

Type: — MEXICO. Jalisco: mun. Tequila, near the microwave towers atop Volcán Tequila south of Tequila, 13 miles south of Mexico highway 15 and 11 miles south of the railroad in Tequila, 9000 ft [2743 m], 11 October 1975, J. L. Reveal & R. M. Harley 4106 (holotype MEXU-297645!, isotypes CAS, F!, MEXU-297646!, NY!, TEX!).

Perennial herb to subshrub, erect or reclined, (0.2–)0.5–1(–1.7) m tall; stems hispidulous, puberulent and generally covered with retrorse hairs on the ribs. Leaf with petiole 2.5–16.6 mm long, hispidulous; blade ovate to ovate-lanceolate, 1.3–5(–7.5) × 1–4.3(–7.2) cm, apex acute to acuminate, base rounded, truncate to cordate, margin crenate to serrate, upper surface glabrescent, short-pilose to covered with appressed hairs, the hairs concentrated on the veins, lower surface usually white tomentose, both surfaces with sessile glandular dots, more evident beneath. Inflorescence in racemes (6–) 10–28.9 cm long, verticillasters (3–)10–19, each one (2–)4–6(–10)-flowered, 0.8–2.7(–3.9) cm apart from each other toward the base, floral axis hispidulous and covered with short glandular-capitate hairs. Floral bract lanceolate, (1.3–)2.2–4.7 × 0.4–1.4 mm, deciduous, apex acute, base truncate, margin entire, short-pilose, covered with glandular-capitate hairs and sessile glandular dots on the outer surface, ciliated at the margin; bracteoles present, narrow lanceolate to linear, late deciduous. Flowers with pedicels 1.9–3.5(–4.7) mm long, puberulent and covered with short glandular-capitate hairs. Calyx (4–)5–7(–7.7) × (1.8–) 2.5–4.8 mm, green and sometimes dorsally wine tinged, short-pilose, with glandular-capitate hairs, covered with sessile glandular dots, glabrous or with short conical hairs inside toward the throat, the upper lip 5 or 7-veined, entire, lobes of the lips acute. Corolla sky blue with white nectar guides on the lower lip; tube (3.3–)4.3–6.5(–7.8) × (1.4–)2–2.8(–3.8) mm, ventricose, slightly constrained at the base, internally epapillate, glabrous to slightly short-pilose toward the lips; upper lip 2.1–4.4(–7.9) mm long, short-pilose and ventrally bordered by tiny glandular-capitate hairs; lower lip (2.5–)4.4–6.2(–9.8) × (3–) 4.2–6.7 mm, deflexed, short-pilose beneath. Stamens included; filament (1.4–) 2–3 mm long; connective 2–3.9 mm long, geniculate (ornate at midportion with a blunt tooth); theca 1.1–1.6 mm long; a pair of staminodes above and behind filament insertion, filiform and usually capitate. Gynobasic horn 0.8–1.4 mm long; style 6.2–10.7 mm long, short-pilose at apex, lower stigmatic branch sigmoid. Mericarp ovoid, 1.3–1.9 × 0.8–1.3 mm, pale brown and irregularly dark brown marbled, smooth, glabrous.

Etymology:—This taxon was named after T. P. Ramamoorthy ( Espejo Serna & Ramamoorthy 1993), a contemporaneous Salvia specialist focused on Mexican species, who made relevant contributions mainly from 1980 to 1993.

Distribution, habitat and phenology: — Salvia ramamoorthyana grows in Distrito Federal, Estado de México, Jalisco, Michoacán and Morelos ( Fig. 3 ). In Tropicos.org (2015) online database specimens from Guerrero and Oaxaca are also registered, though it is probable that these are misidentification of S. protracta Bentham (1848: 309) , a similar species also from S. sect. Sigmoideae , which is very abundant in the Mexican states mentioned above. It inhabits oak, oak-pine, pine-oak and, less frequently, montane cloud forest and tropical deciduous forest, from (1100–)1800– 2800(–3100) m elevation. It shares habitat with Abies religiosa , Alnus jorullensis , Arbutus xalapensis , Clinopodium macrostemum , Comarostaphylis glaucescens , Cunila polyantha , Dahlia barkeriae , D. coccinea , Pinus lumholtzii , P. oocarpa , Quercus candicans , Q. castanea , Q. laeta , Q. resinosa , Q. rugosa , Q. salicifolia , Salvia concolor , S. elegans , S. fulgens , S. gesneriiflora , S. longistyla , S. melissodora and S. mexicana . It might flower and fruit throughout the year, but especially from August to January.

Notes: — The name Salvia nepetoides has been usually applied to designate specimens that exhibit characters of S. sect. Sigmoideae (presence of bracteoles at pedicel base and sigmoid lower stigmatic branch) (e.g., CHAP, HUMO, IBUG, IEB, MEXU, MICH and UC). This is the approach followed by Fernald (1900), Epling (1939), Espejo Serna & Ramamoorthy (1993), Rodríguez-Jiménez & Espinosa-Garduño (1996), Cornejo-Tenorio & Ibarra-Manríquez (2011) and González-Gallegos & Gama-Villanueva (2013). However, the original description by Kunth (1817) makes no mention of the diagnostic characters that would relate it directly to S. sect. Sigmoideae , and the illustration provided therein shows a straight lower stigmatic branch instead of sigmoid. Furthermore, once the type was examined, it became clear that it does not match with the circumscription in use for this species in contemporaneous literature. It lacks bracteoles and lower stigmatic branch is straight ( Fig. 1 ). In fact, morphological characters support it as synonymous of S. amarissima . Bentham (1832 –1836) realized this and submerged S. nepetoides into S. amarissima as a synonym. Hemsley (1881 –1882) and Klitgaard (2012) followed Bentham’s conclusion, but no other author did. Actually, González-Gallegos & Gama-Villanueva (2013) criticized Klitgaard’s (2012) synopsis and reestablished S. nepetoides auct . mult. non Kunth as a valid species distinct from S. amarissima , since they had in mind Epling’s circumscription and did not examine the holotype. Leaves can be similar between S. amarissima and S. nepetoides auct . mult. non Kunth in terms of shape and texture; pubescence of floral axes (glandular-capitate hairs) is also similar though in the latter the hairs are shorter. These similarities may be responsible in part for the misinterpretation of the taxon. Fernald (1900) adds a comment that Bentham (1832 –1836) and Hemsley (1881 –1882) treated S. nepetoides s.str. under S. amarissima , nonetheless he lists S. nepetoides auct . mult. non Kunth as a true species distinct from S. amarissima . Epling (1939) erected section Sigmoideae and provided a complete description of S. nepetoides auct . mult. non Kunth; he also said he had seen the holotype at P herbarium. Epling’s (1939) treatment of Salvia subgenus Calosphace is the most actual and comprehensive work dealing with Mexican sages, so it is not accidental that the majority of Salvia specialists have followed his interpretation of S. nepetoides auct . mult. non Kunth. Recognizing S. nepetoides as synonym with S. amarissima , the taxon in S. sect. Sigmoideae then lacks a name, a new species that has passed unnoticed for a long time under another species name. A solution might be the description of S. nepetoides auct . mult. non Kunth as a new species; however, as stated in the introduction, S. ramamoorthyana holotype belongs to S. nepetoides auct . mult. non Kunth; therefore, the former is a synonym and automatically becomes an available name to be applied to S. nepetoides auct . mult. non Kunth. Hence, these are the reasons supporting the recognition of S. nepetoides auct . mult. non Kunth under S. ramamoorthyana after preparing a re-circumscription of the species in order to discard the morphological characters and specimens of other species that were mixed in the original description and that do not match with the type ( Espejo Serna & Ramamoorthy 1993).

It is noteworthy that Salvia chapalensis Briquet (1898: 145) does not have consistent characters that warrant its distinction from S. nepetoides auct . mult. non Kunth except the absence of glandular-capitate hairs. Moreover, Salvia chapalensis is known exclusively from the type collection gathered in 1892 in mountains at the north of Chapala lake, even though recent botanical exploration has been conducted in that region. Espejo Serna & Ramamoorthy (1993) consider the lack of S. chapalensis collections as the result of the strong anthropogenic pressure that modified the area where it grew due to deforestation and environmental degradation. Another possibility is that S. chapalensis represents an unusual and ephemeral morphological variant of S. nepetoides auct . mult. non Kunth. The last option seems to be more plausible since S. nepetoides auct . mult. non Kunth is abundant in there; and if so, the proper name for the species due to the principle of priority would be S. chapalensis instead of S. ramamoorthyana . However, the presence/ absence of glandular-capitate hairs and their distribution in the plant is a character with great relevance within S. sect. Sigmoideae species delimitation, and until S. chapalensis can be found in the wild and its taxonomic status clarified, it would be better to take a more conservative point of view.

Additional specimens examined: — MEXICO. Distrito Federal: Eslava, 19 November 1902, C. G. Pringle 11125 ( MICH!). Estado de México. Temascaltepec: Tenayac , 1690 m, 18 August 1933, G. B. Hinton 4440 ( GH, MO, NY, UC!) ; Ipericones, 23 July 1935, G. B. Hinton 8084 ( MO). Jalisco. Bolaños: camino al Cerro del Gallo , predio Las Berenjenas, a 10 km al NO de Bolaños , 21º52.6’, 103º51.82’, 27 November 2002, E. Domínguez 29 ( MEXU!) ; 16 km al NO de Bolaños, camino a Tuxpan de Bolaños, arroyo El Nalgazo, 21º53’6’’N, 103º50’35’’E, 2020 m, 24 October 1996, J. Calónico-S. & G. Flores-F. 3216 ( MEXU!) ; Casimiro Castillo: 3–4 km de Puerto Los Mazos por la carretera Autlán a Barra de Navidad , 19º41’, 104º25’, 1100 m, 8 July 1997, L. Guzmán-H. & R. Delgado-C. 1641 ( ZEA!) ; Cuautitlán de García Barragán: on lower shoulders (1 km N) of Cerro El Muñeco, 1–2 km E of and above Los Jardines, E-end of the Sierra of Manantlán Oriental Plateau, 20 km SSE of El Chante, 19º33’30’’N, 104º6’0’’, 2600 m, 8 January 1980, P. Sorensen 7925 ( ENCB!, MEXU!) ; Jocotepec: base del paraje El Quinto, subiendo por la Cañada del Agua de Huejotitán y cerca de El Chiflón , 20º21’39.77’’N, 103º26’48.73’’E, 2441 m, 4 February 2012, J. G. González-G. & J. A. Machuca-N. 1176 ( IBUG!) ; Cerro Viejo, paraje El Rincón-Cresta , Bola del Viejo, Barranca del Agua, 20º24’, 103º25’, 1600 m, 5 February 1987, J. A. Machuca-N. 5479 ( WIS!) ; Cerro Viejo, paraje El Rincón-Cresta , Bola del Viejo, Barranca del Agua, 20º24’, 103º25’, 1800 m, 5 February 1987, J. A. Machuca-N. 5500 ( WIS!) ; El Rincón , ladera S al E de El Molino, 1600 m, 5 February 1987, J. A. Machuca-N. 5437 ( IBUG!) ; Mazamitla: 8 km al SE de Mazamitla y a 1 km al SE de Dos Aguas , 2370 m, 8 April 1988, A. G. Mendoza et al. 3898 ( MEXU!) ; San Gabriel: El Floripondio, carretera Ciudad Guzmán-El Grullo , 19.640323º,103.604429º, 2285 m, 1 November 2012, J. G. González-G. 1187 ( IBUG!) ; km 9 del camino de ascenso al Nevado de Colima, a partir del entronque con la carretera Ciudad Guzmán- Sayulapan , 19º35’57’’N, 103º35’6.5’’E, 3100 m, 7 October 2010, A. Castro-C. & J. L. Villa-V. 2274 ( IBUG!) ; El Floripondio, Nevado de Colima, 19º38’5’’N, 103º37’4’’E, 2347 m, 3 March 1998, F. J. Santana-M. 9047 ( ZEA!) ; 75 km E of El Grullo on paved hwy (Puerto de Floripondio) between El Grullo and Ciudad Guzmán, 12.7 km W of Junction with road to Parque Nacional Volcán de Colima, this along the N shoulders of Nevado de Colima, 19º38’30’’N, 103º35’30’’E, 2145 m, 19 November 1989, M. A. Wetter & R. R. Kowal 1350 ( ZEA) ; along road from El Fresnito to Venustiano Carranza, at km 22.59 marker, in large stand on mountaintop in semishade area in weathered limestone soil, 2347 m, 6 November 1975, K. M. Peterson & C. R. Broome 391 ( IBUG!, MEXU) ; N slopes of Nevado de Colima, about 1 km by lumber road above sawmill called Piedra Ancha , 2400 m, 2 April1949, R. McVaugh 10169 ( MICH!) ; Tapalpa: along gravel road to Juanacatlán and Atemajac de Brizuela , 4.8 km N of junction of this road with paved hwy between Tapalapa and Jalisco hwy 54 at village of San Antonio , junction is 7.8 km E of Tapalpa , this is NE of Tapalpa , 20º0’27’’N, 103º42’42’’E, 2450 m, November 1989, M. A. Wetter & R. R. Kowal 1378 ( ZEA!) ; El Fraile, 10 km al NW de Tapalpa, por el camino a Chiquilistlán , 2300 m, 9 February 1986, L. M. González-V. et al. 2296 ( IBUG!) ; mountains 10–11 km N of Tapalpa, road to Chiquilistlán , 2300 m, 30 January 1975, R. McVaugh 26037 ( MICH!) ; on road to Chiquilistán form Tapalpa, 7.6 miles NW of Tapalpa, 2770 m, 7 November 1975, K. M. Peterson & C. R. Broome 400 ( IBUG!, IEB!, MEXU!) ; El Fraile, Tapalpa , 2400 m, 26 May 1968, L. M. Villarreal de Puga 1708 ( IBUG!) ; Teocuitatlán de Corona: ladera SE de Cerro de García (aprox. 4.7 km en línea recta al SO de San Luis Soyatlán), 20º10’5’’N, 103º20’23’’E, 2610 m, 9 December 2013, P. Carrillo-Reyes et al. 7146 ( IBUG!) ; Tequila: Volcán de Tequila , 9–10 km al S de Tequila por la brecha rumbo a las antenas de microondas, 20º49.255’N, 103º51.366’O, 1980 m, 3 March 2012, J. G. González-G. et al. s.n. ( IBUG!) ; Volcan de Tequila, along the road to the microondas atop the volcan, 7.9 miles S of the junction of the railroad tracks and calle Miguel Hidalgo in the town of Tequila , 2350 m, 8 October 1982, K. M. Peterson et al. 747 ( IBUG!) ; 14–18 m SW of Tequila, on Volcan de Tequila, 2400 m, 7 November 1974, D. E. Breedlove 39269 ( ENCB!, MEXU!) ; Tlajomulco de Zúñiga: cerca de la cima del Cerro Viejo, 5.8 km al SO de San Miguel Cuyutlán y 800–900 m al N de la Mesa de la Gringa, subiendo por vereda a partir de San Miguel, 20º22’20.8’’N, 103º25’12.4’’E, 2783 m, 9 January 2011, J. G. González-G. & A. Castro-C. 817 ( IBUG!) ; base del Cerro Viejo, 5 km al SO de San Miguel Cuyutlán y 2.1 km al N de la Mesa de la Gringa, subiendo por vereda a partir de San Miguel, 20º23’2.5’’N, 103º25’22.9’’E, 2265 m, 9 January 2011, J. G. González-G. & A. Castro-C. 824 ( IBUG!) ; antiguo camino maderero que baja de Cerro Viejo, 9.2 km al sur de Tlajomulco de Zúñiga , 20º23’26.6’’N, 103º28’17.2’’E, 2225 m, 10 January 2011, J. G. González-G. & A. Castro-C. 823 ( IBUG) ; ladera exposición N al W de San Miguel Cuyutlán, en el paraje El Llano, 1950 m, 26 December 1986, J. A. Machuca-N. 5337 ( WIS!) ; Cerro Viejo, ladera de exposición N, enfrente de San Miguel Cuyutlán , brecha hacia el oriente, 1600 m, 30 July 1986, J. A. Machuca-N. 3942 ( WIS!) ; Tuxpan: slopes of the Nevado de Colima, W of summit of the N ridge, near junction of the old pack road to Zapotlán with Atenquique-Jazmín road, 2100 m, 16 October 1952, R. McVaugh 13565 ( MICH!) ; 5 km al SO del Rancho Las Milanés, brecha al Parque Nacional El Nevado, 2310 m, 20 January 1990, J. Villa-C. et al. 544 ( IEB!, MICH!) ; A 10 km sobre la desviación al Nevado de Colima, a partir de El Fresnito, 2510 m, 6 April1988, A. G. Mendoza-A. et al. 3831 ( CIIDIR!, MEXU!) ; NNE-facing slopes of Volcan Colima (just below the very base of the forested steep cone), the flat uplands, all in extensive peach orchards, deeply dissected by narrow 10–40 m deep quebradas, 14–15 km WSW of Ciudad Guzmán (14 km NNE of top Volcan Colima), ca. 2 km of paved road, 0.2 km W of Hda. Piedra Ancha , 19º38’, 103º35’, 2120 m, 31 December 1978, H. H. Iltis et al. 1059 ( ENCB!, WIS!) ; 30 km al SSO de Cd. Guzmán, base N del Volcán de Colima, rancho Piedra Ancha, 1850 m, 28 October 1978, L. M. Villarreal de Puga & J. A. Pérez de la R. 10895 ( IBUG!). Michoacán. Charo: Las Trojes, ca 32 km E de Morelia , 2200 m, 12 November 1985, S. A. Reisfield 1277 ( MEXU!) ; Mil Cumbres , 28 November 1985, T. P. Ramamoorthy 4826 ( MEXU!) ; cerca de Potenzuela, 25 km al E de Morelia, sobre la carretera a Mil Cumbres , 2200 m, 29 January 1987, J. Rzedowski 42419 ( MEXU!) ; cerca de Pontezuela, 22 km al E de Morelia, sobre la carretera a Mil Cumbres, 2200 m, 6 November 1987, J. Rzedowski 45636 ( IBUG!, CIIDIR!, MEXU!) ; Indaparapeo: 11 km al N de Indaparapeo, sobre el camino a Las Peras, 2200 m, 8 November 1987, J. Rzedowski 45660 ( MEXU!, MICH!) ; 10 k al S de Indaparapeo, sobre el camino a Las peras, 2250 m, 23 December 1986, J. Rzedowski 42303 ( MEXU!) ; Morelia: Cerro Azul , 2300 m, 4 February 1909, G. Arsène 2837 ( MEXU!) ; cerca de Durazo, 22 October 1981, T. P. Ramamoorthy & D. Lorence 2577 ( MEXU!) ; Paracho: cerro al N de Ahuitan, 2300 m, 13 December 1990, E. García & E. Pérez 3568 ( MEXU!) ; Queréndaro: 2 km al S de Real de Otzumatlán, sobre el camino a Queréndaro, 2200 m, 20 November 1986, J. Rzedowski 41883 ( IBUG!, CIIDIR!, MEXU!) ; Zitácuaro: Zitácuaro-La Campana , 2025 m, 9 December 1938, G. B. Hinton et al. 13519 ( MICH!) ; Morelos. Cuautla : 20 km NO de Cuautla, 30 July 1960, W. Forbey 68 ( MEXU!) ; 20 km NE de Cuautla , 1981 m, 3 August 1950, C. E. Boyd 84 ( MEXU!) ; Cuernavaca: above Cuernavaca , 2133 m, 9 August 1898, C. G. Pringle 7612 ( MICH!) ; mountains above Cuernavaca , 2286 m, 4 September 1900, C. G. Pringle 8457 ( MEXU!) ; barranca de Atzingo, 1650 m, 13 August 1972, J. Vázquez 3780 ( MEXU!) ; barranca Cilapeña , 2150 m, 16 October 1970, J. Vázquez 2698 ( MEXU!) ; FC de Cuernavaca k-100, Morelos, 2000 m, 17 October 1964, Gold 334 ( MEXU!) ; parte alta de la Barranca de Atzingo, entrando por la Colonia del Bosque , 18º58’17’’N, 99º17’10’’E, 1950 m, 8 September 1996, A. Espejo-S. et al. 5590 ( MEXU!) ; barranca de Chalchihuapan, al NO de Cuernavaca , 1800 m, 24 July 1990, A. R. López-F. et al. 1218 ( MEXU!) ; Cuernavaca 1.3 km después de Tlatempa, rumbo a Buena Vista del Monte , 1900 m, 8 July 2000, A. R. López-F. et al. 2885 ( CIIDIR!, MEXU!) ; Huitzilac: Huitzilac , October 1930, E. Lyonnet 607 ( MEXU) ; Tepoztlán: Sierra de Tepoztlán , 2286 m, 27 September 1904, C. G. Pringle 13166 ( MEXU!, MICH!) ; along Hwy 95, at km 65, 2150 m, 15 August 1985, D. Engle & D. Remington 124 ( MEXU) ; Tepoztlán : a 2 km NO de Tepoztlán, 16 July 1978, J. C. Soto-N. & A. Román-S. 839 ( MEXU!) ; 3 km después de Tepoztlán, rumbo a San Juan Tlacotenco, 2000 m, 10 December 198, A. Espejo-S. & A. Flores-C. 3448 ( CIIDIR!, MEXU!) ; Tlayacapan: faldas del Cerro Cuautzin, 4 km al E de San José de los Laureles , 1900 m, 4 September 1987, A. R. López-F. & A. Espejo-S. 365 ( MEXU!) ; 1 km al NO de San José de los Laureles, rumbo al cerro Cuautzin , 18º59’0’’N, 99º0’30’’E, 1900 m, 19 July 1986, A. Espejo-S. et al. 2562 ( MEXU!) .

Figures

FIGURE 1. A flower from the holotype [F. W. H. A. Humboldt & A. J. A. Bonpland s.n. (P!)] of Salvia nepetoides showing the nonsigmoid lower stigmatic branch; the bar equals 1 mm (picture taken by Elodie Lerat).

FIGURE 3. Distribution maps of Salvia omissa (white triangles) and S. ramamoorthyana (black squares);A. Detail of Jalisco and adjacent areas (box top right), B. Detail of Sierra de Manantlán (shaded area) in Jalisco (box bottom left).