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González et al.- Three new Salvia taxa from Jalisco, Mexico
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Salvia concolor subsp. iltisii, S. meera, S. rogersiana and S. santanae (Lamiaceae), three
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new species and a subspecies from Jalisco, Mexico
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Salvia concolor subsp. iltisii, S. meera, S. rogersiana y S. santanae (Lamiaceae), tres
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nuevas especies y una subespecie de Jalisco, México
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JESÚS GUADALUPE GONZÁLEZ-GALLEGOS1*, JOSÉ ANTONIO VÁZQUEZ-GARCÍA1, FRANCISCO
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JAVIER SANTANA-MICHEL2, RAMÓN CUEVAS-GUZMÁN2 AND LUIS GUZMÁN-HERNÁNDEZ2
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Botánica y Zoología, Universidad de Guadalajara-CUCBA, km 15.5 carr. Guadalajara-
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Nogales, Las Agujas, Nextipac, Zapopan, CP 45110, Jalisco, México; [email protected],
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[email protected]
Herbario Luz María Villarreal de Puga (IBUG), Instituto de Botánica, Departamento de
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Departamento de Ecología y Recursos Naturales, Universidad de Guadalajara-CUCSUR, Av.
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Independencia Nacional 151, Autlán de Navarro, CP 48900, Jalisco, México;
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[email protected], [email protected], [email protected]
Herbario ZEA, Instituto Manantlán de Ecología y Conservación de la Biodiversidad,
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Abstract. We describe and illustrate 4 new taxa of Salvia L. from Jalisco, Mexico. Three of
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them were tentatively proposed as new species in the book Flora de Manantlán. These taxa
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are endemic to the state of Jalisco. Salvia concolor subsp. iltisii J. G. González differs from
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typical populations of S. concolor Lamb. ex Benth. in having thinner and deeply cordate
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leaves, floral bracts 5-6 mm long, yellowish green calyces, up to 15 mm long at fructification,
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and with the upper lip entire. Salvia meera Ramamoorthy ex J. G. González is related to sect.
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Tubiflorae Epling, its long and narrow leaves and white large corollas make this species
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unique. Salvia rogersiana Ramamoorthy ex J. G. González belongs to sect. Briquetia Epling,
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it is distinguished by its short and compact racemes, short calyces (less than 10 mm long), 2-
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flowered verticillasters, and corolla tubes up to 15 mm long. Salvia santanae Ramamoorthy
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ex J. G. González fits well into sect. Angulatae (Epling) Epling and is closely related to Salvia
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longispicata M. Martens & Galeotti; it differs from the latter in having persistent floral bracts,
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2 or 4 (rarely 10) flowered verticillasters and, larger floral characters.
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Key words: Jalisco, sect. Angulatae, sect. Briquetia, sect. Dusenostachys, Sierra de Manantlán
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Resumen. Se describen e ilustran 4 taxa nuevos de Salvia L. de Jalisco, México. Tres de ellos
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fueron tentativamente propuestos como especies nuevas en el libro de Flora de Manantlán.
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Estos taxa son endémicos de Jalisco. Salvia concolor subsp. iltisii J. G. González difiere de
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las poblaciones típicas de S. concolor Lamb. ex Benth. por presentar hojas más delgadas y
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profundamente cordadas, brácteas florales de 5-6 mm de largo, cálices verde amarillentos
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hasta de 15 mm de largo durante la fructificación con el labio superior entero. Salvia meera
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Ramamoorthy ex J. G. González está relacionada con la sect. Tubiflorae Epling, la hacen
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única sus hojas largas y estrechas, y sus corolas grandes y blancas. Salvia rogersiana
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Ramamoorthy ex J. González pertenece a la sect. Briquetia Epling y se distingue por sus
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racimos cortos y compactos, cálices cortos (menores de 10 mm de largo), verticilastros
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bifloros, y corolas que alcanzan como máximo 15 mm de largo. Salvia santanae
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Ramamoorthy ex J. González pertenece a la sect. Angulatae (Epling) Epling, está
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estrechamente relacionada con Salvia longispicata M. Martens & Galeotti, pero se diferencia
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de esta última por tener brácteas florales persistentes, verticilastros con 2 o 4 (rara vez 10)
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flores, y características florales más grandes.
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Palabras claves: Jalisco, sect. Angulatae, sect. Briquetia, sect. Dusenostachys, Sierra de
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Manantlán
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Introduction
The genus Salvia L. as traditionally described is not monophyletic but
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integrated by 3 different clades according to Walker (2004, 2007). One of them corresponds
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to the subgenus Calosphace (Benth.) Benth., which grows from Southern United States to
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Northern Argentina and the Caribbean Islands. This embraces almost all the Mexican species
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including those treated on this paper. Of the 2 remaining clades, one is restricted to Eastern
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Asia and the other one has representatives in North America, Europe, Southern Africa and
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Middle East. Epling and coworkers (Epling, 1939, 1940, 1941, 1944, 1947, 1951; Epling &
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Mathias, 1957; Epling, 1960; Epling & Játiva, 1963, 1966, 1968;) extensively worked on the
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subgenus Calosphace and built the classification proposal that remains in use today. This
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classification often reflects groupings defined more by similarity and convenience than by
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evolutionary assumptions, and is challenging because of the high number of doubtfully
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circumscribed sections. Despite the above, the trend is to keep using it as a support until a
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more natural classification is elaborated (Standley and Williams, 1973; Walker, 2004). We
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agree with this trend.
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In the book Flora de Manantlán (Vázquez et al., 1995), H. H. Iltis and/or T. P.
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Ramamoorthy tentatively listed 6 new Salvia L. species from the Sierra de Manantlán: S.
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brucebenzii, S. cuevasiana, S. mcvaughii, S. meera, S. santanae and S. vazquezii; but the
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names were not validated. It was not stated how to recognize each taxon nor their taxonomic
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position within any of the infrageneric classifications proposed. S. brucebenzii was described
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by Turner (2008) as S. acerifolia B. L. Turner based on a specimen from Coalcomán,
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Michoacán. He probably did not know about the specimens collected from Jalisco, since he
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did not cite them. Salvia vazquezii is being described in a paper with comments about the
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resupination of its flowers (Iltis et al., submitted). Now, as part of the project Flora de Jalisco
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y Áreas Colindantes, we reexamined the Salvia specimens from the Sierra de Manantlán,
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particularly those presumably considered as new taxa. We agree that the corresponding
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populations deserve recognition as new species, except for one case: S. cuevasiana can be
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correctly referred as S. polystachia Cav., according to each of its characters, it corresponds to
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an extreme of variation where the calyces are densely pubescent. In the current paper we
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describe and illustrate the rest of the mentioned taxa, and additionally we propose the
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Manantlán populations of S. concolor Lamb. ex Benth., as a new subspecies.
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Description
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Salvia concolor subsp. iltisii J. G. González subsp. nov. Type: Mexico. Jalisco. Cuautitlán, at
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first stream crossing (and before rapid descent) of lumber road, 2 km S from San Miguel
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“meadows”, E-slope of the Sierra de Manantlán Central, 5.3 km S of Rincón de Manantlán,
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17.5 km S of El Chante, 19º33’00"N, 104º12'15"W, 2400 m, 12 Jan. 1980 (fl, fr), H. H. Iltis,
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P. Sorensen and P. Matekaitis 2623 (holotype: IBUG; isotypes: ENCB, MEXU, WIS not
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seen). Figs. 1 and 2.
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Salvia concolor Lamb. ex Benth. primo adspectu maxime simile, sed foliis tenuibus ad basin
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alte cordatis, calycibus flavovirentibus, calycum labio superiore longe caudato (cauda 3-4 mm
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longa) integro ad apicem.
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Perennial herbs or subshrubs, erect, 1-2(-3) m tall, stems lax and slightly
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inclined, sparsely pilose, mainly on the ribs and with some amber or orange sessile glandular
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dots. Petioles 7-13.2 mm long, glabrescent. Leaves ovate, 10-15 × 6-13 cm, deeply cordate at
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the base, acuminate at the apex, the margin finely serrate and sparsely bordered with straight
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simple hairs, both surfaces green, concolor, adaxial surface sparsely covered with conical tiny
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hairs, abaxial surface glabrous or with some appressed hairs spread on the veins, covered with
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coppery amber sessile glandular dots (not visible to the naked eye), chartaceous texture.
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Inflorescences on terminal racemes, 25-38 cm long, each raceme with 9 to 25 verticillasters,
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these 1.5-2.5 cm apart from each other toward the base, verticillaster 6 to 12-flowered, floral
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axis profusely pilose and with tiny glandular capitate hairs. Floral bracts ovate-lanceolate, 5-6
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× 1-2 mm, rounded or truncate at the base, long attenuate at the apex, yellowish-green,
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deciduous. Pedicels 4-6 mm long in flower, 9-12 mm long during fructification, covered with
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glandular capitate hairs longer than those present on the floral axis. Calyces 6-8 × 3-4 mm in
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flower, accrescent (10-15 × 4-6 mm in fructification), yellowish green, slightly darker on the
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dorsal line, lips acute and unequal particularly during the fructification, the upper one 3.5-5
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mm long, the lower one 2-3 mm long, both aristate at the apex, aristae of the lower lobes 1.8-
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2 mm long, those from the upper lobe 3-4 mm long during fructification, upper lip 5-veined,
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moderately covered with coppery amber sessile glandular dots, occasionally covered with tiny
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conical hairs in the inner surface toward the throat. Corollas dark blue with white nectar
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guides, sparsely pilose throughout its surface, with coppery amber sessile glandular dots
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concentrated toward the lips; tube 20-22 mm long, slightly ventricose, 4.5-5.5 mm wide at its
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widest portion, not invaginated at the base, internally naked (epapillate); the upper corolla lip
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5-6.5 mm long, the lower one 5-6(-8) mm long, 5-6 mm wide. Stamens included, filaments 2
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mm long, connectives 7.5-8.5 mm long, entire, theca 1-2 mm long; with a staminodium pair
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relatively developed behind and above the insertion point of the filaments, 1 mm long,
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globose at the apex. Horn of the gynobase 2-3 mm long; styles 25-26 mm long, abaxially and
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adaxially pilose toward the apex, the upper branch longer and exserted. Nutlets ovate, 2-2.3 ×
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1.3-1.4 mm, tan and dark brown marbled, smooth and glabrous surface.
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Taxonomic summary
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Distribution, habitat and phenology. Salvia concolor subsp. iltisii dwells into montane cloud
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forests at 2400 m altitude, with Abies religiosa (Kunth) Schltdl. & Cham., Alnus jorullensis
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Kunth, Cirsium tolucanum (B. L. Rob. & Seaton) Petr., Montanoa andersonii McVaugh and
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Tillandsia sp. It has been observed in flower and fruit only on January.
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Etymology. This new subspecies honors Hugh H. Iltis, from the University of Wisconsin-
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Madison, who has been instrumental in the establishment of the Reserva de la Biosfera Sierra
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de Manantlán (Jalisco and Colima, Mexico) and in the botanical exploration and knowledge
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of its flora.
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Additional material examined. Mexico. Jalisco. Cuautitlán: 5.3 km S of Rincón de Manantlán,
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2400 m, 12 Jan. 1980 (fl), Iltis 15202 (GUADA).
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Remarks
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Salvia concolor belongs to sect. Dusenostachys (Epling) Epling. This section is
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recognized by means of its leaves rounded or cordate at the base, 6 or more flowers per
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verticillaster, floral bracts deciduous, upper lip of the calyx with 5 to 9 veins, corolla tube
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ventricose, internally naked (epapillate) at the base, stamens included, and style pilose toward
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the apex. Epling (1939) included 9 species within this section. Later, he removed 2 of them:
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S. flaccidifolia to sect. Angulatae (Epling) Epling (Epling, 1941) and S madrensis to sect.
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Longipes Epling (Epling, 1944). Finally, it was added S. divinorum Epling & Játiva (Epling
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and Játiva, 1962). The 8 species of this group are united by the characters delineated by
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Epling; however, their general appearance and distribution (6 species growing in Mexico and
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2 restricted from Southeastern Brazil) make us doubt about its validity as a natural group.
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Specimens of S. concolor subsp. iltisii were previously identified as S.
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concolor, as reported in the book Flora de Manantlán (Vázquez et al. 1995). Nonetheless,
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even when it is evident a close morphological relationship between them, specimens from the
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Sierra de Manantlán differ in having thinner and deeply cordate leaves, floral bracts 5-6 mm
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long, calyces slightly shorter (12-15 mm in fruit), yellowish green, with the upper lobe long
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caudate (3-4 mm long) but entire (Table 1). Meanwhile, S. concolor has thicker leaves and
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with rounded or slightly cordate bases (except for some specimens from the states of Mexico,
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Morelos, and surroundings of Mexico city, which present cordate leaves: Salazar s.n.,
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(MEXU 135793), Engle and Remington 87A (MEXU), Salazar 461 (MEXU)), floral bracts 7-
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23 mm long, calyces 18-22 mm long in fruit, dark blue, with the upper lip long caudate but
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shorter (2-3 mm long) and with 2 lateral mucrones. Furthermore, S. concolor inhabits the
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central region of Mexico: Mexico, D. F., Mexico State, Eastern Michoacán and Morelos.
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Jaliscos´s populations are isolated from those of the typical species.
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KEY FOR SUBSPECIES OF SALVIA CONCOLOR
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1a
Coarse leaves, rounded at the base or slightly cordate; floral bracts 7-23 mm long;
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calyces 18-22 mm long, dark blue, with the upper lip long caudate (2-3 mm long) and
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with 2 lateral mucrones. Endemic to the Valley of Mexico, Puebla and Eastern
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Michoacán ...........................................................................S. concolor subsp. concolor
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Thin leaves, deeply cordate at the base; floral bracts 5-6 mm long; calyces 12-15 mm
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long, yellowish green, with the upper lip long caudate (3-4 mm long) and entire,
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without lateral mucrones. Endemic to the Sierra de Manantlán
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………………………………………………………………..S. concolor subsp. iltisii
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Description
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Salvia meera Ramamoorthy ex J. G. González sp. nov. Type: Mexico. Jalisco. Cuautitlán, en
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el poblado del Aserradero de Manantlán, carretera para Puerta Pesada, Sierra de Manantlán,
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1520 m, 12 Dec. 1982 (fl, fr), J. I. Calzada and G. Nieves H. 9545 (holotype: ZEA; isotype:
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XAL, not found). Figs. 2 and 3.
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Species insignis ob corollas albas, folia anguste lanceolata serrata, flores 2 in verticillastri
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dispositi; bracteas anguste lanceolatas interdum lineares 1.2-1.4 mm longas, 0.1-0.2 mm latas,
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et calycum labiis superiores 3-venis.
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Perennial herbs, erect, (0.7-)1-2.4(-3) m tall, stems with whitish appressed and
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retrorse hairs, abundantly distributed on the ribs and between them. Petioles 3-7 mm long, the
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middle portion of the adaxial surface sunken, pilose. Leaves lanceolate, 7-7.5(-8.5) × 1.4-1.6
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cm, attenuate at the base and the apex, the margin widely serrate, green in both adaxial and
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abaxial surfaces, sparsely covered with appressed hairs on the veins and orange-amber sessile
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glandular dots on both surfaces. Inflorescences arranged on terminal and subterminal axillar
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racemes, 1-4.5 cm long, each one with 2 to 4 verticillasters, 7-9 mm apart from each other
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toward the base, verticillasters 2-flowered, floral axis profusely pilose. Floral bracts narrowly
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lanceolate or almost linear, 1.2-1.4 × 0.1-0.2 mm, attenuate at the base and the apex, margin
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entire, green, densely pilose, soon deciduous. Pedicels 5-8.5 mm long, densely pilose. Calyx
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1.3-1.5 cm long, 4-6 mm wide at the throat, yellowish green, lips equal or subequal in length
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(2.5-5 mm long), the upper one 3-veined, with tiny appressed and antrorse hairs on the veins,
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between them and along the margin of the lips, internally verrucose and with tiny conical
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hairs. Corollas white, glabrous except for the upper lip and the abaxial surface of the lower
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one which are sparsely to densely pilose; tube 2-2.2 cm long, (3-)4-5 mm wide at its widest
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portion, slightly ventricose and arcuate, not invaginated at the base, internally naked
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(epapillate); upper lip 5-6 mm long, the lower one 4.5-6.2 mm long. Stamens included;
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filaments 1.5-2 mm long; connectives 11.5-12.5 mm long, with an acute tooth at the middle;
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theca 2-2.2 mm long; staminodium absent. Horn of the gynobase 0.3-0.4 mm long; styles 2.2-
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2.4 cm long, abaxially and adaxially pilose toward the apex, upper branch of the style longer
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and exserted, the lower one shorter and included. Nutlets ovate, 2-2.5 × (0.3-)9-1.1 mm long,
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pale amber or reddish amber, concolor, smooth surface, glabrous.
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Taxonomic summary
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Distribution, habitat and phenology. Salvia meera grows in pine-oak forests, from 1500-1800
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m altitude. Probably sharing habitat with Abies guatemalensis Rehder, Alnus acuminata
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Kunth, Oreopanax sanderianus Hemsl., Pinus douglasiana Martínez, Quercus candicans
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Née, Q. excelsa Liebm., Q. obtusata Humb. & Bonpl., and Q. scytophylla Liebm., according
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to records made near its location. It flowers and fructifies from late November to middle of
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December.
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Etymology. We conserve the name that was suggested in the book of Flora de Manantlán
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(Vázquez-García et al., 1995) by Ramamoorthy, who identified the labiates collected and
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reviewed for that publication. Meera is the name of a Hindu woman between historical reality
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and myth. She lived during the sixteenth century and there are several songs and poems that
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are attributed to her. Nowadays, she is still a transcendent mystical icon for the Indian people;
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even at least 2 movies exist about her life.
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Additional material examined. Mexico. Jalisco. Cuautitlán: Sierra de Minantlán
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(sic.)[Manantlán], above Haceradero (sic.)[Aserradero], 5900 ft [1800 m], 23 Nov. 1963 (fl,
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fr), Boutin and Brandt 2505 (CAS seen as photograph, MEXU, MICH seen as photograph).
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Remarks
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The characters of Salvia meera fit well between sect. Tubiflorae (Epling) Epling
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except for its striking white corollas with the lower lip as long as the upper one or longer, and
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its narrower leaves from those typical of such group. This section is integrated by shrubs or
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subshrubs with ovate leaves, acuminate at the apex and rounded or attenuate at the base, with
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6 to 12 flowers per verticillaster, floral bracts deciduous, upper lip of the calyces 3-veined or
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sometimes 5-veined, corollas magenta with the tube straight and internally naked (epapillate),
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the upper lip of the corolla as long as the lower one or longer, the connective entire or with a
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short tooth in the middle and the style pilose. The morphologically most related taxa to S.
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meera are: S. pringlei Rob. & Greenm., and S. tubifera Cav. The first one differs because of
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its ovate leaves (4-6 cm wide), inflorescences (3-)6-8(-12) cm long, 4 to 12 flowers per
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verticillaster, magenta corollas with the upper lip 1.2-1.5 cm long and the lower one 0.8-1.0
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cm long. S. tubifera can be recognized by its ovate leaves, 4-6.5 cm wide, inflorescences 7-
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14(-22), 2 to 6 flowers per verticillaster, calyces 0.6-0.9 cm long, corollas magenta with the
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tubes 1.8-2 cm long and the lower lip 0.2-0.4 cm long (table 2). Furthermore, the 3 species
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exhibit different distributional patterns: S. meera is a restricted endemic from the Sierra de
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Manantlán, Jalisco, that grows in montane cloud forests from 1500-1800 m altitude; S.
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pringlei is an endemic with a wider distribution, since it grows in Eastern Jalisco and
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Southern Nayarit from 400-750 m; finally, S. tubifera inhabits high mountains from (1900-
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)2000-2600(-3000) m in Mexico and Guatemala (table 2). The 3 species can be found in
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Jalisco but none of them grow together in the same locality.
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The corollas entirely white are unusual among the mexican Salvia species (
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Ramamoorthy, 1984), from the approximately 300 species that grows in the country, 14
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(4.67%) have entirely white corollas: S. albiflora M. Martens & Galeotti, S. assurgens Kunth,
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S. decora Epling, S. diegoae Martínez-Gordillo & Loazada-Pérez, S. divinorum Epling &
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Játiva, S. durantiflora Epling, S. leninae Epling, S. leucantha Cav., S. meera, S. perblanda
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Epling, S. pericona B. L. Turner, S. pineticola Epling, S. rzedowskii Ramamoorthy and S.
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sphacelifolia Epling. Other species like S. coccinea Buc’hoz ex Etl. (with typical red
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corollas), S. purpurea Cav. (with purple ones) and, S. helianthemifolia Kunth and S. roscida
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Epling (with sky blue ones), can exceptionally present white or very pale blue ones, almost
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white. Hence, white corollas are not highly successful for this genus, and therefore there are
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few species with them.
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These species with white corollas were assigned within different sections by
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Epling and coworkers (1939, 1940, 1941, 1944, 1947, 1951; Epling and Játiva, 1966; Epling
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and Mathias 1957), some of them are morphologically distant: Angulatae (S. albiflora),
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Albolanatae (Epling) Epling (S. leucantha), Carneae (Epling) Epling (S. pineticola),
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Dusenostachys (Epling) Epling (S. divinorum), Nivalis Epling (S. leninae), Polystachyae
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Epling (S. decora, S. durantiflora and S. perblanda), Rudes Epling (S. sphacelifolia) and
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Uliginosae (Epling) Epling (S. assurgens). Three of the species with white corollas
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recognized after Epling’s work were assigned to sect. Nivalis (S. diegoae; Martínez-Gordillo
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and Lozada-Pérez, 2011), Scorodonia (S. pericona; Turner, 2009) and Tubiflorae (S. meera);
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S. rzedowskii were not assigned to any section (Ramamoorthy, 1984). The above indirectly
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indicates, even when Epling’s classification has not proved to be natural, that Salvia
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researches have conceived white corollas as a character that has appeared multiple times in
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the evolutionary history of the subgenus Calosphace.
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Tripp and Manos (2008) postulated that white corollas can act as an
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evolutionary dead-end in Ruellia L. (Acanthaceae) according to the animal species they can
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explode as pollinators. They built a cladogram of 40-55% of the species of Ruellia, and
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mapped floral characters on it, such corolla color. The white corollas of Ruellia attract hawk
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moths and bats; they are the result of a likely irreversible lost of some pigments from species
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with red or purple ones, which are pollinated by hummingbirds and bees, respectively. This
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irreversible lost diminishes the evolutionary potential of the species, since lineages with red
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corollas are originated by lineages with purple ones in the cladrogram and vice versa, but
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lineages with white ones are caused by any of the above but can not give rise to a lineage with
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a different corolla color. It is worth noting the similarity between Ruellia and Salvia in terms
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of its high richness dominated by species with blue to purple corollas, less with red and very
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few with white or yellow ones, and the homoplasy of such character. Moreover, Coberley and
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Rausher (2008) found a probable deleterious pleiotropic effect associated with the allele
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codifying for white corollas in Ipomoea purpurea (L.) Roth. This allele consists of an
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insertion of a transposable element into the exon of the chalcone synthase D gene, which is
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responsible for beginning of the flavonoid pathway; the interruption of this pathway with the
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consequent lack of flavonoids could make the plants vulnerable to drought, UV radiation and
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herbivory. It is presumable that there is operating similar processes in Salvia that limit the
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diversification of species with white corollas.
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KEY FOR SALVIA MEERA AND ITS MORPHOLOGICALLY CLOSEST RELATIVES
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1a
Leaves lanceolate; inflorescences 1-4.5 cm long; floral bracts 1.2-1.4 mm long;
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corollas white. Endemic species from the Sierra de Manantlán Jalisco
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………………………………………………………….……………………... S. meera
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1b
Leaves ovate; inflorescences (3-)6-14(-22) cm long; floral bracts 3-11(-14) mm long;
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corollas magenta. Plants absent in the Sierra de Manantlán.
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2a
Floral bracts 3-4 mm long; calyces 6-9 mm long; upper corolla lips 6-9 mm
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long, lower ones 2-4 mm long. Plants growing in Guatemala and almost
308
throughout the Pacific slope of Mexico, from 2000-2600(-3000) m
309
………………………………………………..………………...…….S. tubifera
310
2b
Floral bracts 6-11(-14) mm long; calyces (8-)11-14 mm long; upper corolla
311
lips 12-15 mm long, lower ones 8-10 mm long. Endemic species to Jalisco
312
and Nayarit, Mexico, which grows from 400-750 m……….……….S. pringlei
313
314
315
KEY FOR THE MEXICAN NATIVE SAGES WITH ENTIRELY WHITE COROLLAS
1a
Leaves adaxially white tomentose.
2a
Petioles 2-3 cm long; leaves ovate-deltoid, cordate at the base; calyces
316
covered with minute glandular capitate hairs between eglandular
317
longer ones. Endemic from Oaxaca…………………...…….S. pericona
318
319
320
2b
Petioles 0.1-1.5 cm long; leaves oblong-lanceolate or elliptic, cuneate
or rounded at the base; calyces without glandular capitate hairs.
3a
Inflorescences in short (2-3 cm long) crowded terminal
321
racemes. Endemic from the state of Mexico
322
………..……………………………………S. rzedowskii
323
324
3a
Inflorescences in long racemes (15-40 cm long ) with
verticillasters 1-2.5 apart from each other.
13
325
4a
Calyces densely purple tomentose; corolla
326
tubes 13-14 mm long. Widespread in
327
Mexico and extensively cultivated
328
worldwide as ornamental
329
…..…………………………..S. leucantha
330
4b
Calyces sparsely pubescent, green; corolla
331
tubes 8-9 mm long. Endemic to Sinaloa
332
…………………………...S. sphacelifolia
333
334
1b
Leaves adaxially glabrous or sparsely covered with appressed or flexible curled hairs.
5a
Upper lip of the calyces 5 to 7-veined. Plants endemic to Michoacán.
335
6a
Petioles 1.5-5 cm long; corolla tubes 12-14 mm long ……S. leninae
336
6b
Petioles 0.2-0.7(-1.3) cm long; corollas tube 5.5-6.5 mm long; floral
337
axis, pedicel and calyx densely covered with glandular capitate hairs
338
……………………..……………………………………….S. assurgens
339
340
341
5b
Upper lip of the calyces 3-veined.
7a
Corolla tubes 1.1 cm or longer.
8a
cm wide …………………………….…………………..S. meera
342
343
Petioles 0.3-0.7 cm long; leaves narrowly lanceolate, 1.4-1.6
8b
Petioles (0.5-)1.5-5 cm long; leaves ovate, (2.1-)3-10 cm
344
wide.
345
9a
Leaves long attenuate at the base; calyces purple with the
346
apices of the lower lobes long caudate; corolla tubes 2-
347
2.2 cm long. Endemic from Oaxaca…..……S. divinorum
348
349
9b
Leaves truncate, rounded or subcordate at the base;
calyces green or red with the apices of the lower lobes
14
350
almost truncate or shortly acute but not long aristate,
351
corolla tubes 1.2-2.9 cm long. Endemic to Michoacán or
352
Guerrero.
353
10a
Leaves glabrous at the upper surface, rounded or
354
subcordate at the base, verticillasters 2-flowered,
355
calyces red, corolla tube 2.4-2.9 cm long,
356
internally epapillate, upper lip 1.2-1.3 cm long
357
and lower one 6.5-10 mm long. Endemic to
358
Guerrero.................................................S. diegoae
359
10b
Leaves covered with long hairse at the upper
360
surface, rounded or truncate at the base,
361
verticillasters 6 to 12-flowered, calyces green and
362
sometimes tingled with purple, corolla tube 1-2
363
cm long, internally ornamented with 4 papillae,
364
upper and lower lips 8-9 mm long. Endemic to
365
Michoacán……………………………..S. leninae
366
367
7b
Corolla tubes shorter than 1 cm long.
11a
Leaves cuneate at the base; corolla tubes internally naked at its
368
base. Plants mainly from the Pacific slope, from Veracruz to
369
Chiapas………………………………………….…...S. albiflora
370
11b
Leaves rounded or truncate at the base; corolla tubes internally
371
ornamented with 1 or 2 papillae pairs at its base.
372
12a
373
Lobes of the calyces connivents at fructification. Plants
from Guerrero and Oaxaca …..…………..S. durantiflora
15
374
12b
Lobes of the calyces straight and separated at
375
fructification.
376
13a
Calyces 0.3-0.4 cm long; corolla tubes 4-5 mm
377
long. Plants from the Pacific slope, from Nayarit
378
to Oaxaca………………………….…....S. decora
379
12b
Calyces 0.5-0.7 mm long; corolla tubes 0.8-1.0
380
cm long.
381
13a
Petioles 5-7 cm long; racemes 5-15 cm
382
long. Endemic to Veracruz
383
………………………..……..S. pineticola
384
13b
Petioles 0.5-1 cm long; racemes 3-5 cm
385
long. Endemic to Guerrero
386
……………………………..S. perblanda
387
388
Description
389
Salvia rogersiana Ramamoorthy ex J. G. González sp. nov. Type: Mexico. Jalisco, Autlán de
390
Navarro, Cerro Las Juntas, Estación Científica Las Joyas, 3 Aug. 1986 (fl), R. Cuevas G.
391
1471 (holotype: ZEA). Figs. 2 and 4.
392
Salviae sectione Briquetia Epling adscribenda, ex species sectionis inflorescentis compactis et
393
brevibus, floribus 2 in verticillastris dispositi, calycibus 10 mm longis vel brevibus, calycum
394
labiis superioribus truncatis ad apicem, corollarum tubis 15-17 mm longis distinguenda.
395
Perennial herbs, erect, (0.5-)0.8-1(-1.5) m tall, stems sparsely to densely pilose
396
on the ribs and between them, mainly on young stems. Petioles (1.4-)1.8-3(-4.5) cm long,
397
sparsely to densely pilose. Leaves ovate or ovate-elliptic, (5-)7-9(-10) × (3.5-)4.5-6.5 cm,
398
rounded and shortly cuneate at the base, acuminate at the apex, margin serrate, sparsely
16
399
covered with appressed hairs on both surfaces. Inflorescences arranged on terminal racemes,
400
(5-)7-12(-17) cm long, 7 to 17 verticillasters per floral axis, 6-12(-15) mm apart from each
401
other toward the base, verticillasters 2-flowered, floral axis densely pilose. Floral bracts ovate,
402
4-8(-9.7) × (2-)3-5 mm, truncate at the base, slightly rounded and abruptly caudate at the
403
apex, margin entire, foliose texture, densely covered with long flexible hairs on the outer
404
surface and along the margin, the inner surface glabrous, the hairs eglandular, pluricellular
405
and uniseriate with red septa that give its yellowish-red appearance, deciduous. Pedicels 2-3.5
406
mm long, thickly pilose. Calyces 7-9.5 × (4-)5-6 mm, yellowish green, concolor, upper lip
407
acute and bent upward, the apex with a truncate appearance, lower lip truncate and with a
408
terminal tiny tooth on each lobe (up to 0.2 mm long), upper lip 3-veined, densely covered
409
with long flexible hairs, inner surface covered with short conical hairs, mainly toward the
410
apex. Corollas dark blue to purple, glabrous except the upper lip which is pilose and in a
411
lesser extent the abaxial surface of the lower lip; tube 15-18.3 long, 5.5-6 mm wide in its
412
widest portion, arcuate, widely ventricose just after the lower lip, straight at the base,
413
internally naked (epapillate); upper lip 5-6.7 mm long, lower lip (4.5-)6-8.6 × 8-8.4 mm.
414
Stamens included; filaments (1.7-)2.3-3.3 mm long; connectives 7.8-13 mm long, non
415
geniculate but with a short ventral tooth; theca 1.4-2 mm long; a pair of filiform staminodium
416
present behind and above the insertion point of the filaments. Horn of the gynobase 0.5 mm
417
long; styles 2.1-2.2 cm long, abaxially and adaxially pilose toward the apex, included except
418
for the upper branch of the style, both branches purple, lower branch acute; horn of the
419
gynobase quadrangular, relatively short (up to 1 mm long), truncate at the apex. Nutlets ovate,
420
1.5-1.6 × 7-1.4 mm, reddish brown and dark brown marbled (concolor when immature),
421
smooth and glabrous (with tiny appressed hairs on immature ones).
422
Distribution, habitat and phenology. Salvia rogersiana grows into montane cloud forests and
423
oak forests within shaded humid ravines, but can grow along dirt roads, from (1100-)1800-
17
424
1900 m altitude. It shares habitat with Alnus jorullensis, Aphananthe monoica (Hemsl.) J.-F.
425
Leroy, Cestrum confertiflorum Schltdl., Clusia salvinii Donn. Sm., Citharexylum mocinnoi D.
426
Don., Juglans major (Torr.) A. Heller, Otatea acuminate (Munro) C. E. Calderón & Soderstr.,
427
Prunus serotina Ehrh. subsp. capuli (Cav.) McVaugh, Symplococarpon purpusii (Brandegee)
428
Kobuski, Synardisia venosa (Mast.) Lundell, Tilia americana L. var. mexicana Schltdl.,
429
Trophis racemosa (L.) Urb., Bursera sp, Ficus sp, Trichilia sp, and Viguiera sp. This species
430
blooms and fructifies from middle June to December.
431
Taxonomic summary
432
Etymology. This species corresponds to what is registered as S. mcvaughii in Flora de
433
Manantlán (Vázquez et al., 1995). However, we did not retain that name because it was
434
already applied for another new species which belongs to sect. Polystachyae Epling (Bedolla-
435
García et al., 2011), and is not morphologically related with the proposed new taxon. Thus,
436
we chose a new specific epithet that still makes reference to Rogers McVaugh, an outstanding
437
botanist who contributed greatly in the study of the Western Mexican flora, and whose most
438
memorable work, Flora Novo-Galiciana (McVaugh 1961, 1974, 1983, 1984, 1985, 1987,
439
1989, 1992, 1993, 1995, 2001), is a crucial reference for any Mexican botanist.
440
Additional material examined. Mexico. Jalisco. Autlán de Navarro: Las Galeras, Estación
441
Científica Las Joyas, 19º37'03"N, 104º16'28"O, 1600-1650 m, 3 Dec. 1996 (fl), Cuevas and
442
Sánchez 5452a (IBUG, ZEA); 500-600 m de Corralitos por la brecha que sube a la Estación
443
Científica Las Joyas, en pared de exposición NO, 19º37'37.4"N, 104º19'16.3"O, 1780 m, 30
444
Oct. 2010 (fl), González et al. 781 (IBUG, IEB, ZEA, MEXU); parte alta de la cañada del
445
Alentrisco, 600 m al S de Corralitos, 1800-1900 m, 19 Jun. 1996 (fl), Sánchez and Cuevas
446
118 (IBUG, ZEA); 11-12 km al SSE de Autlán, 2 km al S de Ahuacapán, 19º39'31"N,
447
104º19'14"O, 1100 m, 24 Jul. 1988 (fl, fr), Santana and De Niz 3669 (ZEA). Talpa de
18
448
Allende: Triunfo, 19 km WSW of Talpa de Allende, along road to La Cuesta and Tomatlán,
449
1555 m, 11 Sep. 1986 (fl), Breedlove and Anderson 64166 (MEXU).
450
Remarks
451
The characters of Salvia rogersiana match those of sect. Briquetia: ovate to ovate
452
lanceolate wide leaves, rounded to attenuate, sometimes truncate or cordate at the base, lips of
453
the calyx subequal in length, the upper one 3-veined, corolla dark blue or purple, tube
454
ventricose and mostly invaginated, internally epapillate, upper corolla lip longer than the
455
lower one, stamens included, connective with an small ventral tooth or subentire, and style
456
pilose; although, it does not seem to be closely related to any of the species of that section. It
457
is distinctive by means of its relatively short and compact inflorescence, 2-flowered
458
verticillasters, calyx 10 mm long or shorter, upper lip truncate and with a tiny terminal tooth,
459
corolla tube 15-18.3 mm long with the upper lip 5-6.7 mm long, the lower one (4.5-)6-8.6 mm
460
long (table 3). None of the other species within the section grows in the same region than S.
461
rogersiana except for S. mexicana; however they have not been found in the same locality.
462
KEY FOR SALVIA MEXICANA AND SALVIA ROGERSIANA
463
1a
Petioles 1-10 cm long, leaves 6-18(-20) cm long; floral bracts 6-12 mm long; calyx
464
18-17(-20) mm long; upper lip of the corolla 13-14(-19) mm long, lower one 12
465
mm long…………………………………………………………………….S. mexicana
466
1b
Petioles 1.8-2.7 cm long, leaves (6-)7-9(-10) cm long; floral bracts 4-5.2 mm long;
467
calyx 7-9.5 mm long; upper lip of the corolla 5-6.5 mm long, lower one
468
(4.5-)6-6.6 mm long ………….…………………………...……………..S. rogersiana
469
470
Description
471
Salvia santanae Ramamoorthy ex J. G. González sp. nov. Type: Mexico. Jalisco. Tolimán, 1-
472
1.5 km al N de El Terrero, 12-13 km al NE de Minatitlán, 19º27'13"N, 103º56'56"O, 2300 m,
19
473
4 Sep. 1990 (fl), L. Guzmán H. and R. Cuevas G. 1091 (holotype: ZEA; isotype: WIS). Figs.
474
2 and 5.
475
Species habitu cum Salvia longispicata M. Martens & Galeotti aemulans optime congruens,
476
sed differt floribus 2-4 in verticillastris dispositi bracteis persistentibus, calycibus 7-9 mm
477
longis et corollarum tubis (9-)1-15 mm longis (vs. flores 6-18 in verticillastris dispositi
478
bracteas caducas, calyces 5-6.5 mm longos, corollarum tubos 5-8 mm longos).
479
Perennial herbs, erect (0.6-)1-1.5 m tall, stems with flexible retrorse hairs between
480
the ribs, puberulent on the ribs. Petioles 3-4.5(-9) cm long, thin, sparsely covered with tiny
481
appressed hairs. Leaves ovate-elliptic, (6-)10.5-13.5 × (3.5-)6-8.5 cm, long cuneate at the
482
base, acuminate at the apex, the margin serrate (the teeth regularly 3-5 mm wide at the base),
483
green, both surfaces barely covered with appressed hairs, concentrated mainly on the veins.
484
Inflorescences an axillar or terminal racemes, (10-)15-20 cm long, each one with 11 to 15
485
verticillasters, these spaced 6-10 mm toward the base, verticillasters 2 or 4-flowered (rarely 5-
486
flowered), floral axis glabrous. Floral bracts narrowly lanceolate, (1.8-)2-4 × 0.5-1.3 mm,
487
green, truncate at the base, aristate at the apex, the margin entire, foliose texture, glabrescent,
488
persistent. Pedicels 2.5-4 mm long, covered with flexible retrorse hairs and light amber sessile
489
glandular dots, the hairs pluricellular and uniseriate with purple septa. Calyces 7-9 mm long,
490
3-4 mm at the throat, the dorsal portion dark green and the ventral one yellowish green, both
491
lips 2-3(-5) mm long, acute and shortly aristate, upper lip 3-veined, veins with the same
492
pubescence than the pedicel and glandular punctate between them, with a scaly appearance
493
due to a thickened at the insertion point of the hairs and its dark purple color. Corollas dark
494
blue or purple with white nectar guides on the lower lip, the upper lip densely pilose, mainly
495
at the ventral surface, the lower lip abaxially pilose, the rest glabrous; tube (9-)1-15 mm long,
496
2-3 mm wide at its widest portion, ventricose and slightly invaginated at the base, internally
497
naked (epapillate); upper lip 6-7 mm long, lower lip equal or subequal in length and 8-8.5 mm
20
498
wide. Stamens included; filaments 2-2.5 mm long; connectives 9-9.5 mm long, straight and
499
with an acute tooth bending backwards at the middle portion; thecae 1.1-1.2 mm long; a pair
500
of staminodium present behind and above the attachment of the filaments to the corolla. Horn
501
of the gynobase 0.5-0.7 mm long; styles 15-16 mm long, abaxially and adaxially pilose (hairs
502
with purple septa), branches purple and exserted. Nutlets ovate, 1.3-1.4 × 0.7-0.8 mm, light
503
brown, almost concolor, smooth and glabrous.
504
Taxonomic summary
505
Distribution, habitat and phenology. Salvia santanae inhabits montane cloud forests and oak
506
forests from (1800-)2100-2300 m, in the karstic topography of Cerro Grande calcareous
507
massif. It grows together with Clethra fragrans L. González & R. Ramírez, Dendropanax
508
arboreus (L.) Decne. & Planch., Oreopanax peltatus Linden, O. xalapensis (Kunth) Decne. &
509
Planch., Quercus candicans Née, Q. castanea Née, Q. crassipes Humb. & Bonpl., Q. laurina
510
Bonpl, Q. obtusata Bonpl., Symplocos citrea Lex. ex La Llave & Lex., Styrax ramirezii
511
Greenm, and Ternstroemia lineata DC. It flowers and fructifies from September to March.
512
Etymology. We conserve the name suggested by Ramamoorthy in the Flora de Manantlán
513
(Vázquez et al., 1995) for this new species, and we are agree to name it in honor to Francisco
514
J. Santana Michel from the Instituto Manantlán de Ecología y Conservación de la
515
Biodiversidad, Universidad de Guadalajara, Mexico, who has contributed continuously to the
516
knowledge of the flora of Jalisco and Colima (Mexico), and of some specific groups like
517
grasses and the genus Aristolochia Juss.
518
Additional material examined. Mexico. Jalisco. Tolimán: 9 km al NE de Minatitlán, 2-4 km al
519
SO de El Terrero, 19º26'00"N, 103º58'38"O, 1750-1900 m, 13 Oct. 1988 (fl, fr), Cuevas and
520
López 3323 (ZEA, WIS); 17-18 km al NNE de Minatitlán, 2 km al S de La Laguna,
521
19º31'28"N, 103º59'00"O, 2100-2200 m, 18 Dec. 1988 (fl, im fr), Cuevas and Núñez 3463
522
(ZEA); Cerro Grande, 19°25’19"N,-103°57’01"O, 1806 m, 7 Dec. 2008 (fl), González and
21
523
Vázquez 231 (IBUG, MEXU); 12-13 km al ENE de Minatitlán, 2 km al NO de El Terrero,
524
19º26'57"N, 103º57'00"O, 2300 m, 16 Mar. 1993 (im fr), Muñoz and Vázquez 35 (ZEA,
525
WIS); El Terrero, 2200 m, 25 Nov. 1992 (im fr), Navarrete 312 (ZEA, WIS); La Ciprecera, 2
526
km al NO de El Terrero, 12-13 km al ENE de Minatitlán, 19º26'57"N, 103º57'00"O, 2280 m,
527
19 Nov. 1993 (fl, im fr), Santana et al. 6286 (ZEA).
528
Remarks
529
The morphology of Salvia santanae is closely related with S. longispicata M.
530
Martens & Galeotti. The shape and size of the vegetative organs, the shape and distribution of
531
pubescence of the floral structures are overlapped between them. However, it differs from the
532
later in having 2 or 4-flowered verticillasters (rarely 10-flowered), persistent floral bracts, and
533
longer floral characters (calyx 7-9 mm long, tube (9-)10-15 mm long, with the upper lip 6-7
534
mm long and style 15-16 mm long) (table 4). Meanwhile, S. longispicata presents usually 10-
535
flowered verticillasters (this may be variable, from 6 to 18-flowered), deciduous floral bracts,
536
and smaller floral characters (calyx 5-6.5 mm and corolla tube 5-8 mm long, with the upper
537
lip (3-)4.5-5 mm long, and the style 8-9(-10) mm long). The denser and more compacted
538
inflorescence of S. longispicata allow us to identify it quickly. They also differ in their
539
ecology: S. santanae grows preferably into montane cloud forests and to a lesser extent into
540
oak forests, above 1800 m; while, S. longispicata inhabits tropical deciduous forest or
541
secondary vegetation along roadsides and dirty roads, it can be rarely found into oak forests or
542
oak-pine forests, generally from 1000-1900 m. Finally, while S. santanae is endemic to Cerro
543
Grande, Jalisco, S. longispicata is one of the widest distributed Salvia species in Mexico and,
544
it can be found in almost all the Mexican states except for those from the California and
545
Yucatán Peninsulas.
546
We have observed an interesting peculiarity in the morphology between S.
547
santanae and S. longispicata: the purple thickened bases at the insertion point of the hairs on
22
548
the calyces, a peculiar character of S. santanae, is also present in some populations of S.
549
longispicata, particularly from those who inhabit the Eastern face of Sierra de Coalcomán
550
(Michoacán), down from Paso Malo to Aguililla, in front of the Balsas Depression.
551
We propose S. santanae as a member of sect. Angulatae, it is distinct from the
552
other species included in that section by its persistent floral bracts and corolla tubes 10 mm or
553
longer.
554
555
KEY FOR SALVIA LONGISPICATA AND SALVIA SANTANAE
1a
Verticillasters 6 to 12 (or rarely 18) flowered, deciduous floral bracts; calyces 5-6.5
556
mm long, corolla tube 5-8 mm long, with the upper lip (3-)4.5-5 mm long; style 8-10
557
mm long………………………………….………………………………S. longispicata
558
1b
Verticillasters 2 or 4 (rarely 10) flowered, persistent floral bracts; calyces 7-9 mm
559
long, corolla tube (9-)10-15 mm long, with the upper lip 6-7 mm long; style 15-16
560
mm long…………………………...……….………………………………..S. santanae
561
562
563
Acknowledgments
We thank the curators and colleagues from the herbaria consulted: ENCB,
564
GUADA, IBUG, IEB, MEXU, ZEA and XAL; and the following herbaria for providing free
565
online photographs of specific specimens we asked for: CAS and MICH, especially to
566
Richard Rabeler, Senior Collection Manager, and Heather Huggins, Imaging Specialist, both
567
from MICH herbarium, and M. A. Wetter, Senior Academic Curator from WIS herbarium.
568
We thank Servando Carvajal from Herbario Luz María Villarreal de Puga, Instituto de
569
Botánica, Universidad de Guadalajara-CUCBA for comments about the latin diagnoses. We
570
are grateful to Oswaldo Zuno-Delgadillo for the 2 illustrations provided (Salvia concolor
571
subsp. iltisii and S. santanae). Finally, we thank the anonymous reviewers who kindly
572
provided us their recommendations. Financial support was provided by CONACYT,
573
COECYTJAL and the Universidad de Guadalajara, Mexico.
23
574
575
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576
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592
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602
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603
McVaugh, R. 1974. Fagaceae. Flora Novo-Galiciana. Contributions from the University of
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McVaugh, R. 1983. Gramineae. Flora Novo-Galiciana 14. The University of Michigan Press,
Ann Arbor.
McVaugh, R. 1984. Compositae. Flora Novo-Galiciana 12. The University of Michigan Press,
Ann Arbor.
McVaugh, R. 1985. Orchidaceae. Flora Novo-Galiciana 16. The University of Michigan
Press, Ann Arbor.
McVaugh, R. 1987. Leguminosae. Flora Novo-Galiciana 5. The University of Michigan
Press, Ann Arbor.
McVaugh, R. 1989. Bromeliaceae to Dioscoreaceae. Flora Novo-Galiciana 15. The
University of Michigan Herbarium, Ann Arbor.
McVaugh, R. 1992. Gymnosperms and Pteridophytes. Flora Novo-Galiciana 17. The
University of Michigan Herbarium, Ann Arbor.
McVaugh, R. 1993. Limnocharitaceae to Typhaceae. Flora Novo-Galiciana 13. The
University of Michigan Herbarium, Ann Arbor.
McVaugh, R. 1995. Euphorbiacearum sertum Novo-Galicianarum. The University of
Michigan Herbarium 20: 173-215.
McVaugh, R. 2001. Ochnaceae to Loasaceae. Flora Novo-Galiciana 3. The University of
Michigan Herbarium, Ann Arbor.
25
623
624
Ramamoorthy, 1984. Notes on Salvia (Labiatae) in Mexico, with three new species. Journal
of the Arnold Arboretum 65: 135-143.
625
Standley, P C. and L. O. Williams. 1973. Labiatae. Flora of Guatemala 24: 237-317.
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Tripp, E. A. and P. S. Manos. 2008. Is floral specialization an evolutionary dead-end?
627
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629
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631
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Walker, J. B., K. J. Sytsma, J. Treulein and M. Wink. 2004. Salvia (Lamiaceae) is not
633
monophyletic: Implications for systematic, radiation, and ecological specializations of
634
Salvia and tribe Mentheae. American Journal of Botany: 921: 1115-1125.
635
Walker, J. B. and K. J. Sytsma. 2007. Staminal evolution in the genus Salvia (Lamiaceae):
636
Molecular phylogenetic evidence for multiple origins of staminal lever. Annals of
637
Botany 100: 375-391.
638
639
640
641
642
643
644
645
646
647
26
648
Tables
649
Table 1. Comparison of morphological and distributional characteristics of Salvia concolor
650
and S. concolor subsp. iltisii.
Character
S. concolor subsp. iltisii
S. concolor subsp.
concolor
Petiole length (cm)
7-13.2
3-8(-12)
10-15 × 6-13
5-13(-20) × 3-12
Inflorescence length (cm)
25-38
14-30
Number of flowers per verticillaster
6-12
6-12
Floral bract length (mm)
7-23
5-6
Calyx length in fruit (mm)
10-15
18-22
Yellowish-green
Blue to purple
Acute and long caudate
Acute, long caudate and
(cauda 3-4 mm long)
with 2 lateral mucrones
Leaf dimensions (cm)
Calyx color
Shape of the upper lip of the calyces
(cauda 2-3 mm long)
Corolla tube length (mm)
20-22
20-25(-32)
Upper corolla lip length (mm)
5-6.5
4-7
Lower corolla lip length (mm)
5-8
(6-)8-10
2400-2500
2650-3300
Montane cloud forests
Montane cloud and pine
Altitudinal range (m)
Habitat
forests
Distribution
Endemic from Sierra de
Mexico D. F., State of
Manantlán, Jalisco
Mexico, Puebla, and
Eastern Michoacán
651
27
652
Table 2. Character comparison between Salvia meera, S. pringlei and S. tubifera.
Character
S. meera
S. pringlei
S. tubifera
Leaf shape
Lanceolate
Ovate
Ovate
1.4-1.6
4-6
4-6.5
1-4.5
(3-)6-8(-12)
7-14(-22)
Flowers per verticillaster
2
4-12
2-6
Floral bract length (mm)
1.2-1.4
6-11(-14)
3-4
Calyx length (mm)
13-15
(8-)11-14
6-9
Corolla color
White
Magenta
Magenta
Corolla tube length
20-22
20-22
18-20
0.5-0.6
1.2-1.5
0.4-0.6
0.45-0.62
0.8-1.0
0.2-0.4
1500-1800
400-750
(1900-)2000-
Leaf width (cm)
Inflorescence length
(cm)
(mm)
Upper corolla lip length
(cm)
Lower corolla lip length
(cm)
Altitudinal range (m)
2600(-3000)
Habitat
Pine-oak forests
Tropical oak forests
Montane cloud
and tropical
forests
deciduous forests
Distribution
Endemic from the
Endemic from
Sparsely
Sierra de Manantlán,
Eastern Jalisco and
distributed in
28
Jalisco, México
Southern Nayarit,
Guatemala and
Mexico
almost throughout
Mexico
653
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
669
670
671
672
673
674
29
675
Table 3. Character comparison between Salvia rogersiana and S. mexicana.
Character
S. rogersiana
S. mexicana
(1.4-)1.8-3(-4.5)
7-13.2
(5-)7-9(-10) × (3.5-)4.5-6.5
10-15×6-13
(5-)7-12(-17)
30-50
2
10-12
4-8(9.7)
6-12(-20)
7-9.5
8-17(-20)
Truncate (the upper lip bent
Acute (lips straight
backward)
forward)
15-18.3
15-25
Upper corolla lip length (mm)
5-6.7
13-14(19)
Lower corolla lip length (mm)
(4.5-)6-8.6
12-16
(1100-)1800-1900
(1300-)1700-2600(-2900)
Montane cloud and oak
Pine-oak and oak forests
Petiole length (cm)
Leaf dimensions (cm)
Inflorescence length (cm)
Number of flowers per
verticillaster
Floral bract length (mm)
Calyx length (mm)
Aspect of the calyx at its apex
Corolla tube length (mm)
Altitudinal range (m)
Habitat
forests
Distribution
Endemic from Western
Widespread in Mexico,
Jalisco
almost in all the states
except for those from the
California and Yucatán
Peninsulas
676
30
677
Table 4. Character comparison between Salvia santanae and S. longispicata
Character
S. santanae
S. longispicata
Petiole length (cm)
3-4.5(-9)
(1-)2-4(-4.5)
(6-)10.5-13.5 × (3.5-) 6-8.5
4-8(-11) × 3-8
Inflorescence length (cm)
10-15(-20)
15-30
Flowers per verticillaster
2-4(-10)
6-12(-18)
(1.8-)2-4 × 0.5-1.3
(2-)4-7 × (1-)2-3
Persistent
Mostly deciduous
7-9
5-6.5
(9-)10-15
5-8
6-7
(3-)4.5-5
15-16
8-9(-10)
(1800-)2100-2300
(700-)1000-1900(-3400)
Montane cloud and oak forests
Tropical deciduous forests
Leaf dimensions
(cm)
Floral bract dimensions (cm)
Floral bract duration
Calyx length (mm)
Corolla tube length (mm)
Upper corolla lip length
(mm)
Style length (mm)
Altitudinal range (m)
Habitat
and secondary vegetation
Distribution
Endemic from the Sierra de
Widespread in Mexico,
Manantlán, Jalisco, México
almost in all the states except
for those from the California
and Yucatán Peninsulas
678
679
680
681
31
682
Figure Legends
683
684
Figure 1. Salvia concolor subsp. iltisii J. G. González. A. General aspect; B. Comparison
685
between the apex of the calyces of S. concolor subsp. concolor and S. concolor subsp. iltisii;
686
upper view; C. Calyx, lateral view; D. Corolla; E. Corolla tube dissection showing the
687
filament and staminodium; F. Stamens; G. Style; H. Nutlets (drawn from the holotype).
688
689
Figure 2. Distribution of the 4 new taxa: Salvia concolor subsp. iltisii J. G. González (stars),
690
S. meera Ramamoorthy ex J. G. González (dots), S. rogersiana Ramamoorthy ex J. G.
691
González (diamonds) and S. santanae Ramamoorthy ex J. G. González (crosses). The shaded
692
area corresponds to the polygon of the Sierra de Manantlán Biosphere Reserve.
693
694
Figure 3. Salvia meera Ramamoorthy ex J. G. González. A. General aspect; B. Floral bract,
695
outer surface; C. Nutlets; D. Calyx; E. Corolla; F. Stamens; G. Style (drawn from the
696
holotype).
697
698
Figure 4. Salvia rogersiana Ramamoorthy ex J. G. González. A. General aspect; B. Floral
699
bract, outer surface; C. Calyx; D. Corolla; E. Corolla tube dissection showing the filament
700
and staminodium; F. Stamens; G. Style; H. Nutlets (drawn from the holotype and González et
701
al. 781).
702
703
Figure 5. Salvia santanae Ramamoorthy ex J. G. González. A. General aspect; B. Leaves; C.
704
Floral node; D. Floral bract; E. Calyx; F. Corolla tube dissection showing the filament and
705
staminodium; G. Stamens; H. Apex of the style; I. Nutlets (drawn from the holotype).
32