Scutellaria cuevasiana and Scutellaria sublitoralis (Lamiaceae), two new species from Jalisco and Nayarit, Mexico

Scutellaria cuevasiana y Scutellaria sublitoralis (Lamiaceae), dos especies nuevas de Jalisco y Nayarit, México

Jesús Guadalupe González-Gallegos* and José Antonio Vázquez-García

Herbario del Instituto de Botánica, de la Universidad de Guadalajara, "Luz María Villarreal de Puga" (IBUG), Departamento de Botánica y Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Km 15.5 carretera Guadalajara-Nogales, Las Agujas, Nextipac, 45110 Zapopan, Jalisco, México. *xanergo@hotmail.com

Abstract

Scutellaria cuevasiana J. G. González et A. Vázquez and Scutellaria sublitoralis J. G. González are described and illustrated. Scutellaria cuevasiana fits best within section Uliginosae (Epling) Epling. It is morphologically close to S. blepharophylla Epling, S. rosei Fernald and S. seleriana Loes. It differs from them by its magenta corollas, small and sessile floral bracts, longer petioles and bigger leaves. Scutellaria sublitoralis does not manifest any clear morphological affinity with species of the sections enumerated by Epling. It could be related to species in sections Pallidiflorae Epling or Uliginosae, particularly with the "Scutellaria ovata" or "Scutellaria caerulea" species groups defined by Paton. It is morphologically similar to S. pallidiflora Epling; however, it can be recognized by wider leaves with usually cordate to subcordate bases, persistent floral bracts, floral axis and calyces without glandular-capitate hairs, longer corollas, and narrower toward the throat.

Key words: Scutellaria caerulea group, Scutellaria ovata group, section Pallidiflorae, section Uliginosae.

Resumen

Se describen e ilustran Scutellaria cuevasiana J. G. González et A. Vázquez y Scutellaria sublitoralis J. G. González. Scutellaria cuevasiana se ajusta mejor a la sección Uliginosae (Epling) Epling; es cercana en su morfología a S. blepharophylla Epling, S. rosei Fernald y S. seleriana Loes. Difiere de las mismas debido a sus corolas magenta, brácteas florales más pequeñas y sésiles, peciolos más largos y hojas más grandes. Scutellaria sublitoralis no muestra una clara afinidad con las especies de las secciones establecidas por Epling; podría estar relacionada con las especies de las secciones Pallidiflorae Epling o Uliginosae, particularmente con los grupos de especies de "Scutellaria ovata" y "Scutellaria caerulea" definidos por Paton. Es morfológicamente similar a S. pallidiflora Epling; sin embargo, puede reconocerse por sus hojas más anchas con bases por lo usual cordadas a subcordadas, brácteas florales persistentes, eje floral y cálices sin tricomas capitado-glandulares, corolas más largas, y estrechas hacia el ápice.

Palabras clave: grupo de Scutellaria caerulea, grupo de Scutellaria ovata, sección Pallidiflorae, sección Uliginosae.

Introduction

The genus Scutellaria includes approximately 360 species, and exhibits a subcosmopolitan distribution: absent or scarcely represented in the arctic, lowland tropical areas, deserts, southern Africa and the Pacific Islands; and present on temperate mountainous areas from the tropics and southern hemisphere, with Central Asia and China as the richest region (Paton, 1990a, 1990b).

The morphology of Mesoamerican species of genus Scutellaria was well synthesized by Pool (2006), her thorough descriptions may be also applicable for the Mexican species. In Mexico, Scutellaria is represented by 34 to 45 species (Ramamoorthy and Elliott, 1998; Domínguez et al., 2003; Villaseñor, 2004). The Mexican species of Scutellaria grow mainly in temperate mountainous areas (Ramamoorthy and Elliott, 1998).

A revision of the Lamiaceae family in the state of Jalisco, western Mexico, as part of the "Flora de Jalisco y Áreas Colindantes" project, brought us to the discovery of 2 forms of the genus Scutellaria L. (Lamiaceae), that do not belong to any of the currently recognized taxa. They are described and illustrated here as 2 new species morphologically similar to sections Pallidiflorae Epling and Uliginosae (Epling) Epling.

Descriptions

Scutellaria cuevasiana J. G. González et A. Vázquez sp. nov.

Figure 1.

Type: Villa Purificación, Villas de Cacoma, 19°49'15" N, 104°33'18" W, 1 607 m, 30 Apr 2010 (fl.), J. L. Rodríguez, J. G. Morales and L. G. Medina 263 (holotype: ZEA; isotypes: IBUG, IEB, MEXU).

Perennial herbs (17-) 30-60 cm tall; erect; roots fasciculate; stems cylindrical or subquadrangular, moderately to densely covered by tiny appressed and retrorse hairs (also present on the floral axis, where occasionally are erect). Petioles (1.5-) 3-3.5 cm long, densely covered with retrorse and erect hairs. Blades variable, lanceolate, ovate-lanceolate to rhomboid, often asymmetric, (3-)5-7.5(-9.2) cm long, (2-)3.5-4.5(-6) cm wide, acute to acuminate at apex, rounded and abrupt and shortly cuneate, or cuneate at base, margin broad and irregularly crenate or sometimes serrate with simple hairs along its border, adaxial surface dark green with some appressed simple hairs, abaxial surface paler with appressed simple hairs only on the veins. Inflorescences in racemes, terminal and lax, 5-6 cm long; flowers alternate or subopposite (sometimes opposite), spirally arranged, nodes 4-6 mm apart at the inflorescence base. Floral bracts lanceolate to ovate, 1-2.7(-5) mm long, 0.2-0.8(-2.1) mm wide, persistent or late deciduous, glabrous or with some tiny hairs, mainly along the margin, rounded to acuminate at apex, truncate at base, margin entire. Pedicels 1.4-2.5 mm long, 3-5.1 mm long in fruit, moderately covered with tiny antrorse hairs. Calyces 2-2.3 mm long, 1.5-2 mm wide at throat (6 mm long, 3.5 mm wide in fruit), pale green, with appressed hairs on the middle vein, the scutellum rib and the base near the pedicel, sparsely covered with tiny translucent glandular dots, the margin of the lips reddish; scutellum 0.3-0.6 mm tall in flower, 3.8-4.5 mm tall in fruit, green with the dorsal margin reddish. Corollas magenta or rarely white; tube 1.9-2.1 cm long, straight or slightly bent basally at 3.7-4 mm, then again straight (1.1-1.2 mm wide), and gradually expanding to 3.2-3.5 mm wide at throat, glabrous inside, sparsely covered with erect tiny hairs outside; upper lip 2.7-3 mm long, lower lip 4-5 mm long, with glandular-capitate hairs along the margin of the lips, mainly on the upper one. Filaments of the upper stamens (3.5-) 4.5-5.2 mm long, filament of the lower stamens 6-6.5 mm long; anthers 1 mm long, white pilose on the slits. Style 2.1-2.2 cm long, glabrous. Nutlets subreniform to piriform, (1-)1.6-1.8 mm long, 1 mm wide, black, surface sparsely spinulose, glabrous.

Taxonomic summary

Distribution, habitat and phenology. Scutellaria cuevasiana is endemic to Jalisco (Fig. 2). It ranges between (1 000-) 1 400-1 800 (-2 090) m. It grows in montane cloud, pine-oak and riparian forests. It shares habitat with Quercus laeta Liebm., Q. resinosa Liebm., Clethra rosei Britton, C. hartwegii Benth., Eugenia culminicola McVaugh, Alnus jorullensis Kunth, Symplocos novogaliciana L. M. González, Magnolia iltisiana A. Vázquez, M. pacifica A. Vázquez, Cedrela odorata L., Ardisia revoluta Kunth, Croton wilburi McVaugh, Euphorbia aff. palmeri Engelm. ex S. Watson, Sideroxylon cartilagineum (Cronquist) T. D. Penn., Trophis racemosa (L.) Urb., Oreopanax peltatus Linden, O. echinops (Cham. et Schltdl.) Decne. et Planch., Ficus velutina Humb. et Bonpl. ex Willd., Fraxinus uhdei (Wenz.) Lingelsh., Clusia salvinii Donn. Sm., Sipatuna thecaphora (Poepp. et Endl.) A. DC., Conostegia sp., Parathesis sp. and Leucaena sp. This plant blooms and fructifies from August to middle December.

Etymology. Scutellaria cuevasiana is named in honor of Ramón Cuevas-Guzmán, from the Instituto Manantlán de Ecología y Conservación de la Biodiversidad, Universidad de Guadalajara, Mexico. He has contributed greatly to the exploration and knowledge of the flora of Jalisco, and to the formation of new botanists and ecologists.

Additional material examined. Mexico. Jalisco. Autlán de Navarro: Las Joyas, Sierra de Manantlán, 1 800 m, 20 Aug 1982 (fl., fr.), Pérez 192 (IBUG); camino de Las Mantequillas a la cascada Las Juntas, Las Joyas, 1 600 m, 8 Dec 1985 (fr.), Vázquez 3766 (ZEA); de Las Galeras a la cascada grande, 1 600-1 650 m, 3 Aug 1986 (fl.), Cuevas 1442 (MEXU); 12-13 km al S de Autlán, 1 500 m, 24 Jul 1988 (fl.), Santana and De Niz 3670 (ZEA). Ayutla: Las Iglesias, 2 090 m, 24 Dec 2002 (fr.), Cuevas et al. 7594 (ZEA). Casimiro Castillo: Sierra de Manantlán Occidental, 2 km E of the microondas tower above puerto Los Mazos, 9 km (by air) NE of Casimiro Castillo, 1 760 m, 16 Dec 1988 (fl., fr.), Iltis and Santana 30109 (MEXU), 30123 (ZEA); Los Mazos, al W de Autlán, 1 800 m, 10 Sep 1989 (fr.), Ramírez and González-T. 1570 (IBUG); Los Mazos, Sierra de Manantlán, 9-10 km al SSW de Autlán, 9-10 km al NNE de Casimiro Castillo, 1 750 m, 15 Jul 1992, Santana et al. 11183 (ZEA); Los Mazos, Sierra de Manantlán, 9-10 km al SSW de Autlán, 9-10 km NNE de Casimiro Castillo, 24 Nov 2008 (fl., fr.), Sigala et al. 23 (ZEA), Mendoza 15 (ZEA). Talpa de Allende: km 19 del camino Talpa-La Cuesta, 1 400 m, 17 Nov 1989 (fl., fr.), Ramírez et al. 1857 (IBUG); brecha Talpa-La Cumbre, 1 400 m, 28 Jul 1990, R. Ramírez D.2121 (IBUG); brecha de Talpa a La Cuesta, en Paso Hondo, 1 550 m, 15 Oct 1995 (fl., fr.), Machuca and Cházaro 7517 (IBUG); TalpaLa Cuesta, 20°12'58.5" N, 104°46'27.4" W, 1 426 m, 8 Aug 2009 (fl.), Quedensely and Arroyo 10160 (CIIDIR).

Remarks. We agree with Pool (2006) and Cuevas (2010) considering the infrageneric classifications proposed by Epling (1939, 1940) and Paton (1990) as unsatisfactory in the circumscriptions of their groups. However, their proposals are still the basis for the study of the genus, until an exhaustive revision and a new natural classification are developed.

Scutellaria cuevasiana J. G. González et A. Vázquez fits within section Uliginosae (Epling) Epling, which is equivalent to section Scutellaria and "Scutellaria uliginosa" species group of Paton (1990), because the magenta corollas (purplish), small (less than 60 cm tall) herbaceous habit, fibrous or fasciculate roots, slightly hirsute leaves on the adaxial surface, and with appressed hairs on the vein of the abaxial one, alternate or subopposite flowers, spirally arranged in terminal racemes (Epling, 1939, 1942; Pool, 1998). However, the corollas are slightly longer than those exhibited by the members of section Uliginosae: 2.2-2.4 cm vs. 1.1-1.8 cm long (measurements including the upper lip length).

It should be noticed that the specimen Cuevas 1442 (MEXU) includes both Scutellaria dumetorum Schltdl., and S. cuevasiana in the same sheet. The first one can be easily distinguished by means of its flowers disposed in the axils of the leaves rather than in terminal racemes. On the other hand, Iltis and Santana collected 2 specimens of S. cuevasiana with the numbers 30109 and 30123, and they wrote on the labels that one plant had purple corollas and the other white ones; apart from this, there are not more differences between the specimens, indicating that this species can have magenta corollas and less frequently white ones (this can be partially observed in the holotype and isotypes, because dried flowers have 2 different colors).

Scutellaria sublitoralis J. G. González sp. nov.

Figure 3.

Type: Mexico. Jalisco. Puerto Vallarta, 3.5 km al SE por la brecha de Puerto Vallarta a Cuale a partir del puente del Remance, 500-700 m al SE del arco del ejido Jorullo, junto a una cascada que vierte en el río Cuale, 20°34'53.8" N, 105°12'28.4" W, 92 m, 24 Oct 2010 (fl., fr.), J. G. González-Gallegos 754 (holotype: IBUG; isotype: MEXU).

Species habitus Scutellariae pallidiflorae Epling optima congruens, sed laminis latioribus (3.5-6 cm vs. 3-3.5 cm longis), bracteis floralibus lanceolatis persistentibus (vs. linearibus deciduis), axe inflorescentiae et calycibus sine pilis glanduliferis, tubis corollarum angustis at apicem (vs. ampliatis at apicem), intus nudis (vs. intus annulis pilorum ornatis) differt.

Perennial herb 20-50 cm tall; erect, roots thin and fasciculate; stem cylindrical to quadrangular, moderately to densely covered with tiny appressed and retrorse hairs, these hairs are present also on the floral axis. Petioles 2.5-4 cm long, covered with appressed and retrorse hairs. Blades ovate, slightly asymmetric, (4-) 5-6.5 cm long, (3.5-) 4-6 cm wide, acute at apex, cordate to subcordate or truncate at base, margin irregularly and sparsely crenate or undulated, adaxial surface dark green, abaxial surface slightly paler, both surfaces glabrous or with some patent hairs along the margin. Inflorescences in racemes, terminal and lax, 6-15 cm long, flowers alternate to opposite, spirally arranged, nodes 7-12 mm apart from each other. Floral bracts linear, 1.5-2.7 mm long, 0.1-0.2 mm wide, persistent, with tiny erect hairs concentrated at base and along the margin, acuminate at apex, truncate at base, margin entire. Pedicels 2-3 mm long, 3.5-4 mm in fruit, covered with tiny retrorse hairs. Calyces (2-) 2.73.2 mm long, 1.4-1.7 mm wide at apex (3.5 mm long, 2.2 mm wide in fruit), green, margin of the lips with a fine purple line invisible to the naked eye, sparsely covered with erect tiny hairs, those concentrated at base and the margin of the lips, scutellum 1 mm tall and wide in flower, 1.5-2 mm tall and 3 mm long in fruit, green with the dorsal margin bordered with a narrow purple line. Corollas white; tube (1.3-) 1.4-2 cm long, slightly bent at 3.5-5 mm long, then straight and consistently narrow (1.1-1.2 mm wide) for 4.7-6 mm long, and gradually expanding to 3-3.3 mm wide at throat, glabrous inside, with sparse glandular-capitate and glandular simple hairs outside; upper lip 3-4 mm long, lower lip 5-6 mm long, tiny glandular-capitate hairs along the margin of the lips. Filaments of the upper stamens 3-4 mm long, filaments of the lower stamens 5.5-6.5 mm long; anther 0.3-1 mm long, white pilose on the slits. Styles 1.4-2.4 cm long, glabrous. Nutlets orbicular to subreniform, 1.5 mm long, 1.2-1.5 mm wide, black, the surface densely tuberculate (tubercules truncate at the apex), glabrous, covered with bright and tiny glandular dots.

Taxonomic summary

Distribution, habitat and phenology. Scutellaria sublitoralis grows in tropical subdeciduous forests, in wet and shady ravines, near the coasts of Jalisco and Nayarit (figure 2). It grows in a narrow altitudinal range, from 50-125 m, together with Hura polyandra Baill., Brosimum alicastrum Sw., Ficus insipida Willd., Euphorbia mexiae Standl., Begonia plebeja Liebm., Achimenes sp. Collected in flower and fruit from August to late November.

Etymology. The name of S. sublitoralis remarks its subcoastal distribution, always growing under 150 m and above 50 m altitude.

Remarks. Scutellaria sublitoralis is morphologically similar to Salvia pallidiflora Epling in sect. Pallidiflorae Epling. It shares with these, perennial herbaceous habit, fasciculate roots, ovate or elliptic petiolate blades, flowers arranged in short racemes, pale or essentially white corollas, and narrow tube. However, it shows some small discrepancies: the bracts are persistent instead of mostly deciduous, and the tube is not internally pilose. Furthermore, S. sublitoralis can be distinguished by its wider leaves (3.5-6 cm vs. 3-3.7 cm wide), generally cordate at base vs. rounded, calyx and floral axis without glandular-capitate hairs, pedicel as long as calyx in flower (vs. twice or more in length than the calyx in flower), and white corolla.

There is also an abrupt difference between the habitat preference of S. pallidiflora and S. sublitoralis. The first one is known exclusively by the type specimen (Rose 2487, seen in a natural sized photograph), where the locality is written as "near Huejuquilla". McVaugh (1972) adds more information to the locality data of this collection: between San Juan Capsitrano (Zacatecas) and Huejuquilla (Jalisco). In this area, according to Vázquez et al. (2004), there are several patches mainly composed of pine-oak and tropical deciduous forests, crassicaule bush and Chihuahuan desert, in an altitudinal range from 1 000-1 700 m. In contrast, S. sublitoralis grows in tropical subdeciduous forest under 150 m altitude. This kind of habitat is remarkably unusual among the Mexican Scutellaria, most of which occupy temperate mountainous areas above 1 500 m altitude.

Scutellaria vitifolia Brandegee also shares some characters with S. sublitoralis. The habit and height, petiole length, blade shape and size, apex and base of the blade, inflorescence length, tube and upper lip length. However, S. sublitoralis can be distinguished by stems and calyces without glandular-capitate hairs, white corolla (vs. blue to rarely purple), corolla tube internally glabrous (vs. pubescent internally in the central middle portion), nutlets ornate with tubercules truncate at the apex (vs. nutlets with tubercules ornate with stellate-peltate scales at apex) (Table 2). Moreover, S. sublitoralis is exclusive of Jalisco and Nayarit lowlands from 50-125 m altitude; while, S. vitifolia is known from Chiapas (Mexico) and adjacent areas of Guatemala, and it grows in a wider altitudinal range, from 60 to 2 100 m altitude (Table 2).

Acknowledgments

We thank Luz María González Villarreal, Ernesto De Castro Arce, Jeanette Hernández Santana and Ramón Cuevas Guzmán, from the University of Guadalajara, and Deborah Vaile, from the library of the Royal Botanical Garden of Edinburgh, for specialized literature acquisition. Servando Carvajal Hernández from Instituto de Botánica (Universidad de Guadalajara, Mexico) reviewed and improved the latin diagnosis. We appreciate the valuable information provided by José Luis Rodríguez Hernández and José Morales Arias about the holotype specimen of S. cuevasiana. We also thank the curators and colleagues from the herbaria consulted: CIIDIR, IBUG, MEXU, ZEA, and XAL, for the help and facilities provided. Ernesto De Castro Arce gave us valuable comments on the paper. Amy Pool (Missouri Botanical Garden) and two anonymous reviewers greatly improved the paper with their suggestions. Financial support was provided by CONACYT and Universidad de Guadalajara, Mexico.

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