Salvia patriciae J.G.González

González-Gallegos, Jesús Guadalupe, Fragoso-Martínez, Itzi, González-Adame, Gabriel, Martínez-Ambriz, Emmanuel & López-Enríquez, Irma Lorena, 2018, Salvia ozolotepecensis, S. patriciae and S. sirenis (Lamiaceae), three new species from Miahuatlán district, Oaxaca, Mexico, Phytotaxa 362 (2), pp. 143-159 : 151-154

Salvia patriciae J.G.González & E.Martínez-A., sp. nov. ( Figs. 5 & 6 )

Salvia stolonifera primo adspectu maxima simile, sed corolla tubis angustatioribus (3.5–6.5 vs. 7–9.5 mm latis), corollarum labiis superis brevioribus [(5.9–)7.8–10 vs. 12–15.5 mm longis] et inferis brevioribus vel longitudo quasi eadem (vs. longioribus vel quasi eadem), papillis erectis (vs. recurvatis) et brevioribus [1–1.4 vs. 6–7(–10) mm longis], staminibus exsertis (vs. inclusis), connectivis longioribus [(18.4–)27.6–40 vs. 21.6–25 mm longis], et stylis prope ad medium pilosulis (vs. pilosulis infra stylos ramarum).

Type:— MEXICO. Oaxaca: Miahuatlán district. Mun. San Juan Ozolotepec: Quiexobra [Cerro del Agua], 3550 m, 15 October 1995, G. B. Hinton 26159 (holotype CIIDIR!, isotype GBH!).

Perennial herb or subshrub, erect, 20–50 cm tall; stems short pilose; developing stolons and tubers. Leaves with petioles (0.3–) 1.2–2.7 cm long, short pilose; blade deltoid to ovate-deltoid, 2.5–5.1 × 1.6–4 cm, acuminate to acute at apex, cuneate at base, margin crenate to crenate-serrate, pilose in both faces and covered with amber glandular dots, pubescence longer and denser beneath. Inflorescence in racemes (5.2–) 9–22 cm long, with 3–8 floral nodes, each with 2–4 flowers, the lowermost 1.4–2.1(–4.4) cm apart from each other; floral axis with sparse simple hairs and more abundant tiny glandular-capitate. Floral bract soon deciduous, rhomboid-elliptic to lanceolate, 3.9–6.1 × 1.2–1.5 mm long, apex short caudate, base cuneate and then truncate, margin entire, pubescence as in the floral axis. Flowers with pedicels (2.9–) 3.6–5.5 mm long, sparsely pilose and with dense tiny glandular-capitate hairs. Calyx (9.3–)10.2–12 × 3.2–5.2 mm, up to 8 mm long in fruit, pubescence outside as in floral axis, short hispidulous inside, lips subequal, (2–)2.5–3.4(–4.5) mm long, the upper one entire and 7-veined. Corolla red to orange-red, sparsely short pilose; tube 22.2–31.7 × 3.5–6.5 mm, thin throughout, straight at base and with two slender linguiform papillae inside (1–1.4 mm long); upper lip (5.9–) 7.8–10 mm long, lower (5.5–)7–11.5 × (4.5–) 7–12.8 mm. Stamens exserted by 3.2–7 mm, filament 3.4–4.5(–6.1) mm long, connective (18.4–) 27.6–40 mm long, slightly curved and with and antrorse short tooth near midportion, theca (1.7–) 2.3–3.1 mm long; a pair of small staminodes above and behind filament insertion. Gynobasic horn 1.3–1.5 mm long, style (30–) 37.7–45 mm long, 7–10.7 mm exserted, sparsely pilose near midportion, lower stigmatic branch acute and shorter than the upper one. Mericarp ovoid, 2.8–3.2 × 1.5–1.8 mm, brownish gray and irregularly punctate with a darker tone, glabrous and smooth

Etymology:—The epithet honors Patricia Hinton, member of an outstanding family of plant collectors in Mexico, which dates back to George B. Hinton (1882–1943). They have contributed greatly to increase the knowledge of the Mexican flora, and in their collecting adventures have explored very remote and inaccessible localities. Their legacy includes 19 type specimens of currently accepted names in Salvia .

Distribution, habitat and phenology:— Salvia patriciae is an endemic species from Oaxaca, Mexico; known from several localities no more than 6 km apart from each other ( Fig. 3 ). It grows in the mountains of Miahuatlán district in Sierra Madre del Sur from 2655–3550 m elevation. It dwells in pine forests. Flowering from June to November, and fruiting probably from late August to February.

Notes:—Following Epling (1939) identification keys for Salvia subgenus Calosphace does not lead to a clear assignment to one of the sections. The most morphologically suitable are Charantia Epling (1939: 141), Flexuosa, Iodophyllae Epling (1939: 141) and Pedicellata Epling (1949: 518) based on sharing exserted stamens, papillae inside corolla tube base, 5–7-veined upper calyx lip, upper corolla lip longer than the tube and deciduous floral bracts. However, Salvia patriciae differs from the species in S. sect. Charantia by having deltoid to ovate-deltoid leaf blades (vs. oblong to oblong-lanceolate) and straight corolla tube at base (vs. invaginated). The species in S. sect. Flexuosae differs by the longer leaves (6–18 vs. 2.5–5.1 cm long), usually longer floral bracts (5–25 vs. 3.9–6.1 mm long) or upper corolla lip longer or as long as the lower; besides, this section is restricted to South America. Salvia sect. Iodophyllae possesses only one species, Salvia iodophylla Epling (1939: 141) , which differs from the new one by its 2-flowered floral nodes (vs. 2–4-flowered), 5-veined upper calyx lip (vs. 7-veined), lower corolla lip shorter than the upper (vs. lower longer than the upper) and entire connective (vs. ornate with an antrorse short tooth). Finally, S. sect. Pedicellata has also only one species, Salvia palealis Epling (1940: 519) , which differs in having bigger leaves (11–13 × 6–9 vs. 2.5–5.1 × 1.6–4 cm), cordate leaf blades (vs. cuneate), 12-flowered floral nodes (vs. 2–4-flowered), 5-veined upper calyx lip (vs. 7-veined), and subequal corolla lips (vs. the upper shorter than de lower).

In general aspect, Salvia patriciae is more similar to the species in section S. sect. Incarnatae. At first glance, it looks like Salvia elegans Vahl (1804: 238) but with bigger flowers and exserted stamens. Nonetheless, more detailed observations reveal there are other deep discrepancies: entire and acute upper calyx lip (vs. trimucronate or caudate), corolla internally ornate with two slender papillae (vs. epapillate), lower corolla lip longer than the upper (vs. shorter than the upper).

If stamen exsertion is ignored, following the identification key leads to S. sect. Fulgentes Epling (1939: 273) when trying to identify Salvia patriciae . Amongst the species of this group, Salvia lineata Bentham (1835: 724) is the morphologically most similar. However, S. patriciae can be distinguished because the pilose leaves (vs. glabrous), shorter floral bracts (3.9–6.1 vs. 6–10 mm long), longer corolla tube (22.2–31.7 vs. 19.6–20.4 mm long), exserted stamens (vs. included below upper corolla lip), antrorse tooth in connective midportion (vs. retrorse) and style short pilose near midportion (vs. pilose towards the apex). If the proportion between corolla lips is also ignored (upper one as long or shorter than the lower, vs. as long or longer), S. patriciae can be related to S. sect. Cardinales Epling (1939: 295), and within this, to S. stolonifera Bentham (1840: 70) . Both species are very similar and hard to be separated considering only vegetative characters, except by the leaves that are cordate, cordate and short cuneate to truncate (vs. cuneate) at base and slightly papillate beneath (vs. smooth) in S. stolonifera . However, once floral characters are compared, stronger evidence arises in order to support the recognition of two different species; S. patriciae differs in having thinner corolla (3.5–6.5 vs. 7–9.5 mm wide), shorter upper corolla lip [(5.9–)7.8–10 vs. 12–15.5 mm long] and this as long or shorter than the lower (vs. as long or longer), straight vertical (vs. recurved and oriented forwards) and shorter papillae [1–1.4 vs. 6–7(–10) mm long] inside corolla tube, exserted stamens (vs. included), longer connective [(18.4–)27.6–40 vs. 21.6–25 mm long], and style short pilose near midportion (vs. with the hairs disposed in the apex just before stigmatic branches).

Because of the similarity between Salvia patriciae and S. stolonifera , this was referred as the latter in the floristic inventory conducted by McDonald (2013) in Cerro Quiexobra. Later, B.L. Turner labeled the specimen McDonald 3004 (TEX!) as Salvia macdonaldii , no date on the label, revealing his intention of publishing this as a new species; however, he did not validate the name.

In summary, Salvia patriciae is a distinctive new species sharing characters with the species of several Epling’s sections, but which cannot be unambiguously assigned to one of these. However, although the results of the phylogenetic analysis ( Fig. 4 ) do not show a clear assignment to any of the sections, it is shown a close relationship with the Fulgentes clade. This group is a heterogeneous assemblage including some species of 4 different sections: Cardinales [as Holwaya Ramamoorthy (1984: 323) according to a more recent proposal], Flocculosae ( Epling 1935: 77) Epling (1939: 153), Fulgentes and Nobiles Epling (1939: 280) [the sampled species from this section also might belong to Holwaya according to dos Santos (1991)]. The species in this clade share 5–7 veined upper calyx lip, red corollas (mostly), ventricose corolla tube and pubescent style [except S. univerticillata Ramamoorthy ex Klitgaard (2007: 208) according to the description in Flora Mesoamericana ( Klitgaard 2012)]. All of these characters are shared with S. patriciae ; however, the remaining species of the clade have stamens that are included in the upper corolla lip. The differences in the level of exsertion of the stamens have been used by Epling (1939) to separate species into contrasting sections. These results could suggest that the transition from exsertion to inclusion is more common and labile, and/or that there are still a number of species that remain to be sampled. Expanding the taxon sampling in the phylogenetic framework would shed light into the evolution of floral morphological characters and their value as taxonomic diagnostic characters, such as stamen exsertion.

Due to the above, and in face that Epling’s classification needs to be rearranged since it is far from natural ( Jenks et al. 2013), no section is here designated.

Additional specimens examined (paratypes):— MEXICO. Oaxaca. Miahuatlán district. Mun. San Juan Mixtepec: San Juan Mixtepec , 5 km SW of town, 16°20’8”N, 96°19’9”W, 2900 m, 23 June 1997, E. Hunn 1259 ( MEXU!). Mun. San Juan Ozolotepec : 35 km ESE of Miahuatlán, 5 km NE of Santo Domingo Ozolotpec , Cerro Quiexobra , 16º10’N 96º15’W, 3500 – 3700 m, 3 October 1990, J. A. McDonald 3004 ( TEX!) ; Quiexobra [Cerro del Agua], 3425 m, 6 August 1996, G. B. Hinton 26787 ( GBH!) ; Zona de aprovechamiento forestal, camino al Cerro del Agua , 16°10’42.77”N 96°14’7.59”W, 3126 m, 27 November 2017, G. González-Adame et al. 2830 (Herbario de la Universidad Sierra Juárez!, SERO!) ; camino de acceso al Cerro del Agua , 16° 9’58.04”N 96°14’52.99”W, 3141 m, 27 November 2017, G. González-Adame et al. 2831 (Herbario de la Universidad Sierra Juárez!, SERO!) ; Cerro del Agua, 3.5–4 km al NE de San Juan Ozolotepec, 16º9’53.9”N 96º14’49.5”W, 3128 m, 27 November 2017, J. G. González-Gallegos et al. 2321 ( CIIDIR!, HUAA!, IEB!, MEXU!, OAX!, ZEA!). Mun. Santo Domingo Ozolotepec: Neverías, 2655 m, 3 August 1996, G. B. Hinton 26651 ( GBH!) ; Las Trancas , 16°11’45”N 96°16’0”W, 3516 m, 8 July 2010, A. Sánchez M. 2906 ( MEXU!, SERO) ; desviación a Santo Domingo Ozoltepec , 16°12’3”N 96°20’47”W, 2920 m 28 November 2015, I. Fragoso-M. et al. 363 ( MEXU!) .

Figures

FIGURE 3. Distribution map of Salvia ozolotepecensis (white dots), S. patriciae (white triangles) and S. sirenis (white squares). Upper right corner showing the location of Oaxaca and Miahuatlán district; the red rectangle highlights the territory magnified in the rest of the figure. Sand lines correspond to municipalities boundaries within Miahuatlán district. Orange shadows represent the towns of, from top to bottom, San Juan Mixtepec, Santo Domingo Ozolotepec, San Juan Ozolotepec and Santiago Xanica.

FIGURE 4. Phylogenetic relationships of Salvia subgenus Calosphace based on three molecular markers (nrITS, trnH-psbA and trnL- trnF) and an extended taxon sampling including the species of Table 1. Bootstrap values ≥80% are shown above the branches. Species names in blue and pink correspond to the new species published earlier and herein, respectively. The clades remaining unchanged with the new taxon additions have been collapsed to ease visualization. An expanded version of the phylogeny can be seen in Supplementary materials.

FIGURE 5. Salvia patriciae. A General habit. B Leaf blade, upperside. C Tuber and roots. D. Flower (Photographs taken by G. González- Adame from Cerro Quiexobra [Cerro del Agua]).

FIGURE 6. Salvia patriciae, floral details. A Calyx. B Ovary and gynobase. C Corolla. D Stamens. E Corolla tube papillae. F Style apex [Photographs taken from I. Fragoso 363 (MEXU!)].