Salvia cajamarcana J.G.González, Uría & M.Roncal, 2023

González-Gallegos, Jesús Guadalupe, Roncal-Rabanal, Manuel & Uría, Rolando, 2023, Salvia cajamarcana (Lamiaceae), a new species from Cajamarca Department, Peru, and lectotypification of S. revoluta, Phytotaxa 597 (1), pp. 15-27 : 20-24

Salvia cajamarcana J.G.González, Uría & M.Roncal , sp. nov., Figures 4 and 5 .

Type:— PERU. Cajamarca department, Celendín province: Oxamarca district , Choctapampa , 3283 m, 23 March 2022, M. R . Roncal-Rabanal & J . Silva-Torres 190 (holotype Herbario Carlos Casanova Lenti 5057!, isotypes (to be distributed) CIIDIR!, CPUN!, MEXU!) .

Salviae revolutae primo adspectu maxime simile, sed indumento arachnoideo trichomatibus plerumque simplicibus vel ramosis sed ramis longioribus (vs. tomentuloso trichomatibus dendriticis), bracteis floralibus lanceolatis, elliptico-lanceolatis, ovato-lanceolatis vel partiale oblanceolatis(vs. nunc ovatis nunc elliptico-ovatis), bracteolis in pedicelli basi dispositis (vs. nullis), pedicellis longioribus (3–9.5 mm vs. 2–4 mm longis), calycis indumentis densis nunc albis nunc ferrugineis (vs. laxis, albis vel interdum violaceis), calycis lobis acutis et mucronatis (vs. acutis sed non mucronatis), corollae tubo invaginato prope basem (vs. prope medium post partem basale cylindraceam et relative longam), corollae labio superiore longiore ((9.8–) 11–14 m vs. 5–8.2 mm longo), filamento longiore ((4.8–) 6–8.5 mm vs 2–3.3 mm longo), connectivo longiore ((12–)16.5– 15 mm vs. 7.5–8.8 mm longo), hoc leniter incrassato ad medium et dentato cum dente minuto retrorso et acuto ad apicem (vs. conspicue incrassato ad medium et non dentato), staminodiis longioribus (1–2.5 mm vs. minus quam 1 mm longis), et stylo longiore ((27–) 29–35 mm vs. 19–22 mm longo) differt.

Shrub, (0.8–)1.4–1.6(–2) m tall, stems basally glabrous, striate and with the bark slightly exfoliating, upper parts white arachnoid to glabrescent, the hairs mostly simple but also with some branched hairs, primary bifurcate or with scarce branches, these less than a third of stalk length; young branches with undeveloped leaves covered with a bright and glutinous resin (except in Lluchubamba population). Leaves aromatic, petioles 1–2.3 cm long, arachnoid as the stems to glabrous, sometimes the hairs concentrated at the basal portion; leaf blade lanceolate to oblong-lanceolate, (3.4–)4–10 × (0.7–) 1.6–4.5 cm, apex acute to attenuate, base rounded, subcordate to cordate, and sometimes oblique, margin crenulate, green, bullate and sparsely pilose to glabrous above, white beneath because of a dense arachnoid pubescence, the hairs uniformly covering the lower surface or restricted between tertiary veins and resembling small cushions, hence the veins evident. Inflorescence in racemes (4–) 7.5–25 cm long, with 3–8 verticillasters, each one (2–)4–6(–8)-flowered, the lowermost floral nodes (0.7–)1.5–2(–4) cm apart from each other, floral axis densely white to ferruginous arachnoid with simple hairs, and some branched with scarce and sparse branches. Floral bracts deciduous, lanceolate, elliptic-lanceolate, ovate-lanceolate to partially oblanceolate, 6–13.5 × 2–5.6 mm, apex acuminate to caudate, base truncate, margin entire, outer surface pubescent as floral axis; bracteoles, soon deciduous (even before the bracts), linear to linear-lanceolate, 5–8 × 0.4–1 mm, apex acute, base truncate, margin entire, also pubescent as floral axis in the outer surface. Pedicel 3–9.5 mm long, pubescent as floral axis; calyx divergent towards the apex, tube (10.8–)13–21 × (6–) 7–10 mm, pubescent as floral axis but usually denser, hispidulous inside with yellowish antrorse hairs, upper lip 7-veined and entire, the lobes mucronate at the tips; corolla lilac, purple or dark blue, paler to white from the middle to the base; corolla indumentum densely pilose in the portion exserted from the calyx, the rest glabrescent to glabrous; tube (16–)17–22.2 × 4.8–8 mm, ventricose, with a ventral invagination near the base, ornate with two folds inside at the invagination; upper lip (9.8–) 11–14 mm long; lower lip (7.5–)9.5–12 × (6–)10–12.5(–16) mm; stamens included within the upper corolla lip; filaments (4.8–) 6–8.5 mm long; connective (12–) 16.5–25 mm long, entire or with a tiny acute tooth near the junction with the filament; thecae 3.5–4.2 mm long; a pair of staminodes placed above and behind filament insertion of each stamen, filiform and capitate at apex, 1–2.5 mm long; gynobasic horn 1–2.2 mm long; style (27–) 29–35 mm long, short pilose along the dorsal and ventral lines towards the apex, upper stigmatic branch subequal or longer than the lower one, the latter acute at apex. Mericarp ovoid, 2.9–3.4 × 1.5–2 mm, brown and irregularly marbled with a darker tone, smooth and glabrous.

Distribution and ecology:— Salvia cajamarcana is a Peruvian endemic species ( Fig. 2 ). It grows on slopes of Andean mountains, in localities relatively humid due mainly to frequent fog, that can be inferred from the abundance of epiphytic bromeliads in the surrounding shrubs ( Fig. 3A–C ). It occupies an elevation range from 2800 to 3300 m. It shares habitat with the herbs and shrubs of Bomarea Mirbel (1802: 71) , Calceolaria Linnaeus (1770: 286) , Clinopodium Linnaeus (1753: 587) , Cortaderia Stapf (1897: 396) , Lantana Linnaeus (1753: 626) , Lupinus Linnaeus (1753: 626) , Oreopanax eriocephalus Harms (1895: 67) , Oxalis Linnaeus (1753: 433) , Pappobolus Blake (1916 : pl. 3057), Rubus robustus Presl (1851: 556) , Salvia oppositiflora , S. punctata Ruiz & Pavón (1798: 27) and S. styphelus Epling (1939: 52) . There are outcrops of sedimentary limestone rocks in the area, and agricultural and cattle raising activities are developed. The seedlings emerged between mosses and lichens.

Phenology:— Salvia cajamarcana flowers and bears fruit from the beginning of March to the end of August.

Etymology:— The new species is dedicated in honor to the Cajamarca culture that developed throughout the department and mainly occupied the valley of the capital city during the years 50 BC to 1400 AD.

Uses:— In Lluchubamba, this new species is known as chacato or chacatos, and in Oxamarca as carquejilla; no uses in folk medicine were reported in these two localities. In Laguna San Nicolás the new species [mistakenly identified as Salvia cuspidata Ruiz & Pavón (1798: 23) ] is known as salvia blanca (white sage) and its leaves are used to prepare a drinking infusion to treat flu and cough ( Cueva-Infante, 2019).

Preliminary conservation status:— Salvia cajamarcana is known only from three populations, all in the Department of Cajamarca, and relatively close, with an EOO of 921 km 2 and an AOO of 20 km 2. The populations from Namora (surroundings of Laguna San Nicolás) and Oxamarca (archeological ruins of Las Chullpas) are small and fragmented, with scattered individuals in the area. The third population, from Lluchabamba, is the largest, spread over about 4 km between Cajabamba and Lluchabamba, although also fragmented. The Namora population is the most deteriorated, showing signs of hydric stress and with scarce young plants and seedlings. The plants in the three populations are under agricultural and animal husbandry pressure, near to the agricultural frontiers. Besides, the plants at Oxamarca grow in archaeological ruins that are visited by local tourists but under no protection or any regulations, which might also constitute a threat. Hence, S. cajamarcana should be treated as an Endangered (EN) species under IUCN (2022) criteria due to the reduced EOO and AOO, the small and deteriorate populations as well as current and future threats derived from farming and tourism: EN B1ab(iii)+2ab(iii). Strategies to conserve this species should integrate its promotion as an ornamental plant in gardens of temperate-cold areas and the protection of the Chullpas at Oxamarca, not only because of its archeological/historical value but also for the biodiversity it embraces.

Additional specimens examined:— PERU. Cajamarca: Cajabamba: Sitacocha district , 3.7 km antes de Lluchubamba desde Cajabamba, 3078 m, 7.5ºN, 77.99ºS, 14 June 2022, R . Uría & F . Lozano ( CIIDIR!, MEXU!). Cajamarca: Namora district , ladera W que converge a la Laguna de San Nicolás , 2800 m, 5 March 2002, J. G . SánchezV. 11274 ( CPUN!); Namora district, caserío San Nicolás , 7º14’14.42”S, 78º19’52.7”W, 2894 m, 11 June 2022, M. R . Roncal-Rabanal & M . Roncal-Ordóñez 191 ( CIIDIR!, CPUN!, Herbario Carlos Casanova Lenti!). Celendín: Celendín , cerro Gelít al S de Celendín, 2 km por terracería del pueblo, 3350 m, 9 November 1984, C. P . Cowan et al. 4340 ( CPUN!, TEX!, USM!); ruta La Chocta – Oxamarca , 3300 m, 30 July 1970, J. G . Sánchez-V. 617 (CPUN!).

Notes:— The new species was collected in the Department of Cajamarca in the spring of 2022. There are previous specimens collected by Sánchez in 1970 and 2002, and by Cowan in 1984, which remained undetermined. It is outstanding for its bicolored leaves (green above and white beneath), the calyces with ferruginous or white densely arachnoid hairs, and the purple to dark blue corollas.

The morphology of Salvia cajamarcana does not fully match with any of the sections of subgenus Calosphace as defined by Epling and coworkers ( Epling 1939, 1940, 1941, 1944, 1947, 1951, Epling & Mathias, 1951, Epling & Játiva 1963, 1966, 1968). Nonetheless, it is closer to S. sect. Pavonia ( Epling 1935: 95) Epling (1939: 279) due to the shrubby habit, oblong-lanceolate leaves, incanous beneath, interrupted inflorescences with six flowers per floral node, deciduous floral bracts, upper calyx lip 5–7-veined, corolla lips subequal in length, included stamens under the upper corolla lip, connective with an acute retrorse tooth, and pilose style ( Epling, 1939, 1947). However, S. cajamarcana deviates from the section definition by the absence of papillae inside corolla tube near the invagination, although it presents a couple of folds, and one of the species of the section also has folds instead of clear papillae.

Among the species of Salvia sect. Pavonia , only S. orbignyi Bentham (1848: 338) has been included in phylogenetic analyses and emerged in a clade with other South American species, but with poorly supported phylogenetic relationships, within the “Core Calosphace ” clade ( Fragoso-Martínez et al. 2018, Kriebel et al. 2019), a group that has not been formally recognized in an infrageneric classification. Because of the current phylogenetic ambiguity in this group, we relied on Epling’s (1939) classification to provisionally place the new species in S. sect. Pavonia, despite acknowledging that this system needs to be restructured to reflect natural groups.

Salvia sect. Pavonia is composed of two species, S. orbignyi and S. revoluta (= S. macbridei Epling (1939: 280)) . It is worth noting that while Epling (1939) described the corollas for the section as magenta (rubro-purpurearum, sic), S. revoluta presents dark blue to purple corollas, as those of S. cajamarcana ( Fig. 1 ). In fact, when following identification keys for Peruvian salvias (MacBride 1960) and that for the species of Salvia subg. Calosphace (Bentham 1833: 198) Epling (1939: 4) ( Epling 1939, 1940, 1941, 1947), S. revoluta is recovered as the morphologically most similar species to S. cajamarcana ; however, it differs in having a tomentulose indument composed by short stalked branched hairs (dendritic) along the stems, leaves, inflorescence axis, floral bracts, pedicels and calyces (vs. arachnoid, mostly with simple hairs and few sparsely branched ones with long stalks, in S. cajamarcana ), floral bracts ovate to elliptic ovate (vs. lanceolate, elliptic-lanceolate, ovate-lanceolate to partially oblanceolate), lacking bracteoles (vs. bracteolate pedicels with soon deciduous linear to linear-lanceolate bracteoles), shorter pedicels (2–4 mm vs. 3–9.5 mm long), calyx indument white or sometimes violet and sparse (vs. calyx white to ferruginous, with hairs covering the surface), calyx lobes acute without a mucro (vs. acute and mucronate at the tip), corolla tube with a ventral invagination near the middle after a relatively long basal cylindrical portion (with the invagination near the base), shorter upper lip (5–8.2 mm vs. (9.8–) 11–14 mm long), shorter filament (2–3.3 mm vs. (4.8–) 6–8.5 mm long), shorter connective (7.5–8.8 mm vs. (12–)16.5– 15 mm long), and the latter widened at the middle with this portion resembling a rhomboid parallelogram (vs. gradually and slightly widened to the middle, but not with a clear rhomboid section, and with a tiny acute retrorse tooth), shorter thecae (2–2.4 mm vs. 3.5–4.2 mm long), shorter staminodes (less than 1 mm vs. 1–2.5 mm long), and shorter style (19–22 mm vs. (27–) 29–35 mm long).

Salvia cajamarcana also recalls, at first sight, a set of Andean shrubby species with tomentose pubescence, bicolored leaves (usually white beneath because of the indumentum), bullate above, and with pubescent calyces. These grow in arid or semiarid habitats and belong to sections Corrugatae ( Bentham 1848: 327) Epling (1939: 50), Discolores ( Epling 1935: 87) Epling (1939: 53) and Leucocephalae ( Epling 1935: 88) Epling (1939: 308). The morphological similarities might respond to similar environmental pressures and could reflect a morphological convergence; however, S. cajamarcana is consistently distinguished from all the above.

The species of Salvia sect. Corrugatae are immediately set apart from S. cajamarcana by the trimucronate upper calyx lip (vs. entire) and a well-developed ventral antrorse tooth at the connective midpoint (vs. entire or with a tiny retrorse tooth) ( Epling 1939). Among section Corrugatae species, S. cajamarcana is most similar to S. corrugata Vahl (1805: 252) due to leaf shape and size. It differs from this species, in addition to the differences in respect to the section, in having lanceolate, elliptic-lanceolate, ovate-lanceolate to partially oblanceolate (vs. ovate) floral bracts, longer pedicels (3–9.5 vs. 2–3 mm long), non-viscid (vs. viscid) calyces, and these longer ((10.8–)13–21 vs. 9–10 mm long), and longer ferruginous or whitish by the pubescence and green underneath (vs. purple to violet), longer corolla tube ((16–)17–22.2 vs. 11–13 mm long), and longer upper ((9.8–)11–14 vs. 7–8 mm long) and lower ((7.5–)9.5–12 vs. 6–9 mm long) corolla lips ( Epling 1939, Wood & Harley 1989).

Salvia discolor Kunth (1818: 294) , an endemic Peruvian species that is the sole representative of Salvia sect. Discolores , differs from S. cajamarcana in presenting coriaceous, entire (vs. papery, crenulate) leaves, calyx covered with white dendritic hairs (vs. white to ferruginous arachnoid simple hairs, and some branched with scarce and long branches, mostly bifurcate), glabrous (vs. hispidulous) inside, corolla tube included inside the calyx as well as half upper corolla lip under calyx lip (vs. upper portion of the corolla tube exserted from the calyx together with corolla lips), shorter corolla tube (13–14 mm vs. (16–)17–22.2 long), and upper corolla lip half the length of the lower one (vs. subequal in length or the upper longer) ( Epling 1939).

Salvia cajamarcana differs from Salvia sect. Leucocephalae in pubescence (white arachnoid hairs and some branched with long branches and mostly bifurcate vs. white dendritic hairs), petiole length (1–2.3 vs. 0.5–1 cm long), leaf shape (lanceolate to oblong-lanceolate vs. ovate to ovate-elliptic) and pubescence of the lower surface (densely white tomentose hiding leaf surface vs. sparsely covered with white branched hairs allowing to see the leaf surface), floral bract shape (lanceolate, elliptic-lanceolate, ovate-lanceolate to partially oblanceolate, acuminate to caudate at apex, vs. orbicular and short acuminate at apex) and width (2–5.6 vs. 6–12 mm), in corolla tube length ((16–)17–22.2 vs. (11–) 20–28 mm long), and in having invaginate corollas near tube base and ornate with a couple folds inside (vs. straight and naked inside) ( Epling 1939, 1960). Section Leucocephalae comprises two Peruvian endemic species, S. consobrina Epling (1960: 149) and S. leucocephala Kunth (1818: 302) . The first also differs from S. cajamarcana in having leaf blades long attenuate at base.

Despite Salvia cajamarcana was misidentified as S. cuspidata in Cuevas-Infante (2019) when documenting folk medicine in Laguna San Nicolás, it can be easily set apart by the following characters: pubescence composed by branched hairs with short branches (vs. simple hairs with some scarce intermixed branched hairs, but with long branches mostly only bifurcate), longer floral bracts (6–13.5 vs. 3–4(–6.1) mm), presence of bracteoles (vs. absent), larger flowers (calyx (10.8–)13–21 vs. (4–)5.4–7.4(–9) mm, corolla tube ((16–) 17–22.2 vs. 4.5–6(–15) mm), with the upper corolla lip longer than the lower one or subequal in length (vs. the lower much longer than the upper), densely arachnoid calyces with ferruginous or white hairs (vs. sparsely white pilose with glandular-capitate hairs intermixed), and upper calyx lip entire (vs. trimucronate) ( Epling 1939, Wood 2007, O’Leary & Moroni 2016).

It is worth noting that Salvia cajamarcana has ornamental potential due to its showy leaves and flowers, as well as for its high tolerance to environmental stress. The last point being relevant in gardening and landscaping after a context of climatic change. It could be used in temperate-cold gardens, planted in garden containers or in rockeries.

Figures

FIGURE 1. Salvia revoluta. A Inflorescence. B Flower. C Calyx. D Corolla. E Ventral corolla dissection showing the stamens and the style (photographs from R. Uría’s private garden in Mar de Plata, Buenos Aires Province,Argentina, originally collected at Callampampa, District Llama, Province Chota, Department Ancash, Perú by R. Uría).

FIGURE 2. Maps showing the geographical distribution of Salvia cajamarcana (white triangles) and S. revoluta (white dots). In the main map, Peru is highlighted with a more intense background; the inset shows the area occupied by both species. Political boundaries of Peruvian Departments are shown (background of the maps taken from Google Earth Pro 7.3.48642).

FIGURE 3. Habitat of different populations of Salvia cajamarcana (A–C) and S. revoluta (D), all in the Department Cajamarca.A Laguna San Nicolás, District Namora, Province Cajamarca. B Lluchubamba, District Sitacocha, Province Cajabamba. C Oxamarca, District Oxamarca, Province Celendín. D Callampampa, District Llama, Province Chota (photographs by R. Uría and F. Lozano).

FIGURE 4. Salvia cajamarcana. A Inflorescence with upper portion of flowering branch. B Flower. C Calyx. D Corolla. E Ventral corolla dissection showing the stamens and the style (lower lip previously removed) (photographs from Oxamarca, District Oxamarca, Province Celendín, Department Cajamarca, by M. Roncal-Rabanal, corresponding to the type collection).

FIGURE 5. Color variation in the flowers of Salvia cajamarcana. Calyces with dense whitish (A) or ferruginous pubescence (B & C), or the hairs not so dense such the green calyx surface is visible (D). Corolla dark blue to purple (A, B & D) or lilac (C) (A & C from Lluchubamba, District Sitachocha, B & D from Oxamarca, District Oxamarca; photographs by R. Uría).