26. Salvia L. (sage, salvia)

Plants annual or perennial herbs (shrubs elsewhere), with taproots or somewhat thickened or woody rootstocks. Stems erect or ascending, bluntly to sharply 4-angled, unbranched or branched, glabrous or variously hairy. Leaves petiolate (sometimes the uppermost leaves sessile or nearly so), the petiole unwinged or winged, with a usually strong fragrance when crushed. Leaf blades variously shaped, unlobed to deeply pinnately lobed, the margins otherwise entire or more commonly toothed, the surfaces hairy, also with inconspicuous or sometimes conspicuous, sessile glands. Inflorescences terminal spikes or spikelike racemes, these sometimes grouped into loose clusters at their bases, the individual axes with more than 5 clusters, these relatively open, more or less continuous along the axis, each with 1–12 flowers, these sessile or short-stalked, subtended by a pair of short bracts, the individual flowers sometimes subtended by inconspicuous bractlets. Calyces zygomorphic, lacking a lateral projection, more or less symmetric at the base, bell-shaped, the tube strongly 10–15-nerved (-ribbed), glabrous in the mouth, hairy externally, 2-lipped, the lobes shorter than the tube, the upper lip unlobed or with 3 shallow, triangular lobes, the lower lip with 2 narrowly triangular lobes, the lobes sharply pointed and sometimes with a short, slender extension of the midnerve, but not spinescent, not or only slightly becoming enlarged (except in S. reflexa) or papery at fruiting. Corollas zygomorphic, white, red, lavender, or blue (pink or purple elsewhere), lacking markings on the lower lip or in some darker-flowered species with white mottling, the outer surface moderately to densely pubescent with minute, spreading hairs, especially on the upper lip, the tube funnelform to nearly cylindric, glabrous in the throat, shallowly to deeply 2-lipped, the upper lip usually shallowly notched at the tip (occasionally entire), either relatively flat (straight to slightly spreading) or hooded, the lower lip noticeably longer and broader than the upper lip, spreading or arched, 3-lobed, the middle lobe longer and broader than the lateral lobes, often shallowly notched at the tip. Stamens 2, not or only slightly exserted (more strongly exserted in S. coccinea), the anthers overall relatively large, the connective slender and elongate, the upper pollen sac fertile and fully formed, the lower sac either absent or reduced and nonfunctional (except sometimes in S. lyrata), the connective appearing attached at or below its midpoint, the fertile sac often yellow. Ovary deeply lobed, the style appearing nearly basal from a deep apical notch. Style not exserted (ascending under the upper lip), unequally 2-branched at the tip. Fruits dry schizocarps, separating into usually 4 nutlets, these 1.5–3.0 mm long, oblong-obovoid, rounded at the tip, the surface brown to dark brown, glabrous, smooth, minutely pebbled, or with dense, minute tubercles. About 900 species, nearly worldwide.

The genus Salvia is usually easily recognized by its characteristic, strongly 2-lipped calyces and often bright, showy corollas. Numerous species are cultivated as ornamentals in gardens (see Compton [1994] for a review of some of the Mexican species that are popular in gardens). The South American S. splendens Sellow ex Wied-Neuw. (scarlet sage) is among the most popular of annual bedding plants in Missouri. A number of salvias also have medicinal uses. Salvia hispanica L., S. officinalis L., and S. fruticosa Mill. are the main species grown as culinary herbs, the sages whose dried or fresh leaves are used to add flavor to cooked dishes. A few other species are used in beverages (mainly in teas. Some of the sages provide a laquerlike oil that has been used in the manufacture of oil paints. Sage is a popular ingredient in soaps, perfumes, toiletry products, sachets, and potpourris; the pleasantly scented S. sclarea L. is commonly used in these products and also is used as a flavorant in vermouth and some other liquors, as well as in an eye wash. Salvia divinorum Epling & Játiva (diviner’s sage) has a long history of cultivation in Latin America as a hallucinogen; that use has spread to the United States in recent years and a number of states are in the process of making its possession illegal.

The unusual structure of the stamens and the associated pollination mechanism was first identified in 1793, but was reviewed by Claβen-Bockhoff et al. (2003). The two stamens develop before the stigmas. They are situated vertically in the throat of the corolla, with the fertile pollen sacs arching under the upper corolla lobe and the sterile arms of the connectives (these sometimes bearing a fertile pollen sac in S. lyrata and a few other species) positioned downward across the tube (and frequently becoming fused into a paddle-shaped structure). A pollinator (variously insects or hummingbirds) entering the flower moves the sterile arms backward, causing a leverlike action that pushes the fertile pollen sacs onto its back (or head), where a load of pollen is deposited. After the pollen has been shed, the stamens wither and the style elongates and arches downward, placing the stigmatic branches in a position to intercept the pollen on a pollinator that previously has visited a different flower during that flower’s functionally staminate phase.

The unique staminal architecture has been used as a key character in defining the genus Salvia taxonomically. However, recent phylogenetic studies using molecular markers (Walker et al., 2004) have suggested that, as traditionally circumscribed, the genus is not a natural group, but instead consists of three independent species groups that are not particularly closely related within the tribe Mentheae. Under this scenario, the remarkable pollination mechanism described above developed independentally at least twice within this tribe of Lamiaceae, and the genus overall likely will be split into three genera in the future. The Missouri species would be classified in the genus Calosphace Raf., but new combinations will need to be published under that generic name.

Steyermark (1963) included S. sclarea L. (clary, clear-eye), based on a single historical collection made in 1910 by Earl Sherff in the city of St. Louis (with no further locality or habitat data). In the introduction to his Flora of Missouri, Steyermark discussed Sherrf’s St. Louis collections, many of which were made from cultivated plants but not indicated as such, and he excluded a number of species from the Missouri flora that were documented only from Sherff’s ambiguous collections. He included S. sclarea based on the anticipation that the species would be discovered by subsequent botanists elsewhere in the state. This optimistic prediction has not been realized. Thus, S. sclarea is here excluded from the Missouri flora. The species is a robust biennial with large ovate leaves that often are 6 cm or more wide and corollas often with yellowish markings in the throat (Pl. 441 g–i).