1. Glandularia
J.F. Gmel. (verbena)
(Umber, 1979)
Plants perennial
herbs (annual elsewhere). Stems few to several from the rootstock, loosely
ascending or creeping with ascending tips, rarely erect, sometimes rooting at
the lower nodes, weakly 4-angled, hairy. Leaves usually with a partially winged
petiole, variously 3-lobed to ternately or pinnately 1 or 2 times deeply
divided, the segments narrowly oblong to linear, pointed at the tip, the
margins sometimes coarsely few-toothed, sparsely to moderately hairy.
Inflorescences terminal on the branches, not associated with slender, elongate
stalks, initially dense, flat-topped clusters, becoming elongated into spikes
with age. Calyces narrowly tubular, 7–13 mm long, 5-lobed, the lobes somewhat
unequal in length, narrowly triangular to nearly linear, erect at flowering,
becoming contorted as the fruits mature, hairy on the outer surface and along
the margins. Corollas 12–30 mm long (shorter elsewhere), trumpet-shaped,
slightly zygomorphic, 5-lobed, pink to lavender or light bluish purple, rarely
white (red elsewhere), usually fading to dark blue, the limb 8–20 mm in
diameter, the lobes often shallowly and broadly notched at the tip. Stamens
inserted at 2 levels toward the tip of the corolla tube, usually with a small
glandular appendage positioned laterally between the anther sacs. Ovary
4-locular, appearing 4-lobed, slightly concave at the tip. Style 15–22 mm long
(shorter elsewhere), the sterile lobe extending noticeably beyond the fertile
lobe (this obscured by the globose stigmatic area), flattened and narrowly
triangular. Fruits consisting of (2–)4 nutlets, these more or less cylindric,
usually rounded at the tip, somewhat asymmetrically concave at the base, the
surface variously wrinkled, lined, and/or with a network of blunt ridges,
sometimes also with small papillae, especially along the inner surface, grayish
black to black (tan to brown elsewhere). About 50 species, North America to
South America, Caribbean Islands; introduced in the Old World.
Traditionally, Glandularia
was regarded as a section within the genus Verbena (Steyermark, 1963).
Umber (1979) and Sanders (2001) have summarized morphological, cytological, and
phytochemical data supporting the separation of the two groups as genera, which
is how most South American botanists have been treating them for many years
(Schnack and Covas, 1944).
Several species
of Glandularia are cultivated as ornamentals in the Midwest, including
both of the taxa growing wild in Missouri. Glandularia canadensis is
popular in native plant gardening because it is easy to grow and flowers over a
long period, but it can be aggressive under some conditions. Most of the other
species that are cultivated in the region are grown as annual groundcovers in
sunny sites. Among these, G. ×hybrida (Groenl. & Rumpler) G.L. Nesom
& Pruski (garden verbena) is the most striking taxon, a complex hybrid of
unconfirmed parentage (Pruski and Nesom, 1992) with relatively large flowers
(calyces 10–15 mm long, corollas 25–40 mm long), with the corollas
peach-colored to pink, blue, or white, but most commonly deep red. This hybrid
has escaped sporadically in a number of southeastern states and eventually may
be recorded as an escape in Missouri, although it does not appear to be very
commonly grown in the Midwest in recent decades.
Cronquist (1991)
cited the existence of moss verbena, G. pulchella (Sweet) Tronc. (as Verbena
tenuisecta Briq.), as an escape in St. Louis, but the specimens upon which
this report apparently was based were gathered from plants cultivated at the
Missouri Botanical Garden. Thus, this widely cultivated South American species
has been excluded for the present from the Missouri flora. It is somewhat
similar to G. bipinnatifida, but has 2 times pinnately deeply lobed
leaves with linear, tapered segments, the floral bracts shorter than the calyx
(thus the spikes appearing more slender), and brown nutlets that are beaked at
the tip.