2. Humulus L. (hops)
Plants annual or
perennial herbs, with long taproots or stout rhizomes respectively, not
strongly aromatic, the staminate and pistillate plants appearing similar
vegetatively. Stems twining, branched (except in very depauperate individuals),
usually finely ridged or angled, solid at maturity, sparsely to densely
rough-pubescent with stiff, prickly 2-branched hairs. Leaves with the petioles
shorter than to longer than the blades, sometimes twining, pubescent with
short-stalked, 2-branched hairs. Stipules lanceolate, sometimes fused laterally
and then appearing solitary rather than paired. Leaf blades ovate to nearly
circular in outline, unlobed or palmately 3–9-lobed, deeply cordate at the
base, the lobes mostly somewhat narrowed toward the base, tapered to a sharply
pointed tip, the margins sharply toothed and sometimes hairy, the surfaces
variously hairy and glandular. Staminate inflorescences appearing as open
axillary or less commonly terminal panicles with small flower clusters at the
branch tips, the flowers with stalks 0.5–3.5 mm long, the branch points and
flowers subtended by small bracts. Pistillate inflorescences short, dense
spikes continuing to elongate as the fruits mature, pendant at maturity,
conelike, the solitary or paired flowers sessile or nearly so, the bracts
elliptic or ovate to broadly ovate, sometimes hairy on the surface and along
the margins, also with glandular dots and/or short-stalked glands cup-shaped at
the tip. Staminate flowers 1.5–3.0 mm long, the sepals lanceolate to ovate or
oblong-ovate, with pale, thin margins, glabrous or less commonly sparsely
hairy, also with glandular dots. Fruits broadly ovoid to nearly spherical,
sometimes slightly flattened, enveloped by the persistent enlarged bracts and
membranous calyx. Three species, North America, Europe, Asia, widely cultivated
and introduced nearly worldwide.
The branched
hairs consist of two short, rigid branches spreading in opposite directions.
These are raised from the surface on short, multicellular bases containing
cystoliths (calcium carbonate inclusions) that appear as small tubercles. This
arrangement apparently is an adaptation to help anchor the plants as they
climb; they impart a strong, prickly roughness to the surfaces of the stems and
leaves.