3. Callitriche
L. (water starwort)
(Philbrick, 1989; N. G. Miller, 2001)
Contributed by Alan E. Brant
Plants annual or
less commonly perennial herbs, monoecious, glabrous but sometimes appearing
minutely scaly or yellowish-glandular spotted at high magnification. Stems
simple to much branched, often rooting at the nodes. Leaves opposite or
sometimes appearing whorled at the stem tip, sessile or short- to
long-petiolate. Leaf blades simple, entire, linear to spatulate, the bases
mostly connected around the stem by a narrow herbaceous membrane. Stipules
absent. Flowers imperfect, tiny, solitary or in small clusters of 2 or 3 per
leaf axil, the staminate and pistillate flowers usually together in the same
axil, in some species subtended by a pair of minute bracts. Perianth absent.
Staminate flower(s) maturing before the adjacent pistillate flower(s),
consisting of 1 stamen, the slender filament attached at the anther base.
Pistillate flower(s) consisting of 1 pistil, the ovary superior, of 2 carpels,
somewhat flattened and 4-lobed, each carpel with 2 ovules. Styles 2, elongate,
often persistent at fruiting, the stigmatic region toward the slender tip.
Fruits depressed-elliptic or more or less heart-shaped in outline, the ovary
separating into 4 achenelike nutlets, these oblong-elliptic to kidney-shaped or
obovate in outline, flattened and sometimes narrowly winged, the surface with a
very fine network of ridges (observable only with magnification), light brown
to brown. About 50 species, nearly worldwide.
Some species of
water starwort are used horticulturally in outdoor pools and ponds. The herbage
is eaten by waterfowl and some species of fish. The seeds are dispersed by
water and by animals, and they reportedly survive passage through the digestive
system of ducks. Studies have shown that species of Callitriche have
developed an unusual breeding system where pollen germinates within intact,
unopened anthers, and the pollen tubes grow downward inside the filament and
then through the vegetative tissue and into adjacent pistillate flowers to
effect pollination (Philbrick, 1984). This presumably helps to ensure abundant
seed production in populations where extreme and rapid environmental
fluctuations occur. Also, this helps to explain observations of seed production
in plants whose stamens appear to be rudimentary and incapable of dehiscing to
release pollen.
As with many
groups of aquatic angiosperms having reduced floral structures, the
relationships of Callitriche have been controversial. Traditionally, the
most widely accepted alignment was as a separate family, Callitrichaceae, close
to the Euphorbiaceae (Steyermark, 1963), presumably because of superficial
similarities in fruit morphology. Studies in ovule and embryo development
suggested a placement as a separate family close to the Lamiaceae and
Verbenaceae, which has been followed more recently by a number of botanists
(Cronquist, 1981, 1991). The most recent evidence from molecular studies has
classified Callitriche into the expanded Plantaginaceae. For further
discussion, see the paragraph under the family description above.