Boraginaceae
Contents
Size
Distribution
Habit
Leaves
Inflorescence
Flower
Fruit
Importance
Relationships

Genera in northern Utah
(genera with native species in green)
Amsinckia
Anchusa
Asperugo
Borago
Cryptantha

Cynoglossum
Echium
Eritrichium
Hackelia
Heliotropium
Lappula
Lithospermum
Mertensia

Mysotis
Plagiobothrys
Symphytum

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The Boraginaceae has around 2,000 species in total but only 50 in our part of the world; of these, 43 are native species. It is worldwide in its distribution, with particularly high concentrations in western North Amera (that includes us) and from the Mediterranean into Asia. 

Globally speaking, the Boraginaceae includes herbs, shrubs, trees and even a few lianas, but all our species are herbaceous.  Most of our species are easily recognized, being covered with rather coarse hairs and having a scorpioid or helicoid cyme for an inflorescence. The flowers are radially symmetric, and the fruit consists of four nutlets. Once familiar with what all these terms mean, the Boraginaceae is easy to recognize.

The strigose hairs are often found throughout the vegetative portion of the plant, and even on the rachis and calyces of the inflorescence. The leaves are simple, estipulate, and alternate.

The inflorescence is usally a scorpioid or helicoid cyme.  This means that it is a determinate inflorescence in which development of the inflorescence is reminiscent of the circinate vernation associated with ferns. What makes the inflorescence of Boraginaceae distinctive (but not unique) is that, as each flower matures, the pedicel supporting it departs at a slight angle from the main stem and elongates until, at flowering, it is often above the flower before it in the flowering sequence.  This makes the inflorescence appear as if it is flowering from the base up.  In other words, it mimics an indeterminate inflorescence.  This, in combination with gynandrous floral development, promotes outcrossing because many pollinating insects work an inflorescence from the base up.

The calyx is 5-merous and usually consists of separate sepals, but it may be united at the base. It is usually radially symmetric. The corolla is also 5-merous and usually radially symmetric (but bilaterally symmetric in a couple of well known genera), and often consists of a short, narrow tube that changes abruptly to the flat, plate-like top. At the base of the corolla lobes and surrounding the entrance to the corolla tube there is often a set of 5 fornices (singular 'fornix') which form a kind of wall around the throat.

There are five epipetalous stamens, all attached to the corolla at the same level. The gynoecium consists of a single, 2 carpellate pistil. The fact that it is 2-carpellate is not always obvious at maturity because of the way the ovary changes as it matures, but may be indicated by the branching of the tip of the style. So what does the ovary do? When very young, the ovary consists of two chambers, each of which has 2 ovules. In most species, the two halves of the ovary are almost completely separate from each other, often connecting to the style at their base. As the fertilized ovules mature, the ovary wall fits more and more closely around each seed so that, at maturity, one sees four 1-seeded nutlets. There are very few families that form such a fruit; in our region, it occurs only in the Boraginaceae, Lamiaceae, and Verbenaceae, so finding it is very useful in determining the family of a plant. A plant with radially symmetric flowers and a fruit of four nutlets is almost bound to be a borage, because flowers of the Lamiaceae are always bilaterally symmetric, and there are not many Verbenaceae around here.

The nutlets are often important for generic identification in the Boraginaceae. I strongly recommend waiting until you can obtain some fruiting specimens before collecting members of this family.

The Boraginaceae has little economic importance, although Borago officinale, borage, used to be used as a herb but is more often grown as a source of nectar for bees nowadays, and Symphytum officinale is often touted as a potherb. In our region, Myosotis or forget-me-not is grown as an ornamental; I cannot remember the derivation of its common name. Mertensia is an attractive genus in montane regions of northern Utah; other genera are most impressive for the ability of their nutlets to cling to one's clothes. Echium plantagineum is known as Paterson's Curse in Australia where it is a widespread weed.  It is not good for horses, but bees love it. 

Cronquist placed the Boraginaceae in his Asteridae.  It is generally considered to be closely related to the Hydrophyllaceae, Verbenaceae, and Lamiaceae.  The last two families have a similar fruit.  The Hydrophyllaceae often has a similar inflorescence.

Amsinckiaretrorsa.jpg (16057 bytes)
Anchusa retrorsa
Note inflorescence
Boragoofficinalis.jpg (14723 bytes)
Borago officinalis
Radially symmetric flower with fornices (light) around protruding (dark) anthers
Cryptantha confertiflora.jpg (36184 bytes)
Cryptantha confertiflora
A widespread, speciose genus in Utah
Echiumplantagineum.jpg (12659 bytes)

Echium plantagineum
A bilaterally symmetric flower
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