Fritillaria

Fritillaria (fritillaries) is a genus of spring flowering herbaceous bulbous perennial plants in the lily family (Liliaceae). These beautiful flowers grow from bulbs and bloom in the spring, often nodding downward with their bell-shaped blossoms. The type species, Fritillaria meleagris, was first described in Europe in 1571 and quickly became a favorite among garden lovers.

Found across the temperate regions of the Northern Hemisphere, from the Mediterranean and North Africa through Eurasia to western North America, fritillaries grow in many different habitats. Sadly, many species are endangered because people pick them too much in the wild.

The name Fritillaria may come from the checkered pattern on F. meleagris, which looks like a box for carrying dice. Today, these plants are important in horticulture as popular ornamental garden plants and are also used in traditional Chinese medicine. Their lovely flowers have inspired artists and have become symbols for various regions and groups.

Description

Fritillaria is a genus of perennial herbaceous bulbous plants that die back after flowering and regrow from underground bulbs each year. They are known for their nodding (drooping) flowers with bell-shaped perianths and unique nectar structures.

These plants have bulbs made of tightly packed scales, and their stems bear leaves arranged in pairs or whorls. Their flowers are usually solitary and droop downward, with six tepals that can be white, yellow, green, purple, or reddish. The flowers have both male and female parts, and they produce fruit in the form of a capsule containing flattened seeds. Some Fritillaria species have strong, unusual scents, while others have sweet fragrances.

Taxonomy

The genus Fritillaria, commonly called fritillaries, consists of about 130–140 species of spring-flowering herbaceous bulbous perennial plants in the lily family, Liliaceae. The type species, Fritillaria meleagris, was first documented in Europe in 1571. Other species from the Middle East and Asia were also introduced to Europe around the same time.

Fritillaria species were known in Persia (modern-day Iran) in the sixteenth century and were later brought to Turkey. European travelers collected these plants along with many other exotic species for the botanical gardens of Europe. By the mid-sixteenth century, there was already a trade in bulbs from Turkey to Europe.

Subgenus Liliorhiza (Kellog) Benth. & Hook.f. This subgenus of North American Fritillaria, centred on California constitutes Clade A and contains more than 20 species. They have distinctive bulbs with several imbricate (overlapping) scales. Their resemblance to the bulbs of Lilium lends this subgenus the name of Liliorhiza, or lily-roots. Most species exhibit loosely attached bulbils, whose size and shape gave them the name of "rice-grain bulbils". These are also seen in subgenus Davidii, but appear to have evolved independently. Stem leaves are whorled.

Subgenus Petilium (L.) Endl. Subgenus Petilium forms a subclade (B1), together with subgenera Korolkowia and Theresia. It is a relatively small subgenus of four species characterized by large (up to 100 cm) sturdy species, with bulbs that are much larger (up to 8 cm) than most Fritillaria with a few large, erect, imbricate and fleshy scales. Flowers are 3–5 in a terminal umbel, in the axils of the lower side of a leaf whorl. They have a trifid (3 lobed) style and winged seeds. The subgenus is found in Turkey, Iraq, Turkestan, Iran, Baluchistan, Afghanistan, and the western Himalayas. The best known example is F. imperialis (crown imperial).

Subgenus Rhinopetalum Fisch. Rhinopetalum together with subgenera Japonica and Fritillaria constitute subclade B2. It is a small subgenus of five species. They are characterized by their nectaries, which are deeply impressed and have a slit-like orifice on the tepals.

Subgenus Japonica Rix Japonica consists of eight species. Rix characterised the subgenus as follows: "Bulb of 2 or 3 solid scales, without rice grains; plants small and delicate; seeds pear-shaped, Erythronium-like, not flat; stem collapsing when seeds are ripe. Japanese woodland plants". The species in this subgenus are dwarf fritillaries, endemic to Japan, bearing a single small campanulate flower on a slender stem with three linear verticillate (in one or more whorls) leaves at its top and two broader, oblong to elliptical and opposite leaves about 1 cm below these. The flower is born on a short pedicel amongst the leaves.

Subgenus Fritillaria Fritillaria is the largest subgenus, with about 100 species, or more than 70% of the total number of species in the genus, and includes the type species, F. meleagris. They are widely distributed from western Europe and the Mediterranean region to eastern Asia. Their characteristic is the Fritillaria-type bulb. This consists of two fleshy more or less tunicated scales that are subglobose. The tunica is formed by the remains of previous years' scales, but sometimes the previous scales persist leading to more than two scales, sometimes three or four. The style may be trifid or undivided, or only trilobulate at the apex (a characteristic that was previously thought to divide the subgenus into sections).

Biogeography and evolution

It is likely that two groups of plants, including Fritillaria, moved across the Bering Straits to reach North America. In Eurasia, different types of bulbs developed in various subgroups. Some Fritillaria bulbs have 2–3 fleshy layers, while others, like Theresia and Korolkowia, have a single large fleshy layer. These special small bulbs, called bulbils, help the plants grow new ones without seeds, giving them an advantage in spreading.

Etymology

When an apothecary named Noël Capperon found F. meleagris in the meadows of the Loire in 1570, he noted that locals called it fritillaria. He thought this was because the flower's checkered pattern looked like the board used for the game checkers. However, another scientist, Carolus Clusius, believed this was a mistake, explaining that the true meaning related to a box for dice, not the game board itself.

In North America, some species of fritillaria are known as "mission bells".

Distribution and habitat

Fritillaria plants grow in many parts of the world, mainly in temperate areas of the Northern Hemisphere. They can be found from western North America through Europe, the Mediterranean, the Middle East, and Central Asia all the way to China and Japan. Some places, like Turkey and the Zagros Mountains in Iran, have many different kinds of Fritillaria.

These plants live in various climates and habitats, but many prefer sunny, open spaces. Some types of Fritillaria have been brought to new places by people. For example, F. meleagris has been grown in British gardens since 1578, and F. imperialis was introduced to Europe in the 1570s.

Ecology

Most fritillaries bloom in the spring. Some insects, like the scarlet lily beetle, like to eat these plants and can sometimes cause problems for gardeners.

These flowers are pollinated by insects. Some are pollinated by wasps, while others are pollinated by bumblebees, depending on the size and sweetness of their nectar.

Conservation

Some Fritillaria plants are in danger because people pick too many of them, their homes are destroyed, and animals eat them. This is especially true in places like Greece, Japan, and China. For example, in China, people take bulbs from Fritillaria cirrhosa to use in medicine, which makes it very rare.

In Iran, F. imperialis and F. persica are also at risk. One species, F. delavayi, is changing its flower color to blend in with rocks to avoid being picked. This shows how some plants try to protect themselves when people take too many of them.

Toxicity

Most fritillaries have poisonous substances in their bulbs. These substances, called steroidal alkaloids, can be harmful if eaten in large amounts. One example of such a substance is imperialin. It's important to handle these plants carefully and never eat them.

Uses

Some species of Fritillaria have bulbs that can be eaten safely if prepared properly. Indigenous peoples along the Pacific Northwest coast of North America used to eat these bulbs. In places like Iran, people visit areas such as the Valley of Roses in late May to see the beautiful flowers of F. imperialis bloom.

Fritillaria plants are also valued for their beauty in gardens and for their medicinal properties. They are popular ornamental plants, and many types are available for gardeners to grow. In traditional medicine, especially in China, parts of these plants have been used for thousands of years to help with respiratory issues. However, the demand for these plants has led to some wild populations being harvested too heavily.

Popular culture

Fritillaries have been loved and written about by famous writers like Shakespeare, Matthew Arnold, George Herbert, and Vita Sackville-West. They were also a favorite subject for Dutch flower painters from around 1600, such as Ambrosius Bosschaert and Jacob de Gheyn II, and appeared in Italian art by Jacopo Ligozzi.

These flowers are used as symbols in many places. For example, F. meleagris is the county flower of Oxfordshire in the UK and the provincial flower of Uppland in Sweden. In Japan, F. camschatcensis is a floral emblem for Ishikawa Prefecture and Obihiro City.