Galanthus

General All species of Galanthus are perennial petaloid herbaceous bulbous (growing from bulbs) monocot plants. The genus is characterised by the presence of two leaves, pendulous white flowers with six free perianth segments in two whorls. The inner whorl is smaller than the outer whorl and has green markings. Vegetative ; Leaves: These are basal, emerging from the bulb initially enclosed in a tubular membranous sheath of cataphylls. Generally, these are two (sometimes three) in number and linear, strap-shaped, or oblanceolate. Vernation, the arrangement of the emerging leaves relative to each other, varies among species. These may be applanate (flat), supervolute (conduplicate), or explicative (pleated). In applanate vernation, the two leaf blades are pressed flat to each other within the bud and as they emerge; explicative leaves are also pressed flat against each other, but the edges of the leaves are folded back (externally recurved) or sometimes rolled; in supervolute plants, one leaf is tightly clasped around the other within the bud and generally remains at the point where the leaves emerge from the soil (for illustration, see Stearn and Davis). In the past, this feature has been used to distinguish between species and to determine the parentage of hybrids, but now has been shown to be homoplasious, and not useful in this regard. The scape (flowering stalk) is erect, leafless, terete, or compressed. Reproductive ;Inflorescence : At the top of the scape is a pair of bract-like spathes (valves) usually fused down one side and joined by a papery membrane, appearing monophyllous (single). From between the spathes emerges a solitary (rarely two), pendulous, nodding, bell-shaped white flower, held on a slender pedicel. The flower bears six free perianth segments (tepals) rather than true petals, arranged in two whorls of three, the outer whorl being larger and more convex than the inner whorl. The outer tepals are acute to more or less obtuse, spathulate or oblanceolate to narrowly obovate or linear, shortly clawed, and erect spreading. The inner tepals are much shorter (half to two thirds as long), oblong, spathulate or oblanceolate, somewhat unguiculate (claw like); tapering to the base and erect. These tepals also bear green markings at the base, the apex, or both, that when at the apex, are bridge-shaped over the small sinus (notch) at the tip of each tepal, which are emarginate. Occasionally, the markings are either green-yellow, yellow, or absent, and the shape and size varies by species. ; Androecium : The six stamens are inserted at the base of the perianth, and are very short (shorter than the inner perianth segments), the anthers basifixed (attached at their bases) with filaments much shorter than the anthers; they dehisce (open) by terminal pores or short slits. ; Gynoecium, fruit and seeds: The inferior ovary is three-celled. The style is slender and longer than the anthers; the stigma is minutely capitate. The ovary ripens into a three-celled capsule fruit. This fruit is fleshy, ellipsoid or almost spherical, opening by three flaps, with seeds that are light brown to white and oblong with a small appendage or tail (elaiosome) containing substances attractive to ants, which distribute the seeds. The chromosome number is 2n=24. Floral formula: \star\; P_{3+3} \; A_{3+3} \; G_{\overline{(3)}} ==Distribution and habitat==

The genus Galanthus is native to Europe and the Middle East, from the Spanish and French Pyrenees in the west through to the Caucasus and Iran in the east, and south to Sicily, the Peloponnese, the Aegean, Turkey, Lebanon, and Syria. The northern limit is uncertain because G. nivalis has been widely introduced and cultivated throughout Europe. G. nivalis and some other species valued as ornamentals have become widely naturalised in Europe, North America, and other regions. In the Udmurt republic of Russia, Galanthus are found even above the 56th parallel. Galanthus nivalis is the best-known and most widespread representative of the genus Galanthus. It is native to a large area of Europe, stretching from the Pyrenees in the west, through France and Germany to Poland in the north, Italy, northern Greece, Bulgaria, Romania, Ukraine, and European Turkey. It has been introduced and is widely naturalised elsewhere. Although it is often thought of as a British native wild flower, or to have been brought to the British Isles by the Romans, it most likely was introduced around the early sixteenth century, and is currently not a protected species in the UK. It was first recorded as naturalised in the UK in Worcestershire and Gloucestershire in 1770. Most other Galanthus species are from the eastern Mediterranean, while several are found in the Caucasus, in southern Russia, Georgia, Armenia, and Azerbaijan. Galanthus fosteri is found in Jordan, Lebanon, Syria, Turkey, and, perhaps, Palestine. Most Galanthus species grow best in woodland, in acid or alkaline soil, although some are grassland or mountain species. ==Taxonomy==

History Early 's Herball, 1597 Snowdrops have been known since early times, being described by the classical Greek author Theophrastus, in the fourth century BCE, in his Περὶ φυτῶν ἱστορία (Latin: Historia plantarum, Enquiry into plants). He gave it, and similar plants, the name λευκόἲον (λευκος, leukos "white" and ἰόν, ion "violet") from which the later name Leucojum was derived. He described the plant as "ἑπεἰ τοῖς γε χρώμασι λευκἂ καἱ οὐ λεπυριώδη" (in colour white and bulbs without scales) Grown as ornamental. • Crimean snowdrop, Galanthus plicatus, 30 cm tall, flowering January/March, white flowers, with broad leaves folded back at the edges (explicative vernation). • Giant snowdrop, Galanthus elwesii, a native of the Levant, 23 cm tall, flowering January/February, with large flowers, the three inner segments of which often have a much larger and more conspicuous green blotch (or blotches) than the more common kinds; supervolute vernation. Grown as ornamental. • Galanthus reginae-olgae, from Greece and Sicily, is quite similar in appearance to G. nivalis, but flowers in autumn before the leaves appear. The leaves, which appear in the spring, have a characteristic white stripe on their upper side; applanate vernation. • G. reginae-olgae subsp. vernalis, from Sicily, northern Greece and the southern part of former Yugoslavia, blooms at the end of the winter with developed young leaves and is thus easily confused with G. nivalis. Etymology Galanthus is derived from the Greek γάλα (gala), meaning "milk" and ἄνθος (anthos) meaning "flower", alluding to the colour of the flowers. The epithet nivalis is derived from the Latin, meaning "of the snow". The word "Snowdrop" may be derived from the German Schneetropfen (snow-drop), the tear drop shaped pearl earrings popular in the sixteenth and seventeenth centuries. Other, earlier, common names include Candlemas bells, Fair maids of February, and White ladies (see Symbols). == Ecology ==

Snowdrops are hardy herbaceous plants that perennate by underground bulbs. They are among the earliest spring bulbs to bloom, although a few forms of G. nivalis are autumn flowering. They naturalise relatively easily forming large drifts. These are often sterile, found near human habitation, and also former monastic sites. The leaves die back a few weeks after the flowers have faded. Galanthus plants are relatively vigorous and may spread rapidly by forming bulb offsets. They also spread by dispersal of seed, animals disturbing bulbs, and water if disturbed by floods. ==Conservation==

Some snowdrop species are threatened in their wild habitats, due to habitat destruction, illegal collecting, and climate change. In most countries collecting bulbs from the wild is now illegal. Under CITES regulations, international trade in any quantity of Galanthus, whether bulbs, live plants, or even dead ones, is illegal without a CITES permit. This applies to hybrids and named cultivars, as well as species. CITES lists all species, but allows a limited trade in wild-collected bulbs of just three species (G. nivalis, G. elwesii, and G. woronowii) from Turkey and Georgia. A number of species are on the IUCN Red List of threatened species, with the conservation status being G. trojanus as critically endangered, four species vulnerable, G. nivalis is near threatened and several species show decreasing populations. G. panjutinii is considered endangered. One of its five known sites, at Sochi, was destroyed by preparations for the 2014 Winter Olympics. ==Cultivation==

Galanthus species and cultivars are extremely popular as symbols of spring and are traded more than any other wild-source ornamental bulb genus. Millions of bulbs are exported annually from Turkey and Georgia. For instance export quotas for 2016 for G. elwesii were 7 million for Turkey. Several gardens open specially in February for visitors to admire the flowers. Sixty gardens took part in Scotland's first Snowdrop Festival (1 Feb–11 March 2007). Several gardens in England open during snowdrop season for the National Gardens Scheme (NGS) and in Scotland for Scotland's Gardens. Colesbourne Park in Gloucestershire is one of the best known of the English snowdrop gardens, being the home of Henry John Elwes, a collector of Galanthus specimens, and after whom Galanthus elwesii is named. Cultivars Numerous single- and double-flowered cultivars of Galanthus nivalis are known, and also of several other Galanthus species, particularly G. plicatus and G. elwesii. Also, many hybrids between these and other species exist (more than 500 cultivars are described in Bishop, Davis, and Grimshaw's book, plus lists of many cultivars that have now been lost, and others not seen by the authors). They differ particularly in the size, shape, and markings of the flower, the period of flowering, and other characteristics, mainly of interest to the keen (even fanatical) snowdrop collectors, known as "galanthophiles", who hold meetings where the scarcer cultivars change hands.  Double-flowered cultivars and forms, such as the extremely common Galanthus nivalis f. pleniflorus 'Flore Pleno', may be less attractive to some people, but they can have greater visual impact in a garden setting. Cultivars with yellow markings and ovaries rather than the usual green are also grown, such as 'Wendy's Gold'. Many hybrids have also occurred in cultivation. Awards , the following have gained the Royal Horticultural Society's Award of Garden Merit: • Galanthus 'Ailwyn' • Galanthus 'Atkinsii' • Galanthus 'Bertram Anderson' • Galanthus elwesii • Galanthus elwesii 'Comet' • Galanthus elwesii 'Godfrey Owen' • Galanthus elwesii 'Mrs Macnamara' • Galanthus elwesii var. monostictus • Galanthus 'John Gray' • Galanthus 'Lady Beatrix Stanley' • Galanthus 'Magnet' • Galanthus 'Merlin' • Galanthus nivalis • Galanthus nivalis f. pleniflorus 'Flore Pleno' • Galanthus nivalis 'Viridapice' • Galanthus plicatus • Galanthus plicatus 'Augustus' • Galanthus plicatus 'Diggory' • Galanthus plicatus 'Three Ships' • Galanthus reginae-olgae subsp. reginae-olgae • Galanthus 'S. Arnott' • Galanthus 'Spindlestone Surprise' • Galanthus 'Straffan' • Galanthus 'Trumps' • Galanthus woronowii Propagation Propagation is by offset bulbs, either by careful division of clumps in full growth ("in the green"), or removed when the plants are dormant, immediately after the leaves have withered; or by seeds sown either when ripe, or in spring. Professional growers and keen amateurs also use such methods as "twin-scaling" to increase the stock of choice cultivars quickly. == Toxicity ==

Snowdrops contain an active lectin or agglutinin named GNA for Galanthus nivalis agglutinin. == Medicinal use ==

In 1983, Andreas Plaitakis and Roger Duvoisin suggested that the mysterious magical herb, moly, that appears in Homer's Odyssey is the snowdrop. One of the active principles present in the snowdrop is the alkaloid galantamine, which, as an acetylcholinesterase inhibitor, could have acted as an antidote to Circe's poisons. Further supporting this notion are notes made during the fourth century BC by the Greek scholar Theophrastus who wrote in Historia plantarum that moly was "used as an antidote against poisons" although which specific poisons it was effective against remains unclear. Galantamine (or galanthamine) may be helpful in the treatment of Alzheimer's disease, although it is not a cure; the substance also occurs naturally in daffodils and other narcissi. ==In popular culture==

Snowdrops figure prominently in art and literature, • In the fairy-tale play The Twelve Months by Russian writer Samuil Marshak, a greedy queen decrees that a basket of gold coins shall be rewarded to anyone who can bring her galanthus flowers in the dead of winter. A young orphan girl is sent out during a snow storm by her cruel stepmother to find the spirits of the 12 months of the year, who take pity on her and not only save her from freezing to death, but also make it possible for her to gather the flowers even in winter. The Soviet traditionally animated film The Twelve Months (1956), Lenfilm film The Twelve Months (1972), and the anime film Twelve Months (1980) (Sekai meisaku dowa mori wa ikiteiru in Japan), are based on this fairy-tale play. • "Snowdrops" was the nickname that the British people gave during the Second World War to the military police of the United States Army (who were stationed in the UK preparatory to the invasion of the continent) because they wore a white helmet, gloves, gaiters, and Sam Browne belt against their olive drab uniforms. • In the German fairy tale, Snow White and the Seven Dwarfs, "Snowdrop" is used as an alternate name for the Princess Snow White. • The short story The Snowdrop by Hans Christian Andersen follows the fate of a snowdrop from a bulb striving toward the light to a picked flower placed in a book of poetry. • Russian composer Tchaikovsky wrote a series of 12 piano pieces, each one named after a month of the year with a second name suggesting something associated with that month. His "April" piece is subnamed "Snow Drop". The Russian climate having a later spring, and winter ending a bit later than in other places. • Johann Strauss II named his very successful waltz Schneeglöckchen (Snowdrops) op. 143 after this flower. The inspiration is especially evident in the cello introduction and in the slow unfurling of the opening waltz. Strauss composed this piece for a Russian Embassy dinner given at the Sperl ballroom in Vienna on 2 December 1853, but did not perform it publicly until the year 1854. The Sperl banquet was given in honour of her Excellency Frau Maria von Kalergis, daughter of the Russian diplomat and foreign minister Count Karl Nesselrode, and Strauss also dedicated his waltz to her. • In the 2007 film, Stardust, a glass snowdrop flower is gifted to the female protagonist, and serves to protect the bearer from all Witches' magic and bring them good luck. Symbolism Early names refer to the association with the religious feast of Candlemas (February 2) – the optimum flowering time of the plant – at which young women, robed in white, would walk in solemn procession in commemoration of the Purification of the Virgin, an alternative name for the feast day. The French name of refers to Candlemas, while an Italian name, , refers to purification. The German name of (little snow bells) invokes the symbol of bells. ==See also==