Snowdrops have been known since the earliest times under various names, but were namedGalanthus in 1753. As the number of recognised species increased, various attempts were made to divide the species into subgroups, usually on the basis of the pattern of the emerging leaves (vernation). In the era ofmolecular phylogenetics this characteristic has been shown to be unreliable and now seven molecularly definedclades are recognised that correspond to thebiogeographical distribution of species. New species continue to be discovered.
These are basal, emerging from the bulb initially enclosed in a tubular membranous sheath ofcataphylls. Generally, these are two (sometimes three) in number and linear, strap-shaped, oroblanceolate.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[4] (for illustration, see Stearn[5] and Davis[6]). 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 behomoplasious, and not useful in this regard.
Thescape (flowering stalk) is erect, leafless,terete, or compressed.[7]
At the top of thescape is a pair ofbract-likespathes (valves) usually fused down one side and joined by a papery membrane, appearingmonophyllous (single). From between the spathes emerges a solitary (rarely two), pendulous, nodding, bell-shaped white flower, held on a slenderpedicel. The flower bears six freeperianth segments (tepals) rather than truepetals, arranged in two whorls of three, the outer whorl being larger and more convex than the inner whorl. The outer tepals areacute to more or lessobtuse,spathulate or oblanceolate to narrowlyobovate or linear, shortly clawed, and erect spreading. The inner tepals are much shorter (half to two thirds as long), oblong, spathulate or oblanceolate, somewhatunguiculate (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 smallsinus (notch) at the tip of each tepal, which areemarginate. Occasionally, the markings are either green-yellow, yellow, or absent, and the shape and size varies by species.[8][7]
Androecium
The sixstamens 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; theydehisce (open) by terminalpores or short slits.[7]
Gynoecium, fruit and seeds
Theinferior ovary is three-celled. Thestyle is slender and longer than the anthers; thestigma is minutelycapitate. The ovary ripens into a three-celledcapsule 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 toants, which distribute the seeds.[7][9]
Distribution map ofGalanthus species in Europe and Western Asia
The genusGalanthus is native to Europe and the Middle East, from the Spanish and FrenchPyrenees in the west through to theCaucasus andIran in the east, and south toSicily, thePeloponnese, theAegean,Turkey,Lebanon, andSyria. The northern limit is uncertain becauseG. nivalis has been widelyintroduced andcultivated throughout Europe.[3]G. nivalis and some other species valued asornamentals have become widelynaturalised in Europe, North America, and other regions.[2] In the Udmurt republic of Russia,Galanthus are found even above the 56th parallel.[citation needed]
Galanthus nivalis is the best-known and most widespread representative of the genusGalanthus. 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, andEuropean Turkey. It has been introduced and is widely naturalised elsewhere.[11] 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.[12] It was first recorded as naturalised in the UK inWorcestershire andGloucestershire in 1770.[13] Most otherGalanthus species are from the eastern Mediterranean, while several are found in theCaucasus, in southern Russia, Georgia, Armenia, and Azerbaijan.[14]Galanthus fosteri is found in Jordan, Lebanon, Syria, Turkey, and, perhaps,Palestine.[15]
MostGalanthus species grow best inwoodland, inacid or alkaline soil,[7] although some aregrassland or mountain species.
Snowdrops have been known since early times, being described by the classicalGreek authorTheophrastus, in the fourth century BCE, in hisΠερὶ 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 nameLeucojum was derived. He described the plant as "ἑπεἰ τοῖς γε χρώμασι λευκἂ καἱ οὐ λεπυριώδη" (in colour white and bulbs without scales)[16] and of their habits "Ἰῶν δ' ἁνθῶν τὀ μἑν πρῶτον ἑκφαἱνεται τὁ λευκόἲον, ὅπου μἑν ό ἀἠρ μαλακώτερος εὐθὑς τοῦ χειμῶνος, ὅπου δἐ σκληρότερος ὕστερον, ἑνιαχοῡ τοῡ ἣρος" (Of the flowers, the first to appear is the white violet. Where the climate is mild, it appears with the first sign of winter, but in more severe climates, later in spring)[17]
Rembert Dodoens, aFlemish botanist, described and illustrated this plant in 1583 as didGerard in England in 1597 (probably using much of Dodoens' material), calling itLeucojum bulbosum praecox (Early bulbous violet). Gerard refers to Theophrastus's description asViola alba orViola bulbosa, usingPliny's translation, and comments that the plant had originated in Italy and had "taken possession" in England "many years past".[18] Thegenus was formally namedGalanthus and described byCarl Linnaeus in 1753,[19] with the singlespecies,Galanthus nivalis, which is thetype species. Consequently, Linnaeus is granted thebotanical authority. In doing so, he distinguished this genus and species fromLeucojum (Leucojum bulbosum trifolium minus), a name by which it previously had been known.[2][20][21]
In 1763Michel Adanson began a system of arranging genera in families. Using thesynonymAcrocorion (also speltAkrokorion),[22] he placedGalanthus in the familyLiliaceae, section Narcissi.[23]Lamarck provided a description of the genus in his encyclopedia (1786),[24] and later,Illustrations des genres (1793).[25] In 1789de Jussieu, who is credited with the modern concept of genera organised infamilies, placedGalanthus and related genera within adivision ofMonocotyledons, using a modified form of Linnaeus' sexual classification, but with the respective topography of stamens to carpels rather than just their numbers. In doing so, he restored the nameGalanthus and retained their placement under Narcissi, this time as a family (known asOrdo, at that time) and referred to the French vernacular name,Perce-neige[26] (Snow-pierce), based on the plants tendency to push through early spring snow (seeEcology for illustration).[27] The modern family ofAmaryllidaceae, in whichGalanthus is placed, dates toJaume Saint-Hilaire (1805) who replaced Jussieu's Narcissi withAmaryllidées.[28] In 1810,Brown proposed that a subgroup of Liliaceae be distinguished on the basis of the position of theovaries and be referred to as Amaryllideae,[29] and in 1813,de Candolle separated them by describing LiliacéesJuss. and AmaryllidéesBrown as two quite separate families.[30] However, in his comprehensive survey of theFlora of France (Flore française, 1805–1815) he divided Liliaceae into a series ofOrdres, and placed Galanthus into the NarcissiOrdre.[31] This relationship of Galanthus to either liliaceous or amaryllidaceaous taxa (seeTaxonomy of Liliaceae) was to last for another two centuries until the two were formally divided at the end of the twentieth century.[32]Lindley (1830) followed this general pattern, placingGalanthus and related genera such asAmaryllis andNarcissus in his Amaryllideae (which he called The Narcissus Tribe in English).[33] By 1853, the number of known plants was increasing considerably and he revised his schema in his last work, placingGalanthus together, and the other two genera in the modernGalantheae in tribe Amarylleae, order Amaryllidaceae, alliance Narcissales.[34] These three genera have been treated together taxonomically by most authors, on the basis of an inferior ovary. As the number of plant species increased, so did the taxonomic complexity. By the timeBentham andHooker published theirGenera plantarum (1862–1883)[35] ordo Amaryllideae[36] contained five tribes, and tribe Amarylleae[37] three subtribes (seeBentham & Hooker system). They placedGalanthus in subtribe Genuinae and included three species.[38]
Cladogram of evolutionary lines inGalanthussensu Ronstedet al.[3]
Galantheae
Galanthus
Acis
Leucojum
Galanthus is one of three closely related genera making up the tribeGalantheae within subfamilyAmaryllidoideae (familyAmaryllidaceae). Sometimes snowdrops are confused with the other two genera,Leucojum andAcis (both called snowflakes).Leucojum species are much larger and flower in spring (or early summer, depending on the species), with all sixtepals in the flower being the same size, although some "poculiform" (goblet- or cup-shaped)Galanthus species may have inner segments similar in shape and length to the outer ones. Galantheae are likely to have arisen in the Caucusus.[39]
The three different forms of leaf shoots from the bulb: flat (applanate), folded (explicative) rolled up (convolute) (left to right)Galanthus elwesiiGalanthus nivalis 'Viridapice'Galanthus plicatusGalanthus woronowii
Galanthus has approximately 20 species, but new species continue to be described.[2]G. trojanus was identified in Turkey in 2001.[40][41]G. panjutinii (Panjutin's snowdrop)[42] was discovered in 2012 in five locations in a small area (estimated at 20 km2 (7.7 sq mi)) of the northernColchis area (westernTranscaucasus) ofGeorgia andRussia.[43][44]G. samothracicus was identified in Greece in 2014. Since it has not been subjected to genetic sequencing, it remains unplaced. It resemblesG. nivalis, but is outside the distribution of that species.[45][46]
Many species are difficult to identify, however, and traditional infrageneric classification based onplant morphology alone, such as those ofStern (1956),[47]Traub (1963)[48] and Davis[49] (1999, 2001),[50][51][7] has not reflected what is known about itsevolutionary history, due to the morphological similarities among the species and relative lack of easily discernible distinguishing characteristics.[52][53][54] Stern divided the genus into three series according to leafvernation (the way the leaves are folded in the bud, when viewed in transverse section, seeDescription);[47]
Stern further utilised characteristics such as the markings of the inner segments, length of the pedicels in relation to the spathe, and the colour and shape of the leaves in identifying and classifying species
By contrast Davis, with much more information and specimens, includedbiogeography in addition to vernation, forming two series. He used somewhat different terminology for vernation, namely applanate (flat), explicative (plicate), and supervolute (convolute). He mergedNivalis andPlicati into seriesGalanthus, and dividedLatifolii into two subseries,Glaucaefolii(Kem.-Nath) A.P.Davis andViridifolii(Kem.-Nath) A.P.Davis.[50]
Earlymolecular phylogenetic studies confirmed the genus wasmonophyletic and suggested fourclades, which were labelled asseries, and showed that Davis' subseries were not monophyletic.[53][54] An expanded study in 2013 demonstrated seven major clades, corresponding tobiogeographical distribution. This study used nuclear encoded nrITS (Nuclear ribosomal internal transcribed spacer), andplastid encoded genesmatK (Maturase K),trnL-F,ndhF, andpsbK–psbI, and examined all species recognised at the time, as well as two naturally occurring putativehybrids. The morphological characteristic of vernation that earlier authors had mainly relied on was shown to be highlyhomoplasious. A number of species, such asG. nivalis andG. elwesii demonstrated intraspecific biogeographical clades, indicating problems withspeciation and there may be a need forrecircumscription. These clades were assigned names, partly according to Davis' previous groupings. In this model clade, the group containingG. platyphyllus issister to the rest of the genus.[3]
By contrast, another study performed at the same time, using bothnuclear andchloroplast DNA, but limited to the 14 species found in Turkey, largely confirmed Davis' series and subseries, and with biogeographical correlation. SeriesGalanthus in this study corresponded to clade nivalis, subseriesGlaucaefolii with clade Elwesii and subseriesViridifolii with clades Woronowii and Alpinus. However, the model did not provide complete resolution.[55]
Cladogram of evolutionary lines inGalanthussensu Margozet al.[55]
Galanthus
Viridifolii
Glaucaefolii
Galanthus
Selected species
Common snowdrop,Galanthus nivalis, grows to around 7–15 cm tall, flowering between January and April in the northern temperate zone (January–May in the wild). Applanate vernation[12] Grown asornamental.
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 theLevant, 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.[56]
Galanthus reginae-olgae, fromGreece andSicily, is quite similar in appearance toG. 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 withG. nivalis.
Galanthus is derived from theGreek γάλα (gala), meaning "milk" and ἄνθος (anthos) meaning "flower", alluding to the colour of the flowers. Theepithetnivalis is derived from theLatin, meaning "of the snow".[57][58] The word "Snowdrop" may be derived from theGermanSchneetropfen (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 (seeSymbols).[21]
Snowdrops arehardyherbaceous plants thatperennate by undergroundbulbs. They are among the earliest spring bulbs to bloom, although a few forms ofG. nivalis are autumn flowering.[9][59] In colder climates, they will emerge through snow (see illustration). To help against the cold and possible damage caused by icecrystals forming on the plant, they are coated in anenzyme that inhibits the creation of these crystals[60]. They naturalise relatively easily forming large drifts. These are often sterile,[61] found near human habitation, and also former monastic sites.[59] The leaves die back a few weeks after the flowers have faded.Galanthus plants are relatively vigorous and may spread rapidly by forming bulboffsets. They also spread by dispersal of seed, animals disturbing bulbs, and water if disturbed by floods.[59][21]
Some snowdrop species are threatened in their wildhabitats, due tohabitat destruction, illegal collecting, andclimate change.[3] In most countries collecting bulbs from the wild is now illegal. UnderCITES regulations, international trade in any quantity ofGalanthus, whether bulbs, live plants, or even dead ones, is illegal without a CITES permit.[62] 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, andG. woronowii) from Turkey and Georgia.[63] A number of species are on theIUCNRed List of threatened species, with theconservation status beingG. trojanus ascritically endangered,[64] four speciesvulnerable,G. nivalis isnear threatened[65] and several species show decreasing populations.[3]G. panjutinii is consideredendangered. One of its five known sites, atSochi, was destroyed by preparations for the2014 Winter Olympics.[43]
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 andGeorgia.[3] For instance export quotas for 2016 forG. elwesii were 7 million for Turkey.[66] Quotas forG. worononowii were 5 million for Turkey and 15 million for Georgia.[67] These figures include both wild-taken and artificially propagated bulbs.
Snowdrop carpet atBank Hall, Bretherton, Lancashire in February 2009
Celebrated as a sign of spring, snowdrops may form impressive carpets of white in areas where they are native or have been naturalised. These displays may attract large numbers of sightseers.[59] There are a number of snowdrop gardens in England, Wales, Scotland, and Ireland.[68] 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).[69] Several gardens in England open during snowdrop season for theNational Gardens Scheme (NGS) and in Scotland for Scotland's Gardens.Colesbourne Park inGloucestershire is one of the best known of the English snowdrop gardens, being the home ofHenry John Elwes, a collector of Galanthus specimens, and after whomGalanthus elwesii is named.[70][71]
Snowdrop with extra tepals (mutation)Galanthus plicatus 'Wendy's Gold' has yellow markings and ovary.
Numerous single- and double-floweredcultivars ofGalanthus nivalis are known, and also of several otherGalanthus species, particularlyG. plicatus andG. 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.[72] Double-flowered cultivars and forms, such as the extremely commonGalanthus nivalisf.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'.[73] Manyhybrids have also occurred in cultivation.[7]
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.
In 1983, Andreas Plaitakis and Roger Duvoisin suggested that the mysterious magical herb,moly, that appears inHomer'sOdyssey is the snowdrop. One of the active principles present in the snowdrop is thealkaloidgalantamine, which, as anacetylcholinesterase inhibitor, could have acted as an antidote toCirce's poisons.[101] Further supporting this notion are notes made during the fourth century BC by the Greek scholarTheophrastus who wrote inHistoria plantarum that moly was "used as an antidote against poisons" although which specific poisons it was effective against remains unclear.[102] Galantamine (or galanthamine) may be helpful in the treatment ofAlzheimer's disease, although it is not a cure;[103] the substance also occurs naturally indaffodils and othernarcissi.[104]
Snowdrops figure prominently in art and literature,[106] often as a symbol in poetry of spring, purity, and religion (seeSymbols), such asWalter de la Mare's poemThe Snowdrop (1929).[107] In this poem, he likened the triple tepals in each whorl ("A triplet of green-pencilled snow") to theHoly Trinity.[59] He used snowdrop imagery several times in his poetry, such asBlow, Northern Wind (1950) – see Box.[105] Another instance is the poemThe Snowdrop. byLetitia Elizabeth Landon in which she asks "Thou fairy gift from summer, Why art thou blooming now?"[108]
In the fairy-tale playThe Twelve Months by Russian writerSamuil 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. TheSoviet traditionally animated filmThe Twelve Months (1956),Lenfilm filmThe Twelve Months (1972), and the anime filmTwelve 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 themilitary 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, andSam Browne belt against their olive drab uniforms.
The short storyThe Snowdrop byHans 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 successfulwaltzSchneeglöckchen (Snowdrops) op. 143 after this flower. The inspiration is especially evident in thecello introduction and in the slow unfurling of the opening waltz. Strauss composed this piece for a Russian Embassy dinner given at theSperl ballroom inVienna on 2 December 1853, but did not perform it publicly until the year 1854. TheSperl banquet was given in honour of her Excellency FrauMaria von Kalergis, daughter of the Russian diplomat and foreign minister Count Karl Nesselrode, and Strauss also dedicated his waltz to her.[109]
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.
Early names refer to the association with thereligious feast ofCandlemas (February 2) – the optimum flowering time of the plant – at which young women, robed in white, would walk in solemn procession in commemoration of thePurification of the Virgin, an alternative name for the feast day. TheFrench name ofviolette de la chandaleur refers to Candlemas, while anItalian name,fiore della purificazione, refers to purification. TheGerman name ofSchneeglöckchen (little snow bells) invokes the symbol of bells.[21][59]
In more recent times, the snowdrop was adopted as a symbol of sorrow and of hope following theDunblane massacre in Scotland, and lent its name to the subsequent campaign to restrict the legal ownership of handguns in the UK.[110][111]
^"AGM Plants – Ornamental"(PDF). Royal Horticultural Society. July 2017. p. 39.Archived(PDF) from the original on 9 October 2022. Retrieved27 February 2018.
Kaplan, Matt (27 October 2015).Science of the magical: from the Holy Grail to love potions to superpowers (First Scribner hardcover ed.). New York.ISBN978-1-4767-7710-8.OCLC904813040.{{cite book}}: CS1 maint: location missing publisher (link)
Hester, Gerko; Kaku, Hanae; Goldstein, Irwin J. & Schubert Wright, Christine (1995). "Structure of mannose-specific snowdrop (Galanthus nivalis) lectin is representative of a new plant lectin family".Nature Structural Biology.2 (6):472–479.doi:10.1038/nsb0695-472.PMID7664110.S2CID45031999.
