Editing Galanthus (section)

==Taxonomy==

=== History ===

==== Early ====
[[File:Leucojum bulbosum praecox Gerard.jpg|thumb|''G. nivalis'' from [[John Gerard]]'s ''Herball'', 1597]]
Snowdrops have been known since early times, being described by the classical [[Greeks|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)<ref name=HP134/> 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)<ref name=HP49/>

[[Rembert Dodoens]], a [[Flanders|Flemish]] botanist, described and illustrated this plant in 1583 as did [[John Gerard|Gerard]] in England in 1597 (probably using much of Dodoens' material), calling it ''Leucojum bulbosum praecox'' (Early bulbous violet). Gerard refers to Theophrastus's description as ''Viola alba'' or ''Viola bulbosa'', using [[Pliny the Elder|Pliny]]'s translation, and comments that the plant had originated in Italy and had "taken possession" in England "many years past".<ref name=Gerard/> The [[genus]] was formally named ''Galanthus'' and described by [[Carl Linnaeus]] in 1753,<ref name=Linnaeus/> with the single [[species]], ''[[Galanthus nivalis]]'', which is the [[type (biology)|type]] species. Consequently, Linnaeus is granted the [[botanical authority]]. In doing so, he distinguished this genus and species from ''[[Leucojum]]'' (''Leucojum bulbosum trifolium minus''), a name by which it previously had been known.<ref name=WCSP/><ref name=APW/><ref name=Churchgal/>

==== Modern ====

In 1763 [[Michel Adanson]] began a system of arranging genera in families. Using the [[synonym (taxonomy)|synonym]] ''Acrocorion'' (also spelt ''Akrokorion''),<ref name=Adansongal/> he placed ''Galanthus'' in the family [[Liliaceae]], section Narcissi.<ref name=Adansonnarc/> [[Lamarck]] provided a description of the genus in his encyclopedia (1786),<ref name=Lamarck/> and later, ''Illustrations des genres'' (1793).<ref name=Lamarckillus/> In 1789 [[Antoine Laurent de Jussieu|de Jussieu]], who is credited with the modern concept of genera organised in [[family (biology)|families]], placed ''Galanthus'' and related genera within a [[division (botany)|division]] of [[Monocotyledons]], 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 name ''Galanthus'' and retained their placement under Narcissi, this time as a family (known as ''Ordo'', at that time) and referred to the French vernacular name, ''[[:fr:Perce-neige|Perce-neige]]''{{sfn|Clos|1862|loc=''Perce-neige'' p.&nbsp;658}} (Snow-pierce), based on the plants tendency to push through early spring snow (see [[#Ecology|Ecology]] for illustration).<ref name=Jussieu/> The modern family of [[Amaryllidaceae]], in which ''Galanthus'' is placed, dates to [[Jaume Saint-Hilaire]] (1805) who replaced Jussieu's Narcissi with ''Amaryllidées''.<ref name=Saint-Hilaire/> In 1810, [[Robert Brown (botanist, born 1773)|Brown]] proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the [[Ovary (botany)|ovaries]] and be referred to as Amaryllideae,<ref name=Brown/> and in 1813, [[de Candolle]] separated them by describing Liliacées <small>Juss.</small> and Amaryllidées <small>Brown</small> as two quite separate families.<ref name=Candolle/> However, in his comprehensive survey of the [[Flora of France]] ([[De Candolle system|Flore française]], 1805–1815) he divided Liliaceae into a series of ''Ordres'', and placed Galanthus into the Narcissi ''Ordre''.<ref name=Candollegal/> This relationship of Galanthus to either liliaceous or amaryllidaceaous taxa (see [[Taxonomy of Liliaceae]]) was to last for another two centuries until the two were formally divided at the end of the twentieth century.{{sfn|Meerow et al.|1999}} [[John Lindley|Lindley]] (1830) followed this general pattern, placing ''Galanthus'' and related genera such as ''[[Amaryllis]]'' and ''[[Narcissus (plant)|Narcissus]]'' in his Amaryllideae (which he called The Narcissus Tribe in English).<ref name=Lindley1830/> By 1853, the number of known plants was increasing considerably and he revised his schema in his last work, placing ''Galanthus'' together, and the other two genera in the modern [[Galantheae]] in tribe Amarylleae, order Amaryllidaceae, alliance Narcissales.<ref name=Lindley53Amarylleae/> 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 time [[George Bentham|Bentham]] and [[Joseph Dalton Hooker|Hooker]] published their ''Genera plantarum'' (1862–1883){{sfn|Bentham|Hooker|1883}} ordo Amaryllideae<ref name=BHAmaryllideae/> contained five tribes, and tribe Amarylleae<ref name=BHAmarylleae/> three subtribes (see [[Bentham & Hooker system]]). They placed ''Galanthus'' in subtribe Genuinae and included three species.<ref name=BHgal/>

=== Phylogeny ===

{{cladogram
| title=Cladogram of evolutionary lines in ''Galanthus'' ''sensu'' Ronsted ''et al.''{{sfn|Ronsted et al.|2013}}
| align=''right''
| cladogram={{clade|style="text-align:right; padding:2.5px; background:#eef"; width:500px;
| label1=Galantheae
| 1={{clade
|   1=''Galanthus''
|   2={{clade
|     1=''Acis''
|     2=''Leucojum''}}
    }}
}}
}}

''Galanthus'' is one of three closely related genera making up the tribe [[Galantheae]] within subfamily [[Amaryllidoideae]] (family [[Amaryllidaceae]]). Sometimes snowdrops are confused with the other two genera, ''[[Leucojum]]'' and ''[[Acis (plant)|Acis]]'' (both called snowflakes). ''Leucojum'' species are much larger and flower in spring (or early summer, depending on the species), with all six [[tepal]]s 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.{{sfn|Meerow et al.|2006a}}

===Subdivision===
[[File:Galanthus vernation types.jpg|thumb|The three different forms of leaf shoots from the bulb: flat (applanate), folded (explicative) rolled up (convolute) (left to right)]]
[[File:Galanthus elwesii, flower.jpg|thumb|''[[Galanthus elwesii]]'']]
[[File:Snowdrop 'Viridi-Apice'.jpg|thumb|''Galanthus nivalis'' 'Viridapice']]
[[File:Підсніжники в урочищі Холодний Яр.jpeg|thumb|''[[Galanthus plicatus]]'']]
[[File:Weiße Galanthusblüten.JPG|thumb|''[[Galanthus woronowii]]'']]

''Galanthus'' has approximately 20 species, but new species continue to be described.<ref name=WCSP/> ''G. trojanus'' was identified in Turkey in 2001.<ref name=WCSPtroj/>{{sfn|Davis|Özhatay|2001}} ''G. panjutinii'' (Panjutin's snowdrop)<ref name=WCSPpanj/> was discovered in 2012 in five locations in a small area (estimated at {{cvt|20|km2}}) of the northern [[Colchis]] area (western [[Transcaucasus]]) of [[Georgia (country)|Georgia]] and [[Russia]].<ref name=RBGpanj/>{{sfn|Zubov|Davis|2012}} ''G. samothracicus'' was identified in Greece in 2014. Since it has not been subjected to genetic sequencing, it remains unplaced. It resembles ''G. nivalis'', but is outside the distribution of that species.<ref name=WCSPsamo/>{{sfn|Tan et al.|2014}}

Many species are difficult to identify, however, and traditional infrageneric classification based on [[plant morphology]] alone, such as those of [[FC Stern|Stern]] (1956),{{sfn|Stern|1956}} [[HP Traub|Traub]] (1963)<ref name=Traub63/> and Davis{{sfn|Davis|2016}} (1999, 2001),<ref name=Davistax/>{{sfn|Davis|2001}}{{sfn|Brickell|2011}} has not reflected what is known about its [[evolutionary history]], due to the morphological similarities among the species and relative lack of easily discernible distinguishing characteristics.{{sfn|Kamenetsky|2012}}{{sfn|Lledo et al.|2004}}{{sfn|Larsen et al.|2010}} Stern divided the genus into three series according to leaf [[vernation]] (the way the leaves are folded in the bud, when viewed in transverse section, see [[#Description|Description]]);{{sfn|Stern|1956}}
* section ''Nivales'' <small>Beck</small> (flat leaves)
* section ''Plicati'' <small>Beck</small> ([[Glossary of botanical terms#plicate|plicate]] leaves)
* section ''Latifolii'' <small>Stern</small> ([[convolute (botany)|convolute]] leaves)
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

Traub considered them as [[subgenera]];
* subgenus ''Galanthus''
* subgenus ''Plicatanthus'' <small>Traub & Moldk.</small>
* subgenus ''Platyphyllanthe'' <small>Traub</small>

By contrast Davis, with much more information and specimens, included [[biogeography]] in addition to vernation, forming two series. He used somewhat different terminology for vernation, namely applanate (flat), explicative (plicate), and supervolute (convolute). He merged ''Nivalis'' and ''Plicati'' into series ''Galanthus'', and divided ''Latifolii'' into two subseries, ''Glaucaefolii'' <small>(Kem.-Nath) A.P.Davis</small> and ''Viridifolii'' <small>(Kem.-Nath) A.P.Davis</small>.<ref name=Davistax/>

Early [[molecular phylogenetic]] studies confirmed the genus was [[monophyletic]] and suggested four [[clades]], which were labelled as [[series (botany)|series]], and showed that Davis' subseries were not monophyletic.{{sfn|Lledo et al.|2004}}{{sfn|Larsen et al.|2010}} An expanded study in 2013 demonstrated seven major clades, corresponding to [[biogeographical]] distribution. This study used nuclear encoded nr[[Internal transcribed spacer|ITS]] (Nuclear ribosomal internal transcribed spacer), and [[plastid]] encoded genes ''matK'' ([[Maturase K]]), ''[[trnL-F]]'', ''[[ndhF]]'', and ''[[psbK–psbI]]'', and examined all species recognised at the time, as well as two naturally occurring putative [[hybrid (biology)|hybrids]]. The morphological characteristic of vernation that earlier authors had mainly relied on was shown to be highly [[homoplasious]]. A number of species, such as ''G. nivalis'' and ''G. elwesii'' demonstrated intraspecific biogeographical clades, indicating problems with [[speciation]] and there may be a need for [[circumscription (taxonomy)|recircumscription]]. These clades were assigned names, partly according to Davis' previous groupings. In this model clade, the group containing ''G. platyphyllus'' is [[sister group|sister]] to the rest of the genus.{{sfn|Ronsted et al.|2013}}

By contrast, another study performed at the same time, using both [[nuclear DNA|nuclear]] and [[chloroplast DNA]], but limited to the 14 species found in Turkey, largely confirmed Davis' series and subseries, and with biogeographical correlation. Series ''Galanthus'' in this study corresponded to clade nivalis, subseries ''Glaucaefolii'' with clade Elwesii and subseries ''Viridifolii'' with clades Woronowii and Alpinus. However, the model did not provide complete resolution.{{sfn|Margoz et al.|2013}}

==== Clades ====

{{cladogram
| title=Cladogram of evolutionary lines in ''Galanthus'' ''sensu'' Ronsted ''et al.''{{sfn|Ronsted et al.|2013}}
| align=''center''
| cladogram={{clade|style="text-align:left; padding:2.5px; background:#eef"; width:500px;
| label1= ''Galanthus''
| 1={{clade
|   1=Platyphyllus
|   2={{clade
|     1=Trojanus
|     2={{clade
|       1={{clade
|         1=Ikariae
|         2=Elwesii}}
|       2={{clade
|         1=Nivalis
|         2={{clade
|           1=Woronowii
|           2=Alpinus}}
          }}
          }}
        }}
      }}
   }}
}}''sensu'' Ronsted et al. 2013{{sfn|Ronsted et al.|2013}}
* Platyphyllus clade ([[Caucasus]], W. [[Transcaucasus]], NE [[Turkey]])
** ''[[Galanthus krasnovii]]'' <small>Khokhr.</small> 1963
** ''[[Galanthus platyphyllus]]'' <small>Traub & Moldenke</small> 1948
** ''[[Galanthus panjutinii]]'' <small>Zubov & A.P.Davis</small> 2012
* Trojanus clade (NW Turkey)
** ''[[Galanthus trojanus]]'' <small>A.P.Davis & Özhatay</small> 2001
* Ikariae clade ([[Aegean Islands]])
** ''[[Galanthus ikariae]]'' <small>Baker</small> 1893
* Elwesii clade (Turkey, Aegean Islands, SE Europe)
** ''[[Galanthus cilicicus]]'' <small>Baker</small> 1897
** ''[[Galanthus elwesii]]'' <small>[[Joseph Dalton Hooker|Hook.f.]]</small> 1875 (2 variants)
** ''[[Galanthus gracilis]]'' <small>Celak.</small> 1891
** ''[[Galanthus peshmenii]]'' <small>A.P.Davis & C.D.Brickell</small> 1994
* Nivalis clade (Europe, NW Turkey)
** ''[[Galanthus nivalis]]'' <small>L.</small> 1753
** ''[[Galanthus plicatus]]'' <small>M.Bieb.</small> 1819 (2 subspecies)
** ''[[Galanthus reginae-olgae]]'' <small>Orph.</small> 1874 (2 subspecies)
* Woronowii clade (Caucasus, E. and NE Turkey, N. [[Iran]])
** ''[[Galanthus fosteri]]'' <small>Baker</small> 1889
** ''[[Galanthus lagodechianus]]'' <small>Kem.-Nath.</small> 1947
** ''[[Galanthus rizehensis]]'' <small>[[Frederick Claude Stern|Stern]]</small> 1956
** ''[[Galanthus woronowii]]'' <small>Losinsk.</small> 1935
* Alpinus clade (Caucasus, NE Turkey, N.Iran)
** [[Galanthus × allenii|''Galanthus'' × ''allenii'']] <small>Baker</small> 1891
** ''[[Galanthus angustifolius]]'' <small>Koss</small> 1951
** ''[[Galanthus alpinus]]'' <small>Sosn.</small> (2 variants) 1911
** ''[[Galanthus koenenianus]]'' <small>Lobin</small> 1993
** ''[[Galanthus transcaucasicus]]'' <small>Fomin</small> 1909
* Unplaced
** ''[[Galanthus bursanus]]'' <small>Zubov, Konca & A.P.Davis</small> 2019 (NW Turkey)
** ''[[Galanthus samothracicus]]'' <small>Kit Tan & Biel</small> 2014 (Greece)

{{cladogram
| title=Cladogram of evolutionary lines in ''Galanthus'' ''sensu'' Margoz ''et al.''{{sfn|Margoz et al.|2013}}
| align=''center''
| cladogram={{clade|style="text-align:left; padding:2.5px; background:#eef"; width:500px;
| label1= ''Galanthus''
| 1={{clade
|   1=''Viridifolii''
|   2={{clade
|     1=''Glaucaefolii''
 |     2=''Galanthus''}}
    }}
}}
}}

;Selected species
* Common snowdrop, ''[[Galanthus nivalis]]'', grows to around 7–15&nbsp;cm tall, flowering between January and April in the northern temperate zone (January–May in the wild). Applanate vernation<ref name="BDG_p17">[[#Bishop|Bishop, Davis & Grimshaw (2002)]], p. 17.</ref> Grown as [[ornamental plant|ornamental]].
* Crimean snowdrop, ''Galanthus plicatus'', 30&nbsp;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&nbsp;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.<ref name=elwesii/>
* ''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 (language)|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".{{sfn|Hyam |Pankhurst |1995}}{{sfn|Hollinger}} The word "Snowdrop" may be derived from the [[German language|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|Symbols]]).<ref name=Churchgal/>