Taxonomy of Narcissus

The taxonomy of ''Narcissus'' is complex, and still not fully resolved. Known to the ancients, the genus name appears in Graeco-Roman literature, although their interest was as much medicinal as botanical. It is unclear which species the ancients were familiar with. Although frequently mentioned in Mediaeval and Renaissance texts it was not formally described till the work of Linnaeus in 1753. By 1789 it had been grouped into a family (Narcissi) but shortly thereafter this was renamed Amaryllideae, from which comes the modern placement within Amaryllidaceae, although for a while it was considered part of Liliaceae.

Many of the species now considered to be ''Narcissus'' were in separate genera during the nineteenth century, and the situation was further confused by the inclusion of many cultivated varieties. By 1875 the current circumscription was relatively settled. By 2004 phylogenetic studies had allowed the place of ''Narcissus'' within its fairly large family to be established, nested within a series of subfamilies (Amaryllidoideae) and tribes (Narcisseae). It shares its position in the latter tribe with ''Sternbergia''.

The infrageneric classification has been even more complex and many schemes of subgenera, sections, subsections and series have been proposed, although all had certain similarities. Most authorities now consider there to be 10 – 11 sections based on phylogenetic evidence. The problems have largely arisen from the diversity of the wild species, frequent natural hybridisation and extensive cultivation with escape and subsequent naturalisation. The number of species has varied anywhere from 16 to nearly 160, but is probably around 50 – 60.

The genus appeared some time in the Late Oligocene to Early Miocene eras, around 24 million years ago, in the Iberian peninsula. While the exact origin of the word ''Narcissus'' is unknown it is frequently linked to its fragrance which was thought to be narcotic, and to the legend of the youth of that name who fell in love with his reflection. In the English language the common name Daffodil appears to be derived from the Asphodel with which it was commonly compared.

History

Early

''Narcissus'' was first described by Theophrastus (, c 371 - c 287 BC) in his ''Historia Plantarum'' ( grc-gre, Περὶ φυτῶν ἱστορία) as , referring to ''N. poeticus'', but comparing it to Asphodelus (). Theophrastus' description was frequently referred to at length by later authors writing in Latin such as Pliny the Elder (Gaius Plinius Secundus, 23 AD – 79 AD) from whom came the Latin form (see also Culture). Pliny's account is from his '' Natural History'' ( la, Naturalis Historia). Like his contemporaries, his interests were as much therapeutic as botanical. Another much-cited Greek authority was Dioscorides (, 40 AD – 90 AD) in his De Materia Medica ( grc-gre, Περὶ ὕλης ἰατρικῆς). Both authors were to remain influential until at least the Renaissance, given that their descriptions went beyond the merely botanical, to the therapeutic (see also Antiquity).

An early European reference is found in the work of Albert Magnus (c. 1200 – 1280), who noted in his the similarity to the leek. William Turner in his ''A New Herball'' (1551) cites all three extensively in his description of the plant and its properties. It was to remain to Linnaeus in 1753 to formally describe and name ''Narcissus'' as a genus in his ''Species Plantarum'', at which time there were six known species (''N. poeticus'', ''N. pseudonarcissus'', ''N. bulbocodium'', ''N. serotinus'', ''N. jonquilla'' and ''N. tazetta''). At that time, Linnaeus loosely grouped it together with 50 other genera into his ''Hexandria monogynia''.

Modern

It was de Jussieu in 1789 who first formally created a 'family' (Narcissi), as the seventh 'Ordo' (Order) of the third class (Stamina epigyna) of Monocots in which ''Narcissus'' and 15 other genera were placed. The use of the term Ordo at that time was closer to what we now understand as Family, rather than Order. The family has undergone much reorganisation since then, but in 1805 it was renamed after a different genus in the family, Amaryllis, as 'Amaryllideae' by Jaume St.-Hilaire and has retained that association since. Jaume St.-Hilaire divided the family into two unnamed sections and recognised five species of ''Narcissus'', omitting ''N. serotinus''.

De Candolle brought together Linnaeus' genera and Jussieau's families into a systematic taxonomy for the first time, but included ''Narcissus'' (together with '' Amaryllis'') in the Liliaceae in his ''Flore française'' (1805-1815) rather than Amaryllidaceae, a family he had not yet recognised. Shortly thereafter he separated the 'Amaryllidées' from 'Liliacées' (1813), though attributing the term to Brown's 'Amaryllideae' in the latter's ''Prodromus'' (1810) rather than St.-Hilaire's 'Amaryllidées'. He also provided the text to the first four volumes of Redouté illustrations in the latter's ''Les liliacées'' between 1805 and 1808 (see illustration here of '' N. candidissimus'').

Historically both wide and narrow interpretations of the genus have been proposed. In the nineteenth century genus splitting was common, favouring the narrow view. Haworth (1831) using a narrow view treated many species as separate genera, as did Salisbury (1866). These authors listed various species in related genera such as ''Queltia'' (hybrids), ''Ajax'' (=''Pseudonarcissus'') and ''Hermione'' (=''Tazettae''), sixteen in all in Haworth's classification. In contrast, Herbert (1837) took a very wide view reducing Harworth's sixteen genera to six. Herbert, treating the Amaryllidacea as an 'order' as was common then, considered the narcissi to be a suborder, the Narcisseae, the six genera being ''Corbularia'', ''Ajax'', ''Ganymedes'', ''Queltia'', ''Narcissus'' and ''Hermione'' and his relatively narrow circumscription of ''Narcissus'' having only three species. Later Spach (1846) took an even wider view bringing most of Harworth's genera into the genus ''Narcissus'', but as separate subgenera. By the time that Baker (1875) wrote his monograph all of the genera with one exception were included as ''Narcissus''. The exception was the monotypic group'' Tapeinanthus'' which various subsequent authors have chosen to either exclude (''e.g.'' Cullen 1986) or include (''e.g.'' Webb 1978, 1980). Today it is nearly always included.

The eventual position of ''Narcissus'' within the Amaryllidaceae family only became settled in the twenty-first century with the advent of phylogenetic analysis and the Angiosperm Phylogeny Group system. The genus ''Narcissus'' belongs to the Narcisseae tribe, one of 13 within the Amaryllidoideae subfamily of the Amaryllidaceae. It is one of two sister clades corresponding to genera in the Narcisseae, being distinguished from ''Sternbergia'' by the presence of a paraperigonium, and is monophyletic.

Subdivision

The infrageneric phylogeny of ''Narcissus'' still remains relatively unsettled. The taxonomy has proved very complex and difficult to resolve, particularly for the ''Pseudonarcissus'' group. This is due to a number of factors, including the diversity of the wild species, the ease with which natural hybridisation occurs, and extensive cultivation and breeding accompanied by escape and naturalisation.

De Candolle, in the first systematic taxonomy of ''Narcissus'', arranged the species into named groups, and those names (''Faux-Narcisse'' or ''Pseudonarcissus'', ''Poétiques'', ''Tazettes'', ''Bulbocodiens'', ''Jonquilles'') have largely endured for the various subdivisions since and bear his name. The evolution of classification was confused by including many unknown or garden varieties, until Baker (1875) made the important distinction of excluding all specimens except the wild species from his system. He then grouped all of the earlier related genera as sections under one genus, ''Narcissus'', the exception being the monotypic ''Tapeinanthus''. Consequently, the number of accepted species has varied widely.

A common modern classification system has been that of Fernandes (1951, 1968, 1975) based on cytology, as modified by Blanchard (1990) and Mathew (2002), although in some countries such as Germany, the system of Meyer (1966) was preferred. Fernandes described two subgenera based on basal chromosome number, ''Hermione'', n = 5 (11) and ''Narcissus'', n = 7 (13). He further subdivided these into ten sections (''Apodanthi'', ''Aurelia'', ''Bulbocodii'', ''Ganymedes'', ''Jonquillae'', ''Narcissus'', ''Pseudonarcissi'', ''Serotini'', ''Tapeinanthus'', ''Tazettae''), as did Blanchard later.

In contrast to Fernandes, Webb's treatment of the genus for the ''Flora Europaea'' (1978, 1980) prioritised morphology over genetics, and abandoned the subgenera ranks. He also restored De Candolle's original nomenclature, and made a number of changes to section ''Jonquilla'', merging the existing subsections, reducing ''Apodanthi'' to a subsection of ''Jonquilla'', and moving ''N. viridiflorus'' from ''Jonquilla'' to a new monotypic section of its own (''Chloranthi''). Finally, he divided ''Pseudonarcissus'' into two subsections. Blanchard (1990), whose ''Narcissus: a guide to wild daffodils'' has been very influential, adopted a simple approach, restoring ''Apodanthae'', and based largely on ten sections alone.

The Royal Horticultural Society (RHS) currently lists ten sections, based on Fernandes (1968), three of which are monotypic (contain only one species), while two others only containing two species. Most species are placed in ''Pseudonarcissus'' While infrageneric groupings within ''Narcissus'' have been relatively constant, their status (genera, subgenera, sections, subsections, series, species) has not. Some authors treat some sections as being further subdivided into subsections, ''e.g.'' ''Tazettae'' (3 subsections). These subdivisions correspond roughly to the popular names for narcissi types, ''e.g.'' Trumpet Daffodils, Tazettas, Pheasant's Eyes, Hoop Petticoats, Jonquils.

While Webb had simply divided the genus into sections, Mathew found this unsatisfactory, implying every section had equal status. He adapted both Fernandes and Webb to devise a more hierarchical scheme he believed better reflected the interrelationships within the genus. Mathew's scheme consists of three subgenera (''Narcissus'', ''Hermione'' and ''Corbularia''). The first two subgenera were then divided into five and two sections respectively. He then further subdivided two of the sections (subgenus ''Narcissus'' section ''Jonquillae'', and subgenus ''Hermione'' section ''Hermione'') into three subsections each. Finally, he divided section ''Hermione'' subsection ''Hermione'' further into two series, ''Hermione'' and ''Albiflorae''. While lacking a phylogenetic basis, the system is still in use in horticulture. For instance the Pacific Bulb Society uses his numbering system (see Table II) for classifying species.

Phylogenetics

The phylogenetic analysis of Graham and Barrett (2004) supported the infrageneric division of ''Narcissus'' into two clades corresponding to the subgenera ''Hermione'' and ''Narcissus'', but does not support monophyly of all sections, with only ''Apodanthi'' demonstrating clear monophyly, corresponding to Clade III of Graham and Barrett (see Cladogram), although some other clades corresponded approximately to known sections. These authors examined 36 taxa of the 65 listed then, and a later extended analysis by Rønsted ''et al.'' (2008) with five additional taxa confirmed this pattern.

A very large (375 accessions) molecular analysis by Zonneveld (2008) utilising nuclear DNA content sought to reduce some of the paraphyly identified by Graham and Barrett. This led to a revision of the sectional structure, shifting some species between sections, eliminating one section and creating two new ones. In subgenus ''Hermione'', ''Aurelia'' was merged with ''Tazettae''. In subgenus ''Narcissus'' section ''Jonquillae'' subsection ''Juncifolii'' was elevated to sectional rank, thus resolving the paraphyly in this section observed by Graham and Barrett in Clade II due to this anomalous subsection, the remaining species being in subsection ''Jonquillae'', which was monophyletic. The relatively large section ''Pseudonarcissi'' was divided by splitting off a new section, ''Nevadensis'' (species from southern Spain) leaving species from France, northern Spain and Portugal in the parent section. At the same time Fernández-Casas (2008) proposed a new monotypic section ''Angustini'' to accommodate ''Narcissus deficiens'', placing it within subgenus ''Hermione''.

While Graham and Barrett (2004) had determined that subgenus ''Hermione'' was monophyletic, using a much larger accession Santos-Gally ''et al.'' (2011) did not. However the former had excluded species of hybrid origins, while the latter included both ''N. dubius'' and ''N. tortifolius''. If these two species are excluded (forming a clade with subgenus ''Narcissus'') then ''Hermione'' can be considered monophyletic, although as a section of ''Hermione'', ''Tazettae'' is not monophyletic. They also confirmed the monophyly of ''Apodanthi''.

Some so-called nothosections have been proposed, predominantly by Fernández-Casas, to accommodate natural ('ancient') hybrids (nothospecies).

Subgenera and sections

Showing revisions by Zonnefeld (2008)

* subgenus ''Hermione'' (Haw.) Spach.
** (''Aurelia'' (Gay) Baker (monotypic) - ''merged with Tazettae (2008)''
** ''Serotini'' Parlatore (2 species)
** ''Tazettae'' de Candolle (16 species) syn. ''Hermione'' (Salisbury) Sprengel, in Fernandes' scheme. Incorporating ''Aurelia'' (2008)
* subgenus ''Narcissus'' L.
** ''Apodanthi'' A. Fernandes (6 species)
** ''Bulbocodium'' de Candolle (11 species)
** ''Ganymedes'' (Haworth) Schultes f. (monotypic)
** ''Jonquillae'' de Candolle (8 species)
** ''Juncifolii'' (A. Fern.) Zonn. ''sect. nov.'' (2008)
** ''Narcissus'' L. (2 species)
** ''Nevadensis'' Zonn. ''sect. nov.'' (2008)
** ''Pseudonarcissus'' de Candolle (36 species) Trumpet daffodils
** ''Tapeinanthus'' (Herbert) Traub (monotypic)

Species

Estimates of the number of species in ''Narcissus'' have varied widely, from anywhere between 16 and nearly 160, even in the modern era. Linnaeus originally included six species in 1753. By the time of the 14th edition of the Systema Naturae in 1784, there were fourteen. The 1819 ''Encyclopaedia Londinensis'' lists sixteen (see illustration here of three species) and by 1831 Adrian Haworth had described 150 species.

Much of the variation lies in the definition of species, and whether closely related taxa are considered separate species or subspecies. Thus, a very wide view of each species, such as Webb's results in few species, while a very narrow view such as that of Fernandes results in a larger number. Another factor is the status of hybrids, given natural hybridisation, with a distinction between 'ancient hybrids' and 'recent hybrids'. The term 'ancient hybrid' refers to hybrids found growing over a large area, and therefore now considered as separate species, while 'recent hybrid' refers to solitary plants found amongst their parents, with a more restricted range.

In the twentieth century Fernandes (1951) accepted 22 species, on which were based the 27 species listed by Webb in the 1980 Flora Europaea. By 1968, Fernandes had accepted 63 species, and by 1990 Blanchard listed 65 species, and Erhardt 66 in 1993. In 2006 the Royal Horticultural Society's (RHS) ''International Daffodil Register and Classified List'' listed 87 species, while Zonneveld's genetic study (2008) resulted in only 36. , the World Checklist of Selected Plant Families accepts 52 species, along with at least 60 hybrids, while the RHS has 81 accepted names in its October 2014 list.

Evolution

Within the Narcisseae, ''Narcissus'' ( western Mediterranean) diverged from ''Sternbergia'' (Eurasia) some time in the Late Oligocene to Early Miocene eras, around 29.3–18.1 Ma, with a best estimate of 23.6 Ma. Later the genus divided into the two subgenera (''Hermione'' and ''Narcissus'') between 27.4 and 16.1 Ma (21.4 Ma). The divisions between the sections of ''Hermione'' then took place during the Miocene period 19.9–7.8 Ma.

''Narcissus'' appears to have arisen in the area of the Iberian peninsula, southern France and northwestern Italy, and within this area most sections of the genus appeared, with only a few taxa being dispersed to North Africa at a time when the African and West European platforms were closer together. Subgenus ''Hermione'' in turn arose in the southwestern mediterranean and north west Africa. However, these are reconstructions, the Amaryllidaceae lacking a fossil record.

Names and etymology

Narcissus

The derivation of the Latin ''narcissus'' ( grc-gre, νάρκισσος) is unknown. It may be a loanword from another language; for instance, it is said to be related to the Sanskrit word , meaning 'hell'. It is frequently linked to the Greek myth of Narcissus described by Ovid in his Metamorphoses, who became so obsessed with his own reflection that as he knelt and gazed into a pool of water, he fell into the water and drowned. In some variations, he died of starvation and thirst. In both versions, the narcissus plant sprang from where he died. Although Ovid appeared to describe the plant we now know as ''Narcissus'' there is no evidence for this popular derivation, and the person's name may have come from the flower's name. The Poet's Narcissus (''N. poeticus''), which grows in Greece, has a fragrance that has been described as intoxicating. This explanation is largely discredited due to lack of proof. Pliny wrote that the plant ('named narcissus from narce, not from the legendary youth'), ''i.e.'' that it was named for its narcotic properties ( , 'I grow numb' in Greek), not from the legend. Furthermore, there were accounts of narcissi growing, such as in the legend of Persephone, long before the story of Narcissus appeared (see Greek culture).Prior here refers to the poet ''Pamphilus'', but it is likely he meant ''Pamphos'' It has also been suggested that daffodils bending over streams evoked the image of the youth admiring his own reflection in the water.

Linnaeus used the Latin name for the plant in formally describing the genus, although Matthias de l'Obel had previously used the name in describing various species of ''Narcissi'' in his ''Icones stirpium'' of 1591, and other publications, as had Clusius in ''Rariorum stirpium'' (1576).

The plural form of the common name ''narcissus'' has caused some confusion. British English sources such as the Shorter Oxford English Dictionary give two alternate forms, ''narcissi'' and ''narcissuses''. In contrast, in American English the Merriam-Webster Dictionary provides for a third form, ''narcissus'', used for both singular and plural. The Oxford dictionaries only list this third form under American English, although the Cambridge Dictionary allows of all three in the same order. However, Garner's Modern American Usage states that ''narcissi'' is the commonest form, ''narcissuses'' being excessively sibilant. For similar reasons, Fowler prefers ''narcissi'' in British English usage. Neither support ''narcissus'' as a plural form. Common names such as ''narcissus'' do not capitalise the first letter in contrast to the person of that name and the Latin genus name.

The name ''Narcissus'' (feminine ''Narcissa'') was not uncommon in Roman times, such as Tiberius Claudius Narcissus, a Roman official in Claudius' time, an early New Testament Christian in Rome and later bishops and saints.

Daffodil

The word ''daffodil'' was unknown in the English language before the sixteenth century. The name is derived from an earlier ''affodell'', a variant of '' asphodel''. In classical Greek literature the narcissus is frequently referred to as the , such as the meadows of the Elysian fields in Homer (see Antiquity). ''Asphodel'' in turn appears to be a loanword coming from French via Mediaeval Latin from Classical Latin and ultimately the Greek ( grc-gre, ἀσφόδελος). The reason for the introduction of the initial ''d'' is not known, although a probable source is an etymological merging from the Dutch article , as in , or English ''the'', as ''th'affodil'' or ''t'affodil'', hence ''daffodil'', and in French and to form and .

From at least the 16th century, ''daffadown dilly'' and ''daffydowndilly'' have appeared as playful synonyms of the name. In common parlance and in historical documents, the term ''daffodil'' may refer specifically to populations or specimens of the wild daffodil, ''N. pseudonarcissus''. H. N. Ellacombe suggests this may be from ''Saffon Lilly'', citing Prior in support, though admittedly conjectural.

Lady Wilkinson (1858), who provides an extensive discussion of the etymology of the various names for this plant, suggests a very different origin, namely the Old English word (that which cometh early), citing a 14th-century (but likely originally much earlier) manuscript in support of this theory, and which appears to describe a plant resembling the daffodil. Ellacombe provides further support for this from a fifteenth century English translation of Palladius that also refers to it.Palladius, the fourth century Roman agricultural writer was best known in English for a 15th-century translation into English, titled ''Palladius on Husbondrie'', where the "affadille" is stated to be a suitable plant for bees to visit. Other editions render this as "Affodyle".

Jonquil

The name ''jonquil'' is said to be a corruption via French from the Latin meaning 'rush-leaf' ( Juncaceae) and its use is generally restricted to those species and cultivars which have rush like leaves, e.g. '' N. juncifolius''.

Other

A profusion of names have attached themselves in the English language, either to the genus as a whole or to individual species or groups of species such as sections. These include ''narcissus'', ''jonquil'', ''Lent lily'', ''Lenten lily'', ''lide lily'', ''yellow lily'', ''wort'' or ''wyrt'', ''Julians'', ''glens'', ''Lent cocks'', ''corn flower'', ''bell rose'', ''asphodel'', ''Solomon's lily'', ''gracy day'', ''haverdrils'', ''giggary'', ''cowslip'', and ''crow foot''.

References

Notes

Bibliography

Antiquity

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Mediaeval and Renaissance

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Modern

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{{Taxonbar, from=Q20818018

Narcissus
Narcissus