Narcissus (plant)

"Daffodil" redirects here. For other uses, see Daffodil (disambiguation).

"Daffodils" redirects here. For the Swedish musical group, see The Daffodils. For the poem by William Wordsworth, see I Wandered Lonely as a Cloud.

This article is about the plant genus (Daffodils). For the mythological hunter who fell in love with his own reflection, see Narcissus (mythology).

Daffodil Temporal range: 24–0Ma PreЄ Є O S D C P T J K Pg N Late Oligocene - Recent
Narcissus poeticus
Scientific classification
Kingdom:	Plantae
Clade:	Angiosperms
Clade:	Monocots
Order:	Asparagales
Family:	Amaryllidaceae
Subfamily:	Amaryllidoideae
Tribe:	Narcisseae
Genus:	Narcissus L.
Type species
Narcissus poeticus L.
Subgenera
See text.

N. poeticus. Thomé: Flora von Deutschland, Österreich und der Schweiz (1885) showing details of plant anatomy

Narcissus /nɑrˈsɪsəs/ is a genus of hardy, spring-flowering, bulbous perennial plants in the Amaryllidaceae (Amaryllis) family, subfamily Amaryllidoideae. Various common names including daffodil, narcissus (plural narcissi), and jonquil are used to describe all or some of the genus.

Narcissi have characteristic showy white to orange flowers with a basal perianth and a trumpet like corona. They were well known to the ancients both medicinally and botanically although not formerly described until Linnaeus' publication in 1753. The exact taxonomy remains relatively unsettled, but generally the genus is considered as having about ten sections with approximately 50 species. The number of defined species has ranged widely depending on the authority, with the number of distinct species depending on how they are classified. The disparity is due to similarity between species and hybridization between species. The genus appears to have arisen some time in the Late Oligocene to Early Miocene eras, in the Iberian peninsula and adjacent areas of south west Europe. The exact origin of the name Narcissus is unknown, but it is often linked to a Greek word for intoxicated (narcotic) and the myth of the youth of that name who fell in love with his own reflection. The English word Daffodil appears to be derived from the Asphodel, with which it was commonly compared.

The Narcissi are native to meadows and woods in Europe, North Africa, and West Asia, with a center of distribution in the Western Mediterranean, particularly the Iberian peninsula. However both wild and cultivated specimens have naturalised widely, and were introduced into the Far East prior to the tenth century. The Narcissi tend to be long lived bulbs which divide, but the flowers are also insect pollinated. Known pests include viruses such as the Narcissus Mosaic virus, fungi, the larvae of flies, mites and nematodes. Some Narcissus species have become extinct, while others are threatened by increasing urbanisation and tourism.

While historical accounts suggest Narcissi have been cultivated from the earliest times, they became increasingly popular in Europe after the sixteenth century, and by the late nineteenth century were an important commercial crop centred primarily on the Netherlands. Nowadays narcissi are popular as cut flowers and as ornamental plants in private and public gardens. There is a long history of Narcissus breeding and this has resulted in thousands of different cultivars. Cultivated Narcissi are found in twelve different types, providing a wide range of shape and colour.

Like other members of their family the Narcissi produce a number of different alkaloids, which while providing some protection for the plant, may be poisonous if accidentally ingested. However, this property has been exploited for medicinal use in traditional healing and has resulted in the production of Galanthamine for the treatment of Alzheimer's dementia.

Long celebrated in art and literature, the Narcissus has become the national flower of Wales and is associated with a number of themes in different cultures ranging from death to good fortune, and as a symbol of Spring. The appearance of the wild flowers in Spring is associated with festivals in many places, but also with fund raising for cancer.

Description

Narcissus shoots emerging, with sheathed leaves

Narcissus floral morphology

'Narcissus flower showing an outer white corolla with a central yellow corona (paraperigonium)

N. cyclamineus showing refexed perianth

N. bulbocodium showing reduced perianth and prominent corona

Solitary Narcissus flower, prior to opening

N. triandrus showing pendant orientation

Narcissus bulb with shoot and roots

Cross section of ovary with ovules

Tazetta cultivar showing stamens and stigma

General

The Narcissi are perennial herbaceous geophytes which grow from pale brown-skinned spherical bulbs with pronounced necks, and reach heights varying from 5–80 cm depending on species. Dwarf forms include N. asturiensis which reaches 5–8 cm, while N. italicus may grow as high as 80 cm.

Specific

Stems 

The leafless stems, appearing from early to late spring depending on the species, bear from 1 to 20 blooms. Stem shape depends on the species, some are highly compressed with a visible seam, while others are rounded. The stems are upright and located at the centre of the leaves. It is hollow in the upper portion but towards the bulb is more solid and filled with a spongy material. In a few species such as N. hedraeanthus the stem is oblique.

The spathe has a singular valve, and is usually scarious. Bracteoles are small or absent.

Flowers 

Narcissi flowers are hermaphrodite, tripartite, solitary to umbellate, the perianth yellow, white or bicoloured, rarely green and sometimes fragrant. There is a hypanthial floral tube, obconic to cylindrical, together with a corona (usually conspicuous) which is free from the stamens and crateriform (cup shaped). Symmetry is actinomorphic to slightly zygomorphic due to declinate-ascending stamens. Tepals spreading, ascending or reflexed (folded back), lanceolate. Paraperigon funnel or cup shaped, rarely a simple callose (hardened, thickened) ring.

Like many monocotyledons, the perianth (perigon or perigonium) is homochlamydeous, that is undifferentiated into separate calyx (sepals) and corolla (petals), but rather has six tepals. In all Narcissus species except N. cavanillesii the perianth consists of three main parts (i) a proximal hypanthial floral tube, formed by fusion of basal segments of the tepals, (ii) free tepals, which are reflexed from the perianth, and (iii) a distal corona. The transition point between the floral tube and corona is marked by insertion of the free tepals on the fused perianth.

While the perianth may point forwards, in some species such as N. cyclamineus it is folded back (reflexed), while in some species such as N. bulbocodium, it is reduced to a few barely visible pointed segments with a prominent corona. The most striking feature of Narcissi flowers is the corona (trumpet) formed during floral development from stamens which fuse into a tubular structure, and the anthers reduced. At the base of the corona the fragrances which attract pollinators are formed. All species produce nectar at the top of the ovary.

In Narcissus the corona (paraperigonium) is a tubular outgrowth, developing from the perianth and situated above the floral tube. The floral tube is formed by the fusion of the six tepal bases to create a narrow cylindrical to funnelform perianth. Flowers are generally showy and some are fragrant. Flower colour ranges from white to pale or deep yellow, although N. viridiflorus is green and night flowering. Flower diameter varies from 12 to 125 mm., and may be solitary such as the Bulbocodii and most Pseudonarcissi, or on racemose inflorescences which appear umbellate, with 2 to 15 or 20 flowers such as N. papyraceus and N. tazetta.

The flowers are notable for their striking corona, a cylindrical central cone which may be bell-, bowl-, or disc-shaped extending beyond the tepals, and their floral tube. The corona is surrounded by a ring of six floral leaves called the perianth which is united into a tube at the forward edge of the 3-locular ovary. The three outer tepal segments are sepals, and the three inner segments are petals. Coronal morphology varies from the tiny pigmented disk of N. serotinus or the rudimentary structure in N. cavanillesii to the elongated trumpets of the Pseudonarcissi (trumpet daffodils). Floral tubes can range from long and narrow in the Apodanthi and Jonquillae to rudimentary (N. cavanillesii).

The other major variable is flower orientation which may be pendant (declinate-ascendant) (N. triandrus, N. alpestris) horizontal (N. gaditanus, N. poeticus) or erect (N. cavanillesii, N. serotinus).

Narcissi demonstrate exceptional floral diversity (and sexual polymorphism), primarily by corona size and floral (corolla) tube length, associated with pollinator groups (see for instance Figs. 1 and 2 in Graham and Barrett). Barrett and Harder (2005) describe a number of different floral patterns. The predominant patterns they name the 'daffodil' and 'paperwhite' forms. A less common form is the 'triandrus' seen in only two species, N. albimarginatus and N. triandrus. Each form corresponds to a different group of pollinators (See Pollination).

The daffodil form which includes Pseudonarcissi and Bulbocodii has a relatively short, broad or highly funnelform tube which grades into an elongate corona, which is large and funnelform (funnel-like), forming a broad, cylindrical or trumpet-shaped perianth.

The paperwhite form including Jonquillae, Apodanthi, and Narcissus has a relatively long, narrow tube and a short, shallow, flaring corona. The flower is horizontal and fragrant.

The triandrus form combines features of both the daffodil and paperwhite forms with a well-developed long, narrow tube and an extended bell-shaped corona of almost equal length. This occurs only in N. albimarginatus a Moroccan endemic, and N. triandrus. The flowers are pendant.

Pseudonarcissi are relatively large flowering with a corolla length of around 50mm, generally solitary but rarely in inflorescences of 2-4 flowers. They have wide greenish floral tubes with funnel shaped bright yellow coronas. The six tepals sometimes differ in colour from the corona and may be cream coloured to a pale yellow.

Androecium 

There are six stamens in one to two rows, attached at the throat or base of the tube, straight or declinate-ascending (curving downwards, then upwards). Anthers basifixed (attached at base).

Gynoecium 

The ovary is inferior and there is a pistil with a small three lobed stigma and style.

Leaves 

The leaves are linear to strap shaped, sometimes channelled adaxially to semiterete, and may (pedicellate) or may not (sessile) have a petiole stalk. The emerging plant generally has two leaves, but the mature plant usually three, rarely four, and they are covered with a cutin containing cuticle, giving them a waxy appearance. Leaf colour is light green to blue-green. In the mature plant the leaves extend higher than the flower stem, but in some species the leaves are low hanging. The leaf base is encased in a colourless sheath. After flowering the leaves turn yellow and die back once the seed pod is ripe.

Fruit 

Narcissus capsule dispersing seed

The fruit consists of loculicidal ellipsoid to subglobose capsules containing numerous seeds which are black, round and swollen with a hard coat. The pericarp is dry. Most species have 12 ovules and 36 seeds, although some species such as N. bulbocodium have more up to a maximum of 60. Seeds take five to six weeks to mature. The seeds of Jonquillae and Bulbocodium are wedge-shaped and matte black, while those of other sections are ovate and glossy black. The seeds are dispersed by wind or by animals, a gust of wind or contact with a passing animal being sufficient to release mature seeds.

Bulbs 

The bulbs have a corky base plate from which arise the root hairs in a ring around the edge, which grow up to 40 cm in length. Once the leaves die back in summer, the roots also wither. After some years, the roots shorten pulling the bulbs deeper into the ground (contractile roots). The bulbs develop from the inside pushing the older layers outwards, becoming brown and dry, forming an outer shell. Up to 60 layers have been counted in some wild species. The flower stalk which will start to grow in the following spring, develops within the bulb surrounded by two to three deciduous leaves and their sheaths. The flower stem lies in the axil of the second true leaf.

Phytochemistry

As with all Amarylidaceae genera, Narcissus contains unique isoquinoline alkaloids. The first alkaloid to be identified was lycorine, from N. pseudonarcissus in 1877. These are considered a protective adaptation, and are utilised in the classification of species. Nearly 100 alkaloids have been identified in the genus, about a third of all known Amaryllidaceae alkaloids, although not all species have been tested. Of the nine alkaloid ring types identified in the family, Narcissus species most commonly demonstrate the presence of alkaloids from within the Lycorine (lycorine, galanthine, pluviine) and Homolycorine (homolycorine, lycorenine) groups. Hemanthamine, Tazettine, Narciclasine, Montanine and Galanthamine alkaloids are also represented. The alkaloid profile of any plant varies with time, location, and developmental stage. Narcissi also contain fructans and low molecular weight glucomannan in the leaves and plant stems.

Taxonomy

History

Genus valde intricatum et numerosissimis dubiis oppressum
Schultes & Schultes fil., Syst. Veg. 1829

Early

Narcissus was first described by Theophrastus (Θεόφραστος, c 371 - c 287 BC) in his Historia Plantarum (Greek: Περὶ φυτῶν ἱστορία) as νάρκισσος, referring to N. poeticus, but comparing it to Asphodelus (ασφοδελωδες). 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 narcissus (see also Culture). Pliny's account comes to us in his Natural History (Latin: Naturalis Historia). Like his contemporaries, his interests were as much therapeutic as botanical. Another much cited Greek authority was Dioscorides (Διοσκουρίδης, 40 AD – 90 AD) in his De Materia Medica (Greek: Περὶ ὕλης ἰατρικῆς). Both authors were to remain influential till at least the Renaisance, 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 De vegetabilibus et plantis 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. jonqulla 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 Narcissus candidissimus).

N. candidissimus by Pierre-Joseph Redouté 1808

Historically both wide and narrow interpretations of the genus have been proposed, Haworth (1831) using a narrow view treated many species as separate genera. 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) had made the important distinction of excluding all specimens except the wild species all of genera with one exception were included as Narcissus. The exception was 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

Tabe I: Comparison of Modern Classifications Schemes of Narcissus

Fernandes 1975			Webb 1978		Blanchard 1990		Mathew 2002				Zonnefeld 2008		RHS 2013
Subgenus	Section	Subsection	Section	Subsection	Section	Subsection	Subgenus	Section	Subsection	Series	Subgenus	Section	Section
Hermione	Serotini		Serotini		Serotini		Hermione	Hermiome	Serotini		Hermione	Serotini	Serotini
	Hermione	Angustifolii	Tazettae	Angustifoliae	Tazettae	Angustifoliae			Angustifoliae			Tazettae	Tazettae
		Hermione		Tazettae		Tazettae			Hermione	Hermione
	Aurelia		Aurelia		Aurelia					Albiflorae
								Aurelia					Aurelia
Narcissus	Apodanthi				Apodanthae		Narcissus	Jonquillae	Apodanthi		Narcissus	Apodanthi	Apodanthi
	Jonquilla	Jonquilla	Jonquillae	Jonquillae	Jonquillae				Jonquillae			Jonquilla	Jonquilla
		Juncifolii		Apodanthi					Chloranthi			Juncifolii
	Tapeinanthus		Tapeinanthus		Tapeinanthus			Tapeinanthus				Tapeinanthus	Tapeinanthus
	Ganymedes		Ganymedes		Ganymedes			Ganymedes				Ganymedes	Ganymedes
	Bulbocodium		Bulbocodium		Bulbocodium							Bulbocodium	Bulbocodium
	Pseudonarcissus		Pseudonarcissi	Pseudonarcissi	Pseudonarcissus			Pseudonarcissus				Pseudonarcissus	Pseudonarcissus
				Reflexi								Nevadensis
	Narcissus		Narcissus		Narcissus			Narcissus				Narcissus	Narcissus
			Chloranthi (N. viridiflorus)				Corbularia syn. Bulbocodium

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. In the nineteenth century when genus splitting was common, many Narcissi were considered as separate genera, such as Queltia (hybrids), Ajax(=Pseudonarcissus) and Hermione(=Tazettae), by authors such as Salisbury and Haworth. For instance Haworth (1831) grouped the species under sixteen separate genera. The situation was further confused by including many unknown or garden varieties, and it was not till the work of Baker (1875) that the wild species were all grouped 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). Finaly, 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 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 interrelatinships 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 Jonqullae, and subgenus Hermione section Hermione) into three subsections each. Finally, he divided section Hermione subsection Hermione further into two series, Hermione and Albiflorae.

Table II: Subdivisions of Narcissus (Mathew 2002)

Subgenus	Section	Subsection	Series	Type species
Narcissus Pax	Narcissus L.			N. poeticus L.
	Pseudonarcissus DC syn. Ajax Spach			N. pseudonarcissus L.
	Ganymedes Salisbury ex Schultes and Schultes fil.			N. triandrus L.
	Jonquillae De Candolle	Jonquillae DC		N. jonquilla L.
		Apodanthi (A. Fernandes) D. A. Webb		N. rupicola Dufour
		Chloranthi D. A. Webb		N. viridiflorus Schousboe
	Tapeinanthus (Herbert) Traub			N. cavanillesii A. Barra and G. López
Hermione (Salisbury) Spach	Hermione syn. Tazettae De Candolle	Hermione	Hermione	N. tazetta L.
			Albiflorae Rouy.	N. papyraceous Ker-Gawler
		Angustifoliae (A. Fernandes) F.J Fernándes-Casas		Link to image N. elegans (Haw.) Spach
		Serotini Parlatore		N. serotinus L.
	Aurelia (J. Gay) Baker			N. broussonetii Lagasca
Corbularia (Salisb.) Pax syn. Bulbocodium De Candolle				N. bulbocodium L.

Phylogenetics

Narcissus Cladogram (Graham and Barrett 2004)

Narcissus I subgen. Hermione Aurelia, Serotinii, Tazettae VI subg. Narcissus II Jonquillae III Apodanthi IV Bulbicodium V Pseudonarcissi, Ganymedes, Narcissus, Tapeinanthus

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. Howerver 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 monophletic. 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

Main article: List of Narcissus species

N. major, N. triandrus and N. jonquilla. Encyclopaedia Londinensis 1819

Estimates of the number of species in Narcissus have varied widely, from anywhere between 16 and 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).

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. There is a distinction between what are referred to as 'ancient hybrids' which are found occurring over a relatively large area, and 'recent hybrids' with a more restricted range and found growing as solitary plants amongst their parents. The former are more often considered as separate species.

Fernandes (1951) originally 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 International Daffodil Register listed 87 species. In contrast, the genetic study by Zonneveld (2008) resulted in only 36 species (for list and comparison with Webb, see Zonneveld Table 4).

As of September 2014, the World Checklist of Selected Plant Families accepts 52 species, along with at least 60 hybrids. Another important source is the Royal Horticultural Society's Botanical Classification and list of botanical names (October 2014) which is the basis of their International Daffodil Register. This is a searchable 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 27.4–16.1 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 north-western 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.

Etymology

Narcissus

N. juncifolius, Carolus Clusius Rariorum stirpium 1576

N. poeticus, Matthias de l'Obel Icones stirpium 1591

The derivation of the Latin narcissus (Greek: νάρκισσος) is unknown. It may be a loanword from another language, for instance it is said to be related to the Sanskrit word nark, 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 (Narcissus poeticus), which grows in Greece, has a fragrance that has been described as intoxicating. Again, this explanation lacks any real proof and is largely discredited. Pliny wrote that the plant ‘narce narcissum dictum, non a fabuloso puero’ (named narcissus from narce, not from the legendary youth), i.e. that it was named for its narcotic properties (ναρκάω narkao, "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). 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). "Narcissus" is the most commonly used plural, but "narcissi" and "narcissuses" are also acceptable plurals in both British and American English usage.

Daffodil

The name "daffodil" is derived from an earlier "affodell", a variant of asphodel. In classical Greek literature the narcissus is frequently, referred to as the asphodel, 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 "affodilus" from Classical Latin "asphodilus" and ultimately the Greek "asphodelos" (Greek: 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 "de", as in "De affodil", or English "the", as th'affodil or t'affodil, hence daffodil, and in French "de" and "affodil" to form fleur d’aphrodille and daphrodille.

From at least the 16th century, "Daffadown Dilly", "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, Narcissus pseudonarcissus.

Distribution and habitat

Distribution

Although the Amaryllidaceae family are predominantly tropical or subtropical as a whole, Narcissi occur primarily in Southwestern Europe and the Mediterranean region, with a centre of diversity in the Iberian Peninsula (Spain and Portugal). A few species extend the range into southern France and Italy, and even few occur in the Balkans (N. poeticus, N. serotinus, N. tazetta) and the Eastern Mediterranean (N. serotinus) including Israel (N. tazetta). The occurrence of N. tazetta in western and central Asia, China and Japan are considered introductions, albeit ancient (see Eastern cultures). While the exact northern limit of the range is unknown, the occurrence of wild N. pseudonarcissus in Great Britain is similarly considered an ancient introduction.

N. elegans occurs on the North West African Coast (Morocco and Libya), as well as the coastline of Corsica, Sardinia and Italy, and N. bulbocodium between Tangier and Algiers and Tangier to Marrakech, but also on the Iberian Peninsula. N. serotinus is found along the entire Mediterranean coast. N. tazetta occurs as far east as Iran and Kashmir. Since this is one of the oldest species found in cultivation, it is likely to have been introduced into Kashmir. N. poeticus and N. pseudonarcissus have the largest distribution ranges. N. poeticus ranges from the Pyrenees along the Romanian Carpathians to the Black Sea and along the Dalmatian coast to Greece. N. pseudonarcissus ranges from the Iberian Peninsula, via the Vosges Mountains to northern France and Belgium, and the United Kingdom where there are still wild stocks in Southern Scotland. The only occurrence in Luxembourg is located near Lellingen, in the municipality of Kiischpelt. In Germany it is found mainly in the nature reserve at Perlenbach-Fuhrtsbachtal and the Eifel National Park, where in the spring at Monschau the meadows are teeming with yellow blooms. One of the most easterly occurrences can be found at Misselberg near Nassau on the Lahn.

N. pseudonarcissus growing in the Perlenbach-Fuhrtsbachtal reserve in Germany

However unlike the above examples most species have very restricted endemic ranges which may overlap resulting in natural hybrids. For instance in the vicinity of the Portuguese city of Porto where both N. pseudonarcissus and N. triandrus occur there are found various intersections of the two species while in a small area along part of the Portuguese Montego river are found intersectional hybrids between other hand, one finds intersections between N. scaberulus and N. triandrus.

The biogeography demonstrates a phylogenetic association, for instance subgenus Hermione having a lowland distribution, but subgenus Narcissus section Apodanthi being montane and restricted to Morocco, Spain and Portugal. The remaining sections within subgenus Narcissus include both lowland and mountain habitats. Section Pseudonarcissus, although widely naturalised is endemic to the Baetic Ranges of the south eastern Iberian peninsula.

Habitats

Their native habitats are very varied, with different elevations, bioclimatic areas and substrates, being found predominantly in open spaces ranging from low marshes to rocky hillsides and montane pastures, and including grassland, scrub, woods, river banks and rocky crevices. Although requirements vary, overall there is a preference for acidic soils, although some species will grow in limestone. Narcissus scaberulus will grow on granite soils where it is moist in the growing season but dry in the summer, while Narcissus dubius thrives best in regions with hot and dry summers. The Pseudonarcissus group in their natural habitat prefer humid situations such as stream margins, springs, wet pastures, clearings of forests or shrublands with humid soils, and moist hillsides. These habitats tend to be discontinuous in the Mediterranean mountains, producing discrete isolated populations. In Germany, which is poor in lime, Narcissus pseudonarcissus grows in small groups on open mountain meadows or in mixed forests of fir, beech, oak, alder, ash and birch trees with well-drained soil.

Ecology

Narcissus are long-lived perennial geophytes with winter-growing and summer-dormant bulbs. While most species flower in late winter to spring, five species are autumn flowering (N. broussonetii, N. cavanillesii, N. elegans, N. serotinus, N. viridiflorus). Plants may spread clonally through the production of daughter bulbs and division producing clumps. Narcissus species hybridise readily, although the fertility of the offspring will depend on the parental relationship.

Pollination

The flowers are insect pollinated, the major pollinators being bees, butterflies, flies, and hawkmoths, while the night flowering N. viridiflorus is pollinated by crepuscular moths. Pollination mechanism fall into three groups corresponding to floral morphology (see Description - Flowers).

1. 'Daffodil' type. Pollinated by bees seeking pollen from anthers within the corona. The broad perianth allows bees (Bombus, Anthophora, Andrena) to completely enter the flower in their search for nectar and /or pollen. In this type the stigma lies in the mouth of the corona, extending beyond the six anthers, whose single whorl lies well within the corona. The bees come into contact with the stigma before their legs, thorax and abdomen contact the anthers, and this approach herkogamy causes cross pollination.
2. 'Paperwhite' type. These are adapted to long-tongued Lepidoptera, particularly sphingid moths such as Macroglossum, Pieridae and Nymphalidae but also some long-tongued bees, and flies, all of which are primarily seeking nectar. The narrow tube admits only the insect's proboscis, while the short corona serves as a funnel guiding the tip of the proboscis into the mouth of the perianth tube. The stigma is placed either in the mouth of the tube, just above two whorls of three anthers, or hidden well below the anthers. The pollinators then carry pollen on their probosci or faces. The long tongued bees cannot reach the nectar at the tube base and so collect just pollen.
3. 'Triandrus' type. Pollinated by long-tongued solitary bees (Anthophora, Bombus) which forage for both pollen and nectar. The large corona allows the bees to crawl into the perianth but then the narrow tube prevents further progres, causing them to probe deeply for nectar. The pendant flowers prevent pollination by lepidoptera. In N. albimarginatus there may be either a long stigma with short and mid length anthers or a short stigma and long anthers (dimorphism). In N. triandrus there are three patterns of sexual organs (trimophism) but all have long upper anthers but vary in stigma position and the length of the lower anthers.

Allogamy (outcrossing) on the whole is enforced through a late-acting (ovarian) self-incompatibility system, but some species such as N. dubius and N. longispathus are self-compatible producing mixtures of selfed and outcrossed seeds.

Pests and diseases

Viruses 

Aphids such as Macrosiphum euphorbiae can transmit viral diseases which affect the colour and shape of the leaves. These include the Narcissus common latent virus (Carlavirus), Narcissus degeneration virus, Narcissus late season yellow virus, Narcissus latent virus, Narcissus mosaic virus (Potexvirus), Narcissus symptomless virus, Narcissus yellow stripe virus (Potyvirus) and Raspberry ringspot virus. These are primarily diseases of commercial nurseries. The growth inhibition caused by viral infection can cause substantial economic damage.

Fungi 

More problematic for non commercial plants is the fungus Fusarium bubigenum which causes rotting of the bulbs and yellowing of the leaves. Since the fungus can remain in the soil for many years it is necessary to remove infected plants immediately, and to avoid planting further narcissi at that spot for a further five years. Not all species and cultivars are equally susceptible. Relatively resistant forms include N. triandrus, N. tazetta and N. jonquilla. Another fungus which attacks the bulbs is Botrytis narcissicola, particularly if improperly stored. Copper sulfate is used to combat the disease, and infected bulbs are burned. Other fungi affect the remainder of the plant. Sclerotinia polyblastis causes brown spots on the flower buds and stems, especially in damp weather and is a threat to the cut flower industry. Ramularia vallisumbrosae is a leaf spot virus found in warmer climates, causing Narcissus white mould disease. Another leaf virus is Stagonospora curtisii.

Animals 

Three types of fly have larvae that attack Narcissus plants, the Narcissus Bulb Fly Merodon equestris, and two species of hoverflies, the Lesser Bulb Flies Eumerus tuberculatus and Eumerus strigatus. The flies lay their eggs at the end of June in the ground around the narcissi, a single female fly being able to lay up to fifty eggs. The hatching larvae then burrow through the soil towards the bulbs and consume their interiors. They then overwinter in the empty bulb shell, emerging in April to pupate in the soil, from which the adult fly emerges in May. Mites infest mainly stored bulbs and multiply particularly at high ambient temperature, but do not attack planted bulbs. These are, however, susceptible to nematodes which cause the leaves to turn yellow and become mishapen. The main offender here is Ditylenchus dipsaci. Infested bulbs have to be destroyed, and where infestation is heavy avoiding planting further narcissi for another five years. Snails are another predator.

Conservation

Many of the smallest species have become extinct, requiring vigilance in the conservation of the wild species. Narcissi are increasingly under threat by urban development and tourism. N. cyclamineus is already considered either extinct or exceedingly rare.

In response a number of protected areas (daffodil meadows) have been established such as the Negraşi Daffodil Meadow in Romania, or Kempley Daffodil Meadow in the UK. These areas often host daffodil festivals in the spring.

Cultivation

History

Narcissi in Hortus Eystettensis, 1613

Early accounts of narcissi referred to wild populations. They have been cultivated from at least as early as the sixteenth century in the Netherlands, when large numbers of bulbs where imported from the field, particularly Narcissus hispanicus which soon became nearly extinct in its native habitat of France and Spain, though still found in the southern part of that country. Cultivation is also documented in Britain at this time. This was a period when the development of exotic formal gardens and parks was becoming popular, particularly in what is known as the 'Oriental Period' (1560 - 1620). In his Hortus Medicus (1588), the first catalogue of a German garden's plants, Joachim Camerarius the Younger states that nine different types of daffodils were represented in his garden in Nuremburg. After his death in 1598, his plants were moved by Basilius Besler to the gardens they had designed at Willibaldsburg, the Bishop's palace at Eichstätt, Upper Bavaria. That garden is described in Besler's Hortus Eystettensis (1613) by which time there were 43 different types present.

N. poeticus, Thomas Hale, Eden: Or, a Compleat Body of Gardening 1757

Although not achieving the sensationalism of tulips, daffodils and narcissi have been much celebrated in art and literature (see The Arts). The most popular narcissi were large trumpet daffodils, N. poeticus and N. bulbocodium, and Istanbul became important in the shipping of bulbs to western Europe. By the early baroque period both tulips and narcissi were an important component of the spring garden. By 1739 a Dutch nursery catalogue listed 50 different varieties. A catalog of a Dutch nursery from 1739 already counted on 50 varieties. In 1757 Hill gave an account of the history and cultivation of the daffodil in his edited version of the works of Thomas Hale, writing "The garden does not afford, in its Kind, a prettier plant than this; nor do we know one that has been so early, or so honorably mention'd by all Kinds of Writers" (see illustration). Interest grew further when varieties that could be grown indoors became available, primarily the bunch flowered (multiple flower heads) N. tazetta. Narcissi became an important floricultural crop in Western Europe in the late nineteenth century and by the beginning of the twentieth century 50 million bulbs of N. Tazetta 'Paperwhite' were being exported annually from the Netherlands to the United States. With the production of triploids such as 'Golden Spur', in the late nineteenth century, and in the beginning of the twentieth century, tetraploids like 'King Alfred' (1899), the industry was well established, with trumpet daffodils dominating the market. The Royal Horticultural Society has been an important factor in promoting narcissi, holding the first Daffodil Conference in 1884, while the Daffodil Society, the first organisation dedicated to the cultivation of narcissi was founded in Birmingham in 1898. Later the American Daffodil Society was founded in 1954, and its quarterly journal is the leading trade publication.

Daffodil trumpets

Narcissi are now popular as ornamental plants for gardens, parks and as cut flowers, providing colour from the end of winter to the beginning of summer in temperate regions. They are one of the major ornamental spring flowering bulb crops, being produced both for their bulbs and cut flowers, though cultivation of private and public spaces is greater than the area of commercial production. Over a century of breeding has resulted in thousands of varieties and cultivars being available from both general and specialist suppliers. They are normally sold as dry bulbs to be planted in late Summer and Autumn (Fall). They are one of the most economically important ornamental plants. Plant breeders have developed some daffodils with double, triple, or ambiguously multiple rows and layers of segments. Much of the breeding programs have concentrated on the corona (trumpet or cup), in terms of its length, shape, and colour, and the surrounding perianth or even as in varieties derived from N. poeticus a very reduced form.

Narcissi as garden plants

While some wild narcissi are specific in terms of their ecological requirements, most garden varieties are relatively tolerant of soil conditions, however very wet soils and clay soils may benefit from the addition of sand to improve drainage. The optimum soil is a neutral pH of 7.0.

Bulbs offered for sale are referred to as either 'round' or 'double nose'. Round bulbs are circular in cross section and produce a single flower stem, while Double Nose bulbs have more than one bulb stem attached at the base and produce two or more flower stems, but bulbs with more than two stems are unusual. Planted narcissi bulbs produce daughter bulbs in the axil of the bulb scales, leading to the dying off of the exterior scales. To prevent planted bulbs forming more and more small bulbs, they can be dug up every 5 – 7 years, and the daughters separated and replanted separately, provided that a piece of the basal plate, where the rootlets are formed, is preserved.

Narcissi are well suited for planting under small thickets of trees, where they can be grouped as 6 - 12 bulbs. They also grow well in perennial borders, especially in association with day lilies which begin to form their leaves as the narcissi flowers are fading. A number of wild species and hybrids such as 'Dutch Master', 'Golden Harvest', 'Carlton', 'Kings Court' and 'Yellow Sun' naturalise well in lawns, but it is important not to mow the lawn till the leaves start to fade, since they are essential for nourishing the bulb for the next flowering season. Blue Scilla and Muscari which also naturalise well in lawns and flower at the same time as narcissus, make an attractive contrast to the yellow flowers of the latter. Unlike Tulips, narcissi bulbs are not attractive to rodents and are sometimes planted near tree roots in orchards to protect them.

Propagation

The commonest form of commercial propagation is by twin-scaling, in which the bulbs are cut into many small pieces in which two scales are still connected by a small fragment of the basal plate. The fragments are disinfected and placed on nutrient media. About 25 - 35 new plants can be produced from a single bulb after four years.

Breeding

For commercial use, varieties with a minimum stem length of 30 cm are sought, making them ideal for cut flowers. Florists require blooms that only open when they reach the retail outlet. For garden plants the objectives are to continually expand the colour palette and to produce hardy forms, and there is a particular demand for miniature varieties.

Classification

Range of Narcissus cultivars

Main article: List of Narcissus horticultural divisions

For horticultural purposes, all Narcissus cultivars are split into 13 divisions as first described by Kington (1998), for the Royal Horticultural Society (RHS), based partly upon flower form (shape and length of corona), number of flowers per stem, flowering period and partly upon genetic background. The classification is a useful tool for planning planting. Most commercially available narcissi come from Divisions 1 (Trumpet), 2 (Large cupped) and 8 (Tazetta). Division 13 which includes the wild daffodil is the exception to this scheme.

Growers register new daffodil cultivars by name and color with the Royal Horticultural Society, which is the international registration authority for the genus. Their International Daffodil Register is regularly updated with supplements available online and is searchable. The most recent supplemt (2014) is the sixth (the fifth was published in 2012). More than 27,000 names were registered as of 2008, and the number has continued to grow. Registered daffodils are given a division number and colour code such as 5 W-W ('Thalia'). In horticultural usage it is common to also find an unofficial Division 14: Miniatures, which although drawn from the other 13 divisions, have their miniature size in common. Over 140 varieties have gained the Royal Horticultural Society's Award of Garden Merit (See List of Award of Garden Merit narcissus).

Colour code

Narcissus 'Geranium' 8 W-O

Daffodils may be self-colored—i.e., both perianth and corona identical in color and shade—or the colors between the perianth and corona may differ widely. Some perianths and some coronas may also contain more than one color or shade. Prevalent colors are all shades and tones of yellow, white, orange, pink, red and green. Pinks vary from apricot to rose in shades from pale to deep, and some more recent cultivars have hints of lavender or lilac. Reds vary from orange-red to salmon red to near scarlet. Pink, red, orange and green tones are mainly confined to the corona. However, breeders are currently working against the genus' natural pigmentation and genetic barriers to create cultivars in which pink, rose, red, orange and green tones suffuse or "bleed" from the more highly colored coronas onto the perianth segments of white or yellow. There is an increasing number of commercially available varieties which display this enhanced coloration.

The color classification lists the perianth color and then the corona color. In the case of multiple colors, the perianth colors are assigned from the outer edge of the perianth segments inward to their juncture with the base of the corona, while the corona colors are assigned from the base of the corona outward to the rim. Thus, 'Accent', a Large Cup (Division 2) Daffodil possessing a white perianth and a pink corona, is officially classifies as 2 W-P, 'Geranium', Tazetta (Division 8) with a white perianth and orange corona is 8 W-O and 'Actaea', Poeticus (Division 9), with white corona and multicoloured corona is 9 W-GYR.

RHS Color Classification

Code	color
W	White
	whitish
G	Green
Y	Yellow
P	Pink
O	Orange
R	Red

Toxicity

Pharmacology

All Narcissus species contain the alkaloid poison lycorine, mostly in the bulb but also in the leaves. Members of the monocot subfamily Amaryllidoideae present a unique type of alkaloids, the norbelladine alkaloids, which are 4-methylcatechol derivatives combined with tyrosine. They are responsible for the poisonous properties of a number of the species. Over 200 different chemical structures of these compounds are known, of which 79 or more are known from Narcissus alone.

The toxic effects of ingesting Narcissus products for both man and animals (such as cattle, goats, pigs and cats) have long been recognised and they have been used in suicide attempts. Ingestion of N. pseudonarcissus or N. jonquilla is followed by salivation, acute abdominal pains, nausea, vomiting, and diarrhea, then neurological and cardiac events, including trembling, convulsions, and paralysis. Death may result if large quantities are consumed. The toxicity of Narcissus varies with species, N. poeticus being more toxic than N. pseudonarcissus, for instance. Nor is the distribution of toxins even within the plant, for instance there is a five times higher concentration of alkaloid in the stem of N. papyraceous than in the bulb, making it dangerous to herbivores more likely to consume the stem than the bulb, and is part of the plant's defence mechanisms.

Not all Narcissus species are equally dangerous. The bulbs of N. poeticus, for example, are more dangerous than those of N. pseudonarcissus. Neither do all plant tissues have the same concentration or profile of alkaloids. Thus, the alkaloid content of N. papyraceus is five times higher in the aerial part than in the bulbs, being toxic for herbivorous mammals. The distribution of the alkaloids in the plant tissues can be related with the plant's defense mechanism such as protection from parasites. The bulbs can also be toxic to other nearby plants, including roses, rice, and cabbages, inhibiting growth. For instance placing cut flowers in a vase alongside other flowers shortens the life of the latter.

Poisoning

Many cases of poisoning or death have occurred when narcissi bulbs have been mistaken for leeks or onions and cooked and eaten. Recovery is usually complete in a few hours without any specific intervention. In more severe cases involving ingestion of large quantities of bulbs activated charcoal, salts, and laxatives may be required, and for severe symptoms intravenous atropine and emetics or stomach pumping may be indicated. However, ingestion of large quantities accidentally is unusual because of a strong unpleasant taste. When narcissi were compared with a nunber of other plants not normally consumed by animals, narcissi were the most repellant, specifically N. pseudonarcissus Consequently Narcissus alkaloids have been used as repellents and may also discourage fungi, molds, and bacteria.

On 1 May 2009 a number of schoolchildren fell ill at Gorseland Primary School in Martlesham Heath, Suffolk, England, after a daffodil bulb was added to soup during a cookery class.

Topical effects

One of the most common dermatitis problems for flower pickers, packers, florists and gardeners, "daffodil itch", involves dryness, fissures, scaling, and erythema in the hands, often accompanied by subungual hyperkeratosis (thickening of the skin beneath the nails). It is blamed on exposure to calcium oxalate, chelidonic acid or alkaloids such as lycorine in the sap, either due to a direct irritant effect or an allergic reaction. It has long been recognised that some cultivars provoke dermatitis more readily than others. N. pseudonarcissus and the cultivars 'Actaea', 'Camparelle', 'Gloriosa', 'Grande Monarque', 'Ornatus', 'Princeps' and 'Scilly White' are known to do so.

If bulb extracts come into contact with wounds, both central nervous system and cardiac symptoms may result. The scent can also cause toxic reactions such as headaches and vomiting from N. bulbocodium.

Uses

Traditional medicine

Despite the lethal potential of Narcissus alkaloids, they have been used for centuries as traditional medicines for a variety of complaints, including cancer. N. poeticus is described in the Bible in the treatment of cancer. In the Classical Greek world Hippocrates (ca. B.C. 460–370) recommended a pessary prepared from Narcissus oil for uterine tumors, a practice continued by Pedanius Dioscorides (ca. A.D. 40–90) and Soranus of Ephesus (A.D. 98–138) in the first and second centuries A.D., while the Roman Pliny the Elder (A.D. 23–79), advocated topical use. The bulbs of N. poeticus contain the antineoplastic agent narciclasine. This usage is also found in later Arabian, North African, Central American and Chinese medicine during the Middle Ages. In China N. tazetta var. chinensis was grown as an ornamental plant but the bulbs were applied topically to tumors in traditional folk medicine. These bulbs contain pretazettine, an active antitumor compound.

Narcissus products have received a variety of other uses. The Roman physician Aulus Cornelius Celsus listed narcissus root in De Medicina among medical herbs, described as emollient, erodent, and "powerful to disperse whatever has collected in any part of the body". N. tazetta bulbs were used in Turkey as a remedy for abscesses in the belief they were antiphlogistic and analgesic. Other uses include the application to wounds, strains, painful joints and various local ailments as an ointment called ‘Narcissimum’. Powdered flowers have also been used medically, as an emetic, a decongestant and for the relief of dysentery, in the form of a syrup or infusion. The French used the flowers as an antispasmodic, the Arabs the oil for baldness and also an aphrodisiac. In the eighteenth century the Irish herbal of John K’Eogh recommended pounding the roots in honey for use on burns, bruises, dislocations and freckles, and for drawing out thorns and splinters. N. tazetta bulbs have also been used for contraception, while the flowers have been recommended for hysteria and epilepsy. A homeopathic medicine made from bulbs was prescribed for bronchitis and whooping cough. In the traditional Japanese medicine of kampo, wounds were treated with narcissus root and wheat flour paste; the plant, however, does not appear in the modern kampo herb list.

There is also a long history of the use of Narcissus as a stimulant and to induce trance like states and hallucinations. Sophocles referred to the narcissus as the ‘Chaplet of the infernal Gods’, a statement frequently wrongly attributd to Socrates (see Antiquity).

Biological properties

Extracts of Narcissus have demonstrated a number of potentially useful biological properties including antiviral, prophage induction, antibacterial, antifungal, antimalarial, insecticidal, cytotoxic, antitumor, antimitotic, antiplatelet, hypotensive, emetic, acetylcholine esterase inhibitory, antifertility, antinociceptive, chronotropic, pheromone, plant growth inhibitor, and allelopathic. An ethanol extract of Narcissus bulbs was found effective in one mouse model of nociception, para-benzoquinone induced abdominal constriction, but not in another, the hot plate test. Most of these properties are due to alkaloids, but some are also due to mannosa-binding lectins. The most-studied alkaloids in this group are galanthamine, lycorine, narciclasine, and pretazettine.

It is likely that the traditional use of narcissi for the treatment of cancer was due to the presence of isocarbostyril constituents such as narciclasine, pancratistatin and their congeners. N. poeticus contains about 0.12g of narciclasine per kg of fresh bulbs.

Acetylcholine esterase inhibition has attracted the most interest as a possible therapeutic intervention, with activity varying by a thousandfold between species, and the greatest activity seen in those that contain galanthamine or epinorgalanthamine.

The rodent repellant properties of Narcissus alkaloids have been utilised in horticulture to protect more vulnerable bulbs.

Therapeutics

Daffodils growing in Wales

Of all the alkaloids, only galanthamine has made it to therapeutic use in man, as the drug galantamine for Alzheimer's disease. Galanthamine is an acetylcholine esterase inhibitor which crosses the blood brain barrier and is active within the central nervous system. Daffodils are grown commercially near Brecon in Powys, Wales, to produce galantamine, a drug used to combat Alzheimer's disease.

Commercial uses

Throughout history the scent of narcissi has been an important ingredient of perfumes, a quality that comes from essential oils rather than alkaloids. Narcissi are also an important horticultural crop, and source of cut flowers (floriculture).

Daffodil production in the Netherlands

The Netherlands, which is the most important source of flower bulbs worldwide is also the centre of narcissus production. Of 16,700 hectares under cultivation for flower bulbs, narcissi account for about 1,800 hectares. In the 1990s narcissus bulb production was at 260 million, sixth in size after tulips, gladioli, irises, crocuses and lilies. About two thirds of the area under cultivation is dedicated to about 20 of the most popular varieties, particularly 'Carlton' (yellow) and 'Ice Follies' (white), both of which are large-cupped narcissi (Division Two). These two varieties have a long history of cultivation, together with 'Dutch Master' and 'Golden Harvest'. 'Carlton' and 'Golden Harvest' were introduced in 1927, and 'Ice Follies' in 1953. 'Carlton' is the commonest cultivar with over 9000 million bulbs (350 000 tons), probably the largest individual plant production in the world. The other major area of production is the United Kingdom.

Narcissi growing at Keukenhof

Bulbs are harvested for market in the summer, sorted, stored for 2 – 3 weeks, and then disinfected by a hot (43.5 °C) bath. This eliminates infestations by narcissus fly and nematodes. The bulbs are then dried at a high temperature, and then stored at 15.5 °C. Traditionally, sales took place in the daffodil fields prior to harvesting the bulbs, but today sales are handled by Marketing Boards although still before harvesting. In the Netherlands there are special exhibition gardens for major buyers to view flowers and order bulbs, some larger ones may have more than a thousand narcissus varieties on display. While individuals can visit these gardens they cannot buy bulbs at retail, which are only available at wholesale, usually at a minimum of several hundredweight. The most famous display is at Keukenhof, although only about 100 narcissus varieties are on display there.

Forcing

There is also a market for both forced blooms in pots and cut flowers through the winter from Christmas to Easter, the long season requiring special preparation by growers. To bloom in December, bulbs are harvested in June, dried, stored for four days at 34 °C, two weeks at 30 and two weeks at 17 and then placed in cold storage at 9 degrees. The bulbs are then planted in light compost in crates in a greenhouse and the blooms appear in 19 – 30 days. For later blooming, the higher temperatures are omitted, being stored a 17 degrees after harvesting and placed in cold storage in September. The bulbs can then be planted in cold frames, and then forced in a greenhouse according to requirements.

Culture

Symbols

N. triandrus 'Thalia', considered a grave flower

The daffodil is the national flower of Wales, where it is traditional to wear a daffodil or a leek on Saint David's Day (March 1). In Welsh the daffodil is known as "Peter's Leek", cenhinen Bedr or cenin Pedr). The narcissus is also a national flower symbolising the new year or Newroz in the Kurdish culture.

The narcissus is perceived in the West as a symbol of vanity, in the East as a symbol of wealth and good fortune (see Eastern cultures). In classical Persian literature, the narcissus is a symbol of beautiful eyes, together with other flowers that equal a beautiful face with a spring garden, such as roses for cheeks and violets for shining dark hair. In some countries the yellow variation is associated with Easter. The German for daffodil is Osterglocke, that is "Easter bell."

Although prized as an ornamental flower, some people consider narcissi unlucky, because they hang their heads implying misfortune, and hence refuse to have them in the house. White narcissi are especially associated with death, especially the pure white N triandrus 'Thalia', and hence are considered grave flowers. Indeed, in Ancient Greece narcissi were planted near tombs. Robert Herrick, describes them as portents of death, an association which also appears in the myth of Persephone and the underworld (see The Arts, below).

The Arts

Antiquity

N. tazetta growing in Israel

Demeter and Persephone surrounded by Narcissi

Narcissi have been used decoratively for a long time, a wreath of white-flowered N. tazetta having been found in an ancient Egyptian grave, and in frescoes on the excavated walls of Pompeii. It is thought to have been mentioned in the Bible, for instance in the Book of Isaiah. The rose mentioned here being the original translation into English from the Biblical Hebrew word chabatstsileth (Hebrew: חבצלת‎). This so-called "Rose of Sharon" being actually a bulbous plant, probably N. tazetta which grows in Israel on the Plain of Sharon, where it is a protected plant. They make a frequent appearance in classical literature.

Greek culture

The narcissus has also frequently appeared in literature and the visual arts, and forms part of two important Graeco-Roman myths, that of the youth Narcissus who was turned into the flower of that name, and of the Goddess Kore, or Persephone (Greek: Περσεφόνη, Latin: Proserpina) daughter of the goddess Demeter, snatched into the Underworld by the god Hades while picking narcissi. Hence, the narcissus is listed as having been sacred to both Hades and Persephone, and to grow along the banks of the river Styx in the underworld.

The Greek poet Stasinos (flourished ca. 800-900 BC) mentioned them in the Cypria in which he sings of the flowers of the island of Cyprus "She clothed herself with garments which the Graces and Hours had made for her and dyed in flowers of spring -- such flowers as the Seasons wear -- in crocus and hyacinth and flourishing violet and the rose's lovely bloom, so sweet and delicious, and heavenly buds, the flowers of the narcissus and lily. In such perfumed garments is Aphrodite clothed at all seasons. " (ll. 1-7). The legend of Persephone comes to us primarily in the anonymous Homeric Hymn (Seventh century BC) to Demeter, which describes the narcissus as being grown as a lure to trap the young Persephone, the last flower she reached for l. 428 "νάρκισσόν θ᾽, ὃν ἔφυσ᾽ ὥς περ κρόκον εὐρεῖα χθών" (and the narcissus which the wide earth caused to grow yellow as a crocus).

Other Greek authors making reference to the narcissus include Sophocles (c. 497 - 406 BC) and Plutarch (c. 46 AD – 120 AD). Sophocles, in his Oedipus at Colonus utilises narcissus in a highly symbolic manner, implying fertility, and allying it with the cults of Demeter and her daughter Kore (Persephone) (μεγάλαιν θεαίν, the Great Goddesses), but by extension through the Persephone association, a symbol of death. Jebb comments here that νάρκισσος is the flower of imminent death with its fragrance being νάρκη or narcotic, emphasised by its pale white colour. Just as Persephone reaching for the flower heralded her doom, the youth Narcissus gazing at his own reflection portended his death.

θάλλει δ ουρανίας υπ άχνας
ο καλλίβοτρυς κατ ημαρ αει
νάρκισσος, μεγάλαιν θεαίν
αρχαιον στεφάνωμ
"And, fed on heavenly dew,
the narcissus blooms day by day with its fair clusters;
it is the ancient crown of the Great Goddesses."
Sophocles. Oedipus at Colonus, 681 - 684

Plutarch refers to this in his Symposiacs as follows, "and the daffodil, because it benumbs the nerves and causes a stupid narcotic heaviness in the limbs, and therefore Sophocles calls it the ancient garland flower of the great (that is, the earthy) gods." This reference to Sophocles' "crown of the great Goddesses", here is the source of the commonly quoted phrase in the English literature "Chaplet of the infernal Gods" incorrectly attributed to Socrates.

Another Greek writer was Homer (ca. 7th century BC), who in his Odyssey in several places (e.g. Od. 11:539; 24.14) described the underworld as having Elysian meadows carpeted with flowers, though using the term asphodel (ἀσφοδελὸν) rather than narcissus. This may well have actually been narcissus, with its associations with the underworld, as described by Theophrastus, and frequently used in later literature to refer to daffodils. A similar account is provided by Lucian (c. 125 – 180 AD) in his Necyomantia or Menippus (Μένιππος ἢ Νεκυομαντεία), describing Asphodel in the underworld (Nec. 11:2; 21:10).

The myth of the youth Narcissus is also taken up by Pausanias (Greek: Παυσανίας, c. 110 – 180AD) in his Description of Greece. Pausanias, deferring to Pamphos, believed that the myth of Persephone long antedated that of Narcissus, and hence discounts the idea the flower was named after the youth.

νάρκισσον δὲ ἄνθος ἡ γῆ καὶ πρότερον ἔφυεν ἐμοὶ δοκεῖν, εἰ τοῖς Πάμφω τεκμαίρεσθαι χρή τι ἡμᾶς ἔπεσι: γεγονὼς γὰρ πολλοῖς πρότερον ἔτεσιν ἢ Νάρκισσος ὁ Θεσπιεὺς Κόρην τὴν Δήμητρός φησιν ἁρπασθῆναι παίζουσαν καὶ ἄνθη συλλέγουσαν, ἁρπασθῆναι δὲ οὐκ ἴοις ἀπατηθεῖσαν ἀλλὰ ναρκίσσοις.
"The flower narcissus grew, in my opinion, before this, if we are to judge by the verses of Pamphos. This poet was born many years before Narcissus the Thespian, and he says that the Maid, the daughter of Demeter, was carried off when she was playing and gathering flowers, and that the flowers by which she was deceived into being carried off were not violets, but the narcissus."
Pausanias. Description of Greece. 9 Boeotia. 31:9

Roman culture

Virgil (Publius Vergilius Maro, 70 BC – 19 AD), the first known Roman writer to refer to the narcissus, does so in several places, for instance twice in the Georgics, Book four, l. 122 "nec sera comantem Narcissum" (nor had I passed in silence the late-flowering narcissus) and l. 159 "pars intra septa domorum, Narcissi lacrymam" (some within the enclosure of their Hives, lay Narcissus' tears). Virgil refers to the cup shaped corona of the narcissus flower, allegedly containing the tears of the youth Narcissus. Milton makes a similar analogy in his Lycidas "And Daffodillies fill their Cups with Tears". Virgil also mentions narcissi three times in the Eclogues. In the second book l. 48 "Narcissum et florem jungit bene olentis anenthi" (joins the narcissus and flower of sweet-smelling Anise), also the fifth book, l. 38 "pro purpureo narcisso" (in lieu of the empurpled narcissus). For the idea that Narcissus could be purple, see also Dioscorides (επ ενίων δε πορφυροειδές) and Pliny (sunt et purpurea lilia). This is thought to be an allusion to the purple rimmed corona of N. poeticus. Finally, in the eighth book of the Eclogues, Virgil writes, l. 53 "narcisso floreat alnus" (the alder with narcissus bloom).

Ovid (Publius Ovidius Naso, 43 BC – 17 AD) was also familiar with narcissi, in his recounting of the self-loving youth who is turned into the flower, in the third book of his Metamorphoses l. 509 "croceum pro corpore florem inveniunt, foliis medium cingemtibus albis"(They came upon a flower, instead of his body, with white petals surrounding a yellow heart) and also the fifth book of his Fasti l. 201 "Tu quoque nomen habes cultos, Narcisse, per hortos" (You too, Narcissus, were known among the gardens). This theme of metamorphosis was broader than just Narcissus, for instance see Crocus and Hyacinth.

Western culture

Narcissus in art

Poussin: Death of Narcissus, 1630

Waterhouse: Echo and Narcissus, 1903

Dali: Metamorphosis of Narcissus, 1937

Jan van Scorel: Madonna of the Daffodils with the Child and Donors, 1535

Vincent van Gogh: Undergrowth with Two Figures, 1890

Waterhouse: Narcissus, 1912

Although there is no clear evidence that the flower's name derives directly from the Greek myth this link between the flower and the myth became firmly part of western culture.

"When a daffadill I see, Hanging down his head t'wards me, Guesse I may, what I must be: First, I shall decline my head; Secondly, I shall be dead: Lastly, safely buryed."

Herrick, 'Divination by a Daffadill', Hesperides 1648.

The narcissus or daffodil appears frequently in British poetry, and no flower has received more poetic description except the Rose and the Lily, with poems by authors including Shakespeare, Spenser, Addison and Thomson, together with Milton (see above), Keats and Shelley. Frequently the poems deal with self-love derived from Ovid's account. Shakespeare, who frequently uses flower imagery, refers to daffodils twice in his The Winter's Tale (e.g. "Daffodils, That come before the swallow dares, and take The winds of March with beauty"). Robert Herrick, in his Hesperides (1648) alludes to their association with death in a number of poems such as To Daffadils and Divination by a Daffadill. Amongst the English romantic movement writers none is better known than William Wordsworth's short 1804 poem I Wandered Lonely as a Cloud (The Daffodils) which has become linked in the popular mind with the daffodils that form its main image, here associated with vitality and pleasure. Wordsworth also included the daffodil in other poems, such as Foresight.

I wandered lonely as a Cloud

I wandered lonely as a Cloud
That floats on high o'er Vales and Hills,
When all at once I saw a crowd
A host of dancing Daffodils;
Along the Lake, beneath the trees,
Ten thousand dancing in the breeze.

The waves beside them danced, but they
Outdid the sparkling waves in glee: –
A poet could not but be gay
In such a laughing company:
I gaz'd – and gaz'd – but little thought
What wealth the shew to me had brought:

For oft when on my couch I lie
In vacant or in pensive mood,
They flash upon that inward eye
Which is the bliss of solitude,
And then my heart with pleasure fills,
And dances with the Daffodils.

William Wordsworth (1804 version)

More recently Cecil Day-Lewis wrote "Now the full throated daffodils, Our trumpeters in gold, Call resurrection from the ground, And bid the year be bold" in his From Feathers to Iron (1931).

In German literature, Paul Gerhardt, a pastor and hymn writer wrote his Geh aus, mein Herz, und suche Freud in 1653 in which the second verse states:

"Narzissus und die Tulipan
Die ziehen sich viel schöner an,
Als Salomonis Seide"

(Daffodil and Tulip are dressed more beautifully than Solomon's silk)

The plant name was used in the title of the 1939 novel by Rumer Godden, a story about the disorientation of European nuns in India. The book was later adapted into the 1947 award-winning British film Black Narcissus.

In the visual arts, narcissi are depicted in three different contexts, mythological (Narcissus, Persephone), floral art, or landscapes. The Narcissus story has been popular with painters and he is frequently depicted with flowers to indicate this association, for instance François Lemoyne, John William Waterhouse, Poussin's with flowers sprouting around the dying youth, or Salvador Dalí's painting Metamorphosis of Narcissus. The Persephone theme is typified by Waterhouse's Narcissus, the floral motif by van Scorel and the landscape by Van Gogh's Undergrowth. Narcissi first started to appear in western art in the late middle ages, in panel paintings, particularly those depicting crucifixion. For instance there is a crucifixion scene in the Wallraf-Richartz-Museum (Cologne), where daffodils symbolise not only death but also hope in the resurrection, and also because they are perennial and bloom at Easter. Another example from this period is the altar panel Noli me tangere by the Meister des Göttinger Barfüßeraltars (c. 1410) showing daffodils growing between the hand of Jesus and Mary Magdalene.

Eastern cultures

In Chinese culture interest in narcissi centres on Narcissus tazetta which can be grown indoors. N. tazetta subsp. chinensis, best known outside China as N. tazetta 'Chinese Sacred Lily' is widely grown in China as an ornamental plant. Tazetta daffodils were probably introduced to China by Arab traders travelling the Silk Road at some time prior to the beginning of the Song Dynasty, presumably for their claimed medicinal properties, where they became naturalised. Flowering in the spring, they became associated with the Chinese New Year, signifying good fortune, prosperity and good luck. If the narcissus blooms on Chinese New Year, it is said to bring extra wealth and good fortune throughout the year. Its sweet fragrances are also highly revered in Chinese culture. The flower has many names in Chinese culture. These include water narcissus (since they can be grown in water), seui sin faa (water immortal flowers), or joss flowers. In ancient Chinese culture the narcissus is referred to as water goddess of the Xiang River (Chinese: 水仙, shuǐ xiān), or the "Goddess standing above the waves" (lingbo xianzi). There is also a legend about a poor but good man who was brought many cups of gold and wealth by this flower.

Chinese decorative carved Narcissus bulb

As noted Chinese Garden Art expert Marianne Beuchert writes, in contrast to the west, narcissi have not played as significant part in Chinese Garden art, but have become a symbol of good luck, in which the multi headed inflorescence of N. tazetta symbolised a hundred headed water spirit. However, Zhao Mengjian (趙孟堅, c. 1199 – 1267), in the Southern Song Dynasty was noted for his portrayal of narcissi, and Zhao's love of the flower is celebrated by the poet Qiu Yuan (c. 1247 – 1327). Narcissus bulb carving and cultivation has become an art akin to Japanese bonsai. The bulbs are carved to create curling leafs (crab claw culture). The bulbs are often divided, to produce many flower stems from a single bulb (twin-scaling).

The Japanese visual novel Narcissu contains many references to the narcissus.

Islamic culture

N. poeticus symbolising the eye in Islamic culture

Islamic scholar Annemarie Schimmel states that the narcissi are one of the most popular garden plants in Islamic culture. The Persian ruler Chusrau Anuschirwan, or Khosrau I (531 - 579) is said to have not been able to tolerate them at feasts because they reminded him of eyes, an association that persists to this day, as described by the poet Ghalib (1797 – 1869), "God has given the eye of the narcissus the power of seeing". The imagery could also be negative, such as blindness (white eye), sleepless or longing for love. The eye imagery is also found in a number of poems by Abu Nuwas (756–814). In one of his most famous poems about narcissi he writes "eyes of silver with pupils of molten gold united with an emerald stalk". Schimmel describes an Arab legend that despite the apparent sinfulness of much of his poetry, his narcissus poems alone would earn him a place in Paradise. Another poet who refers to narcissi, is Rumi (1207 – 1273). Even the prophet Mohammed is said to have praised the narcissus, "Whoever has two loaves of bread, sell one and buy narcissi, for while bread nourishes the body, the narcissus feeds the soul".

Festivals

Narcissi (May Snow) growing near Montreux in May

In some areas where wild narcissi are particularly prevalent, their blooming in spring is celebrated in festivals. The slopes around Montreux, Switzerland and its associated riviera come alive with blooms each May (May Snow), and are associated with the Narcissi Festival. However, the narcissi are now considered threatened. Festivals are held in many other countries and regions including Fribourg (Switzerland), Austria and in the United States, including Hawaii (Chinese New Year) and Washington State's Daffodil Festival.

Cancer

Various cancer charities around the world, including the American Cancer Society, New Zealand Cancer Society, Cancer Council Australia, the Irish Cancer Society, and Marie Curie (UK)'s Great Daffodil Appeal use the daffodil emblem as a fundraising symbol. "Daffodil Days", first instituted in Toronto in 1957 by the Canadian Cancer Society, are organized to raise funds by offering the flowers in return for a donation.

See also

• List of AGM narcissus - list of narcissus species and cultivars which have gained the Royal Horticultural Society's Award of Garden Merit
• List of Narcissus species

Bibliography

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Flora

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Narcissus

Articles

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Phylogeny

• Graham, S. W.; Barrett, S. C. H. (1 July 2004). "Phylogenetic reconstruction of the evolution of stylar polymorphisms in Narcissus (Amaryllidaceae)". American Journal of Botany 91 (7): 1007–1021. doi:10.3732/ajb.91.7.1007. Retrieved 30 October 2014. 
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• Rønsted, Nina; Savolainen, Vincent; Mølgaard, Per; Jäger, Anna K. (May–June 2008). "Phylogenetic selection of Narcissus species for drug discovery". Biochemical Systematics and Ecology 36 (5-6): 417–422. doi:10.1016/j.bse.2007.12.010. Retrieved 30 October 2014. 
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• Fernández-Casas, Francisco Javier (2008). "Narcissorum notulae, X". Fontqueria 55: 547–558. Retrieved 20 October 2014. 
• Jiménez, Juan F.; Sánchez-Gómez, Pedro; Guerra, Juan; Molins, Arántzazu; Rosselló, Josep A. (12 June 2009). "Regional Speciation or Taxonomic Inflation? The Status of Several Narrowly Distributed and Endangered Species of Narcissus Using ISSR and Nuclear Ribosomal ITS Markers". Folia Geobotanica 44 (2): 145–158. doi:10.1007/s12224-009-9040-2. Retrieved 29 October 2014.  (Full text on Research Gate)
• Zonneveld, B. J.M. (8 July 2010). "The involvement of Narcissus hispanicus Gouan in the origin of Narcissus bujei and of cultivated trumpet daffodils (Amaryllidaceae)". Anales del Jardín Botánico de Madrid 67 (1): 29–39. doi:10.3989/ajbm.2219. Retrieved 5 October 2014. 
• Fernández-Casas, Francisco Javier (2011). "Narcissorum Notulae, XXXI". Fontqueria 56 (26): 239–248. Retrieved 5 October 2014. 
• Gage, Ewan; Wilkin, Paul; Chase, Mark W.; Hawkins, Julie (2011). "Phylogenetic systematics of Sternbergia (Amaryllidaceae) based on plastid and ITS sequence data". Botanical Journal of the Linnean Society 166 (2): 149–162. doi:10.1111/j.1095-8339.2011.01138.x. Retrieved 30 October 2014. 
• Marques, I.; Nieto Feliner, G.; Martins-Loucao, M. A.; Fuertes Aguilar, J. (11 November 2011). "Genome size and base composition variation in natural and experimental Narcissus (Amaryllidaceae) hybrids". Annals of Botany 109 (1): 257–264. doi:10.1093/aob/mcr282. Retrieved 30 October 2014. 
• Santos-Gally, Rocío; Vargas, Pablo; Arroyo, Juan (April 2012). "Insights into Neogene Mediterranean biogeography based on phylogenetic relationships of mountain and lowland lineages of Narcissus (Amaryllidaceae)". Journal of Biogeography 39 (4): 782–798. doi:10.1111/j.1365-2699.2011.02526.x. Retrieved 30 October 2014. 
• Medrano, Mónica; López-Perea, Esmeralda; Herrera, Carlos M. (June 2014). "Population Genetics Methods Applied to a Species Delimitation Problem: Endemic Trumpet Daffodils (Narcissus Section Pseudonarcissi) from the Southern Iberian Peninsula". International Journal of Plant Sciences 175 (5): 501–517. doi:10.1086/675977. Retrieved 5 October 2014. 

Pharmacology

• John M. Riddle, John M. (September 1985). "Ancient and Medieval Chemotherapy for Cancer". Isis 76 (3): 319–330. Retrieved 31 October 2014. 
• Bastida, Jaume; Lavilla, Rodolfo; Viladomat, Francesc Viladomat (2006). Cordell, G. A., ed. "Chemical and biological aspects of "Narcissus" alkaloids". The Alkaloids: Chemistry and Biology (Amsterdam: Elsevier Inc) 63: 87–179. doi:10.1016/S1099-4831(06)63003-4. Retrieved 31 October 2014. 
• Kornienko, Alexander; Evidente, Antonio (2008). "Chemistry, Biology, and Medicinal Potential of Narciclasine and its Congeners". Chemical Reviews 108 (6): 1982–2014. doi:10.1021/cr078198u. PMC 2856661. Retrieved 31 October 2014. 
• Pettit, George R.; Tan, Rui; Bao, Guan-Hu; Melody, Noeleen; Doubek, Dennis L.; Gao, Song; Chapuis, Jean-Charles; Williams, Lee (27 April 2012). "Antineoplastic Agents. 587. Isolation and Structure of 3-Epipancratistatin from Narcissus cv. Ice Follies". Journal of Natural Products 75 (4): 771–773. doi:10.1021/np200862y. PMID 22413911. Retrieved 31 October 2014. 
• Pigni, Natalia B.; Ríos-Ruiz, Segundo; Martínez-Francés, Vanessa; Nair, Jerald J.; Viladomat, Francesc; Codina, Carles; Bastida, Jaume (28 September 2012). "Alkaloids from Narcissus serotinus". Journal of Natural Products 75 (9): 1643–1647. doi:10.1021/np3003595. Retrieved 5 October 2014. 
• Pigni, Natalia B.; Ríos-Ruiz, Segundo; Luque, F. Javier; Viladomat, Francesc; Codina, Carles; Bastida, Jaume (November 2013). "Wild daffodils of the section Ganymedes from the Iberian Peninsula as a source of mesembrane alkaloids". Phytochemistry 95: 384–393. doi:10.1016/j.phytochem.2013.07.010. Retrieved 5 October 2014. 
• Takos, Adam; Rook, Fred (31 May 2013). "Towards a Molecular Understanding of the Biosynthesis of Amaryllidaceae Alkaloids in Support of Their Expanding Medical Use". International Journal of Molecular Sciences 14 (6): 11713–11741. doi:10.3390/ijms140611713. Retrieved 5 October 2014. 
• Berkov, Strahil; Martínez-Francés, Vanessa; Bastida, Jaume; Codina, Carles; Ríos, Segundo (March 2014). "Evolution of alkaloid biosynthesis in the genus Narcissus". Phytochemistry 99: 95–106. doi:10.1016/j.phytochem.2013.11.002. Retrieved 31 October 2014. 

Books

• Haworth, A.H. (1831). Narcissinearum Monographia (2 ed.). London: Ridgway. Retrieved 20 October 2014. 
• Burbidge, Frederick William (1875). The Narcissus: Its History and Culture: With Coloured Plates and Descriptions of All Known Species and Principal Varieties.. London: L. Reeve & Company. Retrieved 28 September 2014.  (also available as pdf)
• Barr, Peter; Burbidge, F.W. (1884). Ye Narcissus Or Daffodyl Flowere, Containing Hys Historie and Culture, &C., With a Compleat Liste of All the Species and Varieties Known to Englyshe Amateurs. London: Barre & Sonne. ISBN 978-1104534271. Retrieved 2 November 2014. 
• Blanchard, J. W. (1990). Narcissus: a guide to wild daffodils. Surrey, UK: Alpine Garden Society. ISBN 0900048530. Retrieved 2 October 2014. 
• Erhardt, Walter (1993). Narzissen : Osterglocken, Jonquillen, Tazetten. Stuttgart (Hohenheim): E. Ulmer. ISBN 3800164892. Retrieved 18 October 2014. 
• Hanks, Gordon R (2002). Narcissus and Daffodil: The Genus Narcissus. London: Taylor and Francis. ISBN 0415273447. Retrieved 2 October 2014. 

Websites

• Libertiny, Susan (5 February 2013). "Peter Barr – The King of Daffoldils". All about my garden. Retrieved 2 November 2014. 
• Trinklein, David (2007). "Spring Flowering Bulbs: Daffodils". University of Missouri Extension. Retrieved 25 October 2014. 
• Van Beck, John; Christman, Steve (3 Oct 2005). "Narcissus". Floridata. Retrieved 25 October 2014. 
• Vigneron, Pascal. "Narcissus". Amaryllidaceae.org (in French). Retrieved 23 October 2014. 

Historical research

• Anonymous (May–October 1887). "Homer the botanist". Macmillan's Magazine (London: Macmillan and Company) 56: 428–436. Retrieved 3 November 2014. 
• Arber, Agnes (1912). Herbals, Their Origin and Evolution: A Chapter in the History of Botany 1470-1670. Cambridge University Press 1912. Retrieved 2 November 2014. 
• Krausch, Heinz D (2012). 'Kaiserkron und Päonien rot?': Entdeckung und Einführung unserer Gartenblumen. Munich: Dölling und Galitz Verlag G. ISBN 3862180220. Retrieved 16 October 2014. 

Literature and art

Antiquity

• Singer, Samuel Weller, ed. (1846). Hesperides: or, The works both humane and divine of Robert Herrick, Volume 1. London: William Pickering. Retrieved 1 October 2014. 
• Jebb, Sir Richard Claverhouse, ed. (1889). Sophocles: the plays and fragments with critical notes, commentary, and translation in English prose. Part II. The Oedipus Coloneus (2 ed.). Cambridge University Press. 
• McClintock, John; Strong, James (1889). Cyclopaedia of Biblical, Theological, and Ecclesiastical Literature, Vol. IX RH-ST. New York: Harper & Brothers. Retrieved 8 October 2014. 
• Zimmerman, Clayton (1994). The pastoral Narcissus : a study of the first idyll of Theocritus. Lanham: Rowman & Littlefield. ISBN 0847679624. Retrieved 19 October 2014. 
• Jashemski, Wilhelmina Feemster; Meyer, Frederick G., eds. (2002). The natural history of Pompeii. New York: Cambridge University Press. ISBN 0521800544. Retrieved 19 October 2014. 
• Reece, Steve (2007). "Homer’s Asphodel Meadow". Greek, Roman, and Byzantine Studies 47 (4): 389–400. Retrieved 7 October 2014. 

Renaissance

• Shakespeare, William (1623). "The Winter's Tale". The Complete Works of William Shakespeare. Retrieved 6 November 2014. 
• D'Ancona, Mirella Levi (1977). Garden of the Renaissance: Botanical Symbolism in Italian Painting. Firenze: Casa Editrice Leo S.Olschki. ISBN 9788822217899. 
• Sparrow, Lady Amie (November 2007). "Flowers and Their Renaissance Symbolism". The Bull, Newsletter for the Barony of Stierbach, Vol. 10 Issue XI. Retrieved 6 October 2014. 

Modern

• Wordsworth, William (1807). Poems in Two Volumes, VOL. II. Longman, Hurst, Rees and Orme. Retrieved 12 October 2014. 
• Lewis, C. Day (1992). The Complete Poems. Stanford, California: Stanford University Press. ISBN 0804725853. Retrieved 5 November 2014. 

Islam

• Schimmel, Annemarie (1984). Stern und Blume : die Bilderwelt der persischen Poesie. Wiesbaden: O. Harrassowitz. ISBN 3447024348. Retrieved 16 October 2014. 
• Schimmel, Annemarie (1992). Two-colored brocade : the imagery of Persian poetry.. Univ Of North Carolina. ISBN 0807856207. Retrieved 15 October 2014. 
• Schimmel, Annemarie (1998). Die Träume des Kalifen : Träume und ihre Deutung in der islamischen Kultur. München: Beck. ISBN 3406440568. Retrieved 16 October 2014. 
• Schimmel, Annemarie (2001). Kleine Paradiese : Blumen und Gärten im Islam. Freiburg im Breisgau: Herder. ISBN 3451051923. 
• Meisami, Julie Scott; Starkey, Paul, eds. (1998). Encyclopedia of Arabic literature, vol. 2. London: Routledge. ISBN 0415185726. Retrieved 16 October 2014. 

Databases

• "Narcissus". The Plant List (2013). Version 1.1. 2013. Retrieved 3 November 2014. 
• "Narcissus L.". International Plant Names Index. Retrieved 16 October 2014. 
• "Narcissus". World Checklist of Selected Plant Families. Royal Botanic Gardens, Kew. Retrieved 16 October 2014. 

Societies and organisations

• "The American Daffodil Society". 2014. Retrieved 4 November 2014. 
  • "DaffLibrary – books, articles, and journals about daffodils". American Daffodil Society. Retrieved 4 November 2014. 
  • "DaffSeek - Daffodil Database with Photos". American Daffodil Society. Retrieved 4 November 2014. 
• "Daffodils". Royal Horticultural Society. Retrieved 4 November 2014. 
• "IBS Gallery". Gallery of the world's bulbs. International Bulb Society. 2012. Retrieved 4 November 2014. 
• "National Daffodil Society of New Zealand". 2014. Retrieved 23 October 2014. 
• "Narcissus". Plant Encyclopaedia. Alpine Garden Society. 2011. Retrieved 29 October 2014. 

Cultivation

• "Flower Bulb Research Program". Department of Horticulture, College of Agriculture and Life Sciences, Cornell University. Retrieved 3 November 2014. 
  • "Research Newsletter". Retrieved 3 November 2014. 
• "International Flower Bulb Centre". Retrieved 3 November 2014. 
  • "Bulb flower production: Narcissus". Retrieved 3 November 2014. 
  • "Narcissus". Narcissus, Hyacinth & Special Flower Bulbs Picture Book. Retrieved 3 November 2014. 

Other

• Gold, Sara; Eshel, Amram; Plotnizki, Abraham (2014). "Wild Flowers of Israel". Retrieved 7 October 2014. 
• "Daffodil Festivals and Fields". Retrieved 3 November 2014.