Narcissus (plant)

''Narcissus'' is a genus of predominantly spring flowering perennial plants of the amaryllis family, Amaryllidaceae. Various common names including daffodil,The word "daffodil" is also applied to related genera such as '' Sternbergia'', ''Ismene'' and ''Fritillaria meleagris''. It has been suggested that the word "Daffodil" be restricted to the wild species of the British Isles, '' N. pseudonarcissus''. narcissus, and jonquil are used to describe all or some members of the genus. ''Narcissus'' has conspicuous flowers with six petal-like tepals surmounted by a cup- or trumpet-shaped corona. The flowers are generally white and yellow (also orange or pink in garden varieties), with either uniform or contrasting coloured tepals and corona.

''Narcissus'' were well known in ancient civilisation, both medicinally and botanically, but formally described by Linnaeus in his ''Species Plantarum'' (1753). The genus is generally considered to have about ten sections with approximately 50 species. The number of species has varied, depending on how they are classified, due to similarity between species and hybridisation. The genus arose some time in the Late Oligocene to Early Miocene epochs, in the Iberian peninsula and adjacent areas of southwest 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 " asphodel", with which it was commonly compared.

The species are native to meadows and woods in southern Europe and North Africa with a centre of diversity in the Western Mediterranean, particularly the Iberian peninsula. Both wild and cultivated plants have naturalised widely, and were introduced into the Far East prior to the tenth century. Narcissi tend to be long-lived bulbs, which propagate by division, but are also insect-pollinated. Known pests, diseases and disorders include viruses, fungi, the larvae of flies, mites and nematodes. Some ''Narcissus'' species have become extinct, while others are threatened by increasing urbanisation and tourism.

Historical accounts suggest narcissi have been cultivated from the earliest times, but became increasingly popular in Europe after the 16th century and by the late 19th century were an important commercial crop centred primarily in the Netherlands. Today narcissi are popular as cut flowers and as ornamental plants in private and public gardens. The long history of breeding has resulted in thousands of different cultivars. For horticultural purposes, narcissi are classified into divisions, covering a wide range of shapes and colours. Like other members of their family, narcissi produce a number of different alkaloids, which provide some protection for the plant, but may be poisonous if accidentally ingested. This property has been exploited for medicinal use in traditional healing and has resulted in the production of galantamine for the treatment of Alzheimer's dementia. Long celebrated in art and literature, narcissi are associated with a number of themes in different cultures, ranging from death to good fortune, and as symbols of spring.

The daffodil is the national flower of Wales and the symbol of cancer charities in many countries. The appearance of the wild flowers in spring is associated with festivals in many places.

Description

General

''Narcissus'' is a genus of perennial herbaceous bulbiferous geophytes, which die back after flowering to an underground storage bulb. They regrow in the following year from brown-skinned ovoid bulbs with pronounced necks, and reach heights of depending on the species. Dwarf species such as '' N. asturiensis'' have a maximum height of , while ''Narcissus tazetta'' may grow as tall as .

The plants are scapose, having a single central leafless hollow flower stem (scape). Several green or blue-green, narrow, strap-shaped leaves arise from the bulb. The plant stem usually bears a solitary flower, but occasionally a cluster of flowers (umbel). The flowers, which are usually conspicuous and white or yellow, sometimes both or rarely green, consist of a perianth of three parts. Closest to the stem (proximal) is a floral tube above the ovary, then an outer ring composed of six tepals (undifferentiated sepals and petals), and a central disc to conical shaped corona. The flowers may hang down (pendant), or be erect. There are six pollen bearing stamens surrounding a central style. The ovary is inferior (below the floral parts) consisting of three chambers (trilocular). The fruit consists of a dry capsule that splits ( dehisces) releasing numerous black seeds.

The bulb lies dormant after the leaves and flower stem dieback and has contractile roots that pull it down further into the soil. The flower stem and leaves form in the bulb, to emerge the following season. Most species are dormant from summer to late winter, flowering in the spring, though a few species are autumn flowering.

Specific

Vegetative

; Bulbs :
The pale brown-skinned ovoid tunicate bulbs have a membranous tunic and a corky stem (base or basal) plate from which arise the adventitious root hairs in a ring around the edge, which grow up to 40 mm in length. Above the stem plate is the storage organ consisting of bulb scales, surrounding the previous flower stalk and the terminal bud. The scales are of two types, true storage organs and the bases of the foliage leaves. These have a thicker tip and a scar from where the leaf lamina became detached. The innermost leaf scale is semicircular only partly enveloping the flower stalk (semisheathed).(see Hanks Fig 1.3). The bulb may contain a number of branched bulb units, each with two to three true scales and two to three leaf bases. Each bulb unit has a life of about four years.

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 which become brown and dry, forming an outer shell, the tunic or skin. Up to 60 layers have been counted in some wild species. While the plant appears dormant above the ground 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.

; Stems :
The single leafless stem or scape, appearing from early to late spring depending on the species, bears 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. In a few species such as '' N. hedraeanthus'' the stem is oblique (asymmetrical). The stem is hollow in the upper portion but towards the bulb is more solid and filled with a spongy material.

; Leaves :
''Narcissus'' plants have one to several basal leaves which are linear, ligulate or strap-shaped (long and narrow), sometimes channelled adaxially to semiterete, and may (pedicellate) or may not (sessile) have a petiole stalk. The leaves are flat and broad to cylindrical at the base and arise from the bulb. 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 (fruit) is ripe.

Jonquils usually have dark green, round, rush-like leaves.

Reproductive

; Inflorescence :
The inflorescence is scapose, the single stem or scape bearing either a solitary flower or forming an umbel with up to 20 blooms. Species bearing a solitary flower include section ''Bulbocodium'' and most of section ''Pseudonarcissus''. Umbellate species have a fleshy racemose inflorescence (unbranched, with short floral stalks) with 2 to 15 or 20 flowers, such as '' N. papyraceus'' (see illustration, left) and '' N. tazetta'' (see Table I). The flower arrangement on the inflorescence may be either with (pedicellate) or without (sessile) floral stalks.

Prior to opening, the flower buds are enveloped and protected in a thin dry papery or membranous (scarious) spathe. The spathe consists of a singular bract that is ribbed, and which remains wrapped around the base of the open flower. As the bud grows, the spathe splits longitudinally. Bracteoles are small or absent.

; Flowers :
The flowers of ''Narcissus'' are hermaphroditic (bisexual), have three parts (tripartite), and are sometimes fragrant (see Fragrances). The flower symmetry is actinomorphic (radial) to slightly zygomorphic (bilateral) due to declinate-ascending stamens (curving downwards, then bent up at the tip). ''Narcissus'' flowers are characterised by their, usually conspicuous, corona (trumpet).

The three major floral parts (in all species except '' N. cavanillesii'' in which the corona is virtually absent - Table I: Section ''Tapeinanthus'') are;
* (i) the proximal floral tube (hypanthium),
* (ii) the surrounding free tepals, and
* (iii) the more distal corona (paraperigon, paraperigonium).

All three parts may be considered to be components of the perianth (perigon, perigonium). The perianth arises above the apex of the inferior ovary, its base forming the hypanthial floral tube.

The floral tube is formed by fusion of the basal segments of the tepals (proximally connate). Its shape is from an inverted cone (obconic) to funnel-shaped (funneliform) or cylindrical, and is surmounted by the more distal corona. Floral tubes can range from long and narrow in sections ''Apodanthi'' and ''Jonquilla'' to rudimentary (''N. cavanillesii'').

Surrounding the floral tube and corona and reflexed (bent back) from the rest of the perianth are the six spreading tepals or floral leaves, in two whorls which may be distally ascending, reflexed (folded back), or lanceolate. Like many monocotyledons, the perianth is homochlamydeous, which is undifferentiated into separate calyx (sepals) and corolla (petals), but rather has six tepals. The three outer tepal segments may be considered sepals, and the three inner segments petals. The transition point between the floral tube and the corona is marked by the insertion of the free tepals on the fused perianth.

The corona, or paracorolla, is variously described as bell-shaped (funneliform, trumpet), bowl-shaped (cupular, crateriform, cup-shaped) or disc-shaped with margins that are often frilled, and is free from the stamens. Rarely the corona is a simple callose (hardened, thickened) ring. The corona is formed during floral development as a tubular outgrowth from stamens which fuse into a tubular structure, the anthers becoming reduced. At its base, the fragrances which attract pollinators are formed. All species produce nectar at the top of the ovary. Coronal morphology varies from the tiny pigmented disk of '' N. serotinus'' (see Table I) or the rudimentary structure in '' N. cavanillesii'' to the elongated trumpets of section ''Pseudonarcissus'' (trumpet daffodils, Table I).

While the perianth may point forwards, in some species such as '' N. cyclamineus'' it is folded back (reflexed, see illustration, left), while in some other species such as '' N. bulbocodium'' ( Table I), it is reduced to a few barely visible pointed segments with a prominent corona.

The colour of the perianth is white, yellow or bicoloured, with the exception of the night flowering '' N. viridiflorus'' which is green. In addition the corona of '' N. poeticus'' has a red crenulate margin (see Table I). Flower diameter varies from 12 mm ('' N. bulbocodium'') to over 125 mm (''N. nobilis''='' N. pseudonarcissus'' subsp. ''nobilis'').

Flower orientation varies from pendent or deflexed (hanging down) as in '' N. triandrus'' (see illustration, left), through declinate-ascendant as in ''N. alpestris'' = ''N. pseudonarcissus'' subsp. ''moschatus'', horizontal (patent, spreading) such as '' N. gaditanus'' or '' N. poeticus'', erect as in ''N. cavanillesii'', ''N. serotinus'' and '' N. rupicola'' ( Table I), or intermediate between these positions (erecto-patent).

The flowers of ''Narcissus'' demonstrate exceptional floral diversity and sexual polymorphism, primarily by corona size and floral tube length, associated with pollinator groups (see for instance Figs. 1 and 2 in Graham and Barrett). Barrett and Harder (2005) describe three separate floral patterns;
* "Daffodil" form
* "Paperwhite" form
* "Triandrus" form.

The predominant patterns are the 'daffodil' and 'paperwhite' forms, while the "triandrus" form is less common. Each corresponds to a different group of pollinators (See Pollination).

The "daffodil" form, which includes sections ''Pseudonarcissus'' and ''Bulbocodium'', has a relatively short, broad or highly funnelform tube (funnel-like), which grades into an elongated corona, which is large and funnelform, forming a broad, cylindrical or trumpet-shaped perianth. Section ''Pseudonarcissus'' consists of relatively large flowers 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 pale yellow.

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

The "triandrus" form is seen in only two species, '' N. albimarginatus'' (a Moroccan endemic) and '' N. triandrus''. It 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. The flowers are pendent.

; Androecium :
There are six stamens in one to two rows (whorls), with the filaments separate from the corona, attached at the throat or base of the tube (epipetalous), often of two separate lengths, straight or declinate-ascending (curving downwards, then upwards). The anthers are basifixed (attached at their base).

; Gynoecium :
The ovary is inferior (below the floral parts) and trilocular (three chambered) and there is a pistil with a minutely three lobed stigma and filiform (thread like) style, which is often exserted (extending beyond the tube).

; Fruit :
The fruit consists of dehiscent loculicidal capsules (splitting between the locules) that are ellipsoid to subglobose (almost spherical) in shape and are papery to leathery in texture.

; Seeds :
The fruit contains numerous subglobose seeds which are round and swollen with a hard coat, sometimes with an attached elaiosome. The testa is black and the pericarp 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 sections ''Jonquilla'' and ''Bulbocodium'' are wedge-shaped and matte black, while those of other sections are ovate and glossy black. A gust of wind or contact with a passing animal is sufficient to disperse the mature seeds.

Chromosomes

Chromosome numbers include 2n=14, 22, 26, with numerous aneuploid and polyploid derivatives. The basic chromosome number is 7, with the exception of '' N. tazetta'', '' N. elegans'' and ''N. broussonetii'' in which it is 10 or 11; this subgenus (''Hermione'') was in fact characterised by this characteristic. Polyploid species include '' N. papyraceus'' (4x=22) and '' N. dubius'' (6x=50).

Phytochemistry

Alkaloids

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 galantamine alkaloids are also represented. The alkaloid profile of any plant varies with time, location, and developmental stage. ''Narcissus'' also contain fructans and low molecular weight glucomannan in the leaves and plant stems.

Fragrances

Fragrances are predominantly monoterpene isoprenoids, with a small amount of benzenoids, although '' N. jonquilla'' has both equally represented. Another exception is '' N. cuatrecasasii'' which produces mainly fatty acid derivatives. The basic monoterpene precursor is geranyl pyrophosphate, and the commonest monoterpenes are limonene, myrcene, and ''trans''-β-ocimene. Most benzenoids are non-methoxylated, while a few species contain methoxylated forms (ethers), ''e.g.'' ''N. bugei''. Other ingredient include indole, isopentenoids and very small amounts of sesquiterpenes. Fragrance patterns can be correlated with pollinators, and fall into three main groups (see Pollination).

Taxonomy

History

Early

The genus ''Narcissus'' was well known to the ancients. In Greek literature Theophrastus and Dioscorides described ''νάρκισσος'', probably referring to '' N. poeticus'', although the exact species mentioned in classical literature cannot be accurately established. Pliny the Elder later introduced the Latin form ''narcissus''. These early writers were as much interested in the plant's possible medicinal properties as they were its botanical features and their accounts remained influential until at least the Renaissance (see also Antiquity). Mediaeval and Renaissance writers include Albert Magnus and William Turner, but it remained to Linnaeus to formally describe and name ''Narcissus'' as a genus in his Species Plantarum (1753) at which time there were six known species.

Modern

De Jussieu (1789) grouped ''Narcissus'' into a "family", which he called Narcissi. This was renamed Amaryllideae by Jaume Saint-Hilaire in 1805, corresponding to the modern Amaryllidaceae. For a while ''Narcissus'' was considered part of Liliaceae (as in the illustration seen here of ''Narcissus candidissimus''), but then the Amaryllidaceae were split off from it.

Various authors have adopted either narrow (e.g. Haworth, Salisbury) or wide (e.g.Herbert, Spach ) interpretations of the genus. The narrow view treated many of the species as separate genera. Over time the wider view prevailed with a major monograph on the genus being published by Baker (1875). One of the more controversial genera was ''Tapeinanthus'', but today it is included in ''Narcissus''.

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

Subdivision

The infrageneric phylogeny of ''Narcissus'' still remains relatively unsettled, the taxonomy having proved complex and difficult to resolve, due to the diversity of the wild species, the ease with which natural hybridization occurs, and extensive cultivation and breeding accompanied by escape and naturalisation. Consequently, the number of accepted species has varied widely.

De Candolle, in the first systematic taxonomy of ''Narcissus'', arranged the species into named groups, and those names have largely endured for the various subdivisions since and bear his name as their authority. The situation was confused by the inclusion of many unknown or garden varieties, and it was not till the work of Baker that the wild species were all grouped as sections under one genus, ''Narcissus''.

A common classification system has been that of Fernandes based on cytology, as modified by Blanchard (1990) and Mathew (2002). Another is that of Meyer (1966). Fernandes proposed two subgenera based on basal chromosome number, and then subdivided these into ten sections as did Blanchard.

Other authors (e.g. Webb) prioritised morphology over genetics, abandoning subgenera, although Blanchard's system has been one of the most influential. While infrageneric groupings within ''Narcissus'' have been relatively constant, their status (genera, subgenera, sections, subsections, series, species) has not. The most cited system is that of the Royal Horticultural Society (RHS) which simply lists ten sections. Three of these are monotypic (contain only one species), while two others contain only two species. Most species are placed in section ''Pseudonarcissus''. Many of these subdivisions correspond roughly to the popular names for daffodil types, ''e.g.'' Trumpet Daffodils, Tazettas, Pheasant's Eyes, Hoop Petticoats, Jonquils.

The most hierarchical system is that of Mathew, illustrated here -

Phylogenetics

The phylogenetic analysis of Graham and Barrett (2004) supported the infrageneric division of ''Narcissus'' into two clades corresponding to Fernandes' subgenera, but did not support monophyly of all sections. A later extended analysis by Rønsted ''et al.'' (2008) with additional taxa confirmed this pattern.

A large molecular analysis by Zonneveld (2008) sought to reduce some of the paraphyly identified by Graham and Barrett. This led to a revision of the sectional structure.
While Graham and Barrett (2004) had determined that subgenus ''Hermione'' was monophyletic, Santos-Gally ''et al.'' (2011) did not. If two species excluded in the former study are removed from the analysis, the studies are in agreement, the species in question instead forming a clade with subgenus ''Narcissus''. Some so-called nothosections have been proposed, to accommodate natural ('ancient') hybrids (nothospecies).

Species

Estimates of the number of species in ''Narcissus'' have varied widely, from anywhere between 16 and almost 160, even in the modern era. Linnaeus originally included six species in 1753, by 1784 there were fourteen by 1819 sixteen, and by 1831 Adrian Haworth had described 150 species.

Much of the variation lies in the definition of species. 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, with a distinction between "ancient hybrids" and "recent hybrids". The term "ancient hybrid" refers to hybrids found growing over a large area, and therefore now considered as separate species, while "recent hybrid" refers to solitary plants found amongst their parents, with a more restricted range.

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

Evolution

Within the Narcisseae, ''Narcissus'' (western Mediterranean) diverged from '' Sternbergia'' (Eurasia) some time in the Late Oligocene to Early Miocene eras, around 29.3–18.1 Ma. Later the genus divided into the two subgenera (''Hermione'' and ''Narcissus'') between 27.4 and 16.1 Ma. The divisions between the sections of ''Hermione'' then took place during the Miocene period 19.9–7.8 Ma. ''Narcissus'' appears to have arisen in the area of the Iberian peninsula, southern France and northwestern Italy. Subgenus ''Hermione'' in turn arose in the southwestern mediterranean and north west Africa.

Names and etymology

Narcissus

The derivation of the Latin ' is from Greek ''narkissos''.. According to Plutarch ''narkissos'' has been connected because of the plant's narcotic properties, with ''narkē'' "numbness"; it may also be connected with hell. On the other hand, its etymology is considered to be clearly Pre-Greek by Beekes.

It is frequently linked to the myth of Narcissus, who became so obsessed with his own reflection in water that he drowned and the narcissus plant sprang from where he died. There is no evidence for the flower being named after Narcissus. ''Narcissus poeticus'', which grows in Greece, has a fragrance that has been described as intoxicating. Pliny wrote that the plant was named for its fragrance ( ''narkao'', "I grow numb" ), rather than Narcissus. Furthermore, there were accounts of narcissi growing long before the story of Narcissus appeared (see Greek culture).Prior here refers to the poet ''Pamphilus'', but it is likely he meant ''Pamphos''. It has also been suggested that narcissi bending over streams represent the youth admiring his reflection. Linnaeus used the Latin name "narcissus" for the plant but was preceded by others such as Matthias de l'Obel (1591) and Clusius (1576). The name Narcissus was not uncommon for men in Roman times.

The plural form of the common name "narcissus" has been the cause of some confusion. Dictionaries list "narcissi", "narcissuses" and "narcissus". However, texts on usage such as Garner and Fowler state that "narcissi" is the preferred form. The common name narcissus should not be capitalised.

Daffodil

The name "daffodil" is derived from "affodell", a variant of '' asphodel''. The narcissus was frequently referred to as the asphodel (see Antiquity). Asphodel in turn appears to come from the Greek "asphodelos" ( grc-gre, ἀσφόδελος). The reason for the introduction of the initial "d" is not known. From at least the 16th century, "daffadown dilly" and "daffydowndilly" have appeared as alternative names. Other names include "Lent lily".

Distribution and habitat

Distribution

Although the family Amaryllidaceae are predominantly tropical or subtropical as a whole, ''Narcissus'' occurs primarily in Mediterranean region, with a centre of diversity in the Iberian Peninsula (Spain and Portugal). A few species extend the range into southern France, Italy, 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 as well as East Asia are considered introductions, albeit ancient (see Eastern cultures). While the exact northern limit of the natural range is unknown, the occurrences of wild ''N. pseudonarcissus'' in Great Britain, middle and northern Europe are similarly considered ancient introductions.

While the Amaryllidaceae are not native to North America, ''Narcissus'' grows well in USDA hardiness zones 3B through 10, which encompass most of the United States and Canada.

'' 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.

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 Mondego river are found intersectional hybrids 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 southeastern 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 on 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 has relatively little limestone, ''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

Life cycle

''Narcissus'' are long-lived perennial geophytes with winter-growing and summer-dormant bulbs that are mainly synanthous (leaves and flowers appearing at the same time). While most species flower in late winter to spring, five species are autumn flowering (''N. broussonetii'', '' N. cavanillesii'', '' N. elegans'', '' N. serotinus'', '' N. viridiflorus''). By contrast these species are hysteranthous (leaves appear after flowering).

Flower longevity varies by species and conditions, ranging from 5–20 days. After flowering leaf and root senescence sets in, and the plant appears to be 'dormant' till the next spring, conserving moisture. However, the dormant period is also one of considerable activity within the bulb primordia. It is also a period during which the plant bulb may be susceptible to predators . Like many bulb plants from temperate regions, a period of exposure to cold is necessary before spring growth can begin. This protects the plant from growth during winter when intense cold may damage it. Warmer spring temperatures then initiate growth from the bulb. Early spring growth confers a number of advantages, including relative lack of competition for pollinators, and lack of deciduous shading. The exception to requiring cold temperatures to initiate flowering is '' N. tazetta''.

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 highly scented night-flowering '' N. viridiflorus'' is pollinated by crepuscular moths. Pollination mechanism fall into three groups corresponding to floral morphology (see Description - Flowers).

# 'Daffodil' form. 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.
# 'Paperwhite' form. 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.
# 'Triandrus' form. 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 progress, 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

Diseases of ''Narcissus'' are of concern because of the economic consequences of losses in commercial cultivation. Pests include viruses, bacteria, and fungi as well as arthropods and gastropods. For control of pests, see Commercial uses.

; Viruses :
Aphids such as ''Macrosiphum euphorbiae'' can transmit viral diseases which affect the colour and shape of the leaves, as can nematodes. Up to twenty-five viruses have been described as being able to infect narcissi. These include the '' Narcissus common latent virus'' (NCLV, ''Narcissus mottling-associated virus''),This ''Carlavirus'' should not be confused with the similarly named'' Narcissus latent virus'' which is a ''Macluravirus''. '' Narcissus latent virus'' (NLV, ''Narcissus mild mottle virus'') which causes green mottling near leaf tips, '' Narcissus degeneration virus'' (NDV), '' Narcissus late season yellows virus'' (NLSYV) which occurs after flowering, streaking the leaves and stems, '' Narcissus mosaic virus'', '' Narcissus yellow stripe virus'' (NYSV, ''Narcissus yellow streak virus''), ''Narcissus tip necrosis virus'' (NTNV) which produces necrosis of leaf tips after flowering and '' Narcissus white streak virus'' (NWSV).

Less host specific viruses include '' Raspberry ringspot virus'', '' Nerine latent virus'' (NeLV) ='' Narcissus symptomless virus'', '' Arabis mosaic virus'' (ArMV), '' Broad Bean Wilt Viruses'' (BBWV) ''Cucumber mosaic virus'' (CMV), '' Tomato black ring virus'' (TBRV), '' Tomato ringspot virus'' (TomRSV) and ''Tobacco rattle virus leaf striping in '' N. tazetta''. Infection with NYSV produces light or grayish-green, or yellow stripes or mottles on the upper two-thirds of the leaf, which may be roughened or twisted. The flowers which may be smaller than usual may also be streaked or blotched. NWSV produces greenish-purple streaking on the leaves and stem turning white to yellow, and premature senescence reducing bulb size and yield. These viruses are primarily diseases of commercial nurseries. The growth inhibition caused by viral infection can cause substantial economic damage.

; Bacteria :
Bacterial disease is uncommon in ''Narcissus'' but includes ''Pseudomonas'' (bacterial streak) and ''Pectobacterium carotovorum'' sp. ''carotovorum'' (bacterial soft rot).

; Fungi :
More problematic for non-commercial plants is the fungus, ''Fusarium oxysporum'' f. sp. ''narcissi'', which causes basal rot (rotting of the bulbs and yellowing of the leaves). This is the most serious disease of ''Narcissus''. 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, causing narcissus smoulder, is ''Botrytis narcissicola'' (''Sclerotinia narcissicola'') and other species of '' Botrytis'', including ''Botrytis cinerea'', particularly if improperly stored. Copper sulfate is used to combat the disease, and infected bulbs are burned. Blue mould rot of bulbs may be caused by infection with species of ''Penicillium'', if they have become damaged either through mechanical injury or infestation by mites (see below). Species of Rhizopus (''e.g.'' ''Rhizopus stolonifer'', ''Rhizopus nigricans'') cause bulb soft rot and ''Sclerotinia bulborum'', black slime disease. A combination of both ''Peyronellaea curtisii'' (''Stagonosporopsis curtisii'') and ''Botrytis narcissicola'' causes neck rot in the bulbs.

Fungi affecting the roots include '' Nectria radicicola'' (''Cylindrocarpon destructans''), a cause of root rot and ''Rosellinia necatrix'' causing white root rot, while others affect root and bulb, such as ''Aspergillus niger'' (black mold), and species of ''Trichoderma'', including '' T. viride'' and '' T. harzianum'' (=''T. narcissi'') responsible for green mold.

Other fungi affect the remainder of the plant. Another ''Botrytis'' fungus, ''Botrytis polyblastis'' (''Sclerotinia polyblastis'') causes brown spots on the flower buds and stems (narcissus fire), especially in damp weather and is a threat to the cut flower industry. '' Ramularia vallisumbrosae'' is a leaf spot fungus found in warmer climates, causing narcissus white mould disease. ''Peyronellaea curtisii'', the Narcissus leaf scorch, also affects the leaves as does its synanamorph, ''Phoma narcissi'' (leaf tip blight). '' Aecidium narcissi'' causes rust lesions on leaves and stems.

; Animals :
Arthropods that are ''Narcissus'' pests include insects such as three species of fly that have larvae that attack the plants, 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. The larvae of some moths such as ''Korscheltellus lupulina'' (the common swift moth) attack ''Narcissus'' bulbs.

Other arthropods include Mites such as ''Steneotarsonemus laticeps'' (Bulb scale mite), ''Rhizoglyphus'' and '' Histiostoma'' infest mainly stored bulbs and multiply particularly at high ambient temperature, but do not attack planted bulbs.

Planted bulbs are susceptible to nematodes, the most serious of which is ''Ditylenchus dipsaci'' (Narcissus eelworm), the main cause of basal plate disease in which the leaves turn yellow and become misshapen. Infested bulbs have to be destroyed; where infestation is heavy avoiding planting further narcissi for another five years. Other nematodes include ''Aphelenchoides subtenuis'', which penetrates the roots causing basal plate disease and ''Pratylenchus penetrans'' (lesion nematode) the main cause of root rot in narcissi.
Other nematodes such as the longodorids ('' Longidorus'' spp. or needle nematodes and ''Xiphinema'' spp. or dagger nematodes) and the stubby-root nematodes or trichodorids