Chapter 86 - The Daffodils Speak (Part 2)

"The flowers of Narcissus are hermaphroditic (bisexual), have three parts (tripartite), and are sometimes fragrant. 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 characterized by their, usually conspicuous, corona (trumpet)."

"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, that is undifferentiated into a 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."

"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."

"The flowers of Narcissus demonstrate exceptional floral diversity and sexual polymorphism, primarily by corona size and floral tube length, associated with pollinator groups. There are 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.

"The "daffodil" form, 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, 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 color from the corona and maybe cream-colored 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 like a pendant."

"The Androecium comprises 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)."

"The Gynoecium comprises the ovary which 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)."

"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."

"The fruit contains numerous subglobose seeds which are round and swollen with a hard coat, sometimes with an attached elaiosome. The testa is black[9] and the pericarp dry."

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 utilized 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 number 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 ingredients include indole, isopentenoids and very small amounts of sesquiterpenes. Fragrance patterns can be correlated with pollinators, and fall into three main groups.

History

The genus Narcissus was well known to the ancients. In Greek literature, Theophrastus and Dioscorides described 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. 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, but then the Amaryllidaceae was split off from it.

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 naturalization. Consequently, the number of accepted species has varied widely.

It was not till the work of Baker that the wild species were all grouped as sections under one genus, Narcissus. There were many others like Webb, Graham and Barrett (2004) who tried to classify them.