Chapter 87 - The Daffodils Speak (Part 3)

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 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.

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 27.4–16.1 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 north-western Italy. Subgenus Hermione, in turn, arose in the southwestern Mediterranean and northwest Africa.

The derivation of the Latin narcissus is unknown but is frequently linked to the myth of Narcissus, who became so obsessed with his own reflection in the water that he drowned and the narcissus plant sprang from where he died. There is no evidence for the flower being named for the youth. 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 (ναρκάω narkao, "I grow numb" ), not the 123456 name Narcissus was not uncommon for men in Roman times.

Daffodil

The name "daffodil" is derived from "affodell", a variant of asphodel. The narcissus was frequently referred to as the asphodel. Asphodel in turn appears to come from the Greek "asphodelos" (Greek: 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

Although the family Amaryllidaceae are predominantly tropical or subtropical as a whole, Narcissus occurs primarily in the Mediterranean region, with a center 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, China and Japan and N. pseudonarcissus in Great Britain, are considered introductions,

Habitats

Their native habitats are varied, with different elevations, bioclimatic areas. Found in open areas from low marshes to rocky hillsides, grassland, woods, riverbanks, and rocky crevices. They preference 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.

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 survive from 5–20 days. After flowering the plant appears to be 'dormant' till the next spring, conserving moisture. However, the dormant period is also one of considerable activity. 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.

Plants may spread clonally through the production of daughter bulbs and division producing clumps. Narcissus species hybridize 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 falls into three groups corresponding to floral morphology.

In 'Daffodil' form the pollination is 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 causes cross-pollination.

In the 'Paperwhite' form pollination is by 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.

In the 'Triandrus' form pollination is 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).