Amaryllidaceae

Amaryllidaceae
Amaryllis belladonna
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Monocots
Order:	Asparagales
Family:	Amaryllidaceae J.St.-Hil.
Genera
see list of genera

The Amaryllis family or Amaryllidaceae are a family of herbaceous, perennials and bulbous flowering plants included in the monocot order Asparagales, taking its name from the genus Amaryllis. The family consists of about sixty genera, with over eight hundred species with a worldwide distribution.

All members of this family, which may be described as 'amaryllids', can be recognised by their rather fleshy and two-ranked leaves and their large and showy flowers, with an inferior ovary and six stamens, arranged in umbels at the apex of leafless flowering stems, or scapes. The somewhat similar-looking members of the onion family (Alliaceae) differ in having a superior ovary and the presence of allyl sulfide compounds which give them their characteristic smell. From another close relative, the agapanthus family (Agapanthaceae), differs also in the position of the ovary and in lacking steroidal saponins. ^[1]

Description

Pink Lily

Members of Amaryllidaceae are perennial, mostly deciduous, rarely shrubby or treelike plants, often with bulbs or, rarely, with rhizomes, as in the genera Clivia, Cryptostephanus and Scadoxus. While growing, the bulb is kept deep below ground by a special type of root that lengthen and contract. Leaves are arranged in a basal rosette or fan. They are often narrow, with an entire or spiny margin, and without marked olours, in particular they are not onion-scented. Some genera, like Eucrosia and Scadoxus, which occupy habitats with low light-intensity, have leaves that are broad and flattened, whereas in semi-arid regions like southern Africa, species of Brunsvigia, Crossyne, Gethyllis and Haemanthus have leaves covered with variously shaped hairs. The leaves in Crossyne and some Haemanthus species are also attractively spotted with dark green or red.^[2][3][4]

The flowers are large and showy, bisexual, trimerous, actinomorphic (less often slightly zygomorphic, as in Sprekelia). Shape of the flowers varies from star-like to trumpet-shaped or tubular and Colours range from red, orange, yellow and pink to white, whereas bluish flowers are only found in Griffinia, Worsleya and Lycoris. The perianth present 6 segments arranged in two whorls, free or fused to form a short tube. The flowers of Narcissus (the popular daffodil) characteristically have a large, cup-shaped corona, which is an outgrowth of the tepals. The androecium is composed by six stamens, inserted at the perianth throat or at the base of each tepal. Some Griffinia species have five stamens and some Gethyllis species have multiple stamens with about 60 anthers. In Pancratium and Hymenocallis the stamens are fused to form a large cup, known as "staminal cup" which resembles the corona in Narcissus. The ovary is inferior (located below the tepals), with three locules and few to many ovules per locule. All the members of the family produce nectar and are often heavily scented. The inflorescence can be of one or many flowers in a terminal spike, raceme, panicle or, more often, in an umbel-like inflorescence, subtended by an involucre of one to many bracts and with ephemeral hyaline bracts between the flowers. The inflorescence appear at the end of a leafless stem, called a scape. In unusual genera like Gethyllis, however, the scape carries only one flower and remains subterranean. ^[2][3][4]

The fruit is a dry capsule, or more rarely, a fleshy berry and contain dry, dark and often flattened, or fleshy, round, and greenish seeds. The seeds have an oily endosperm and usually with a black or brown phytomelanous testa, sometimes with a caruncular elaiosome at the chalazal end. ^[2][3][4]

The most common basic chromosome number in the family is x = 11, although during its evolution a vast arrange of different basic chromosome numbers arose, from x= 6 to x= 23. ^[3]

Members of this family present a unique type of alkaloids, the norbelladine alkaloids, which are tyrosine derivatives (combined with 4-methylcatechol). 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.^[5]

Distribution and habitat

Holarctic, Paleotropical, Neotropical, Cape, and Australian. World distribution: widespread In the Neotropics, the family occurs from Mexico through Central America and the West Indies to Chile and Argentina in South America. Notable areas of diversity throughout this range include eastern Brazil, north-central Chile (outside of the tropical zone, however), and the central Andes of Ecuador and Peru. Hippeastrum is primarily found in the Andes and eastern Brazil, Hymenocallis occurs mostly in Mesoamerica, Clinanthus is largely endemic to Peru, and Zephyranthes is broadly distributed. The greatest generic diversity is found in Peru.

Taxonomy

Acis autumnalis

Amaryllis belladonna,.

A paraperigonium or corona is characteristic of the genus Narcissus, as can be seen in the image, the tepals in white and the corona in yellow colour.

Clivia

Hymenocallis

Lycoris radiata

Sternbergia lutea.

Sprekelia formosissima, an example of zygomorphic flowers in the family.

Scadoxus.

Zephyranthes candida .

Haemanthus albiflos.

Hippeastrum.

Habranthus tubispathus

Several members of Amaryllidaceae show a "staminal cup" formed by a membrane that fuse the six stames. In the image, it can be observed the staminal cup in the flower of the "Alligator Lilly" (Hymenocallis palmeri), note that the staminal cup in this species is greater than the tepals.

Pancratium parvum

Crinum species

|} First described by the French naturalist Jean Henri Jaume Saint-Hilaire in 1805, the family was named after Amaryllis, a beautiful shepherdess mentioned by Theocritus, Virgil and Ovid, and hence an allegory of beauty. Close relatives to the Amaryllis family are the onion family (Alliaceae) and the agapanthus family (Agapanthaceae).

Amaryllidaceae has been recognized, with varying definitions, by most classification systems of the 20th Century, although the Cronquist system included it within a very broadly defined Liliaceae. The two families have traditionally been separated by including species with inferior ovaries in Amaryllidaceae and those with superior ovaries in Liliaceae. The APG II system (2003) advocates recognizing all Agapanthaceae, Alliaceae, and Amaryllidaceae as three subtaxa of a single family in the order Asparagales, citing Alliaceae Batsch (1786) as the name of earliest priority. They do, however, leave open the option for recognizing three families. The APG system, of 1998, accepted Amaryllidaceae as a separate family. Despite these proposals to combine the three families in Alliaceae sensu lato, their differences regarding chemical compounds and morphology are nevertheless sufficiently important to keep them apart. ^[3] In fact, Amaryllidaceae are one of the few families of the higher Asparagales well defined by other than molecular characters, namely the combination of umbellate cymes, inferior ovaries, and unique alkaloid chemistry.^[6]

Evolution

Amaryllidaceae and its sister group, Agapanthaceae, originated in western Gondwanaland. ^[7] Africa has also been the site of considerable innovation in the family's history as well, as typified by the Afrocentric tribes Amaryllideae, Haemantheae, and Cyrtantheae. Most of the diversity within those three tribes is, however, centered in South Africa, and thus may reflect radiation engendered by the more recent paleoclimatic and geological history of Africa encompassing Neogene and later times.^[7][8] The increased aridity of the African climate and the uplift of the continental mass beginning near the end of the Oligocene, further abetted by Quaternary climatic fluctuations, were catastrophic to many elements of the African flora, but it may have been a selective pressure for diversity among groups of geophytes capable of adapting to increasing drought. The geophyte richness of South Africa is well documented,^[9] and the Cape region has been suggested as a possible refuge for certain African plant and animal groups as the tropical flora of the continent was impoverished.^[7] On the other hand, the three genera of the baccate-fruited Haemantheae, Clivia, Cryptostephanus, and Scadoxus, are all forest understory adapted taxa, do not form bulbs, and are at least in part (Scadoxus, Cryptostephanus) elements of tropical vegetation farther north.^[10]

The Calostemmateae, the only exclusively Australasian element of the family, may have been isolated from the African lineages as Australia separated from western Gondwanaland.^[7] Direct migration between Africa and Australia may have persisted up through the close of the early Cretaceous, although India and Madagascar may have provided a less direct corridor up until the late Cretaceous.^[7] Crinum is the only amaryllid that is known to occur on Madagascar, whereas indigenous Indian amaryllids are restricted to Crinum and two to three species of Pancratium. The adaptations of Crinum for long-distance dispersal have been demonstrated and Pancratium may have been able to directly enter India from either Africa or Eurasia during the late Cretaceous or early Eocene.^[7][10]

To summarize, molecular data indicate an African origin for the family, with later dispersal to Australasia and ultimately Eurasia and the New World, the latter two geographic groups sharing an ostensibly African ancestor. The family evolved in Africa and subsequently spread to other continents, further suggesting that South America is the center of secondary diversification.^[11] The sister relationship of the Eurasian/Mediterranean clade to the American genera raises the interesting question of when and where the Amaryllidaceae, in the main, entered the New World. Given the extant distribution of Amaryllidaceae in North America and the generic richness south of the equator, a northern latitude entry into the New World for the family would necessitate massive extinction in North America sometime after migration to South America took place. Glaciation would be the likely factor involved. Little migration of plants from North America to South America probably took place before the Eocene.^[7] All indications are that the movement of extant Amaryllidaceae has been northward from South America. This does not necessarily preclude an earlier, initial arrival in North America, migration to South America, and a more recent, but secondary, return of some elements of the family to North America long after glaciation extirpated the founder populations.^[10]

Phylogeny

The Amaryllidaceae has so far defied precise understanding of its intrafamilial phylogeny, and, its relationships to other families of higher Asparagales. Combined analysis of three plastid DNA sequences (rbcL, trnL intron, and trnL-F spacer) for 50 genera of Amaryllidaceae analyzed together with members of Alliaceae, Behniaceae, Convallariaceae, Hyacinthaceae, Themidaceae, and Hemerocallidaceae, resolves Agapanthaceae as sister to Amaryllidaceae with weak support and places Agapanthus-Amaryllidaceae as a sister clade to a monophyletic Alliaceae.

The African tribe Amaryllideae is sister to the remainder of the Amaryllidaceae. Amaryllis and Boophone forms the most basal branches of the phylogeny inside this tribe. Two major lineages are subsequently resolved in all the analyses. The most diverse of them is the southern African lineage that encompasses Strumaria and its allies. The other is the predominantly sub-Saharan African group that includes Crinum and related genera. ^[12] The African baccate-fruited Haemantheae and the Australasian Calostemmateae are sister tribes, and the African endemic Cyrtantheae is sister to them both. ^[13] The most completely resolved and best supported tree for Haemantheae, divides the tribe into two main clades. The smaller clade, uniting Clivia and Cryptostephanus, represents entirely rhizomatous genera that never form bulbs. Cryptostephanus is also the only genus of the tribe that retains the plesiomorphic character of a phytomelanous testa. The second clade contains all of the genera that form true bulbs, though Scadoxus is polymorphic for this character. This second clade contains two subclades that can be characterized morphologically as well. The sister relationship of Haemanthus and Scadoxus is well supported by the morphological synapomorphy of the brush-like inflorescence, facilitated by the reduction in perianth size and the dominance of the spathe bracts during anthesis. The gethyllid subclade is characterized by a suite of morphological characters, such as uniflory, obsolete scape, and the long, aromatic, cylindrical, many-seeded fruit of both recognized genera, in contrast to the one or few seeded berry of the other genera in the tribe.^[14]

The Eurasian and neotropical genera of the Amaryllidaceae are a well-supported clade, and are sister groups.^[15] Lycorideae are basal in the Eurasian clade and begin a grade that continues with Hannonia, then Pancratium, then Lapiedra. The genera Galanthus, Narcissus, and Sternbergia are resolved as monophyletic with strong support. Leucojum sensu lato is paraphyletic and recognition of Acis for the mostly autumn-flowering Mediterranean species is supported by molecular data.^[16]

The American genera of the family form two major clades. The first, or ‘‘hippeastroid’’ clade, are diploid (x = 11), primarily the extra-Andean element of the family (though several of the genera do have Andean representatives), comprising the genera treated as the tribe Hippeastreae. The second clade constitutes the tribes centered in the Andes whose basic chromosome number is derived by polyploidy (x = 23). Several genera within the hippeastroid clade resolve as polyphyletic (such as Rhodophiala and Zephyranthes) and the possibility of reticulate evolution (i.e., early hybridization) in these lineages was hypothesized. A petiolate-leafed Andean subclade, containing elements of both Eucharideae and Stenomesseae, was resolved. Within the Andean subclade, Eustephieae resolves as sister to all other tribes; a distinct petiolate-leafed group is resolved, combining the tribe Eucharideae and the petiolate Stenomesseae; and a distinct Hymenocallideae is supported. It was inferred from the molecular data that a great deal of the diversity of the family in the Americas is recent, and that the American Amaryllidaceae may have been reduced to peripheral isolates some time after its initial entry and spread through the Americas.^[17] In both of the major American clades, there is a small tribe that is sister to the rest of the clade, Eustephieae in the Andean group, and Griffineae in the hippeastroid clade. These two small tribes may represent either ancestral or merely very isolated elements of their respective clades.

A cladogram which summarizes the phylogenetic relationships among tribes and subtribes of Amaryllidaceae is given below.

Tribes, subtribes and genera

The four modern infrafamilial classifications of Amaryllidaceae are those of Traub (1963),^[18] Dahlgren, Clifford, and Yeo (1985),^[19] Müller-Doblies and Müller-Doblies (1996),^[20] and Meerow and Snijman (1998).^[21] Traub's clasiification included Alliaceae, Hemerocallidaceae, and Ixioliriaceae as subfamilies, following partially the scheme proposed by Hutchinson (1934).^[22] Within his subfamily Amarylloideae, he erected two informal categories: "infrafamilies" Amarylloidinae and Pancratioidinae, both of which were polyphyletic.^[23] Dahlgren, Clifford, and Yeo (1985) recognized eight tribes, and treated as Amaryllidaceae only those genera in Traub's Amarylloideae.^[23] Müller-Doblies and Müller-Doblies (1996) recognized ten tribes and 19 subtribes, many of them with a single genus. Meerow and Snijman (1998) recognized 14 tribes, with two subtribes only in one of them, resurrected Eustephieae from Stenomesseae and recognized two new tribes, Calostemmateae and Hymenocallideae. Later on, they recognized four subtribes in Amaryllidae,^[12] and three subtribes for Haemantheae, changing the taxonomic rank of Gethyllidinae.^[14]

Basally African and Australasian clades

• Amaryllideae J.St.-Hil. (13 genera, x= 10, 11). Members of this tribe are endemic to Africa, with the exception of the pantropical genus Crinum. The Amaryllideae is sister to the rest of the Amaryllidaceae and is marked by a large number of diagnostic characters, such as extensible fibers in the bulb tunics, the type of pollen grains, the scapes with a sclerenchymatous sheath, and nondormant, water-rich seeds with chlorophyllous embryos and without phytomelanin. A few of the genera extend outside of South Africa, but only Crinum, has the seeds well adapted to oceanic dispersal ranges through Asia, Australia, and America. ^[12][24]
  • Subtribe Amaryllidinae.^[25] The single member of this subtribe (Amarylis, two species) has leaves with a prominent midrib, zygomorphic flowers with free tepals, dehiscent fruits, and large, pink or colorless seeds. It is endemic to the winter rainfall region of southern Africa.
  • Subtribe Boophoninae.^[26] Leaves spreading into an erect fan. Inflorescence of numerous helicoid cymes; pedicels elongating and radiating after anthesis; flowers actinomorphic, with a perigone tube; stamens free; fruit indehiscent, trigonal, 3-ribbed; fruiting head detaching from top of scape during seed dispersal; seeds endosperm-rich, partially chlorophyllous, cork-covered. Widespread in sub-Saharan Africa. It consists in a single genus (Boophone) with two species.
  • Subtribe Crininae.^[27] Leaves often with an intercalary meristem, usually fringed with cartilaginous teeth, apex often truncate. Flowers actinomorphic to zygomorphic, with a perigone tube; stamens free; fruit indehiscent, irregular, often rostellate; scape not abscising during seed dispersal except in Cybistetes where it detaches at ground level; seeds lacking an integument, endosperm-rich, partially chlorophyllous, cork-covered. Widespread in the tropics and sub-Saharan Africa. It consists in three genera: Crinum (65 species), Ammocharis (5 species) and Cybistetes (1 species).
  • Subtribe Strumariinae.^[28] Leaves often prostrate. Flowers zygomorphic or actinomorphic, with or without a perigone tube; stamens connate into a tube proximally (except in Strumaria where one whorl of stamens is fused to the style); fruit dehiscent; seeds with a well-developed chlorophyllous integument and stomatose testa. The subtribe is distributed in Southern Africa and consists in five genera Crossyne (2 species), Strumaria (24 species), Nerine (23 species), Hessea (13 species), Namaquanula (2 species), and Brunsvigia (23 species). Still, there exist controversy about the delimitation of genera in this subtribe.^[29]

• Cyrtantheae Salisb.^[30](1 genus, x=8). Consisting of a single African endemic genus, Cyrtanthus (56 species), this tribe is the only African taxon with the flat, winged phytomelanous seed characteristic of many American genera. Also, it is the most diverse in floral morphology of any genus in the family.

• Haemantheae (Pax) Hutchinson (6 genera, X= 6, 8, 9, 11 and 12). This tribe is characterized by its type of fruit, a berry, which differentiate all its members from the rest of the family. It includes three subtribes. ^[14]
  • Cliviinae, ^[31] are bulbless, rhizomatous perennials and include two genera, Cryptostephanus (2 species) and Clivia (5 species).
  • Haemanthineae,^[32] contains all of the genera of the tribe that form true bulbs, though Scadoxus is polymorphic for this character and has been misdiagnosed as being entirely rhizomatous. It is yet unclear whether bulbs form in Scadoxus only under certain environmental conditions or if bulb formation is limited to just certain species. Scadoxus (9 species) and Haemanthus (22 species) belong to this subtribe. Both genera have brush-like inflorescences, in which the bracts often form part of the pollinator attraction system.
  • Gethyllidinae.^[33] This subtribe consists of two closely related South African endemic genera, Apodolirion (6 species) and Gethyllis (30 species). Both genera maintain their scapes inside the bulb and have long, fragrant baccate fruits that have many seeds.

• Calostemmateae (two genera, x=10)^[34] The tribe consists of two Australasian genera. Proiphys (4 species) are forest understory herbs of Malaysia, Indonesia, the Philippines and tropical Australia, and Calostemma (3 species), endemic to Australia. The indehiscent capsules of both genera are similar in appearance to the unripe berry-fruits of Scadoxus and Haemanthus (Haemantheae), but early in the development of the seed, the embryo germinates precociously, and a bulbil forms within the capsule and functions as the mature propagule.

The Eurasian clades

• Lycorideae Traub, (two genera, x = 11) is a small tribe which represents the more or less temperate Asian component of the family, with Lycoris (22 species) ranging from Korea, through China, Myanmar, and Japan, and Ungernia (10 species) restricted to the mountains of central Asia. ^[35]

• Narcisseae Lam. & DC.,^[36] (two genera, x = 7, 10, 11). Together with the closely related Galantheae, this tribe represents the southwest Laurasian element of the family. At least some species exhibit 22 chromosomes in their cells, but much more chromosome evolution has occurred in the largest genus Narcissus (56 species) which is widely distributed from Macaronesia to Afghanistan, and South Eastern China to Japan. Sternbergia (8 species), the other member of the tribe, is distributed from Central and Southern Europe to Central Asia. Characteristically, the scape is solid and the spathe bracts are fused into a tube. Seeds are black, round and angular. A "paraperigone" is also characteristic of the tribe but absent from Sternbergia. No relationships with any other tribe outside of Galantheae have been proposed.^[37]

• Galantheae (Herb.) Parl.,^[38] (5 genera, x= 7, 8, 9, 11, 12). Of similar distribution to that of Narcisseae, the Galantheae are distinguished from the former by the type of anther dehiscence and leaf anatomy. Acis (9 species) is distributed in Western and Central Mediterranean region. Galanthus (20 species) and Leucojum (3 species) occur from Europe to Northern Iran. The other two members of the tribe are monotypic, Hannonia from Morocco and Lapiedra from Western Mediterranean. The delimitation of these two last genera are not definitive. Indeed, some botanists treated this tribe as a subtribe of Narcisseae. ^[31][39]

• Pancratieae Dumort.,^[40] (2 genera, x= 11). This Old World tribe has uncertain limits and a controversial phylogenetic position. Pancratium (21 species) is the largest genus and the most widespread, from South Africa to the Mediterranean and into Asia, and the only genus with stamens fused into a staminal cup. The flowers bear a remarkable resemblance to those of Hymenocallis. Vagaria (2 species), the other member of ths tribe, has a more restricted distribution: the coastal regions of Mediterranean countries, from Morocco to Lebanon and Israel. ^[41]

The American clades

• Griffineae.^[42] It is a tribe composed by two genera endemic to Brazil, Griffinia (21 species) and the monotypic Worsleya, which are characterized by their zygomorphic, pedicellate to nearly sessile flowers in shades of blue. ^[17][39]

• Hippeastreae.^[31] Species in this tribe are distributed in South America. Flowers are large and showy, zygomorphic, with the stamens in varying lengths, inflorescence bracts are often fused basally (along one side). The seeds are flattened, winged or D-shaped. Reported basic chromosome numbers are x= 8-13, 17, and higher. All the species in this tribe presents a remarkable aesthetic interest and horticultural value.^[17][43] It includes two subtribes:
  • subtribe: Hippeastrineae, includes species of medium height and often with many flowers in each inflorescence and inflorescence bracts are different in size and fused basally. Genera in this subtribe are Placea (6 species), Hippeastrum (91 species), Griffiniopsis (incl.: Eithea), Rhodophiala (28 species), Phycella (5 species) and Traubia (1 species).
  • subtribeZephyranthinea,^[31] includes species of small height with solitary flowers. Inflorescence bracts are fused forming a tube surrounding the pedicel of the flower. Genera in this subtribe are Sprekelia (2 species), Habranthus (74 apecies), and Zephyranthes (94 species).

• Eucharideae (4 genera, x = 23). Members of this neotropical tribe are recognized by their phytomelanous seeds, petiolate leaves, and white flowers. They are found in the understory of primary tropical rain forest. Genera in this tribe are Eucharis (17 species), Caliphruria (4 species), Urceolina (43 species), and the monotypic Plagiolirion.^[17][44]

• Stenomesseae^[45] (5 genera, x= 23). This Andean tribe, like the Neotropical Eucharidae, is characterized by its basic chromosome number and a well developed false corona or staminal cup in most of the genera. The seeds are uniformly dry, flattened and obliquely winged. In its original circumscription, the tribe was polyphyletic, for this reason a new tribe, Clinanteheae, has ben proposed for the members with linear leaves. Stenomesson (16 species) is distributed from Colombia to Northern Chile, Phaedranassa (9 species) ranges from Costa Rica to Ecuador, Eucrosia (8 species) ranges from Ecuador to Peru, and Rauhia (4 species) and the monotypic Mathieua are endemic to Peru.^[17][46][47]

• Eustephieae^[48] (4 genera, x= 23). This tribe represents a southern central andean clade. All the members have leaves with well developed palisade layers, exhibit a reduction series in staminal fusion and have trifid stigmas. The seeds are dry, flatetened and discoid. The species also exhibit changes in chrmosome number from the ancestral 2n =46. Eustephia (6 species) is endemic to Peru, Chlidanthus is distributed from Peru to Bolivia, Pyrolirion (6 species) from Peru to Northern Chile, and Hieronymiella (8 species) ranges from Southern Bolivia to North Western Argentina. ^[17][49]

• Hymenocallideae (3 genera, x = 23).^[50] Leptochiton (2 species) and Ismene (10 species) are two strictly central Andean genera, whereas the third genus of the tribe, Hymenocallis (63 species), is poorly represented in South America since it is primarily distributed in South Eastern USA, Mexico and West Indias. The flower of the members of this tribe have their stamens fused forming a characteristic corona. The fleshy seeds of Hymenocallidae are comprised of a thick outer integument formed by chlorenchyma with a well-developed vascular system, and an embryo which stores starch.^[17][51]

• Clinantheae^[52] (3 genera, x=23) Members of this tribe have linear or lorate leaves linear, often glaucous and lacking palisade in the mesophyll. The perianth is often brightly colored, consisting of six tepals in two series fused below into a tube of varying length. The filaments of the stamens are fused into a staminal cup. The fruit is a papery or woody loculicidal capsule with dry, flattened, obliquely winged seeds with a black or brown phytomelanous testa. The tribe includes the genera Clinanthus (22 species) distributed from Ecuador to North Western Argentina, Pamianthe (3 species) and Paramongaia (2 species) from Peru, Ecuador and Bolivia.^[17][53]

Hybrids

Several members of different genera of Amaryllidaceae hybridize readily, and the resulting hybrids are often sterile but can be propagated asexually. A hybrid name is usually reserved for these horticulturally arising hybrids which is indicated by a multiplication sign "×" placed before the name or epithet, as the case may be. Some nothogenera (a taxonomic rank given to artificial hybrids between species from different genera, from the Greek νόθος (“bastard”) and “genus”) of Amaryllidaceae are:

• × Amarcrinum Coutts (syn.: × Crindonna; × Crinodonna; × Amarcrinaflora): is the result of crossing Amaryllis belladonna with species of the genus Crinum.^[54]

• × Amarine Sealy, is the name applied to the hybrids between Amaryllis belladonna with members of Nerine.^[55]

• × Hippeastrelia is the result of crossing species of Hippeastrum with members of the genus Sprekelia.^[56]

Haemanthus × clarkei W. Wats., is the product of crossing H. albiflos with H. coccineus, it was raised in the UK by 1891.^[57][58] Clivia cyrtanthifora was raised by Charles Raes in Ghent, Belgium in the late 1850s and it was suggested to be a hybrid between Clivia miniata and Clivia nobilis,^[59][60] which was confirmed later by means of cytogenetics tools. ^[61] By means of DNA analysis, it was also confirmed that Lycoris straminea originated from hybridization between Lycoris chinensis and Lycoris radiata, and Lycoris caldwellii and Lycoris albiflora derived from hybridization between L. chinensis and Lycoris sprengeri.^[62]

In the genus Narcissus, high frequencies of natural hybrids have been reported and hybridization has been suggested as an explanation for the phenotypic variability within and between populations.^[63] Narcissus cavanillesii is a rare species, while N. serotinus is widely distributed across the Mediterranean. The hybrid, N. x perezlarae, is quite frequent in southeastern Spain but is scarce in Portugal.^[64]

Genera

• Acis Salisb.
• Amaryllis L.
• Ammocharis Herb. (Incl.: Cybistetes Milne-Redh. & Schweick.)
• Apodolirion Baker
• Boophone Herb.
• Brunsvigia Heist.
• Caliphruria Herb.
• Calostemma R.Br.
• Chlidanthus Herb. (incl.: Castellanoa Traub)
• Clinanthus Herb. (syn.: Anax Ravenna)
• Clivia Lindl.
• Crinum L.
• Crossyne Salisb.
• Cryptostephanus Welw. ex Baker
• Cyrtanthus Aiton (syn: Anoiganthus Baker; Vallota Salisb. ex Herb.)
• Eucharis Planch. & Linden
• Eucrosia Ker Gawl. (syn: Callipsyche Herb.)
• Eustephia Cav.
• Galanthus L.
• Gethyllis L. (syn.: Klingia Schönl.)
• Griffinia Ker Gawl. (incl. Hyline Herb. as a subgenus)
• Griffiniopsis Dutilh & Meerow (sin.: Eithea Ravenna)
• Habranthus Herb. (sin.: Zephyranthella (Pax) Pax; Haylockia Herb.)
• Haemanthus L.
• Hannonia Braun-Blanq. & Maire
• Hessea Herb. (syn.: Kamiesbergia Snijman)
• Hieronymiella Pax (syn.: Eustephiopsis R.E.Fr.)
• Hippeastrum Herb. (syn: Moldenkea Traub)
• Hymenocallis Salisb.
• Ismene Salisb. ex Herb (incl. Elisena Herb. and Pseudostenomesson Velarde as subgenera)
• Lapiedra Lag.
• Leptochiton Sealy
• Leucojum L.
• Lycoris Herb.
• Mathieua Klotzsch
• Namaquanula D.Müll.-Doblies & U.Müll.-Doblies
• Narcissus L. (incl.: Braxireon Raf. and Tapeinanthus Herb.)
• Nerine Herb.
• Pamianthe Stapf
• Pancratium L. (syn.: Mizonia A.Chev.; Chapmanolirion Dinter )
• Paramongaia Velarde
• Phaedranassa Herb. (sin: Neostricklandia Rauschert; Stricklandia Baker)
• Phycella Lyndl. (incl.:Famatina Ravenna)
• Placea Miers
• Plagiolirion Baker
• Proiphys Herb. (syn.: Eurycles Salisb. ex Schult. & Schult.)
• Pyrolirion Herb.
• Rauhia Traub
• Rhodophiala C.Presl (syn.: Rhodolirium Phil.; Rhodolirion Dalla Torre & Harms)
• Scadoxus Raf. (syn: Choananthus Rendle)
• Sprekelia Heist.
• Stenomesson Herb. (syn: Anax Ravenna; Callithauma Herb.; Crocopsis Pax, Pucara Ravenna)
• Sternbergia Waldst. & Kit.
• Strumaria Jacq. ex Willd. (Sin.: Bokkeveldia D.Müll.-Doblies & U.Müll.-Doblies; Carpolyza Salisb.;Gemmaria Salisb.; Carpolyza Salisb.; Tedingea D.Müll.-Doblies & U.Müll.-Doblies)
• Traubia Moldenke
• Ungernia Bunge
• Urceolina Rchb. (sin: Collania Schult. & Schult.f.; Pseudourceolina Vargas)
• Vagaria Herb.
• Worsleya (W.Watson ex Traub) Traub
• Zephyranthes Herb. (syn: Cooperia Herb.)

Ecology

Most species are adapted to seasonal climates that have a pronounced dry or cold period unfavourable for plant growth and during which the plants remain dormant. As a result most species are deciduous. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos. The aboveground parts (leaves and stems) of deciduous species die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavourable for growth by retreating underground. This is particularly useful in grasslands and fynbos, which are adapted to regular burning in the dry season. At that point the plants are dormant and their bulbs or corms are able to survive underground. The Neotropical genera of Amaryllidaceae are chiefly adapted for seasonally dry habitats and some prefer truly xeric environments in which their bulbs may remain dormant for a period longer than they are in active growth (e.g., Leptochiton, Paramongaia, some Eucrosia). At the other extreme, species have colonized the understory of rain forests (Eucharis, Griffinia) and aquatic habitats (a number of Hymenocallis, Hippeastrum angustifolium, Crinum). The family has also adapted to the high montane tropical climates of the Andes. Certain genera are primarily found at elevations in excess of 2000 meters; and Clinanthus humilis is found above 4000 meters. This species has adapted to high elevations by retaining the scape (and developing fruit) inside the bulb until the seeds are ripe. ^[65][66]

Veld fires clear the soil surface of competing vegetation and fertilise it with ash. With the arrival of the first rains, the dormant bulbs are ready to burst into growth, sending up flowers and stems before they can be shaded out by other vegetation. Many Amaryllidaceae species are adapted to cope with wildfires in their natural habitats, and those that depend on fire to flower are appropriately known as "fire lilies". In South Africa, several members of the genus Cyrtanthus are noted for their extremely rapid flowering response to natural bush fires. Indeed, species like Cyrtanthus contractus, widespread in the eastern half of South Africa, as well as Cyrtanthus ventricosus from the south western Cape, and Cyrtanthus odorus from the southern Cape, only flower after fires. The flowers of C. ventricosus are known to reach full flowering stage in just nine days following a fire.^[67] At least in C. ventricosus, this dependency on fire for initiating the flowering process is regulated by the smoke. ^[68] Aspects of the life history of Haemanthus pubescens were examined in the fire-prone Mediterranean climate zone of Lowland Coastal Fynbos in South Africa. The juvenile period of the species spans 9 years. The young reproductive period starts at 10 to 13 years and the reproductive maturity peaked at 16 years. Plants older than 17 years showed a marked reduction in reproductive potential. These age states indicated that the life-history strategy in terms of age states was similar to that of other fynbos plants and showed remarkable synchronization with the suggested fire frequency of about 15 to 20 years for this area. The phenological study indicated that the species is well adapted to the putative fire season. Flowers are only produced toward the end of the fire season, and leaves appear only in the cooler, wetter months when fires are unlikely to occur. Seeds are not dormant and germinate on the soil surface at the start of the cool, wet season.^[69]

Reproductive biology

Members of Amaryllidaceae possess extraordinary diversity in reproductive traits, even among closely related species. This variation is the result of the evolutionary lability of reproductive characters and implies that there are diverse functional solutions for achieving mating and fertility. The floral diversification that has accompanied the coevolution of flowers and animal pollinators is particularly striking and has resulted in contrasting suites of floral characters associated with different pollinator groups ^[70][71]

Heterostyly is a genetically controlled floral polymorphism with two morphological forms, namely, distyly and tristyly, that are characterized by two or three morphs, respectively. These morphs show variations in style and filament length, pollen grain size and production, and in the size of the stigmatic papillae and corolla. In addition, the morphs have an self-incompatibility system by which seeds are only produced in intermorph crosses. Reciprocal herkogamy forces pollinators to contact same-level floral organs with the same region of their body, thereby producing "legitimate" (intermorph) pollination. This morphological (floral parts) and physiological (incompatibility system) convergence in distylous species provides an efficient mechanism that avoids selfing and maximizes male and female fitness.^[72] Narcissus, a small genus of animal-pollinated species, has four major classes of stylar condition: stylar monomorphism, stigma-height dimorphism, distyly, and tristyly are represented among the 10 sections recognized in the genus. No other heterostylous taxon displays this range of stylar variation. Monomorphism is the ancestral condition in Narcissus and stigma-height dimorphism appears to have evolved multiple times. Floral morphology and pollinator relationships played an important role in the evolution of stylar polymorphisms in the genus. Long, narrow floral tubes (correlated with relatively precise depth-probed pollination by Lepidoptera) probably promoted the evolution of stigma-height dimorphism. Finally, the unusual conjunction of long, narrow floral tubes and deep coronas, part of a suite of floral features associated with pollination by long-tongued solitary bees, likely facilitated the convergent origins of heterostyly in Narcissus.^[73]

Zephyranthes is a rather versatile genus and contains self-incompatible and self-compatible species coupled with positional barrier between stigma and anthers. Furthermore, the genus also contains sexual and agamospermous species. The latter are often self-pollinated and pseudogamous. Zephyranthes atamasca is capable of producing seeds both by self-pollination and by outcrossing with other individuals of the species. Because the styles project beyond the anthers, however, self pollination in nature is probably less frequent than outcrossing.^[74] In contrast, it was documented a process of asexual seed production (apomixis) in Zephyranthes texensis. Ovules of Z. texensis produce embryos without any cells ever undergoing meiosis and without fusion of sperm and egg. So, whereas the seeds of most plant species are produced by sexual processes and bear a shuffled allotment of genes from both parents, the asexual seeds of Z. texensis are all genetically identical to the parent plant from which they were produced. ^[75] Z. sulphurea (2n=48) when pollinated with Z. candida (2n=40 and 41) has consistently given rise to seedlings with maternal chromosome number and morphology. On the other hand, the crosses involving sexual species like Z. candida (2n=41) as the female parent have generated a large heterogeneous progeny ranging in chromosome number from 2n=33 to 48 depending upon the number in the male parent. Such versatility of the breeding system together with chromosomal repatterning, hybridization, polyploidy and vegetative multiplication/apomixis explains the origin and preservation of an astonishing range in chromosome numbers from 2n=18 to 96 in this genus. ^[76]

Especially in Cyrtanthus the flowers are so diverse that they attract sunbirds, bees, long-tongued flies, butterflies and moths. Members of the genus Brunsvigia appear to show considerable variation in pollination syndromes. ^[77] A total of 22 species of southern African genera Crinum, Cyrtanthus and Pancratium conform to the syndrome of sphingophily, which includes a long-tubed, pale-coloured perianth that expands more fully at night, a strong sweet fragrance dominated by the acyclic terpenoid alcohol, linalool and abundant nectar.^[78] Southern Africa also has several Amaryllidaceae with remarkable dispersal abilities. Species of Brunsvigia, Boophone and Crossyne in particular have large, light, spherical fruiting heads that tumble along the ground in the wind, shedding their seeds as they move.

Economic and cultural value

Worldwide the Amaryllidaceae have greatest economic value as ornamentals. In addition, huge numbers of plants are traded for traditional medicines. Africans use the bulbs and leaves as poultices and decoctions for treating sores and digestive disorders, but in large dosages they are extremely poisonous. The Zulu people of South Africa also use rhizomes of clivias as protective charms. In Peru, the Inca people frequently depicted flowers of Amaryllidaceae (Ismene, Pyolirion and Stenomesson) on ceremonial drinking vessels. In southern Africa, however, indigenous art portraying plants is rare. The single known rock painting of a Brunsvigia species in Lesotho probably emphasizes how much the San people valued the bulbs for their psychoactive effects. ^[3]

In cool temperate climates, Narcissus (daffodils), Leucojum (snowflakes) and Galanthus (snowdrops) are among the most important spring-flowering bulbs in commerce. Elsewhere, in warm temperate and subtropical climates, species of Amaryllis, Clivia, Hippeastrum, Nerine, and Zephyranthes are the most popular choices for gardens and containers.^[3]

Conservation status

Habitat loss is currently the greatest threat to the Amaryllidaceae in South Africa, where 59 species are endangered or vulnerable and 58 species are near threatened.^[3]

References

Bibliography

• Michael F. Fay and Mark W. Chase. 1996. Resurrection of Themidaceae for the Brodiaea alliance, and Recircumscription of Alliaceae, Amaryllidaceae and Agapanthoideae. Taxon 45: 441-451 (abstract)

External links

Wikimedia Commons has media related to: Amaryllidaceae

• Images of Amaryllidaceae species in Topwalks
• links at CSDL, Texas
• International Bulb Society
• Images and information of Amaryllidaceae in Israel
• Images of Amarilids from Chile