Amaryllidoideae is the subfamily of
flowering plantThe flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...
s that takes its name from the
genusIn biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
AmaryllisAmaryllis is a small genus of flowering bulbs, with two species. The better known of the two, Amaryllis belladonna, is a native of South Africa, particularly the rocky southwest region near the Cape...
. It is part of the family
AmaryllidaceaeAmaryllidoideae is the subfamily of flowering plants that takes its name from the genus Amaryllis. It is part of the family Amaryllidaceae, in order Asparagales...
, in order
AsparagalesAsparagales is the name of an order of plants, used in modern classification systems such as the APG III system . The order takes its name from the family Asparagaceae and is placed in the monocots. The order has only recently been recognized in classification systems...
. The most recent
APGThe Angiosperm Phylogeny Group, or APG, refers to an informal international group of systematic botanists who came together to try to establish a consensus on the taxonomy of flowering plants that would reflect new knowledge about plant relationships discovered through phylogenetic studies., three...
classification,
APG IIIThe APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy...
, takes a broad view of the Amaryllidaceae, which then has three subfamilies, one of which is Amaryllidoideae (the old Amaryllidaceae family), and the others are Allioideae (the old Alliaceae family) and Agapanthoideae (the old Agapanthaceae family). The subfamily consists of about sixty
generaIn biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
, with over eight hundred
speciesIn biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
, and a worldwide distribution.
The Amaryllidoideae are
herbaceousA herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...
,
perennialA perennial plant or simply perennial is a plant that lives for more than two years. The term is often used to differentiate a plant from shorter lived annuals and biennials. The term is sometimes misused by commercial gardeners or horticulturalists to describe only herbaceous perennials...
flowering plants, usually with
bulbsA bulb is a short stem with fleshy leaves or leaf bases. The leaves often function as food storage organs during dormancy.A bulb's leaf bases, known as scales, generally do not support leaves, but contain food reserves to enable the plant to survive adverse conditions. At the center of the bulb is...
(some are rhizomatous). Their fleshy leaves are arranged in two vertical columns, and their flowers are large.
Description
Members of Amaryllidoideae are perennial, mostly
deciduousDeciduous means "falling off at maturity" or "tending to fall off", and is typically used in reference to trees or shrubs that lose their leaves seasonally, and to the shedding of other plant structures such as petals after flowering or fruit when ripe...
, rarely shrubby or treelike plants, often with
bulbA bulb is a short stem with fleshy leaves or leaf bases. The leaves often function as food storage organs during dormancy.A bulb's leaf bases, known as scales, generally do not support leaves, but contain food reserves to enable the plant to survive adverse conditions. At the center of the bulb is...
s or, rarely, with
rhizomeIn botany and dendrology, a rhizome is a characteristically horizontal stem of a plant that is usually found underground, often sending out roots and shoots from its nodes...
s, as in the genera
CliviaClivia is a genus of monocot flowering plants native to southern Africa. They are from the family Amaryllidaceae, subfamily Amaryllidoideae. Common names include Kaffir lily and bush lily....
, Cryptostephanus and
ScadoxusScadoxus is a genus of 9 species native to tropical Africa in the family Amaryllidaceae, subfamily Amaryllidoideae. The genus has close affinities with Haemanthus from which it has only recently been separated.Species in the genus are...
. While growing, the bulb is kept deep below ground by a special type of
rootIn vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...
that lengthen and contract.
Leaves-History:Vocalist Arnar Gudjonsson was formerly the guitarist with Mower, and he was joined by Hallur Hallsson , Arnar Ólafsson , Bjarni Grímsson , and Andri Ásgrímsson . Late in 2001 they played with Emiliana Torrini and drew early praise from the New York Times...
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
EucrosiaEucrosia is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family distributed from Ecuador to Peru. The name is derived from the Greek eu, beautiful, and krossos, a fringe, referring to the long stamens. The genus contains eight species...
and
ScadoxusScadoxus is a genus of 9 species native to tropical Africa in the family Amaryllidaceae, subfamily Amaryllidoideae. The genus has close affinities with Haemanthus from which it has only recently been separated.Species in the genus are...
, which occupy habitats with low light-intensity, have leaves that are broad and flattened, whereas in semi-arid regions like southern
AfricaAfrica is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...
, species of
BrunsvigiaBrunsvigia is a flowering plant genus in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains about 20 species native to South Africa....
, Crossyne, Gethyllis and
HaemanthusHaemanthus is a Southern African genus of Amaryllidaceae, subfamily Amaryllidoideae, with some 22 known species, endemic to South Africa, Namibia and the kingdoms of Lesotho and Swaziland...
have leaves covered with variously shaped hairs. The leaves in
Crossyne and some
HaemanthusHaemanthus is a Southern African genus of Amaryllidaceae, subfamily Amaryllidoideae, with some 22 known species, endemic to South Africa, Namibia and the kingdoms of Lesotho and Swaziland...
species are also attractively spotted with dark green or red.
The flowers are large and showy, bisexual, trimerous, actinomorphic (less often slightly zygomorphic, as in
SprekeliaSprekelia is a genus of at least three bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are native to Central America. Like Hippeastrum, these plants were known as Amaryllis. Sprekelia plants are sometimes called "Aztec lilies", although they are not true lilies...
). 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
GriffiniaGriffinia is a genus of monocotyledonous plants in the Amaryllidaceae family, subfamily Amaryllidoideae. It includes 21 species which are endemic to South America, Brazil. The most closely related genus to it is the monotypic Worsleya. The members of the genus Griffinia are tropical, bulbous plants...
, WorsleyaThe genus Worsleya contains only one species, Worsleya procera, previously known as Worsleya rayneri. It is one of the largest and rarest members of the subfamily Amaryllidoideae . Worsleya is a tropical plant. This species is also known as the empress of Brazil because of its origin in South...
and
Lycoris. The
perianthThe term perianth has two similar but separate meanings in botany:* In flowering plants, the perianth are the outer, sterile whorls of a flower...
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
tepalTepals are elements of the perianth, or outer part of a flower, which include the petals or sepals. The term tepal is more often applied specifically when all segments of the perianth are of similar shape and color, or undifferentiated, which is called perigone...
s. The androecium is composed by six stamens, inserted at the perianth throat or at the base of each
tepalTepals are elements of the perianth, or outer part of a flower, which include the petals or sepals. The term tepal is more often applied specifically when all segments of the perianth are of similar shape and color, or undifferentiated, which is called perigone...
. Some
Griffinia species have five stamens and some
Gethyllis species have multiple stamens with about 60 anthers. In
PancratiumPancratium may be:* Pankration, a sport or martial art introduced in the Olympic games in 648 BC, and its modern version* Pancratium , a genus of flowering plants in the family Amaryllidaceae...
and
HymenocallisHymenocallis is a genus of plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains more than 60 species native to tropical and subtropical America. Hymenocallis are bulbous perennial herbs. The flowers have their stamens united to a characteristic corona...
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 subfamily produce nectar and are often heavily scented. The
inflorescenceAn inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
can be of one or many flowers in a terminal spike,
racemeA raceme is a type of inflorescence that is unbranched and indeterminate and bears pedicellate flowers — flowers having short floral stalks called pedicels — along the axis. In botany, axis means a shoot, in this case one bearing the flowers. In a raceme, the oldest flowers are borne...
,
panicleA panicle is a compound raceme, a loose, much-branched indeterminate inflorescence with pedicellate flowers attached along the secondary branches; in other words, a branched cluster of flowers in which the branches are racemes....
or, more often, in an
umbelAn umbel is an inflorescence which consists of a number of short flower stalks which are equal in length and spread from a common point, somewhat like umbrella ribs....
-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
scapeIn biology, the term scape may refer to:* The first segment of an insect antenna* A finger-like appendage of the epigyne of a female spider* Scape , a flowering stemScape may also refer to:...
. In unusual genera like
Gethyllis, however, the scape carries only one flower and remains subterranean.
The
fruitIn broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
is a dry
capsuleIn botany a capsule is a type of simple, dry fruit produced by many species of flowering plants. A capsule is a structure composed of two or more carpels that in most cases is dehiscent, i.e. at maturity, it splits apart to release the seeds within. A few capsules are indehiscent, for example...
, or more rarely, a fleshy
berryThe botanical definition of a berry is a fleshy fruit produced from a single ovary. Grapes are an example. The berry is the most common type of fleshy fruit in which the entire ovary wall ripens into an edible pericarp. They may have one or more carpels with a thin covering and fleshy interiors....
and contain dry, dark and often flattened, or fleshy, round, and greenish
seedA seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
s. The seeds have an oily
endospermEndosperm is the tissue produced inside the seeds of most flowering plants around the time of fertilization. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This makes endosperm an important source of nutrition in human diet...
and usually with a black or brown phytomelanous testa, sometimes with a caruncular
elaiosomeElaiosomes are fleshy structures that are attached to the seeds of many plant species. The elaiosome is rich in lipids and proteins, and may be variously shaped. Many plants have elaiosomes to attract ants, which take the seed to their nest and feed the elaiosome to their larvae...
at the chalazal end.
The most common basic
chromosomeA chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
number in the subfamily is
x = 11, although during its evolution a vast arrange of different basic chromosome numbers arose, from
x= 6 to
x= 23.
Members of this subfamily present a unique type of alkaloids, the norbelladine alkaloids, which are
tyrosineTyrosine or 4-hydroxyphenylalanine, is one of the 22 amino acids that are used by cells to synthesize proteins. Its codons are UAC and UAU. It is a non-essential amino acid with a polar side group...
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.
Distribution and habitat
Members of the subfamily are widespread, being found in the Holarctic, Paleotropical, Neotropical, Cape, and Australian regions. In the Neotropics, the family occurs from
MexicoThe United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
through
Central AmericaCentral America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...
and the West Indies to
ChileChile ,officially the Republic of Chile , is a country in South America occupying a long, narrow coastal strip between the Andes mountains to the east and the Pacific Ocean to the west. It borders Peru to the north, Bolivia to the northeast, Argentina to the east, and the Drake Passage in the far...
and
ArgentinaArgentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...
in
South AmericaSouth America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
. Notable areas of diversity throughout this range include eastern
BrazilBrazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
, north-central Chile (outside of the tropical zone, however), and the central
AndesThe Andes is the world's longest continental mountain range. It is a continual range of highlands along the western coast of South America. This range is about long, about to wide , and of an average height of about .Along its length, the Andes is split into several ranges, which are separated...
of
EcuadorEcuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...
and Peru.
HippeastrumHippeastrum is a genus of about 90 species and 600+ hybrids and cultivars of bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to tropical and subtropical regions of the Americas from Argentina north to Mexico and the Caribbean. Some species are grown for their large...
is primarily found in the Andes and eastern Brazil,
HymenocallisHymenocallis is a genus of plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains more than 60 species native to tropical and subtropical America. Hymenocallis are bulbous perennial herbs. The flowers have their stamens united to a characteristic corona...
occurs mostly in Mesoamerica,
Clinanthus is largely endemic to Peru, and
ZephyranthesZephyranthes is a genus of 71 species in the Amaryllis family . There are numerous hybrids and cultivars. Common names for species in this genus include fairy lily, rainflower, zephyr lily, magic lily, Atamasco lily, and rain lily.The name is derived from Ζέφυρος , the Greek god of the west...
is broadly distributed. The greatest generic diversity is found in Peru.
Taxonomy
The group was first described (as the family Amaryllidaceae) by the French naturalist
Jean Henri Jaume Saint-HilaireJean Henri Jaume Saint-Hilaire was a French naturalist and artist, born in Grasse, France.-Biography:Born as Jaume, he added Saint-Hilaire later. Some biographers indicate that this addition was to distinguish himself from a family member, Henri-Honore Jaume, a Jacobin who had been involved in...
in 1805. The type genus,
Amaryllis, is named after Amaryllis, a beautiful shepherdess mentioned by
TheocritusTheocritus , the creator of ancient Greek bucolic poetry, flourished in the 3rd century BC.-Life:Little is known of Theocritus beyond what can be inferred from his writings. We must, however, handle these with some caution, since some of the poems commonly attributed to him have little claim to...
,
VirgilPublius Vergilius Maro, usually called Virgil or Vergil in English , was an ancient Roman poet of the Augustan period. He is known for three major works of Latin literature, the Eclogues , the Georgics, and the epic Aeneid...
and
OvidPublius Ovidius Naso , known as Ovid in the English-speaking world, was a Roman poet who is best known as the author of the three major collections of erotic poetry: Heroides, Amores, and Ars Amatoria...
, and hence an allegory of beauty.
An Amaryllidaceae family has been recognized, with varying definitions, by most classification systems of the 20th Century, although the
Cronquist systemThe Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in his texts An Integrated System of Classification of Flowering Plants and The Evolution and Classification of Flowering Plants .Cronquist's system places flowering plants into two...
included it within a very broadly defined
LiliaceaeThe Liliaceae, or the lily family, is a family of monocotyledons in the order Liliales. Plants in this family have linear leaves, mostly with parallel veins but with several having net venation , and flower arranged in threes. Several have bulbs, while others have rhizomes...
. The two families were traditionally separated by including species with inferior ovaries in Amaryllidaceae and those with superior ovaries in Liliaceae.
The
APGThe Angiosperm Phylogeny Group, or APG, refers to an informal international group of systematic botanists who came together to try to establish a consensus on the taxonomy of flowering plants that would reflect new knowledge about plant relationships discovered through phylogenetic studies., three...
initially (
1998The APG system of plant classification is the first, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in 1998 by the Angiosperm Phylogeny Group. It was superseded in 2003 by a revision, the APG II system, and then in 2009 by a further...
) recognized the Agapanthaceae, Alliaceae, and Amaryllidaceae
sensu stricto as separate families. The
2003The APG II system of plant classification is the second, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Group. It was a revision of the first APG system, published in 1998, and was superseded in 2009...
revision tentatively combined the three as subtaxa of a single family in the order
AsparagalesAsparagales is the name of an order of plants, used in modern classification systems such as the APG III system . The order takes its name from the family Asparagaceae and is placed in the monocots. The order has only recently been recognized in classification systems...
, citing Alliaceae Batsch (1786) as the name of earliest priority. The most recent revision (
2009The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy...
) is clear that all three families should be combined as separate subfamilies of a broadly defined Amaryllidaceae, the approach followed here.
The APG III approach has been disputed. Differences between the subfamilies regarding chemical compounds and morphology are considered sufficiently important by some authors to keep them as three separate families. If the Amaryllidoideae is treated as a family, it is 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.
Evolution
Amaryllidoideae and its sister genus
AgapanthusAgapanthus is the only genus in the subfamily Agapanthoideae of the flowering plant family Amaryllidaceae. The family is in the monocot order Asparagales....
(the subfamily Agapanthoideae), originated in western Gondwanaland, and Africa has been the site of considerable innovation in the subfamily's history as well.
Crinum is unusual in the subfamily because of its dispersal ability that comes from seeds that are adapted for oceanic dispersal and it has a pantropical distribution. The most recent data indicate that genetic relationships within the subfamily closely mirror their geographic distributions.
The Calostemmateae, the only exclusively Australasian element of the subfamily, may have been isolated from the African lineages as Australia separated from western Gondwanaland.
The Eurasian/Mediterranean members of the subfamily are the sister group of the American genera, but it is unclear when and how the major dispersal between these areas occurred. There are considerably more species in South America than on the Northern Continent, and there has been some recent movement northward, but this does not imply that the subfamily reached South America earlier.
Phylogeny
The Amaryllidoideae has so far defied precise understanding of its internal phylogeny, and its relationships to other higher Asparagales. Combined analysis of three
plastidPlastids are major organelles found in the cells of plants and algae. Plastids are the site of manufacture and storage of important chemical compounds used by the cell...
DNADeoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
sequences (
rbcL,
trnL intronAn intron is any nucleotide sequence within a gene that is removed by RNA splicing to generate the final mature RNA product of a gene. The term intron refers to both the DNA sequence within a gene, and the corresponding sequence in RNA transcripts. Sequences that are joined together in the final...
, and
trnL-F spacer) for 50 genera of Amaryllidoideae analyzed together with members of Allioideae, Behniaceae, Convallariaceae, Scillioideae,
ThemidaceaeBrodiaeoideae is a monocot subfamily of flowering plants in the family Asparagaceae, order Asparagales. It has been treated as a separate family, Themidaceae. They are native to Central America and western North America, from British Columbia to Guatemala...
, and Hemerocallidoideae, resolves Agapanthoideae as sister to Amaryllidoideae with weak support and places
Agapanthus-Amaryllidoideae as a sister clade to a monophyletic Allioideae.
The African tribe
AmaryllideaeAmaryllideae is a tribe of perennial, herbaceous and bulbous plants in the Amaryllis family . Members of this tribe are distributed in Africa, with the exception of the pantropical genus Crinum...
is sister to the remainder of the Amaryllidoideae.
AmaryllisAmaryllis is a small genus of flowering bulbs, with two species. The better known of the two, Amaryllis belladonna, is a native of South Africa, particularly the rocky southwest region near the Cape...
and
BoophoneBoophone is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of two species distributed in Tropical and Southern Africa...
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
StrumariaStrumaria is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Strumaria barbarae, Oberm.* Strumaria hardyana, D.& U. Müll.-Doblies...
and its allies. The other is the predominantly sub-Saharan African group that includes
CrinumCrinum is a genus of about 180 species of perennial plants that have large showy flowers on leafless stems, and develop from bulbs. They are found along the sides of streams and lakes in tropical and subtropical areas worldwide, including South Africa....
and related genera. The African baccate-fruited Haemantheae and the Australasian Calostemmateae are sister tribes, and the African endemic Cyrtantheae is sister to them both. The most completely resolved and best supported tree for Haemantheae, divides the tribe into two main clades. The smaller clade,
uniting
CliviaClivia is a genus of monocot flowering plants native to southern Africa. They are from the family Amaryllidaceae, subfamily Amaryllidoideae. Common names include Kaffir lily and bush lily....
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.
The Eurasian and neotropical genera of the Amaryllidoideae are a well-supported clade, and are sister groups. Lycorideae are basal in the Eurasian clade and begin a grade that continues with
Hannonia, then
Pancratium, then
Lapiedra. The genera
Galanthus,
Narcissus, and
SternbergiaSternbergia is a genus in the family Amaryllidaceae, subfamily Amaryllidoideae, of around 8 species that show a broad distribution throughout Mediterranean Europe and Asia. It was first described by Clusius in 1601 as Narcissus, before being redescribed by Carl Linnaeus as Amaryllis in 1753...
are resolved as monophyletic with strong support.
Leucojum sensu lato is paraphyletic and recognition of
AcisThe 3D ACIS Modeler is a 3D modelling kernel owned by Spatial Corporation . ACIS is used by many software developers in industries such as computer-aided design , Computer-aided manufacturing , Computer-aided engineering , Architecture, engineering and construction , Coordinate-measuring machine...
for the mostly autumn-flowering Mediterranean species is supported by molecular data.
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
HippeastreaeHippeastreae is a tribe of plants belonging to the subfamily Amaryllidoideae of the Amaryllis family . 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 . The seeds are...
. The second clade constitutes the tribes centered in the Andes whose basic chromosome number is derived by
polyploidyPolyploid is a term used to describe cells and organisms containing more than two paired sets of chromosomes. Most eukaryotic species are diploid, meaning they have two sets of chromosomes — one set inherited from each parent. However polyploidy is found in some organisms and is especially common...
(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 (Eucharideae has nomenclatural priority); 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 Amaryllidoideae may have been reduced to peripheral isolates some time after its initial entry and spread through the Americas. 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
GriffineaeThe tribe Griffineae includes 2 genera with 22 species from South America which are actually endemic to Brazil. A typical character of the representatives of the tribe are the flowers - They are with blue or lilac color collected into an umbel...
in
the hippeastroid clade. These two small tribes may represent either ancestral or merely very isolated elements of their
respective clades.
A
cladogramA cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...
which summarizes the phylogenetic relationships among tribes and subtribes of Amaryllidoideae is given below.
Tribes, subtribes and genera
Two modern sub-classifications of the Amaryllidoideae are those of Müller-Doblies and Müller-Doblies (1996) and Meerow and Snijman (1998). Müller-Doblies and Müller-Doblies recognized ten tribes and 19 subtribes, many of them with a single genus. Meerow and Snijman 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, and three subtribes for Haemantheae, changing the taxonomic rank of Gethyllidinae.
Basally African and Australasian clades:
- Amaryllideae
Amaryllideae is a tribe of perennial, herbaceous and bulbous plants in the Amaryllis family . Members of this tribe are distributed in Africa, with the exception of the pantropical genus Crinum...
J.St.-Hil. (13 genera, x= 10, 11). Members of this tribe are endemic to Africa, with the exception of the pantropical genus CrinumCrinum is a genus of about 180 species of perennial plants that have large showy flowers on leafless stems, and develop from bulbs. They are found along the sides of streams and lakes in tropical and subtropical areas worldwide, including South Africa....
. The Amaryllideae is sister to the rest of the Amaryllidoideae 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 AsiaAsia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...
, AustraliaAustralia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
, and America.
- Subtribe Amaryllidinae. 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. 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
Boophone is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of two species distributed in Tropical and Southern Africa...
) with two species.
- Subtribe Crininae. 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
Crinum is a genus of about 180 species of perennial plants that have large showy flowers on leafless stems, and develop from bulbs. They are found along the sides of streams and lakes in tropical and subtropical areas worldwide, including South Africa....
(65 species), AmmocharisAmmocharis is a genus in the Amaryllidaceae family which includes 6 species distributed in Africa. The plant grows as a succulent, above-ground bulb, preferring seasonally wet, hot, sandy soils and full sun.-Species:...
(5 species) and Cybistetes (1 species).
- Subtribe Strumariinae. 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
Strumaria is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Strumaria barbarae, Oberm.* Strumaria hardyana, D.& U. Müll.-Doblies...
(24 species), NerineNerine is a genus of plants belonging to the Amaryllidaceae family, subfamily Amaryllidoideae Native to South Africa, there are about 30 different species in the genus. Nerine have been widely cultivated and much hybridized and are now spread world wide....
(23 species), Hessea (13 species), Namaquanula (2 species), and BrunsvigiaBrunsvigia is a flowering plant genus in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains about 20 species native to South Africa....
(23 species). Still, there exists controversy about the delimitation of genera in this subtribe.
- Cyrtantheae Salisb.
Richard Anthony Salisbury FRS was a British botanist. While he is remembered as a valuable worker in horticultural and botanical sciences, several bitter disputes caused him to be ostracised by his contemporaries.-Life:...
(1 genus, x=8). Consisting of a single African endemic genus, CyrtanthusCyrtanthus is a genus of perennial, herbaceous and bulbous plants in the Amaryllis family . This highly ornamental genus encompasses about 60 species...
(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 subfamily.
- Haemantheae (Pax
Ferdinand Albin Pax was a German botanist and entomologist. He specialised in Lepidoptera, Diptera, as well as in spermatophytes, describing several species....
) HutchinsonJohn Hutchinson, OBE, FRS was a renowned English botanist, taxonomist and author.-Life and career:...
(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 subfamily. It includes three subtribes.
- Cliviinae, are bulbless, rhizomatous perennials and include two genera, Cryptostephanus (2 species) and Clivia
Clivia is a genus of monocot flowering plants native to southern Africa. They are from the family Amaryllidaceae, subfamily Amaryllidoideae. Common names include Kaffir lily and bush lily....
(5 species).
- Haemanthineae, 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
Scadoxus is a genus of 9 species native to tropical Africa in the family Amaryllidaceae, subfamily Amaryllidoideae. The genus has close affinities with Haemanthus from which it has only recently been separated.Species in the genus are...
(9 species) and HaemanthusHaemanthus is a Southern African genus of Amaryllidaceae, subfamily Amaryllidoideae, with some 22 known species, endemic to South Africa, Namibia and the kingdoms of Lesotho and Swaziland...
(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. This subtribe consists of two closely related South African endemic genera, Apodolirion
Apodolirion is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of 6 species distributed in South Africa...
(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) The tribe consists of two Australasian genera. Proiphys
Proiphys is a genus of herbaceous, perennial and bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It consists of 4 species that are native to Southeast Asia and Australia.-Description:...
(4 species) are forest understory herbs of Malaysia, Indonesia, the Philippines and tropical Australia, and CalostemmaCalostemma is a small genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists in 3 species endemic to Australia, where they are distributed in arid regions with summer precipitations...
(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 subfamily, with Lycoris
Lycoris is a genus of 13–20 species of flowering plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are native to eastern and southern Asia in Japan, southern Korea, eastern and southern China, northern Vietnam, northern Laos, northern Thailand, northern Burma, Nepal, northern...
(22 species) ranging from KoreaKorea ) is an East Asian geographic region that is currently divided into two separate sovereign states — North Korea and South Korea. Located on the Korean Peninsula, Korea is bordered by the People's Republic of China to the northwest, Russia to the northeast, and is separated from Japan to the...
, through ChinaChinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
, MyanmarBurma , officially the Republic of the Union of Myanmar , is a country in Southeast Asia. Burma is bordered by China on the northeast, Laos on the east, Thailand on the southeast, Bangladesh on the west, India on the northwest, the Bay of Bengal to the southwest, and the Andaman Sea on the south....
, and JapanJapan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
, and Ungernia (10 species) restricted to the mountains of central Asia.
- Narcisseae Lam.
Jean-Baptiste Pierre Antoine de Monet, Chevalier de la Marck , often known simply as Lamarck, was a French naturalist...
& DC., (two genera, x = 7, 10, 11). Together with the closely related Galantheae, this tribe represents the southwest LaurasiaIn paleogeography, Laurasia was the northernmost of two supercontinents that formed part of the Pangaea supercontinent from approximately...
n element of the subfamily. 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 MacaronesiaMacaronesia is a modern collective name for several groups of islands in the North Atlantic Ocean near Europe and North Africa belonging to three countries: Portugal, Spain, and Cape Verde...
to AfghanistanAfghanistan , officially the Islamic Republic of Afghanistan, is a landlocked country located in the centre of Asia, forming South Asia, Central Asia and the Middle East. With a population of about 29 million, it has an area of , making it the 42nd most populous and 41st largest nation in the world...
, and South Eastern China to Japan. SternbergiaSternbergia is a genus in the family Amaryllidaceae, subfamily Amaryllidoideae, of around 8 species that show a broad distribution throughout Mediterranean Europe and Asia. It was first described by Clusius in 1601 as Narcissus, before being redescribed by Carl Linnaeus as Amaryllis in 1753...
(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.
- Galantheae (Herb.) Parl.
Filippo Parlatore was an Italian botanist.Italian botanist, b. at Palermo, 8 Aug., 1816; d. at Florence, 9 Sept., 1877, a devout and faithful Catholic. He studied medicine at Palermo, but practiced only for a short time, his chief activity being during the cholera epidemic of 1837...
, (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. AcisAcis is a genus of perennial, herbaceous and bulbous plants in the Amaryllis family . The genus consists in 9 species distributed in Europe and Northern Africa.The genus was reinstated in 2004, after it was determined on morphological and molecular grounds that these species should be segregated...
(9 species) is distributed in Western and Central Mediterranean region. Galanthus (20 species) and Leucojum (3 species) occur from Europe to Northern IranIran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...
. 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.
- Pancratieae Dumort.
Barthélemy Charles Joseph, Baron Dumortier was a Belgian politician and botanist. Some consider him to be the true discoverer of cell division, although he is rarely credited as such.-Works:...
, (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 MoroccoMorocco , officially the Kingdom of Morocco , is a country located in North Africa. It has a population of more than 32 million and an area of 710,850 km², and also primarily administers the disputed region of the Western Sahara...
to LebanonLebanon , officially the Republic of LebanonRepublic of Lebanon is the most common term used by Lebanese government agencies. The term Lebanese Republic, a literal translation of the official Arabic and French names that is not used in today's world. Arabic is the most common language spoken among...
and IsraelThe State of Israel is a parliamentary republic located in the Middle East, along the eastern shore of the Mediterranean Sea...
.
The American clades:
- Griffineae
The tribe Griffineae includes 2 genera with 22 species from South America which are actually endemic to Brazil. A typical character of the representatives of the tribe are the flowers - They are with blue or lilac color collected into an umbel...
. It is a tribe composed by two genera endemic to Brazil, GriffiniaGriffinia is a genus of monocotyledonous plants in the Amaryllidaceae family, subfamily Amaryllidoideae. It includes 21 species which are endemic to South America, Brazil. The most closely related genus to it is the monotypic Worsleya. The members of the genus Griffinia are tropical, bulbous plants...
(21 species) and the monotypic WorsleyaThe genus Worsleya contains only one species, Worsleya procera, previously known as Worsleya rayneri. It is one of the largest and rarest members of the subfamily Amaryllidoideae . Worsleya is a tropical plant. This species is also known as the empress of Brazil because of its origin in South...
, which are characterized by their zygomorphic, pedicellate to nearly sessile flowers in shades of blue.
- Hippeastreae
Hippeastreae is a tribe of plants belonging to the subfamily Amaryllidoideae of the Amaryllis family . 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 . The seeds are...
. 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. 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
Placea is a genus of about 6 species of herbaceous, perennial and bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are endemic to Chile.-List of species:...
(6 species), HippeastrumHippeastrum is a genus of about 90 species and 600+ hybrids and cultivars of bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to tropical and subtropical regions of the Americas from Argentina north to Mexico and the Caribbean. Some species are grown for their large...
(91 species), Griffiniopsis (incl.: Eithea), RhodophialaRhodophiala is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of 28 South American species distributed in southern Brazil, Argentina, and, specially, in Chile....
(28 species), Phycella (5 species) and Traubia (1 species).
- subtribeZephyranthinea
Zephyranthinea is a subtribe of plants classified under the tribe Hippeastreae. It belongs to the subfamily Amaryllidoideae of the Amaryllis family . They are generally small plants with solitary flowers. Spathes are fused forming a tube surrounding the pedicel of the flower. Most of its members...
, 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 SprekeliaSprekelia is a genus of at least three bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are native to Central America. Like Hippeastrum, these plants were known as Amaryllis. Sprekelia plants are sometimes called "Aztec lilies", although they are not true lilies...
(2 species), HabranthusHabranthus is a genus of tender herbaceous flowering bulbs in the Amaryllidaceae, subfamily Amaryllidoideae. The genus was first identified by pioneering bulb enthusiast William Herbert in 1824.-Description:...
(74 apecies), and ZephyranthesZephyranthes is a genus of 71 species in the Amaryllis family . There are numerous hybrids and cultivars. Common names for species in this genus include fairy lily, rainflower, zephyr lily, magic lily, Atamasco lily, and rain lily.The name is derived from Ζέφυρος , the Greek god of the west...
(94 species).
- Eucharideae (9 genera, x = 23). Members of this neotropical tribe are recognized by their phytomelanous seeds, petiolate leaves, and variously colored flowers, most of which have the stamens connate below into a staminal cup. The seed are either dry, flattened and obliquely winged (the majority) or globose (sausage-shaped in Urceolina), turgid, and with oily endosperm. They are found in the understory of primary tropical rain forest, and seasonally dry vegetation of Central America (Eucharis
Eucharis is a genus of about 15-20 species of monocotyledonous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to Central America and South America from Guatemala south to Bolivia.-Description:...
)to Andean South America. Genera in this tribe are EucharisEucharis is a genus of about 15-20 species of monocotyledonous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to Central America and South America from Guatemala south to Bolivia.-Description:...
(17 species), CaliphruriaCaliphruria is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of four species distributed in tropical regions of South America, three of them are endemic to Colombia. This genus is closely related with the genera Eucharis and Urceolina, all of them known...
(4 species), Urceolina (4-5 species), the monotypic Plagiolirion, Stenomesson (ca. 16 species), distributed from Colombia to northern Peru, PhaedranassaPhaedranassa is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Phaedranassa brevifolia, Meerow* Phaedranassa cinerea, Ravenna...
(9 species), from Colombia to Ecuador (a poorly known species described from Costa Rica may be adventive), EucrosiaEucrosia is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family distributed from Ecuador to Peru. The name is derived from the Greek eu, beautiful, and krossos, a fringe, referring to the long stamens. The genus contains eight species...
(8 species), from Ecuador to Peru, Rauhia (4 species), endemic to Peru, and the monotypic Peruvian endemic Mathieua, which may ultimately be referred to Stenomesson. .
- Eustephieae (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
Chlidanthus is a genus that consists of 10 species of tender bulbs from tropical South America, mostly natives to the Andes. The botanical name comes from the Greek, meaning "delicate flower". The plants have large spherical bulbs with gray-green, strap-shaped leaves 30cm long arising from the base...
is distributed from Peru to Bolivia, PyrolirionPyrolirion, commonly known as fire lilies or flame lilies, is a small genus of herbaceous flowering bulbs native to Chile, Peru, and Bolivia.-Description:...
(6 species) from Peru to Northern Chile, and Hieronymiella (8 species) ranges from Southern Bolivia to North Western Argentina.
- Hymenocallideae (3 genera, x = 23). Leptochiton
Leptochiton is an extinct of polyplacophoran mollusc. Leptochiton became extinct during the Pliocene period....
(2 species) and IsmeneIsmene, or Peruvian daffodil, is a genus of tender perennial bulbs bearing a strong resemblance to Hymenocallis, a genus into which Ismene had often been grouped in the past...
(10 species) are two strictly central Andean genera, whereas the third genus of the tribe, HymenocallisHymenocallis is a genus of plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains more than 60 species native to tropical and subtropical America. Hymenocallis are bulbous perennial herbs. The flowers have their stamens united to a characteristic corona...
(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 seedA seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
s of Hymenocallidae are made up of a thick outer integument formed by chlorenchyma with a well-developed vascular system, and an embryo which stores starch.
- Clinantheae (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
Pamianthe is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Pamianthe parviflora, Meerow...
(3 species) and Paramongaia (2 species) from Peru, Ecuador and Bolivia.
Hybrids
Several members of different genera of Amaryllidoideae hybridize readily, and the resulting hybrids are often sterile but can be propagated asexually. A
hybrid nameIn botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name. The ICBN provides the following options in dealing with a hybrid:...
is usually reserved for these
horticulturallyHorticulture is the industry and science of plant cultivation including the process of preparing soil for the planting of seeds, tubers, or cuttings. Horticulturists work and conduct research in the disciplines of plant propagation and cultivation, crop production, plant breeding and genetic...
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 Amaryllidoideae are:
- × Amarcrinum
× Amarcrinum is the scientific name applied to those hybrid plants obtained from the cross between the genera Amaryllis and Crinum...
CouttsCoutts & Co. is one of the UK's private banking houses, now wholly owned by the Royal Bank of Scotland . RBS acquired Coutts and all of its overseas subsidiaries when it bought NatWest. On 1 January 2008, Coutts' international businesses were renamed RBS Coutts, aligning them more closely with...
(syn.: × Crindonna; × Crinodonna; × Amarcrinaflora): is the result of crossing Amaryllis belladonna with species of the genus CrinumCrinum is a genus of about 180 species of perennial plants that have large showy flowers on leafless stems, and develop from bulbs. They are found along the sides of streams and lakes in tropical and subtropical areas worldwide, including South Africa....
.
- × Amarine Sealy
Sealy is a surname, and may refer to:* Allan Sealy, author nominated for the Booker Prize* Alison Sealy-Smith, Canadian actress* Derek Sealy, West Indian cricketer* Glenroy Sealy, Canadian cricketer...
, is the name applied to the hybrids between Amaryllis belladonna with members of NerineNerine is a genus of plants belonging to the Amaryllidaceae family, subfamily Amaryllidoideae Native to South Africa, there are about 30 different species in the genus. Nerine have been widely cultivated and much hybridized and are now spread world wide....
.
- × Hippeastrelia is the result of crossing species of Hippeastrum
Hippeastrum is a genus of about 90 species and 600+ hybrids and cultivars of bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to tropical and subtropical regions of the Americas from Argentina north to Mexico and the Caribbean. Some species are grown for their large...
with members of the genus SprekeliaSprekelia is a genus of at least three bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are native to Central America. Like Hippeastrum, these plants were known as Amaryllis. Sprekelia plants are sometimes called "Aztec lilies", although they are not true lilies...
.
Haemanthus × clarkei W. Wats., is the product of crossing
H. albiflos with
H. coccineus, it was raised in the UK by 1891.
Clivia cyrtanthifora was raised by Charles Raes
in Ghent, Belgium in the late 1850s and it was suggested to be a hybrid between
Clivia miniataClivia miniata is a species of clivia, from South Africa. It grows to a height of about 45cm in the shade of trees and shrubs, and flowers are red, orange or yellow, with a faint, but very sweet perfume...
and
Clivia nobilisClivia nobilis, also called Drooping clivia, is a plant belonging to the genus Clivia. It grows to about 38 cm of height. It has pendent flowers with scarlet red and some green.-External links:**...
, which was confirmed later by means of cytogenetics tools. 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.
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.
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.
Genera
- Acis
Acis is a genus of perennial, herbaceous and bulbous plants in the Amaryllis family . The genus consists in 9 species distributed in Europe and Northern Africa.The genus was reinstated in 2004, after it was determined on morphological and molecular grounds that these species should be segregated...
Salisb.Richard Anthony Salisbury FRS was a British botanist. While he is remembered as a valuable worker in horticultural and botanical sciences, several bitter disputes caused him to be ostracised by his contemporaries.-Life:...
- Amaryllis
Amaryllis is a small genus of flowering bulbs, with two species. The better known of the two, Amaryllis belladonna, is a native of South Africa, particularly the rocky southwest region near the Cape...
L.
- Ammocharis
Ammocharis is a genus in the Amaryllidaceae family which includes 6 species distributed in Africa. The plant grows as a succulent, above-ground bulb, preferring seasonally wet, hot, sandy soils and full sun.-Species:...
Herb. (Incl.: Cybistetes Milne-Redh. & Schweick.)
- Apodolirion
Apodolirion is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of 6 species distributed in South Africa...
BakerJohn Gilbert Baker was an English botanist.Baker was born in Guisborough, the son of John and Mary Baker and educated at Quaker schools in Ackworth and York....
- Boophone
Boophone is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of two species distributed in Tropical and Southern Africa...
Herb.
- Brunsvigia
Brunsvigia is a flowering plant genus in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains about 20 species native to South Africa....
Heist.Lorenz Heister was a German anatomist, surgeon and botanist born in Frankfurt am Main....
- Caliphruria
Caliphruria is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of four species distributed in tropical regions of South America, three of them are endemic to Colombia. This genus is closely related with the genera Eucharis and Urceolina, all of them known...
Herb.
- Calostemma
Calostemma is a small genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists in 3 species endemic to Australia, where they are distributed in arid regions with summer precipitations...
R.Br.Robert Brown was a Scottish botanist and palaeobotanist who made important contributions to botany largely through his pioneering use of the microscope...
- Chlidanthus
Chlidanthus is a genus that consists of 10 species of tender bulbs from tropical South America, mostly natives to the Andes. The botanical name comes from the Greek, meaning "delicate flower". The plants have large spherical bulbs with gray-green, strap-shaped leaves 30cm long arising from the base...
Herb. (incl.: Castellanoa Traub)
- Clinanthus Herb. (syn.: Anax Ravenna)
- Clivia
Clivia is a genus of monocot flowering plants native to southern Africa. They are from the family Amaryllidaceae, subfamily Amaryllidoideae. Common names include Kaffir lily and bush lily....
Lindl.John Lindley FRS was an English botanist, gardener and orchidologist.-Early years:Born in Catton, near Norwich, England, John Lindley was one of four children of George and Mary Lindley. George Lindley was a nurseryman and pomologist and ran a commercial nursery garden...
- Crinum
Crinum is a genus of about 180 species of perennial plants that have large showy flowers on leafless stems, and develop from bulbs. They are found along the sides of streams and lakes in tropical and subtropical areas worldwide, including South Africa....
L.
- Crossyne Salisb.
- Cryptostephanus Welw.
Friedrich Martin Josef Welwitsch was an Austrian explorer and botanist who in Angola discovered the plant Welwitschia mirabilis...
ex Baker
- Cyrtanthus
Cyrtanthus is a genus of perennial, herbaceous and bulbous plants in the Amaryllis family . This highly ornamental genus encompasses about 60 species...
AitonWilliam Aiton was a Scottish botanist.Aiton was born near Hamilton. Having been regularly trained to the profession of a gardener, he travelled to London in 1754, and became assistant to Philip Miller, then superintendent of the Chelsea Physic Garden...
(syn: Anoiganthus Baker; Vallota Salisb. ex Herb.)
- Eucharis
Eucharis is a genus of about 15-20 species of monocotyledonous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to Central America and South America from Guatemala south to Bolivia.-Description:...
Planch.Jules Émile Planchon was a French botanist born in Ganges, Hérault.-Biography:After receiving his Doctorate of Science at the University of Montpellier in 1844, he worked for a while at the Royal Botanical Gardens in London, and for a few years was a teacher in Nancy and Ghent...
& LindenJean Jules Linden , was a Belgian botanist and explorer, horticulturist and businessman, specialising in orchids, on which subject he wrote a number of books....
- Eucrosia
Eucrosia is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family distributed from Ecuador to Peru. The name is derived from the Greek eu, beautiful, and krossos, a fringe, referring to the long stamens. The genus contains eight species...
Ker Gawl.John Bellenden Ker, originally John Gawler was an English botanist born about 1764 in Ramridge, Andover, Hampshire and died in June 1842 in the same town. On 5 November 1804 he changed his name to Ker Bellenden, but continued to sign his name as Bellenden Ker until his death...
(syn: Callipsyche Herb.)
- Eustephia Cav.
- Galanthus L.
- Gethyllis L.
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...
(syn.: Klingia Schönl.)
- Griffinia
Griffinia is a genus of monocotyledonous plants in the Amaryllidaceae family, subfamily Amaryllidoideae. It includes 21 species which are endemic to South America, Brazil. The most closely related genus to it is the monotypic Worsleya. The members of the genus Griffinia are tropical, bulbous plants...
Ker Gawl.John Bellenden Ker, originally John Gawler was an English botanist born about 1764 in Ramridge, Andover, Hampshire and died in June 1842 in the same town. On 5 November 1804 he changed his name to Ker Bellenden, but continued to sign his name as Bellenden Ker until his death...
(incl. Hyline Herb. as a subgenus)
- Griffiniopsis Dutilh & Meerow
Alan W. Meerow is an American botanist, born in New York in 1952. He specializes in the taxonomy of the family Amaryllidaceae and the horticulture of palms and tropical ornamental plants....
(sin.: Eithea Ravenna)
- Habranthus
Habranthus is a genus of tender herbaceous flowering bulbs in the Amaryllidaceae, subfamily Amaryllidoideae. The genus was first identified by pioneering bulb enthusiast William Herbert in 1824.-Description:...
Herb. (sin.: Zephyranthella (Pax) Pax; Haylockia Herb.)
- Haemanthus
Haemanthus is a Southern African genus of Amaryllidaceae, subfamily Amaryllidoideae, with some 22 known species, endemic to South Africa, Namibia and the kingdoms of Lesotho and Swaziland...
L.
- Hannonia Braun-Blanq.
Josias Braun-Blanquet was an influential phytosociologist and botanist. Braun-Blanquet was born in Chur, Switzerland and died in Montpellier, France.-Phytosociology:...
& Maire
- Hessea Herb. (syn.: Kamiesbergia Snijman)
- Hieronymiella Pax
Ferdinand Albin Pax was a German botanist and entomologist. He specialised in Lepidoptera, Diptera, as well as in spermatophytes, describing several species....
(syn.: Eustephiopsis R.E.Fr. Robert Elias Fries , the youngest son of Thore M. Fries and grandson of Elias Magnus Friesand an expert on mushrooms...
)
- Hippeastrum
Hippeastrum is a genus of about 90 species and 600+ hybrids and cultivars of bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to tropical and subtropical regions of the Americas from Argentina north to Mexico and the Caribbean. Some species are grown for their large...
Herb. (syn: Moldenkea Traub)
- Hymenocallis
Hymenocallis is a genus of plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains more than 60 species native to tropical and subtropical America. Hymenocallis are bulbous perennial herbs. The flowers have their stamens united to a characteristic corona...
Salisb.
- Ismene
Ismene, or Peruvian daffodil, is a genus of tender perennial bulbs bearing a strong resemblance to Hymenocallis, a genus into which Ismene had often been grouped in the past...
Salisb. ex Herb (incl. Elisena Herb. and Pseudostenomesson Velarde as subgenera)
- Lapiedra Lag.
Mariano la Gasca y Segura was a Spanish botanist, writer and doctor. He was the director of Royal Botanical Garden of Madrid.- Early life :...
- Leptochiton Sealy
- Leucojum L.
- Lycoris Herb.
- Mathieua Klotzsch
Johann Friedrich Klotzsch was a German pharmacist and botanist.His principal work was in the field of mycology, with the study and description of many species of mushroom.-Selected works:...
- Namaquanula D.Müll.-Doblies
Dietrich Muller-Doblies is a German systematic botanist. His main areas of interest are the Bryophyta, the Spermatophyta, the Monocotyledons, Amaryllidaceae, Colchicaceae and Hyacinthaceae. He is currently at the Herbarium of the Technische Universität Berlin -References:...
& U.Müll.-Doblies
- Narcissus L. (incl.: Braxireon Raf. and Tapeinanthus Herb.)
- Nerine
Nerine is a genus of plants belonging to the Amaryllidaceae family, subfamily Amaryllidoideae Native to South Africa, there are about 30 different species in the genus. Nerine have been widely cultivated and much hybridized and are now spread world wide....
Herb.
- Pamianthe
Pamianthe is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Pamianthe parviflora, Meerow...
StapfOtto Stapf FRS was an Austrian born botanist and taxonomist.Stapf trained in Vienna, moving to Royal Botanic Gardens, Kew in 1890. He was keeper of the Herbarium from 1909 to 1920...
- Pancratium
Pancratium is a genus of about 21 species of perennial, herbaceous and bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, which latter also includes the genera Narcissus and Galanthus . The genus is found along the coastline of the Mediterranean area extending to the Canary...
L. (syn.: Mizonia A.Chev.; Chapmanolirion Dinter )
- Paramongaia Velarde
- Phaedranassa
Phaedranassa is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Phaedranassa brevifolia, Meerow* Phaedranassa cinerea, Ravenna...
Herb. (sin: Neostricklandia Rauschert; Stricklandia Baker)
- Phycella Lyndl. (incl.:Famatina Ravenna)
- Placea
Placea is a genus of about 6 species of herbaceous, perennial and bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are endemic to Chile.-List of species:...
MiersJohn Miers, FRS FLS, knight grand cross of the Order of the Rose, was a British botanist and engineer, best known for his work on the flora of Chile and Argentina....
- Plagiolirion Baker
- Proiphys
Proiphys is a genus of herbaceous, perennial and bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It consists of 4 species that are native to Southeast Asia and Australia.-Description:...
Herb. (syn.: EuryclesProiphys is a genus of herbaceous, perennial and bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It consists of 4 species that are native to Southeast Asia and Australia.-Description:...
Salisb. ex Schult. & Schult.)
- Pyrolirion
Pyrolirion, commonly known as fire lilies or flame lilies, is a small genus of herbaceous flowering bulbs native to Chile, Peru, and Bolivia.-Description:...
Herb.
- Rauhia Traub
- Rhodophiala
Rhodophiala is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of 28 South American species distributed in southern Brazil, Argentina, and, specially, in Chile....
C.PreslKarel Bořivoj Presl was a Bohemian botanist.He lived all his life in Prague, and was a professor at the University of Prague. He made an expedition to Sicily in 1817, and published a Flora bohemica in 1820....
(syn.: Rhodolirium Phil.Rodolfo Amando Philippi was a German-Chilean paleontologist and zoologist....
; Rhodolirion Dalla Torre & Harms)
- Scadoxus
Scadoxus is a genus of 9 species native to tropical Africa in the family Amaryllidaceae, subfamily Amaryllidoideae. The genus has close affinities with Haemanthus from which it has only recently been separated.Species in the genus are...
Raf. (syn: Choananthus Rendle)
- Sprekelia
Sprekelia is a genus of at least three bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae. They are native to Central America. Like Hippeastrum, these plants were known as Amaryllis. Sprekelia plants are sometimes called "Aztec lilies", although they are not true lilies...
Heist.Lorenz Heister was a German anatomist, surgeon and botanist born in Frankfurt am Main....
- Stenomesson Herb. (syn: Anax Ravenna; Callithauma Herb.; Crocopsis Pax, Pucara Ravenna)
- Sternbergia
Sternbergia is a genus in the family Amaryllidaceae, subfamily Amaryllidoideae, of around 8 species that show a broad distribution throughout Mediterranean Europe and Asia. It was first described by Clusius in 1601 as Narcissus, before being redescribed by Carl Linnaeus as Amaryllis in 1753...
Waldst.Franz de Paula Adam Norbert Wenzel Ludwig Valentin von Waldstein was an Austrian soldier, explorer and naturalist....
& Kit.Pál Kitaibel was a Hungarian botanist and chemist.He was born at Mattersburg and studied botany and chemistry at the undersity of Buda. He became professor taught these subjects at Pest in 1794...
- Strumaria
Strumaria is a genus of plant in family Amaryllidaceae, subfamily Amaryllidoideae. It contains the following species :* Strumaria barbarae, Oberm.* Strumaria hardyana, D.& U. Müll.-Doblies...
Jacq.Nikolaus Joseph Freiherr von Jacquin or Baron Nikolaus von Jacquin. was a scientist who studied medicine, chemistry and botany....
ex Willd.Carl Ludwig Willdenow was a German botanist, pharmacist, and plant taxonomist. He is considered one of the founders of phytogeography, the study of the geographic distribution of plants...
(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.
Heinrich Gottlieb Ludwig Reichenbach was a German botanist and ornithologist.He was the son of Johann Friedrich Jakob Reichenbach, the author in 1818 of the first Greek-German dictionary. He was the father of Heinrich Gustav Reichenbach, equally a botanist and an eminent orchid...
(sin: Collania Schult.Josef August Schultes 1773-1831 was an Austrian botanist and professor in Vienna. Together with Johann Jacob Roemer, he published the 16th edition of Linnaeus' Systema Vegetabilium. In 1821, he was elected a foreign member of the Royal Swedish Academy of Sciences.Father of Julius Hermann...
& Schult.f. Julius Hermann Schultes was an Austrian botanist in Vienna. He co-authored volume 7 of the Roemer & Schultes edition of the Systema Vegetabilium with his father Josef August Schultes.-Notes:...
; Pseudourceolina Vargas)
- Vagaria Herb.
- Worsleya
The genus Worsleya contains only one species, Worsleya procera, previously known as Worsleya rayneri. It is one of the largest and rarest members of the subfamily Amaryllidoideae . Worsleya is a tropical plant. This species is also known as the empress of Brazil because of its origin in South...
(W.WatsonWilliam Watson may refer to:*W. Marvin Watson , U.S. Postmaster General*William E. Watson, military historian*William H. Watson , Mexican-American War soldier from Maryland*William J. Watson...
ex Traub) Traub
- Zephyranthes
Zephyranthes is a genus of 71 species in the Amaryllis family . There are numerous hybrids and cultivars. Common names for species in this genus include fairy lily, rainflower, zephyr lily, magic lily, Atamasco lily, and rain lily.The name is derived from Ζέφυρος , the Greek god of the west...
Herb. (syn: CooperiaCooperia is a genus of tender herbaceous perennials native to South America and the southern reaches of North America. They are closely related to Zephyranthes and Habranthus, all members of the Amaryllidaceae, or Amaryllis family. All three genera are commonly known as rain lilies because of...
Herb.)
Ecology
Most species are adapted to
seasonA season is a division of the year, marked by changes in weather, ecology, and hours of daylight.Seasons result from the yearly revolution of the Earth around the Sun and the tilt of the Earth's axis relative to the plane of revolution...
al 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
deciduousDeciduous means "falling off at maturity" or "tending to fall off", and is typically used in reference to trees or shrubs that lose their leaves seasonally, and to the shedding of other plant structures such as petals after flowering or fruit when ripe...
. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist
fynbosFynbos is the natural shrubland or heathland vegetation occurring in a small belt of the Western Cape of South Africa, mainly in winter rainfall coastal and mountainous areas with a Mediterranean climate...
. The aboveground parts (leaves and stems) of deciduous species die down when the bulb or corm enters
dormancyDormancy is a period in an organism's life cycle when growth, development, and physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be closely associated with environmental conditions...
. The plants thus survive periods that are unfavourable for growth by retreating underground. This is particularly useful in
grasslandGrasslands are areas where the vegetation is dominated by grasses and other herbaceous plants . However, sedge and rush families can also be found. Grasslands occur naturally on all continents except Antarctica...
s 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 Amaryllidoideae are chiefly adapted for seasonally dry habitats and some prefer truly xeric environments in which their bulbs may remain
dormantDormant means lacking activity. It can refer to:*Dormancy in an organism's life cycle*Dormant volcano, a volcano that is inactive but may become active in the future...
for a period longer than they are in active growth (e.g.,
LeptochitonLeptochiton is an extinct of polyplacophoran mollusc. Leptochiton became extinct during the Pliocene period....
, Paramongaia, some
EucrosiaEucrosia is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family distributed from Ecuador to Peru. The name is derived from the Greek eu, beautiful, and krossos, a fringe, referring to the long stamens. The genus contains eight species...
). At the other extreme, species have colonized the understory of rain forests
(Eucharis-Organisms:* Eucharis , a genus of ants* Eucharis , a genus of monocotyledons* a lapsus for the bee genus Epicharis* an invalid name for the bivalves genus Basterotia* an invalid name for the ctenophore genus Cabira...
, GriffiniaGriffinia is a genus of monocotyledonous plants in the Amaryllidaceae family, subfamily Amaryllidoideae. It includes 21 species which are endemic to South America, Brazil. The most closely related genus to it is the monotypic Worsleya. The members of the genus Griffinia are tropical, bulbous plants...
) and aquatic habitats (a number of
HymenocallisHymenocallis is a genus of plants in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains more than 60 species native to tropical and subtropical America. Hymenocallis are bulbous perennial herbs. The flowers have their stamens united to a characteristic corona...
,
Hippeastrum angustifolium, CrinumCrinum is a genus of about 180 species of perennial plants that have large showy flowers on leafless stems, and develop from bulbs. They are found along the sides of streams and lakes in tropical and subtropical areas worldwide, including South Africa....
). The subfamily 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.
VeldThe term Veld refers primarily to the wide open rural spaces of South Africa or southern Africa and in particular to certain flatter areas or districts covered in grass or low scrub...
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 Amaryllidoideae 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
CyrtanthusCyrtanthus is a genus of perennial, herbaceous and bulbous plants in the Amaryllis family . This highly ornamental genus encompasses about 60 species...
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. At least in
C. ventricosus, this dependency on fire for initiating the flowering process is regulated by the smoke. Aspects of the life history of
Haemanthus pubescensHaemanthus pubescens is an endemic South African bulbous geophyte in the genus Haemanthus....
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.
Reproductive biology
Members of Amaryllidoideae possess extraordinary diversity in reproductive traits, even among closely related species. The floral diversification that has accompanied the coevolution of flowers and animal pollinators is particularly striking.
HeterostylyHeterostyly is a unique form of polymorphism and herkogamy in flowers. In a heterostylous species, two or three different morphological types of flowers, termed morphs, exist in the population. On each individual plant, all flowers share the same morph. The flower morphs differ in the lengths of...
is a genetically controlled floral polymorphism; distyly and tristyly are characterized by two or three morphs, respectively, that differ in the style length of the carpel and the filament length of the
stamenThe stamen is the pollen producing reproductive organ of a flower...
. Pollen grain size and production, and the size of the stigmatic papillae and the corolla also vary. The positions of the stigma and anthers in the different morphs are such that pollinators contact same-level but different floral organs with the same region of their body. Thus pollination occurs and seeds are produced only in intermorph crosses. This is reciprocal
herkogamyHerkogamy is a common strategy employed by hermaphroditic angiosperms to reduce sexual interference between male and female function. Herkogamy differs from other such strategies by supplying a spatial separation of the anthers and stigma.Two forms of herkogamy are most common:1...
, a self-incompatibility system. A physiological incompatibility system is also present in distylous species that contributes to an efficient mechanism that avoids selfing and maximizes male and female fitness.
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
LepidopteraLepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
) 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.
ZephyranthesZephyranthes is a genus of 71 species in the Amaryllis family . There are numerous hybrids and cultivars. Common names for species in this genus include fairy lily, rainflower, zephyr lily, magic lily, Atamasco lily, and rain lily.The name is derived from Ζέφυρος , the Greek god of the west...
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 atamasco 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. In contrast, it was documented a process of asexual seed production (
apomixisIn botany, apomixis was defined by Winkler as replacement of the normal sexual reproduction by asexual reproduction, without fertilization. This definition notably does not mention meiosis...
) in
Zephyranthes texensis.
Zephyranthes 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,
polyploidyPolyploid is a term used to describe cells and organisms containing more than two paired sets of chromosomes. Most eukaryotic species are diploid, meaning they have two sets of chromosomes — one set inherited from each parent. However polyploidy is found in some organisms and is especially common...
and vegetative multiplication/apomixis explains the origin and preservation of an astonishing range in chromosome numbers from 2n=18 to 96 in this genus.
Especially in
CyrtanthusCyrtanthus is a genus of perennial, herbaceous and bulbous plants in the Amaryllis family . This highly ornamental genus encompasses about 60 species...
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. A total of 22 species of southern African genera
Crinum,
Cyrtanthus and
Pancratium conform to the syndrome of "sphingophily", attracting
SphingidaeSphingidae is a family of moths , commonly known as hawk moths, sphinx moths and hornworms, that includes about 1,200 species . It is best represented in the tropics but there are species in every region . They are moderate to large in size and are distinguished among moths for their rapid,...
moths, 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.
Southern Africa also has several Amaryllidoideae with remarkable dispersal abilities. Species of
BrunsvigiaBrunsvigia is a flowering plant genus in the family Amaryllidaceae, subfamily Amaryllidoideae. It contains about 20 species native to South Africa....
, BoophoneBoophone is a genus of herbaceous, perennial and bulbous plants in the Amaryllis family . It consists of two species distributed in Tropical and Southern Africa...
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 Amaryllidoideae 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
PeruPeru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....
, the Inca people frequently depicted flowers of Amaryllidaceae
(IsmeneIsmene is the name of two women of Greek mythology. The more famous is a daughter and half-sister of Oedipus, daughter and granddaughter of Jocasta, and sister of Antigone, Eteocles, and Polynices. She appears in several plays of Sophocles: at the end of Oedipus the King, in Oedipus at Colonus and...
, 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
LesothoLesotho , officially the Kingdom of Lesotho, is a landlocked country and enclave, surrounded by the Republic of South Africa. It is just over in size with a population of approximately 2,067,000. Its capital and largest city is Maseru. Lesotho is a member of the Commonwealth of Nations. The name...
probably emphasizes how much the San people valued the bulbs for their psychoactive effects.
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
AmaryllisAmaryllis is a small genus of flowering bulbs, with two species. The better known of the two, Amaryllis belladonna, is a native of South Africa, particularly the rocky southwest region near the Cape...
, CliviaClivia is a genus of monocot flowering plants native to southern Africa. They are from the family Amaryllidaceae, subfamily Amaryllidoideae. Common names include Kaffir lily and bush lily....
, HippeastrumHippeastrum is a genus of about 90 species and 600+ hybrids and cultivars of bulbous plants in the family Amaryllidaceae, subfamily Amaryllidoideae, native to tropical and subtropical regions of the Americas from Argentina north to Mexico and the Caribbean. Some species are grown for their large...
, NerineNerine is a genus of plants belonging to the Amaryllidaceae family, subfamily Amaryllidoideae Native to South Africa, there are about 30 different species in the genus. Nerine have been widely cultivated and much hybridized and are now spread world wide....
, and
ZephyranthesZephyranthes is a genus of 71 species in the Amaryllis family . There are numerous hybrids and cultivars. Common names for species in this genus include fairy lily, rainflower, zephyr lily, magic lily, Atamasco lily, and rain lily.The name is derived from Ζέφυρος , the Greek god of the west...
are the most popular choices for gardens and containers.
Conservation status
Habitat loss is currently the greatest threat to the Amaryllidoideae in South Africa, where 59 species are endangered or vulnerable and 58 species are near threatened.
External links