Scilloideae (named after the genus ''
Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and sea ...
'', "squill") is a
subfamily of
bulbous plants within the family ''
Asparagaceae''. Scilloideae is sometimes treated as a separate family Hyacinthaceae, named after the genus ''
Hyacinthus''. Scilloideae or Hyacinthaceae include many familiar garden plants such as ''
Hyacinth
Hyacinth or Hyacinthus may refer to:
Nature Plants
* Hyacinth (plant), genus ''Hyacinthus''
** '' Hyacinthus orientalis'', common hyacinth
* Grape hyacinth, '' Muscari'', a genus of perennial bulbous plants native to Eurasia
* Hyacinth bean, ''L ...
us'' (hyacinths), ''
Hyacinthoides'' (bluebells), ''
Muscari'' (grape hyacinths) and ''
Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and sea ...
'' and ''
Puschkinia
''Puschkinia'' is a genus of four known species of bulbous perennials in the family Asparagaceae, subfamily Scilloideae. It is native to the Caucasus and the Middle East. ''Puschkinia scilloides'' is grown as an ornamental bulbous plant.
Descri ...
'' (squills or scillas). Some are important as
cut flowers
Cut flowers are flowers or flower buds (often with some stem and leaf) that have been cut from the plant bearing it. It is usually removed from the plant for decorative use. Typical uses are in vase displays, wreaths and garlands. Many garde ...
.
Scilloideae are distributed mostly in
Mediterranean climate
A Mediterranean climate (also called a dry summer temperate climate ''Cs'') is a temperate climate sub-type, generally characterized by warm, dry summers and mild, fairly wet winters; these weather conditions are typically experienced in the ...
s, including
South Africa
South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. It is bounded to the south by of coastline that stretch along the Atlantic Ocean, South Atlantic and Indian Oceans; to the ...
,
Central Asia
Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
and
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
. Their flowers have six tepals and six stamens with a
superior ovary
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule(s) and is located above or below or at the point of connection with the ...
, which previously placed them within the
lily family (Liliaceae), and their leaves are fleshy,
mucilaginous
Mucilage is a thick, gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion. The direction of their movement is always opposite to that of the secretion of ...
, and arranged in a
basal rosette
In botany, a rosette is a circular arrangement of leaves or of structures resembling leaves.
In flowering plants, rosettes usually sit near the soil. Their structure is an example of a modified stem in which the internode gaps between the lea ...
.
The Scilloideae, like most
lily-like monocots, were at one time placed in a very broadly defined lily family (Liliaceae). The subfamily is recognized in modern classification systems such as the
APG III system of 2009. It is also treated as the separate family Hyacinthaceae, as it is by many researchers and was in earlier APG systems. Determining the boundaries between genera within the Scilloideae is an active area of research. The number of genera varies widely from source to source, from about 30 to about 70. The situation has been described as being in a "state of flux".
[
]
Description
The subfamily contains many popular spring-flowering garden bulbs, such as hyacinth
Hyacinth or Hyacinthus may refer to:
Nature Plants
* Hyacinth (plant), genus ''Hyacinthus''
** '' Hyacinthus orientalis'', common hyacinth
* Grape hyacinth, '' Muscari'', a genus of perennial bulbous plants native to Eurasia
* Hyacinth bean, ''L ...
s (''Hyacinthus''), grape hyacinths ('' Muscari''), bluebells ('' Hyacinthoides'') and squills (''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and sea ...
''). Other members are summer- and autumn-flowering, including ''Galtonia
''Galtonia'' is a genus of plants in the family Asparagaceae, subfamily Scilloideae. Native to Southern Africa, the genus is named after Sir Francis Galton. According to some authorities it has been subsumed into '' Ornithogalum'' as a subgenu ...
'' and ''Eucomis
''Eucomis'' is a genus of flowering plants in the family Asparagaceae, subfamily Scilloideae, native to southern Africa. Most species of this genus are commonly referred to as pineapple flowers or pineapple lilies. They are bulbous perennials wit ...
'' ('pineapple lilies'). Most are native to Mediterranean climate
A Mediterranean climate (also called a dry summer temperate climate ''Cs'') is a temperate climate sub-type, generally characterized by warm, dry summers and mild, fairly wet winters; these weather conditions are typically experienced in the ...
zones and neighboring areas in the Mediterranean Basin and South Africa
South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. It is bounded to the south by of coastline that stretch along the Atlantic Ocean, South Atlantic and Indian Oceans; to the ...
. Others are found in Central Asia
Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
, the Far East
The ''Far East'' was a European term to refer to the geographical regions that includes East and Southeast Asia as well as the Russian Far East to a lesser extent. South Asia is sometimes also included for economic and cultural reasons.
The ter ...
and South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
.
Morphologically the subfamily is characterised by having 6 tepals
A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
and 6 stamens
The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filame ...
with a superior ovary, a characteristic which placed them within the older order of Liliales in many older classification systems, such as the Cronquist system
The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) ...
, but they now separate from them within the Asparagales
Asparagales (asparagoid lilies) is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in t ...
order. They have also been included in the family Liliaceae
The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fair a ...
.
Roots: contractile and mucilaginous.
Leaves: fleshy and mucilaginous arranged in a basal rosette, alternate and spiral, simple, margin entire, with parallel venation, sheathing at the base, without stipules and hair simple.
Flowers: arranged in scapiflorous inflorescences (in racemes, in spikes, and in heads). The peduncles are articulated. The flowers are hermaphroditic
In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes.
Many taxonomic groups of animals (mostly invertebrates) do not have s ...
, actinomorphic
Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts.
Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
, often showy.
Perianths: six tepals divided into two whorls, free or joined ( connate). When joined, the perianth forms a tubular bell. The tepals are imbricate
Aestivation or estivation is the positional arrangement of the parts of a flower within a flower bud before it has opened. Aestivation is also sometimes referred to as praefoliation or prefoliation, but these terms may also mean vernation: the ar ...
and petaloid
Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corolla''. Petals are usually ...
. The corolla may be white, yellow, violet, blue, brown and even black (see images).
Androecium
The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the fila ...
: composed of 6 stamens (exceptionally 3, as in Albuca
''Albuca'' is a genus of flowering plants in the family Asparagaceae, subfamily Scilloideae. The genus is distributed mainly in southern and eastern Africa, with some species occurring in northern Africa and the Arabian Peninsula. Plants of the g ...
, for example), with the filaments free or adnate
Adnate may refer to:
* Adnation, in botany, the fusion of two or more whorls of a flower
* Adnate, in mycology, a classification of lamellae (gills)
* Conjoined twins
Conjoined twins – sometimes popularly referred to as Siamese twins – are ...
to the tube, often appendiculate. The anthers are dorsifixed and pollen dehiscence occurs by longitudinal openings. The pollen is monosulcate (having a linear furrow).
Gynoecium
Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils' ...
: superior ovary, tricarpelate, connate and trilocular. Single stigma, capitate to 3-lobed. May contain from one to several ovules in each locule. They have nectaries
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
at the septa of the ovaries.
Fruit
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
: dehiscence loculicidal.
Seed
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
: Seed morphology is diverse, from globular to flattened, and occasionally aril. The seed coat usually contains phytomelan (phytomelanin), one of the defining characteristics of the order, a black pigment present in the seed coat, creating a dark crust.
Chromosomes
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
: Chromosome size varies widely, from 1.2 to 18 µm in length, karyotype bimodal or trimodal. The basic chromosome number is also very variable (X = 2, 6, 7, 10, 15, 17, etc.).[
]
Systematics
When treated as a subfamily, the name Scilloideae is derived from the generic name of the type genus
In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name.
Zoological nomenclature
According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal ...
, ''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and sea ...
'', and is attributed to Gilbert Thomas Burnett in 1835.[ When treated as a family, the name Hyacinthaceae is derived from the type genus '' Hyacinthus'', and is usually attributed to ]August Batsch
August Johann Georg Karl Batsch (28 October 1761 – 29 September 1802) was a German naturalist. He was a recognised authority on mushrooms, and also described new species of ferns, bryophytes, and seed plants.
Life and career
Batsch was born ...
from ("ex") a 1797 publication by Moritz Borkhausen.[
]
Phylogeny
The monophyly
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
of Scilloideae is well supported by studies based on molecular data.[ These studies also give support to the exclusion of '']Camassia
''Camassia'' is a genus of plants in the asparagus family native to North America. Common names include camas, quamash, Indian hyacinth, camash, and wild hyacinth.
It grows in the wild in great numbers in moist meadows. They are perennial p ...
'', ''Chlorogalum
The common names soap plant, soaproot and amole refer to the genus ''Chlorogalum''. They are native to western North America, with some species in Oregon but they are mostly found in California. Common names of the genus and several species ...
'' and related genera, i.e. the former Hyacinthaceae subfamily Chlorogaloideae, now placed in the subfamily Agavoideae
Agavoideae is a subfamily of monocot flowering plants in the family Asparagaceae, order Asparagales. It has previously been treated as a separate family, Agavaceae. The group includes many well-known desert and dry-zone types, such as the agaves ...
.[
The exact position of the Scilloideae within the broadly defined Asparagaceae is less clear. One possible phylogeny for the seven subfamilies recognised within the family is shown below.][
Although generally agreeing on the main division of the Asparagaceae into two clades, studies have produced slightly different relationships among the Agavoideae, Aphyllanthoideae, Brodiaeoideae and Scilloideae. For example, Seberg et al. (2012) present analyses based on parsimony and on maximum likelihood. In the first, the Scilloideae are sister to the Agavoideae; in the second, they are sister to the Brodiaeoideae.][
]
Early classifications
Detailed historical accounts of taxonomic issues relating to the modern subfamily Scilloideae have been provided by Pfosser & Speta (1999)[ and Chase et al. (2009).][ The lilioid monocots have long created classification problems. At one extreme, e.g. in the ]Cronquist system
The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) ...
of 1968, they have been regarded as one large family (Liliaceae
The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fair a ...
''sensu lato
''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
''). At the other extreme, e.g. in the Dahlgren system
One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 19 ...
of 1985, they have been divided between orders and split into many often small families. Dahlgren divided the lilioid monocots in search of monophyly
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, but in practice he was unsuccessful. His major contribution was to split the Liliaceae into two families, the true Liliaceae, Liliaceae ''sensu stricto'', and the Hyacinthaceae (families which are now placed in separate orders, Liliales and Asparagales
Asparagales (asparagoid lilies) is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in t ...
).
Splitting off the Hyacinthaceae from the Liliaceae was originally suggested by Batsch
August Johann Georg Karl Batsch (28 October 1761 – 29 September 1802) was a German naturalist. He was a recognised authority on mushrooms, and also described new species of ferns, bryophytes, and seed plants.
Life and career
Batsch was born i ...
in 1786.[ Batsch's version of the family only superficially resembles the modern version, but did include '']Hyacinth
Hyacinth or Hyacinthus may refer to:
Nature Plants
* Hyacinth (plant), genus ''Hyacinthus''
** '' Hyacinthus orientalis'', common hyacinth
* Grape hyacinth, '' Muscari'', a genus of perennial bulbous plants native to Eurasia
* Hyacinth bean, ''L ...
us'' and ''Lachenalia
''Lachenalia'' is a genus of bulbous perennial plants in the family Asparagaceae, subfamily Scilloideae, which are usually found in Namibia and South Africa. Most of them have a dormancy period, but new roots will always grow every year.
''Lache ...
''. The group was reduced to a tribe by Endlicher
Stephan Ladislaus Endlicher also known as Endlicher István László (24 June 1804, Pressburg, Bratislava (Pozsony) – 28 March 1849, Vienna) was an Austrian Empire, Austrian botanist, numismatist and Sinologist. He was a director of the Botan ...
in 1836, and included ''Camassia
''Camassia'' is a genus of plants in the asparagus family native to North America. Common names include camas, quamash, Indian hyacinth, camash, and wild hyacinth.
It grows in the wild in great numbers in moist meadows. They are perennial p ...
''. In 1866 Salisbury
Salisbury ( ) is a cathedral city in Wiltshire, England with a population of 41,820, at the confluence of the rivers Avon, Nadder and Bourne. The city is approximately from Southampton and from Bath.
Salisbury is in the southeast of ...
redistributed the genera into several families.[
In the 1870s, Baker used tribes to divide up the Liliaceae '' s.l.''. introducing the Hyacintheae, Scilleae, Massonieae, and Chlorogaleae.][ In 1887 Engler divided the Liliaceae ''s.l.'' into two tribes, Lilieaoe and Scilleae.][ In the twentieth century, Fritsch proposed the division of Liliaceae ''s.l.'' into smaller more homogeneous families.][ In the 1930s the Viennese school elevated Engler's tribes to subfamilies.][ They questioned the inclusion of such different groups as Lilioideae and Scilloideae within the same family, and even Scilloideae was considered to be composed of at least three groups.][ By 1969, ]Huber
Huber is a German-language surname. It derives from the German word ''Hube'' meaning hide, a unit of land a farmer might possess, granting them the status of a free tenant. It is in the top ten most common surnames in the German-speaking world, ...
was recognizing the Scilloideae as the family Hyacinthaceae, and dividing it into tribes.[ How many tribes were recognised and how the genera were distributed within those tribes depended on the diagnostic characters chosen. Huber used seeds, while Schulze in 1980 used pollen.][ Morphology and chromosome analysis were supplemented by chemotaxonomy, due to the presence of cardiac steroids, such as the bufadienolids in the Urgineoideae and cardenolids in Ornithogaloideae. Even Linnaean genera such as ''Hyacinthus'', ''Scilla'' and ''Ornithoglum'' proved heterogeneous and characters useful in other families failed to define satisfactory taxa.
]
Modern classifications
Modern classification systems for plants are largely derived from molecular phylogenetic analysis. The initial molecular analysis of the Liliaceae ''s.l.'' was based on the Dahlgren system, as for example in the work by Chase et al. in 1995.[ When it was discovered that the Dahlgren families were not monophyletic, the tendency was to create new families out of each identified clade, as in the first ]Angiosperm Phylogeny Group
The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disc ...
system of 1998, the APG system
The APG system (Angiosperm Phylogeny Group system) of plant classification is the first version of a modern, mostly molecular-based, system of plant taxonomy. Published in 1998 by the Angiosperm Phylogeny Group, it was replaced by the improved A ...
. This placed many lilioid families and genera in the order Asparagales
Asparagales (asparagoid lilies) is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in t ...
(a term derived from Dahlgren, and the largest monocot order). One of the 29 families into which the Asparagales were divided was the Hyacinthaceae.[
With further work it was evident that these 29 families, some of which had few genera, could be grouped into larger clades. The ]APG II system
The APG II system (Angiosperm Phylogeny Group 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 Gr ...
of 2003 was a compromise. It divided the Asparagales into 14 broadly defined families, while allowing an alternative system in which some of the larger families could be replaced by smaller ones. The Hyacinthaceae was one of these optional smaller families, which could alternatively be sunk into a broadly defined Asparagaceae.[
This compromise approach was abandoned in the APG III system of 2009, which allowed only the broader families. The paper presenting the system states "The area around Asparagaceae is difficult from the standpoint of circumscription. Although Asparagaceae s.l. are heterogeneous and poorly characterized, Asparagaceae s.s., Agavaceae, Laxmanniaceae, Ruscaceae and even Hyacinthaceae have few if any distinctive features."] At the same time, Chase et al. provided subfamilies to replace the alternative narrowly defined families of APG II. The Hyacinthaceae became the subfamily Scilloideae of the family Asparagaceae.[
Many sources have adopted the APG III system; for example, the ]World Checklist of Selected Plant Families
The World Checklist of Selected Plant Families (usually abbreviated to WCSP) is an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected pla ...
places genera such as ''Hyacinthus'' only in the broadly defined Asparagaceae. Other sources prefer to retain the narrower families of APG II; for example, Seberg et al. say that it "remains a moot point whether the difficult-to-recognize bracketed families of APG II are a worse or a better choice than the equally difficult-to-recognize subfamilies of APG III", and in their analyses of the phylogeny of the Asparagales they continue to use families such as Hyacinthaceae.[
]
Tribes
In 1990, Pfosser and Speta stated that their earlier classification of the Hyacinthaceae into the subfamilies Hyacinthoideae, Ornithogaloideae, Oziroeoideae and Urgineoideae continued to be supported by ongoing studies. (They further divided the subfamilies Hyacinthoideae and Ornithogaloideae into tribes.)[ A part of reducing the Hyacinthaceae to the subfamily Scilloideae, Chase et al. (2009) suggested dividing it into four tribes, corresponding to Pfosser and Speta's four subfamilies: Hyacintheae Dumort., Ornithogaleae Rouy, Oziroëeae M.W.Chase, Reveal & M.F.Fay and Urgineeae Rouy.][ The possible relationship of the four tribes is represented in the following ]cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
,[ which has, however, only "moderate" statistical support.][
The exact boundaries between genera within these tribes remains controversial;][ the situation has been described as being in a "state of flux".][
]
Oziroëeae
Species are found only in western South America. They have flowers with stamens which are joined to the petals, rounded seeds and the embryo as long as the seed. The basic chromosome numbers are ''n'' = 15, 17. The tribe contains only the genus '' Oziroë''.[
]
Ornithogaleae
In terms of the number of species, this is the largest tribe. Its species are distributed in Europe, western Asia and Africa. They have flowers with three stamens which have flattened filaments. Their seeds are flattened and angular. The basic chromosome numbers range from ''n'' = 2 to ''n'' = 10.[ In the treatment by Manning et al. (2009) and Stevens at the ]Angiosperm Phylogeny Website
The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy.
The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis ...
, the tribe contains four genera, ''Albuca
''Albuca'' is a genus of flowering plants in the family Asparagaceae, subfamily Scilloideae. The genus is distributed mainly in southern and eastern Africa, with some species occurring in northern Africa and the Arabian Peninsula. Plants of the g ...
'' (about 110–140 species), ''Dipcadi
''Dipcadi'' is a genus of bulbous flowering plants in the family Asparagaceae, subfamily Scilloideae (also treated as the family Hyacinthaceae).Stevens, P.F. (2001 onwards),Asparagales: Scilloideae, ''Angiosperm Phylogeny Website'', retrieved 2 ...
'', ''Ornithogalum
''Ornithogalum'' is a genus of perennial plants mostly native to southern Europe and southern Africa belonging to the family Asparagaceae. Some species are native to other areas such as the Caucasus. Growing from a bulb, species have linear bas ...
'' (about 160 species, including ''Galtonia'' and ''Neopatersonia'') and '' Pseudogaltonia''.[ By contrast, Martínez-Azorín et al. (2011) divide the tribe into 19 genera.][
]
Urgineeae
Species within this tribe contain bufadienolides and are distributed mainly in Africa, Madagascar, and the Mediterranean through to India. The seeds are flattened and winged with the head barely attached to the endosperm. The basic chromosome numbers are ''n'' = 6, 7 and 10.[ Depending on the source, the tribe may include the genera '' Bowiea'', '']Drimia
''Drimia'' is a genus of flowering plants. In the APG IV classification system, it is placed in the family Asparagaceae, subfamily Scilloideae (formerly the family Hyacinthaceae). When broadly circumscribed, the genus includes a number of other ...
'' (including ''Urginea''), '' Schizobasis'' (sometimes included in ''Drimia'') and '' Fusifilum'' (also sometimes included in ''Drimia'').[
]
Hyacintheae
In terms of the number of species, this is the second largest tribe. Its species have leaves with pustules or spots, rounded seeds and contain homoisoflavanone
Homoisoflavonoids (3-benzylidenechroman-4-ones) are a type of phenolic compounds occurring naturally in plants.
Chemically, they have the general structure of a 16-carbon skeleton, which consists of two phenyl rings (A and B) and heterocyclic rin ...
s. The tribe can in turn be divided into three clades (subtribes):[
* '' Pseudoprospero'' Speta
: The only species in the genus, ''Pseudoprospero firmifolium'', is from eastern South Africa. It has two ovules per carpel with one seed per locule and a basic chromosome number ''n'' = 9.][
* Massoniinae Bentham & Hooker f.
: Species are distributed in Africa south of the Sahara and India. There are two or more ovules per carpel. The seeds have ]elaiosome
Elaiosomes ( grc, ἔλαιον ''élaion'' "oil" + ''sóma'' "body") 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 elaio ...
s. The basic chromosome number is 5 to 10+ (many 20).[ The subtribe contains about 13–20 genera (depending on the treatment), including '' Daubenya'', '' Drimiopsis'', '']Eucomis
''Eucomis'' is a genus of flowering plants in the family Asparagaceae, subfamily Scilloideae, native to southern Africa. Most species of this genus are commonly referred to as pineapple flowers or pineapple lilies. They are bulbous perennials wit ...
'', ''Lachenalia
''Lachenalia'' is a genus of bulbous perennial plants in the family Asparagaceae, subfamily Scilloideae, which are usually found in Namibia and South Africa. Most of them have a dormancy period, but new roots will always grow every year.
''Lache ...
'' (about 110 species), ''Ledebouria
''Ledebouria'' is a genus of African bulbous perennial herbs in the Asparagus family, Asparagaceae, subfamily Scilloideae. Most members were previously part of the genus ''Scilla''. A number of species are grown by cacti and succulent enthusi ...
'' (about 80 species), '' Massonia'' (including ''Whiteheadia''), '' Merwilla'', '' Schizocarphus'' and '' Veltheimia''.[
* Hyacinthinae Parlatore
: Species are distributed in Europe, the Mediterranean and North Africa and the Middle East, and then again in the Far East. There are two to eight ovules per carpel; elaiosomes are present in the seeds; and the basic chromosome number is 4 to 8+.][ The subtribe contains about 14–25 genera (depending on the treatment), including '' Bellevalia'' (about 50 species), '' Brimeura'', '' Hyacinthoides'', '' Muscari'' (about 50 species), '']Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and sea ...
'' (about 30 species) and ''Prospero
Prospero ( ) is a fictional character and the protagonist of William Shakespeare's play '' The Tempest''.
Prospero is the rightful Duke of Milan, whose usurping brother, Antonio, had put him (with his three-year-old daughter, Miranda) to se ...
'' (about 25 species).[
]
Genera and species
Some genera that were formerly placed within the Scillioideae (as Hyacinthaceae), e.g., ''Chlorogalum
The common names soap plant, soaproot and amole refer to the genus ''Chlorogalum''. They are native to western North America, with some species in Oregon but they are mostly found in California. Common names of the genus and several species ...
'' and ''Camassia
''Camassia'' is a genus of plants in the asparagus family native to North America. Common names include camas, quamash, Indian hyacinth, camash, and wild hyacinth.
It grows in the wild in great numbers in moist meadows. They are perennial p ...
'', are currently placed in the Agavoideae.[
Both historically and , there has been "considerable disagreement over generic limits" in the remaining Scilloideae, with different sources listing from 15 to 45 genera for sub-Saharan Africa alone.][ The total number of genera has been given as anything between about 30 (with about 500–700 species)][ and 70 (with about 1000 species).][
]
List of genera
Unless otherwise noted, the list below is based on genera accepted by the World Checklist of Selected Plant Families
The World Checklist of Selected Plant Families (usually abbreviated to WCSP) is an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected pla ...
as in the family Asparagaceae (with synonyms from the same source),[ with assignments to the subfamily Scilloideae based on the Germplasm Resources Information Network.][ As noted above, other sources divide up some of these genera, creating a significantly larger number; thus the genus ''Ornithogalum'' as conceived by Manning et al. (2009) is divided by Martínez-Azorín et al. (2011) into a more narrowly circumscribed ''Ornithogalum'' plus an additional 11 genera.][
]
Distribution and ecology
Scilloideae are widely but discontinuously distributed. The genus '' Oziroe'' is found only in parts of western South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
. Other genera occur in Africa south of the Sahara and parts of the Arabian Peninsula, on both sides of the Mediterranean
The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western Europe, Western and Southern Europe and Anatolia, on the south by North Africa ...
, further north in Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
through the Middle East
The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabia (including the Arabian Peninsula and Bahrain), Asia Minor (Asian part of Turkey except Hatay Province), East Thrace (Europ ...
to India
India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
, and on the east coast of Asia, in China, Korea
Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...
and Japan. Scilloideae are found in temperate
In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
to tropical
The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in
the Southern Hemisphere at S. The tropics are also referred to ...
habitats, but are more diverse in areas of Mediterranean climate
A Mediterranean climate (also called a dry summer temperate climate ''Cs'') is a temperate climate sub-type, generally characterized by warm, dry summers and mild, fairly wet winters; these weather conditions are typically experienced in the ...
(i.e., with a pronounced dry season during the summer).
Scilloideae reproduce
Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual org ...
both sexually and asexually. The showy flowers of many species of the subfamily are pollinated
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
by a wide range of insects
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
including bees, wasps
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
, flies
Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced m ...
and moths, as well as birds. Both nectar and pollen act as incentives to pollinating species. Vegetative reproduction
Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or spec ...
may be by bulbils
A bulbil (also referred to as bulbel, bulblet, and/or pup) is a small, young plant that is reproduced vegetatively from axillary buds on the parent plant's stem or in place of a flower on an inflorescence. These young plants are clones of the p ...
or by seeds through apomixis. The dispersal of seeds may occur by water, wind, or by ants
Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Creta ...
attracted by elaiosome
Elaiosomes ( grc, ἔλαιον ''élaion'' "oil" + ''sóma'' "body") 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 elaio ...
s.
Uses
Cultivation
Many members of the subfamily are popular garden plants, such as '' Hyacinthus'', '' Muscari'', ''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and sea ...
'', ''Puschkinia
''Puschkinia'' is a genus of four known species of bulbous perennials in the family Asparagaceae, subfamily Scilloideae. It is native to the Caucasus and the Middle East. ''Puschkinia scilloides'' is grown as an ornamental bulbous plant.
Descri ...
'', '' Hyacinthoides'', and ''Ornithogalum
''Ornithogalum'' is a genus of perennial plants mostly native to southern Europe and southern Africa belonging to the family Asparagaceae. Some species are native to other areas such as the Caucasus. Growing from a bulb, species have linear bas ...
'' (including those formerly placed in ''Galtonia'').
In South Africa the species of ''Eucomis
''Eucomis'' is a genus of flowering plants in the family Asparagaceae, subfamily Scilloideae, native to southern Africa. Most species of this genus are commonly referred to as pineapple flowers or pineapple lilies. They are bulbous perennials wit ...
'', ''Ornithogalum
''Ornithogalum'' is a genus of perennial plants mostly native to southern Europe and southern Africa belonging to the family Asparagaceae. Some species are native to other areas such as the Caucasus. Growing from a bulb, species have linear bas ...
'', '' Veltheimia'', among others, are grown as ornamentals. '' Ornithogalum thyrsoides'' and the different cultivars of hyacinths are important in the cut flower market.[
]
Medicinal use
''Drimia maritima
''Drimia maritima'' ( syn. ''Urginea maritima'') is a species of flowering plant in the family Asparagaceae, subfamily Scilloideae (formerly the family Hyacinthaceae). This species is known by several common names, including squill, sea squill, ...
'', the sea squill, has been used as a medicinal plant since ancient times. Its use for treatment of edema is mentioned in a papyrus
Papyrus ( ) is a material similar to thick paper that was used in ancient times as a writing surface. It was made from the pith of the papyrus plant, '' Cyperus papyrus'', a wetland sedge. ''Papyrus'' (plural: ''papyri'') can also refer to a ...
from 1554 BC, the Middle Kingdom of Egypt
The Middle Kingdom of Egypt (also known as The Period of Reunification) is the period in the history of ancient Egypt following a period of political division known as the First Intermediate Period. The Middle Kingdom lasted from approximately ...
. Bufadienolides isolated from ''Drimia maritima'' and '' Drimia indica'' are used for the production of substances for the treatment of heart conditions.
Food
The Scilloideae are only occasionally used as food plants for humans. In Italy
Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical ...
the bulbs of '' Leopoldia comosa'' are grown for food and in Greece
Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders ...
they are consumed as pickles. In France
France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of overseas regions and territories in the Americas and the Atlantic, Pacific and Indian Oceans. Its metropolitan area ...
the inflorescence of '' Ornithogalum pyrenaicum'' is consumed as a vegetable. In Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
some tribes consume the bulbs of '' Ledebouria apertiflora'' and '' Ledebouria revoluta''.
Toxicity
Many Scilloideae produce poisonous steroidal saponins
Saponins (Latin "sapon", soap + "-in", one of), also selectively referred to as triterpene glycosides, are bitter-tasting usually toxic plant-derived organic chemicals that have a foamy quality when agitated in water. They are widely distributed ...
such as bufadienolides and cardenolides
A cardenolide is a type of steroid. Many plants contain derivatives, collectively known as cardenolides, including many in the form of cardenolide glycosides (cardenolides that contain structural groups derived from sugars). Cardenolide glycoside ...
, making them inedible.
Several species are toxic. In South Africa, for example, ''Ornithogalum thyrsoides'', and several ''Ledebouria
''Ledebouria'' is a genus of African bulbous perennial herbs in the Asparagus family, Asparagaceae, subfamily Scilloideae. Most members were previously part of the genus ''Scilla''. A number of species are grown by cacti and succulent enthusi ...
'' species ('' Ledebouria cooperi'', ''L. inguinata'', ''L. ovatifolia'', ''L. revoluta''), ''Ornithogalum saundersiae
''Ornithogalum saundersiae'', or giant chincherinchee, is a species of ''Ornithogalum'' (star of Bethlehem) in the subfamily Scilloideae of family Asparagaceae.
Description
''Ornithogalum saundersiae'' is a perennial, herbaceous bulbous p ...
'' and several members of the tribe Urgineeae are poisonous to livestock. Scilliroside (a bufadienolide) is used to poison rats, traditionally by spreading dried chips of ''Drimia maritima
''Drimia maritima'' ( syn. ''Urginea maritima'') is a species of flowering plant in the family Asparagaceae, subfamily Scilloideae (formerly the family Hyacinthaceae). This species is known by several common names, including squill, sea squill, ...
'' bulbs.
References
Bibliography
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*
External links
Hyacinthaceae in Topwalks
''Monocot families'' (USDA)
NCBI Taxonomy Browser
links at CSDL, Texas
{{taxonbar, from=Q133292, from2=Q13833438
Asparagales subfamilies