Scilloideae (named after the genus ''
Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial plant, perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine ...
'', "squill") is a
subfamily
In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
of
bulb
In botany, a bulb is a short underground stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs duri ...
ous plants within the family ''
Asparagaceae
Asparagaceae (), known as the asparagus family, is a family of flowering plants, placed in the order Asparagales of the monocots. The family name is based on the edible garden asparagus, '' Asparagus officinalis''. This family includes both ...
''. 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
''Hyacinthus'' is a genus of bulbous herbs, and spring-blooming Perennial plant, perennials. They are fragrant flowering plants in the family Asparagaceae, subfamily Scilloideae and are commonly called hyacinths (). The genus is native predomin ...
us'' (hyacinths), ''
Hyacinthoides
''Hyacinthoides'' is a genus of flowering plants in the family Asparagaceae, known as bluebells.
Systematics
''Hyacinthoides'' is classified in the subfamily Scilloideae (now part of the family Asparagaceae, but formerly treated as a separate f ...
'' (bluebells), ''
Muscari
''Muscari'' is a genus (biology), genus of perennial bulbous plants native to Eurasia that produce spikes of dense, most commonly blue, urn-shaped flowers resembling bunches of grapes in the spring. The common name for the genus is grape hyacint ...
'' (grape hyacinths) and ''
Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial plant, perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine ...
'' and ''
Puschkinia
''Puschkinia'' is a genus of four known species of bulbous perennial plant, perennials in the family (biology), family Asparagaceae, subfamily Scilloideae. It is native to the Caucasus and the Middle East. ''Puschkinia scilloides'' is grown as an ...
'' (squills or scillas). Some are important as
cut flowers
Cut flowers are flowers and flower buds (often with some Plant stem, stem and leaf) that have been cut from the plant bearing it. It is removed from the plant for decorative use. Cut greens are leaves with or without stems added to the cut flow ...
.
Scilloideae are distributed mostly in
Mediterranean climate
A Mediterranean climate ( ), also called a dry summer climate, described by Köppen and Trewartha as ''Cs'', is a temperate climate type that occurs in the lower mid-latitudes (normally 30 to 44 north and south latitude). Such climates typic ...
s, including
South Africa
South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
,
Central Asia
Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
and
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. 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 ba ...
, 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, with the direction of their movement always opposite to that of the secretion of ...
, and arranged in a
basal rosette.
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
The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a f ...
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
''Hyacinthus'' is a genus of bulbous herbs, and spring-blooming Perennial plant, perennials. They are fragrant flowering plants in the family Asparagaceae, subfamily Scilloideae and are commonly called hyacinths (). The genus is native predomin ...
s (''Hyacinthus''), grape hyacinths (''Muscari
''Muscari'' is a genus (biology), genus of perennial bulbous plants native to Eurasia that produce spikes of dense, most commonly blue, urn-shaped flowers resembling bunches of grapes in the spring. The common name for the genus is grape hyacint ...
''), bluebells (''Hyacinthoides
''Hyacinthoides'' is a genus of flowering plants in the family Asparagaceae, known as bluebells.
Systematics
''Hyacinthoides'' is classified in the subfamily Scilloideae (now part of the family Asparagaceae, but formerly treated as a separate f ...
'') and squills (''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial plant, perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine ...
''). Other members are summer- and autumn-flowering, including '' Galtonia'' and '' Eucomis'' ('pineapple lilies'). Most are native to Mediterranean climate
A Mediterranean climate ( ), also called a dry summer climate, described by Köppen and Trewartha as ''Cs'', is a temperate climate type that occurs in the lower mid-latitudes (normally 30 to 44 north and south latitude). Such climates typic ...
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. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. Others are found in Central Asia
Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
, the Far East
The Far East is the geographical region that encompasses the easternmost portion of the Asian continent, including North Asia, North, East Asia, East and Southeast Asia. South Asia is sometimes also included in the definition of the term. In mod ...
and South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
.
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 ve ...
and 6 stamens
The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filamen ...
with a superior ovary
The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
, a characteristic which placed them within the older order of Liliales
Liliales is an order (biology), order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web List of systems of plant classification, system, within the lilioid monocots. This order of necessity includ ...
in many older classification systems, such as the Cronquist system
The Cronquist system is a list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...
, but they now separate from them within the Asparagales
Asparagales (asparagoid lilies) are a diverse order of flowering plants in the monocots. Under the APG IV system of flowering plant classification, Asparagales are the largest order of monocots with 14 families, 1,122 genera, and about 36,00 ...
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 fai ...
.
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
A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic.
The individuals of many ...
, 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
The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals w ...
: 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 form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''coroll ...
. The corolla may be white, yellow, violet, blue, brown and even black (see images).
Androecium
The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filament ...
: composed of 6 stamens (exceptionally 3, as in Albuca, for example), with the filaments free or adnate to the tube, often appendiculate. The anthers are dorsifixed and pollen dehiscence occurs by longitudinal openings. The pollen
Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
is monosulcate (having a linear furrow).
Gynoecium
Gynoecium (; ; : gynoecia) 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 (botany), whorl of a flower; it consists ...
: superior ovary, tricarpelate, connate and trilocular. Single stigma, capitate to 3-lobed. May contain from one to several ovules in each locule
A locule (: locules) or loculus (; : loculi) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus).
In angiosperms (flowering plants), the term ''locule'' usually refers to a chamber within an ovary ...
. They have nectaries
Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
at the septa of the ovaries.
Fruit
In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering.
Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
: dehiscence loculicidal.
Seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
: Seed morphology is diverse, from globular to flattened, and occasionally aril
An aril (), also called arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode, or false aril, is sometimes distinguished: whereas an aril grows from the attachment point of the seed to the ova ...
. 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 package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most importa ...
: Chromosome size varies widely, from 1.2 to 18 μm
The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
in length, karyotype
A karyotype is the general appearance of the complete set of chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is discerned by de ...
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 (''genus typica'') 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-bearin ...
, ''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial plant, perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine ...
'', 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria:
# the grouping contains its own most recent comm ...
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 pl ...
'', ''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 deri ...
'' 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
Lilioid monocots (lilioids, liliid monocots, petaloid monocots, petaloid lilioid monocots) is an informal name used for a grade (biology), grade (grouping of taxa with common characteristics) of five monocot order (biology), orders (Petrosavial ...
have long created classification problems. At one extreme, e.g. in the Cronquist system
The Cronquist system is a list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...
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 fai ...
''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 co ...
''). At the other extreme, e.g. in the Dahlgren system 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria:
# the grouping contains its own most recent comm ...
, 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
Liliales is an order (biology), order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web List of systems of plant classification, system, within the lilioid monocots. This order of necessity includ ...
and Asparagales
Asparagales (asparagoid lilies) are a diverse order of flowering plants in the monocots. Under the APG IV system of flowering plant classification, Asparagales are the largest order of monocots with 14 families, 1,122 genera, and about 36,00 ...
).
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
''Hyacinthus'' is a genus of bulbous herbs, and spring-blooming Perennial plant, perennials. They are fragrant flowering plants in the family Asparagaceae, subfamily Scilloideae and are commonly called hyacinths (). The genus is native predomin ...
us'' and '' Lachenalia''. The group was reduced to a tribe by Endlicher
Stephan Friedrich Ladislaus Endlicher, also known as Endlicher István László (24 June 1804 – 28 March 1849), was an Austrian botanist, numismatist and Sinologist. He was a director of the Botanical Garden of Vienna.
Biography
Endlicher ...
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 pl ...
''. In 1866 Salisbury
Salisbury ( , ) is a city status in the United Kingdom, cathedral city and civil parish in Wiltshire, England with a population of 41,820, at the confluence of the rivers River Avon, Hampshire, Avon, River Nadder, Nadder and River Bourne, Wi ...
redistributed the genera into several families.[
In the 1870s, ]Baker
A baker is a tradesperson who baking, bakes and sometimes Sales, sells breads and other products made of flour by using an oven or other concentrated heat source. The place where a baker works is called a bakery.
History
Ancient histo ...
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 Fritsch is a German surname. Like Fritsche, Fritzsch and Fritzsche, it is a patronymic derived from Friedrich.
Notable people with the surname include:
* Ahasverus Fritsch (1629–1701), German jurist, poet and hymn writer
* Antonin Fritsch (183 ...
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 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
In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria:
# the grouping contains its own most recent co ...
, the tendency was to create new families out of each identified clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, 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 ...
. This placed many lilioid families and genera in the order Asparagales
Asparagales (asparagoid lilies) are a diverse order of flowering plants in the monocots. Under the APG IV system of flowering plant classification, Asparagales are the largest order of monocots with 14 families, 1,122 genera, and about 36,00 ...
(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 phylogenetics, molecular-based, list of systems of plant taxonomy, system of plant taxonomy that ...
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
The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a f ...
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) was an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected p ...
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.][ Hyacintheae was further divided into three subtribes: Pseudoprosperinae, Massoniinae and Hyacinthinae.][ The possible relationship of the tribes and subtribes is illustrated in the following ]cladogram
A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
,[ 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 second 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 that presents up-to-date research on the phylogeny and taxonomy of flowering plants (angiosperms) in what is intended to be a user-friendly way.
The site is hosted by the Missouri Botanical ...
, the tribe contains four genera, '' Albuca'' (about 110–140 species), ''Dipcadi
''Dipcadi'' is a genus of bulbous flowering plants in the family (biology), family Asparagaceae, subfamily Scilloideae (also treated as the family Hyacinthaceae).Stevens, P.F. (2001 onwards),Asparagales: Scilloideae, ''Angiosperm Phylogeny Webs ...
'', ''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. Some species are classified as noxious invas ...
'' (about 160 species, including ''Galtonia'' and ''Neopatersonia'') and ''Pseudogaltonia
''Pseudogaltonia'' 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'', retrie ...
''.[ By contrast, Martínez-Azorín et al. (2011) divide the tribe into 19 genera.][
]
Urgineeae
Species within this tribe contain bufadienolide
Bufadienolide is a chemical compound with steroid structure. Its derivatives are collectively known as bufadienolides, including many in the form of bufadienolide glycosides (bufadienolides that contain structural groups derived from sugars). These ...
s 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 endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the Embryo#Pla ...
. The basic chromosome numbers are ''n'' = 6, 7 and 10.[ Depending on the source, the tribe may include the genera '']Bowiea
''Bowiea'', commonly known as climbing-onion, is a genus of bulbous, perennial, succulent plants which thrive in dry and desert regions of eastern and southern Africa, ranging from Uganda to South Africa. It is native to a region stretching from ...
'', '' Drimia'' (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 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 ri ...
s. The tribe can in turn be divided into three subtribes:[
* Pseudoprosperinae Speta
: A monotypic subtribe genus with a single species, '' 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 ( ''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 elaiosomes that attract ...
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'', '' Lachenalia'' (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 enthusias ...
'' (about 80 species), ''Massonia
''Massonia'' is a genus (biology), genus of bulbous perennial plant, perennial flowering plants in the family (biology), family Asparagaceae, subfamily Scilloideae (also treated as the family Hyacinthaceae). It is native to southern Africa, and i ...
'' (including ''Whiteheadia''), '' Merwilla'', '' Schizocarphus'' and ''Veltheimia
''Veltheimia'' is a genus of perennial plants native to the Cape Provinces of South Africa in the family Asparagaceae, subfamily Scilloideae
Scilloideae (named after the genus ''Scilla'', "squill") is a subfamily of bulbous plants within the f ...
''.[
* 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
''Brimeura'' is a genus of bulb-forming, monocotyledonous perennial plants. They have narrow leaves and bear bluebell-like flowers in Spring. ''Brimeura'' is treated in the family Asparagaceae, subfamily Scilloideae, tribe Hyacintheae, subtribe H ...
'', ''Hyacinthoides
''Hyacinthoides'' is a genus of flowering plants in the family Asparagaceae, known as bluebells.
Systematics
''Hyacinthoides'' is classified in the subfamily Scilloideae (now part of the family Asparagaceae, but formerly treated as a separate f ...
'', ''Muscari
''Muscari'' is a genus (biology), genus of perennial bulbous plants native to Eurasia that produce spikes of dense, most commonly blue, urn-shaped flowers resembling bunches of grapes in the spring. The common name for the genus is grape hyacint ...
'' (about 50 species), ''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial plant, perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine ...
'' (about 30 species) and ''Prospero
Prospero ( ) is a fictional character and the protagonist of William Shakespeare's ''The Tempest''.
Character
Twelve years before the play begins, Prospero is usurped from his position as the rightful Duke of Milan by his brother Antonio, ...
'' (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 deri ...
'' 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 pl ...
'', 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
Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
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) was an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected p ...
as in the family Asparagaceae
Asparagaceae (), known as the asparagus family, is a family of flowering plants, placed in the order Asparagales of the monocots. The family name is based on the edible garden asparagus, '' Asparagus officinalis''. This family includes both ...
(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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. Other genera occur in Africa south of the Sahara
The Sahara (, ) is a desert spanning across North Africa. With an area of , it is the largest hot desert in the world and the list of deserts by area, third-largest desert overall, smaller only than the deserts of Antarctica and the northern Ar ...
and parts of the Arabian Peninsula
The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world.
Geographically, the ...
, 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 east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
, further north in Europe
Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
through the Middle East
The Middle East (term originally coined in English language) is a geopolitical region encompassing the Arabian Peninsula, the Levant, Turkey, Egypt, Iran, and Iraq.
The term came into widespread usage by the United Kingdom and western Eur ...
to India
India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
, and on the east coast of Asia, in China
China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, Korea
Korea is a peninsular region in East Asia consisting of the Korean Peninsula, Jeju Island, and smaller islands. Since the end of World War II in 1945, it has been politically Division of Korea, divided at or near the 38th parallel north, 3 ...
and Japan
Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
. Scilloideae are found in temperate
In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
to tropical
The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
habitats, but are more diverse in areas of Mediterranean climate
A Mediterranean climate ( ), also called a dry summer climate, described by Köppen and Trewartha as ''Cs'', is a temperate climate type that occurs in the lower mid-latitudes (normally 30 to 44 north and south latitude). Such climates typic ...
(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. There are two forms of reproduction: asexual and sexual.
In asexual reprod ...
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. Pollinating agents can be animals such as insects, for example bees, beetles or butterflies; bird ...
by a wide range of insects
Insects (from Latin ') are 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 jointed ...
including bees
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
, 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. Th ...
, flies
Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
and moths
Moths are a group of insects that includes all members of the order Lepidoptera that are not butterflies. They were previously classified as suborder Heterocera, but the group is paraphyletic with respect to butterflies (suborder Rhopalocera) a ...
, as well as birds
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
. Both nectar and pollen act as incentives to pollinating species. Vegetative reproduction
Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is a form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or specializ ...
may be by bulbils
A bulbil (also referred to as a 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 ...
or by seeds through apomixis
In botany, apomixis is asexual development of seed or embryo without fertilization. However, other definitions include replacement of the seed by a plantlet or replacement of the flower by bulbils.
Apomictically produced offspring are geneti ...
. The dispersal of seeds may occur by water, wind, or by ants
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
attracted by elaiosome
Elaiosomes ( ''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 elaiosomes that attract ...
s.
Uses
Cultivation
Many members of the subfamily are popular garden plants, such as '' Hyacinthus'', ''Muscari
''Muscari'' is a genus (biology), genus of perennial bulbous plants native to Eurasia that produce spikes of dense, most commonly blue, urn-shaped flowers resembling bunches of grapes in the spring. The common name for the genus is grape hyacint ...
'', ''Scilla
''Scilla'' () is a genus of about 30 to 80 species of bulb-forming perennial plant, perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine ...
'', ''Puschkinia
''Puschkinia'' is a genus of four known species of bulbous perennial plant, perennials in the family (biology), family Asparagaceae, subfamily Scilloideae. It is native to the Caucasus and the Middle East. ''Puschkinia scilloides'' is grown as an ...
'', ''Hyacinthoides
''Hyacinthoides'' is a genus of flowering plants in the family Asparagaceae, known as bluebells.
Systematics
''Hyacinthoides'' is classified in the subfamily Scilloideae (now part of the family Asparagaceae, but formerly treated as a separate f ...
'', 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. Some species are classified as noxious invas ...
'' (including those formerly placed in ''Galtonia'').
In South Africa the species of '' Eucomis'', ''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. Some species are classified as noxious invas ...
'', ''Veltheimia
''Veltheimia'' is a genus of perennial plants native to the Cape Provinces of South Africa in the family Asparagaceae, subfamily Scilloideae
Scilloideae (named after the genus ''Scilla'', "squill") is a subfamily of bulbous plants within the f ...
'', among others, are grown as ornamentals. ''Ornithogalum thyrsoides
''Ornithogalum thyrsoides'' is a bulbous plant species that is endemic to the Cape Province in South Africa. It is also known by the common names of chinkerinchee or chincherinchee, star-of-Bethlehem or wonder-flower. It produces long-lasting f ...
'' 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, s ...
'', the sea squill, has been used as a medicinal plant since ancient times. Its use for treatment of edema
Edema (American English), also spelled oedema (British English), and also known as fluid retention, swelling, dropsy and hydropsy, is the build-up of fluid in the body's tissue (biology), tissue. Most commonly, the legs or arms are affected. S ...
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'' or ''papyruses'') can 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 of Egypt, First Intermediate Period. The Middl ...
. 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, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
the bulbs of ''Leopoldia comosa
''Leopoldia comosa'' (syn. ''Muscari comosum'') is a perennial bulbous flowering plant in the family Asparagaceae. Usually called the tassel hyacinth or tassel grape hyacinth, it is one of a number of species and genera also known as grape hyacin ...
'' are grown for food and in Greece
Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...
they are consumed as pickles. In France
France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
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. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
some tribes consume the bulbs of ''Ledebouria apertiflora
''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 enthusiasts ...
'' and ''Ledebouria revoluta
''Ledebouria revoluta'', the south Indian squill, is a flowering plant species in the genus ''Ledebouria'' found in Southern Africa and India.
Description
The leaves of ''Ledebouria revoluta'' are smooth and fleshy, and unlike those of some othe ...
''.
Toxicity
Some members of Scilloideae produce toxic steroids known as cardiac glycosides
Cardiac glycosides are a class of organic compounds that increase the output force of the heart and decrease its rate of contractions by inhibiting the cellular sodium-potassium ATPase pump. Their beneficial medical uses include treatments for ...
and thus should not be ingested.
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 enthusias ...
'' species (''Ledebouria cooperi
''Ledebouria cooperi'' is a species of bulbous flowering plant in the family Asparagaceae
Asparagaceae (), known as the asparagus family, is a family of flowering plants, placed in the order Asparagales of the monocots. The family name is ...
'', ''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 plan ...
'' and several members of the tribe Urgineeae are poisonous to livestock. Scilliroside
Scilliroside is a toxic compound derived from the plant '' Drimia maritima'' ( syn. ''Urginea maritima''), which is sometimes used as a rodenticide
Rodenticides are chemicals made and sold for the purpose of killing rodents. While commonly re ...
(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, s ...
'' bulbs.
References
Bibliography
*
*
*
*
*
External links
Hyacinthaceae in Topwalks
''Monocot families'' (USDA)
NCBI Taxonomy Browser
links at CSDL, Texas
{{Authority control
Asparagales subfamilies