Themidaceae
Encyclopedia
Brodiaeoideae is a monocot subfamily of flowering plants in the family
Asparagaceae
, order
Asparagales
. It has been treated as a separate family, Themidaceae. They are native
to Central America
and western North America
, from British Columbia
to Guatemala
. The name of the subfamily is based on the type genus Brodiaea
.
In molecular phylogenetic analyses, Brodiaeoideae is strongly supported
as monophyletic. It is probably sister to Scilloideae. Recent treatments
have divided Brodiaeoideae (or Themidaceae) into 12 genera
. The monophyly of several of the genera remains in doubt. As currently circumscribed
, the largest genera are Triteleia, with 15 species
, and Brodiaea, with 14. Nine of the 12 genera are known in cultivation, but only species of Brodiaea and Triteleia are commonly grown.
Perennial
herbs
arising from a starch
y corm
; a new corm arising each year from the old one.
Leaves
linear
, often fleshy, forming a closed sheath at their base. Veins parallel.
Inflorescence
an umbel
, or rarely a single flower, at the apex of a solitary scape
. Flower
s bisexual, actinomorphic. Tepal
s all similar, in 2 whorls
of 3.
Fertile stamen
s 6, or 3 and alternating with 3 staminode
s. Stamens and staminodes inserted on tepals. Anthers basifixed and introrse.
Ovary
superior and trilocular
.
Fruit
a loculicidal capsule
. Seed
covered with phytomelan.
was recognized at all, it was usually at tribal
rank
and usually called Brodiaeeae. Most authors assigned it to Liliaceae
, Alliaceae
, or Amaryllidaceae
. In 1985, Dahlgren
, Clifford, and Yeo treated it as tribe Brodiaeeae of Alliaceae.
Toward the end of the 20th century, it became increasingly evident that the heterogeneous Liliaceae recognized by most authors was several times polyphyletic and that Brodiaea and its relatives were closer to Asparagus
than to Allium
or Amaryllis
. For these reasons, the family Themidaceae was resurrected in an article in Taxon in 1996. The name
'Themidaceae' was first used by Richard Salisbury in 1866. The name was based
on the now-defunct genus Themis, which was established by Salisbury along with the family. The only species ever assigned to Themis was Themis ixioides. Its name was changed to Brodiaea ixioides by Sereno Watson
in 1879, then to Triteleia ixioides by Edward Lee Greene
in 1886. It is known as Triteleia ixioides in Flora of North America
.
When the Angiosperm Phylogeny Group
published the APG II system
in 2003, Themidaceae was treated as an optional circumscription
for those who thought that Asparagaceae
sensu lato should be divided into smaller segregate
families. When the APG III system
was published in 2009, Themidaceae was not accepted. In an accompanying article, it was treated as Brodiaeoideae, one of 7 subfamilies in Asparagaceae.
, the genera included in the subfamily are:
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Asparagaceae
Asparagaceae
Asparagaceae is the botanical name of a family of flowering plants, placed in the order Asparagales of the monocots.In earlier classification systems, the species involved were often treated as belonging to the family Liliaceae...
, order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
Asparagales
Asparagales
Asparagales is the name of an order of plants, used in modern classification systems such as the APG III system . The order takes its name from the family Asparagaceae and is placed in the monocots. The order has only recently been recognized in classification systems...
. It has been treated as a separate family, Themidaceae. They are native
Indigenous (ecology)
In biogeography, a species is defined as native to a given region or ecosystem if its presence in that region is the result of only natural processes, with no human intervention. Every natural organism has its own natural range of distribution in which it is regarded as native...
to Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...
and western North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
, from British Columbia
British Columbia
British Columbia is the westernmost of Canada's provinces and is known for its natural beauty, as reflected in its Latin motto, Splendor sine occasu . Its name was chosen by Queen Victoria in 1858...
to Guatemala
Guatemala
Guatemala is a country in Central America bordered by Mexico to the north and west, the Pacific Ocean to the southwest, Belize to the northeast, the Caribbean to the east, and Honduras and El Salvador to the southeast...
. The name of the subfamily is based on the type genus Brodiaea
Brodiaea
Brodiaea is a monocot genus of flowering plants in the family Asparagaceae, subfamily Brodiaeoideae, also known by the common name cluster-lilies...
.
In molecular phylogenetic analyses, Brodiaeoideae is strongly supported
Resampling (statistics)
In statistics, resampling is any of a variety of methods for doing one of the following:# Estimating the precision of sample statistics by using subsets of available data or drawing randomly with replacement from a set of data points # Exchanging labels on data points when performing significance...
as monophyletic. It is probably sister to Scilloideae. Recent treatments
Treatise
A treatise is a formal and systematic written discourse on some subject, generally longer and treating it in greater depth than an essay, and more concerned with investigating or exposing the principles of the subject.-Noteworthy treatises:...
have divided Brodiaeoideae (or Themidaceae) into 12 genera
Genera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
. The monophyly of several of the genera remains in doubt. As currently circumscribed
Circumscription (taxonomy)
In taxonomy, circumscription is the definition of the limits of a taxonomic group of organisms. One goal of taxonomy is to achieve a stable circumscription for every taxonomic group. Achieving stability can be simple or difficult....
, the largest genera are Triteleia, with 15 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
, and Brodiaea, with 14. Nine of the 12 genera are known in cultivation, but only species of Brodiaea and Triteleia are commonly grown.
Description
The following description is derived from two sources.Perennial
Perennial plant
A perennial plant or simply perennial is a plant that lives for more than two years. The term is often used to differentiate a plant from shorter lived annuals and biennials. The term is sometimes misused by commercial gardeners or horticulturalists to describe only herbaceous perennials...
herbs
Herbaceous plant
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...
arising from a starch
Starch
Starch or amylum is a carbohydrate consisting of a large number of glucose units joined together by glycosidic bonds. This polysaccharide is produced by all green plants as an energy store...
y corm
Corm
A corm is a short, vertical, swollen underground plant stem that serves as a storage organ used by some plants to survive winter or other adverse conditions such as summer drought and heat ....
; a new corm arising each year from the old one.
Leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
linear
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...
, often fleshy, forming a closed sheath at their base. Veins parallel.
Inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
an umbel
Umbel
An umbel is an inflorescence which consists of a number of short flower stalks which are equal in length and spread from a common point, somewhat like umbrella ribs....
, or rarely a single flower, at the apex of a solitary scape
Scape (botany)
In botany, scapes are leafless flowering stems that rise from the ground. Scapes can have a single flower or many flowers, depending on the species....
. Flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...
s bisexual, actinomorphic. Tepal
Tepal
Tepals are elements of the perianth, or outer part of a flower, which include the petals or sepals. The term tepal is more often applied specifically when all segments of the perianth are of similar shape and color, or undifferentiated, which is called perigone...
s all similar, in 2 whorls
Whorl (botany)
In botany, a whorl is an arrangement of sepals, petals, leaves, or branches in which all the parts are attached at the same point and surround or wrap around the stem.There are four whorls in a general flower...
of 3.
Fertile stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...
s 6, or 3 and alternating with 3 staminode
Staminode
In botany, a staminode is an often rudimentary, sterile or abortive stamen. This means that it does not produce pollen. Staminodes are frequently inconspicuous and stamen-like, usually occurring at the inner whorl of the flower, but are also sometimes long enough to protrude from the...
s. Stamens and staminodes inserted on tepals. Anthers basifixed and introrse.
Ovary
Ovary (plants)
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 and is located above or below or at the point of connection with the base of the petals and sepals...
superior and trilocular
Locule
A locule is a small cavity or compartment within an organ or part of an organism ....
.
Fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
a loculicidal capsule
Capsule (fruit)
In botany a capsule is a type of simple, dry fruit produced by many species of flowering plants. A capsule is a structure composed of two or more carpels that in most cases is dehiscent, i.e. at maturity, it splits apart to release the seeds within. A few capsules are indehiscent, for example...
. Seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
covered with phytomelan.
History
For most of the 19th and 20th centuries, when the groupTaxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
was recognized at all, it was usually at tribal
Tribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...
rank
Taxonomic rank
In biological classification, rank is the level in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, and class. Each rank subsumes under it a number of less general categories...
and usually called Brodiaeeae. Most authors assigned it to Liliaceae
Liliaceae
The Liliaceae, or the lily family, is a family of monocotyledons in the order Liliales. Plants in this family have linear leaves, mostly with parallel veins but with several having net venation , and flower arranged in threes. Several have bulbs, while others have rhizomes...
, Alliaceae
Alliaceae
Allioideae is the botanical name of a monocot subfamily of flowering plants in the family Amaryllidaceae, order Asparagales. It was formerly treated as a separate family, Alliaceae...
, or Amaryllidaceae
Amaryllidaceae
Amaryllidoideae is the subfamily of flowering plants that takes its name from the genus Amaryllis. It is part of the family Amaryllidaceae, in order Asparagales...
. In 1985, Dahlgren
Rolf Dahlgren
Rolf Martin Theodor Dahlgren was a Swedish-Danish botanist, professor at the University of Copenhagen from 1973 to his death....
, Clifford, and Yeo treated it as tribe Brodiaeeae of Alliaceae.
Toward the end of the 20th century, it became increasingly evident that the heterogeneous Liliaceae recognized by most authors was several times polyphyletic and that Brodiaea and its relatives were closer to Asparagus
Asparagus
Asparagus officinalis is a spring vegetable, a flowering perennialplant species in the genus Asparagus. It was once classified in the lily family, like its Allium cousins, onions and garlic, but the Liliaceae have been split and the onion-like plants are now in the family Amaryllidaceae and...
than to Allium
Allium
Allium is a monocot genus of flowering plants, informally referred to as the onion genus. The generic name Allium is the Latin word for garlic....
or Amaryllis
Amaryllis
Amaryllis is a small genus of flowering bulbs, with two species. The better known of the two, Amaryllis belladonna, is a native of South Africa, particularly the rocky southwest region near the Cape...
. For these reasons, the family Themidaceae was resurrected in an article in Taxon in 1996. The name
Botanical name
A botanical name is a formal scientific name conforming to the International Code of Botanical Nomenclature and, if it concerns a plant cultigen, the additional cultivar and/or Group epithets must conform to the International Code of Nomenclature for Cultivated Plants...
'Themidaceae' was first used by Richard Salisbury in 1866. The name was based
Basionym
Basionym is a term used in botany, regulated by the International Code of Botanical Nomenclature...
on the now-defunct genus Themis, which was established by Salisbury along with the family. The only species ever assigned to Themis was Themis ixioides. Its name was changed to Brodiaea ixioides by Sereno Watson
Sereno Watson
Sereno Watson was an American botanist.Graduating from Yale in 1847, he drifted through various occupations until, in California, he joined the Clarence King Expedition and eventually became its expedition botanist...
in 1879, then to Triteleia ixioides by Edward Lee Greene
Edward Lee Greene
Edward Lee Greene, Ph.D., was an American botanist known for his numerous publications including the two-part Landmarks of Botanical History and the naming or redescribing of over 4,400 species of plants in the American West.- Early Life :Edward Lee Greene was born on August 20, 1843 in...
in 1886. It is known as Triteleia ixioides in Flora of North America
Flora of North America
The Flora of North America North of Mexico is a multivolume work describing the native plants of North America. These days much of the Flora is available . The work is expected to fill 30 volumes when completed...
.
When the Angiosperm Phylogeny Group
Angiosperm Phylogeny Group
The Angiosperm Phylogeny Group, or APG, refers to an informal international group of systematic botanists who came together to try to establish a consensus on the taxonomy of flowering plants that would reflect new knowledge about plant relationships discovered through phylogenetic studies., three...
published the APG II system
APG II system
The APG II system of plant classification is the second, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Group. It was a revision of the first APG system, published in 1998, and was superseded in 2009...
in 2003, Themidaceae was treated as an optional circumscription
Circumscription (taxonomy)
In taxonomy, circumscription is the definition of the limits of a taxonomic group of organisms. One goal of taxonomy is to achieve a stable circumscription for every taxonomic group. Achieving stability can be simple or difficult....
for those who thought that Asparagaceae
Asparagaceae
Asparagaceae is the botanical name of a family of flowering plants, placed in the order Asparagales of the monocots.In earlier classification systems, the species involved were often treated as belonging to the family Liliaceae...
sensu lato should be divided into smaller segregate
Segregate (taxonomy)
In taxonomy, a segregate, or a segregate taxon is created when a taxon is split off, from another taxon. This other taxon will be better known, usually bigger, and will continue to exist, even after the segregate taxon has been split off...
families. When the APG III system
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...
was published in 2009, Themidaceae was not accepted. In an accompanying article, it was treated as Brodiaeoideae, one of 7 subfamilies in Asparagaceae.
Genera
According to the Angiosperm Phylogeny WebsiteAngiosperm Phylogeny Website
Angiosperm Phylogeny Website is a well-known web site dedicated to research on Angiosperm phylogeny and taxonomy.The site is hosted by the Missouri Botanical Garden website and maintained by researcher Peter F. Stevens, who is a member of the Angiosperm Phylogeny Group...
, the genera included in the subfamily are:
- AndrostephiumAndrostephiumThe genus Androstephium belongs to the family Asparagaceae, subfamily Brodiaeoideae. It contains three species.Species include:*Androstephium breviflorum - pink funnel lily*Androstephium caeruleum - blue funnel lily-External links:*...
Torr. - BesseraBesseraBessera is a genus of plants in the family Asparagaceae, subfamily Brodiaeoideae. A small genus of mostly herbaceous flowering plants with corms, they are perennials native to Mexico...
Schult.f. (including Behria) - BloomeriaBloomeriaBloomeria, a geophyte in the family Asparagaceae, subfamily Brodiaeoideae,, was named for H. G. Bloomer an early San Francisco botanist. It consists of three species of California and Baja California:* Bloomeria clevelandii S. Watson...
Kellogg - BrodiaeaBrodiaeaBrodiaea is a monocot genus of flowering plants in the family Asparagaceae, subfamily Brodiaeoideae, also known by the common name cluster-lilies...
Sm. - DandyaDandya (plant)Dandya is a genus of about four species of flowering plants found in Mexico. In the APG III classification system, it is placed in the family Asparagaceae, subfamily Brodiaeoideae ....
H.E.Moore - DichelostemmaDichelostemmaDichelostemma is a genus of seven species of flowering plants in the order Asparagales. The genus is native to the western United States, especially in northern California, but also east to Utah and north to Oregon and southern Washington....
Kunth (including Brevoortia, Dipterostemon, Stropholirion) - MillaMillaMilla is a genus of plant belonging to the family Asparagaceae, subfamily Brodiaeoideae, and contains the following species of plants.* Milla biflora, Mexican star...
Cav. (including Diphalangium) - MuillaMuillaThe genus Muilla includes three to four species of flowering plants. In the APG III classification system, it is placed in the family Asparagaceae, subfamily Brodiaeoideae. They are native to western North America...
S.Watson ex Benth. - PetronymphePetronymphePetronymphe is a genus of one species of flowering plant found in Mexico. In the APG III classification system, it is placed in the family Asparagaceae, subfamily Brodiaeoideae ....
H.E.Moore - Triteleia Douglas ex Lindl. (including Hesperoscordium, Themis)
- TriteleiopsisTriteleiopsisTriteleiopsis is a genus of one species of flowering plant found in in southern Arizona and north-west Mexico. In the APG III classification system, it is placed in the family Asparagaceae, subfamily Brodiaeoideae ....
Hoover
External links
- Brodiaeoideae At: Angiosperm Phylogeny Website At: Missouri Botanical Garden Website
- Liliaceae At: Flora of North America At: eFloras
- A fragment containing Liriogamae At: Google Books
- Sereno Watson 1879 At: Biodiversity Heritage Library
- Edward Lee Greene 1886 At: Biodiversity Heritage Library
- Resurrection of Themidaceae