Limnanthaceae

Limnanthaceae
Limnanthes douglasii
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Rosids
Order:	Brassicales
Family:	Limnanthaceae R.Br. [1]
Genera
Floerkea Limnanthes

The Limnanthaceae are a small family of annual herbs occurring throughout temperate North America. There are eight species and nineteen taxa currently recognized. Members of this family are prominent in vernal pool communities of California. Some taxa have been domesticated for use as an oil seed crop. Some members are listed as threatened or endangered and have been the focus of disputes over development plans (e.g. Limnanthes floccosa subsp. californica, ^[2] Limnanthes vinculans ^[3] )

Contents

• Taxonomy
• Morphology
• Systematics of Limnanthaceae
• References

The Limnanthaceae are members of a recently identified clade (Brassicales) of mustard oil producing plants. They have a sharp flavor similar to mustard greens, radish or capers. ^[4]

Two genera are recognized in the family. The monotypic genus Floerkea inhabits shaded, vernally wet habitats in eastern North American deciduous forests, high montane islands in the great basin and humid coniferous forests along the northwest coast. All seven species of Limnanthes , with one British Columbian exception, occur solely in the California Floristic Province, most commonly occupying grassland or savanna vernal pool habitats. Various taxa of this genus are prominent elements in the flora of the Great Central Valley "hogwallow" communities, the coastal prairie, and wet meadows of the Coast Ranges and the Sierra Nevada/Cascade foothills up to 1800 meters. Disjunct populations occur in the Peninsular Ranges just north of the Mexican border and in the Umpqua River valley of central Oregon. In favorable years Limnanthes can cover large areas with white flowers (hence the common name Meadowfoam) and in hogwallow habitats sometimes forms spectacular rings surrounding the deepest parts of the pools.

Two sorts of flowers are found in the family, reflecting different breeding systems: some taxa have inconspicuous perianths and reproduce largely by self-pollination. Others have large, showy flowers, usually pentamerous and white, some with varying amounts of yellow or ultraviolet, others with prominent rose or brown veins and anthers. These are fully to predominantly outcrossing, usually pollinated by species of solitary bees in the genera Panurginus, Andrena and Hesperandrina.

The Flora of North America Project has chosen a line drawing of Floerkea to serve as its logo because of this taxon's ubiquitous (but obscure) occurrence in many areas of North America, and the diverse aspects of the family including economic and horticultural value, endangered species status and fruitful subject of scientific research.

Taxonomy

Limnanthaceae currently consist of two genera, eight species and nineteen taxa.

Morphology

Limnanthaceae are all herbaceous annuals. Leaves are alternate, simple or compound with pinnate venation. Flowers are produced singly in the axils of leaves. They have 3, 4 or 5 petals, mostly white. Fruits are ridged or tuberculate schizocarps (nutlets).

Systematics of Limnanthaceae

When Robert Brown created the family Limnanthaceae in 1833 he declared that, "the place of this new family is not absolutely determined; but it is suggested that in two remarkable points of its structure, namely, the presence of glands subtending the alternate filaments, and the existence of a gynobase, it more nearly approaches to Hypogynous families than to Perigynous, with which it has hitherto been associated". ^[5] Ultimately he demurred from assigning a systematic position to his family. Subsequent taxonomists promoted various divergent placements. Bentham and Hooker (1862) ^[6] decided that given the great similarity of Limnanthes and Floerkea to taxa in the Geraniaceae, family status was not warranted; accordingly, they placed both genera in the tribe Limnantheae in that family. Engler and Prantl (1896), ^[7] however, disagreed, noting that the position of the seed indicated the family should be placed in the Sapindales.

Most recent authors of taxonomic treatises have maintained the family Limnanthaceae but placed it in the order Geraniales, including Thorne (1976) Cronquist (1988), who nevertheless states that Limnanthaceae are, "without...obvious affinities..." and Takhtajan (1980). Hutchinson (1973), proposed that Limnanthaceae should be included in Geraniales, but its similarity to Caryophyllales (which he believed is derived from Geraniales) suggested that Limnanthaceae form a link between these two groups.

A number of studies over the years have noted various morphological, developmental and embryological characters at variance with the Geraniales (as well as other groups to which Limnanthaceae have been assigned). Maheshwari and Johri (1956) conducted an extensive investigation of the morphology of Floerkea noting that, among other things, the herbaceous habit, gynobasic style, unusual type of tetrasporic embryo sac and basal parietal placentation of the unitegmic, tenuinucellate ovules differ from the Geraniales which has among its woody to herbaceous members, (at most) lobed syncarpous gynoecia, monosporic embryo sacs, generally axile placentation and bitegmic, crassinucellate ovules. Additionally the fruit type of Limnanthaceae, a schizocarpic nutlet, is unlike anything in the Geraniales most of which produce capsules. They also found a number of key differences between Limnanthaceae and Sapindales, and concluded that Limnanthaceae should be given their own order. ^[8]

Hofmann and Ludewig (1985) undertook a similar detailed study of the morphology of Limnanthes douglasii and made systematic inferences from their findings. They concluded that there is nothing in the morphology to suggest a relationship to the Coriariaceae (suggested by Chatin in 1856), Geraniales or the Sapindales (sensu Cronquist), though they allow that the, "level in evolution is about the same...." They also rejected any meaningful similarity between Limnanthaceae and Tropaeolaceae, as suggested by Dahlgren, who noted the shared possession of certain phytochemicals (glucosinolate, myrosinase, erucic acid and eicosenoic acid). They rejected this connection on the grounds that these phytochemicals also occur in the Brassicaceae which is assumed to be unrelated and suggest that the secondary compounds must be convergent. They concluded that the systematic position of Limnanthaceae is uncertain and cannot be presently determined.

Dahlgren (1975), who believed strongly in the systematic value of secondary chemicals, placed Limnanthaceae in Capparales with Brassicaceae. He later (1980) placed Limnanthaceae together with Tropaeolaceae in the order Tropaeolales.

Buchner, Halbritter, Prundner and Hesse (1990) investigated the pollen morphology of Limnanthaceae and discovered that the zonosulcate morphology is unlike the pollen of any known angiosperm and therefore relationships cannot be inferred from it. They reiterated Maheshwari and Johri's suggestion that a new order, the Limnanthales, should be created to contain the family.

Rodman (1991a, b) ^{[9] [10]} included Limnanthaceae in a twin phenetic and cladistic analysis of all 15 taxa then known to produce glucosinolates. In UPGMA phenograms, Limnanthaceae clustered with Balsaminaceae. Similarly, cladistic analysis showed Limnanthaceae either in a clade with Balsaminaceae and sometimes Pentadiplandraceae or in a polytomy with Balsaminaceae, Pentadiplandraceae, Caricacaeae, Centrospermae, and a clade which includes all the rest of the glucosinolate taxa (except Drypetes).

Rodman et al. (1993, 1997) ^{[11] [12]} assembled additional DNA sequence data sets for glucosinolate taxa and applied cladistic methods to generate hypotheses about relationship. They discovered that all the glucosinolate-producing taxa save Drypetes, form a clade. Despite the paucity of morphological synapomorphies (and therefore, to the surprise of many taxonomists), rbcL sequences have provided well supported evidence for the group's monophyly. 18S nuclear ribosomal DNA sequences have similarly, though less certainly, indicated a common ancestor for all the glucosinolate containing plants except Drypetes.

The Angiosperm Phylogeny Group places Limnanthaceae in Brassicales, which is in the Malvid/Rosid II lineage. ^[13]

Related Research Articles

Asparagales 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 the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.

The Brassicales are an order of flowering plants, belonging to the eurosids II group of dicotyledons under the APG II system. One character common to many members of the order is the production of glucosinolate compounds. Most systems of classification have included this order, although sometimes under the name Capparales.

The Dioscoreales are an order of monocotyledonous flowering plants, organized under modern classification systems, such as the Angiosperm Phylogeny Group or the Angiosperm Phylogeny Web. Among monocot plants, Dioscoreales are grouped with the lilioid monocots, wherein they are a sister group to the Pandanales. In total, the order Dioscoreales comprises three families, 22 genera and about 850 species.

Rosales is an order of flowering plants. It is sister to a clade consisting of Fagales and Cucurbitales. It contains about 7,700 species, distributed into about 260 genera. Rosales comprise nine families, the type family being the rose family, Rosaceae. The largest of these families are Rosaceae (90/2500) and Urticaceae (54/2600). The order Rosales is divided into three clades that have never been assigned a taxonomic rank. The basal clade consists of the family Rosaceae; another clade consists of four families, including Rhamnaceae; and the third clade consists of the four urticalean families.

Sapindales is an order of flowering plants. Well-known members of Sapindales include citrus; maples, horse-chestnuts, lychees and rambutans; mangos and cashews; frankincense and myrrh; mahogany and neem.

The Cornales are an order of flowering plants, early diverging among the asterids, containing about 600 species. Plants within the Cornales usually have four-parted flowers, drupaceous fruits, and inferior to half-inferior gynoecia topped with disc-shaped nectaries.

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Burmanniaceae is a family of flowering plants, consisting of 99 species of herbaceous plants in eight genera.

Moringa is the sole genus in the plant family Moringaceae. It contains 13 species, which occur in tropical and subtropical regions of Africa and Asia and that range in size from tiny herbs to massive trees. Moringa species grow quickly in many types of environments.

The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons.

Peridiscaceae is a family of flowering plants in the order Saxifragales. Four genera comprise this family: Medusandra, Soyauxia, Peridiscus, and Whittonia., with a total of 12 known species. It has a disjunct distribution, with Peridiscus occurring in Venezuela and northern Brazil, Whittonia in Guyana, Medusandra in Cameroon, and Soyauxia in tropical West Africa. Whittonia is possibly extinct, being known from only one specimen collected below Kaieteur Falls in Guyana. In 2006, archeologists attempted to rediscover it, however, it proved unsuccessful.

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 by a further revision, the APG III system.

The Kubitzki system is a system of plant taxonomy devised by Klaus Kubitzki, and is the product of an ongoing survey of vascular plants, entitled The Families and Genera of Vascular Plants, and extending to 15 volumes in 2018. The survey, in the form of an encyclopedia, is important as a comprehensive, multivolume treatment of the vascular plants, with keys to and descriptions of all families and genera, mostly by specialists in those groups. The Kubitzki system served as the basis for classification in Mabberley's Plant-Book, a dictionary of the vascular plants. Mabberley states, in his Introduction on page xi of the 2008 edition, that the Kubitzki system "has remained the standard to which other literature is compared".

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References