The Cleomaceae are a small family of
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s in the order
Brassicales
The Brassicales (or Cruciales) are an order (biology), order of flowering plants, belonging to the malvid group of eudicotyledons under the APG IV system. Well-known members of Brassicales include cabbage, cauliflower, Brussels sprout, broccoli, ...
, comprising about 220 species in two genera, ''
Cleome'' and ''
Cleomella''.
[ These genera were previously included in the family ]Capparaceae
The Capparaceae (or Capparidaceae), commonly known as the caper family, are a family of plants in the order Brassicales. As currently circumscribed, the family contains 15 genera and about 430 species. The largest genera are ''Capparis'' (about 1 ...
, but were raised to a distinct family when DNA
Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
evidence suggested the genera included in it are more closely related to the Brassicaceae
Brassicaceae () or (the older but equally valid) Cruciferae () is a medium-sized and economically important Family (biology), family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous pla ...
than they are to the Capparaceae. 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 ...
allows for Cleomaceae to be included in Brassicaceae.[Stevens, P. F. (2001 onwards) Angiosperm Phylogeny Website]
Brassicales
Cleomaceae includes C3, C3–C4, and C4 photosynthesis species.
Taxonomy
In 1994, a group of scientists including Walter Stephen Judd suggested to merge the Capparaceae
The Capparaceae (or Capparidaceae), commonly known as the caper family, are a family of plants in the order Brassicales. As currently circumscribed, the family contains 15 genera and about 430 species. The largest genera are ''Capparis'' (about 1 ...
(which at that time included the Cleomaceae) with the Brassicaceae. Early DNA-analysis showed that the Capparaceae - as defined at that moment - were paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, and others suggested to assign the genera closest to the Brassicaceae to the Cleomaceae. The Cleomaceae and Brassicaceae diverged approximately 41 million years ago. All three families have consistently been placed in one order (variably called Capparales or Brassicales
The Brassicales (or Cruciales) are an order (biology), order of flowering plants, belonging to the malvid group of eudicotyledons under the APG IV system. Well-known members of Brassicales include cabbage, cauliflower, Brussels sprout, broccoli, ...
). The APG II system merged Cleomaceae and Brassicaceae. Other classifications have continued to recognise the Capparaceae, but with a more restricted circumscription, either including ''Cleome'' and its relatives in the Brassicaceae or recognizing them in the segregate family Cleomaceae. 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 ...
has recently adopted this last solution, but this may change as a consensus arises on this point. Current insights in the relationships of the Brassicaceae, based on a 2012 DNA-analysis, are summarized in the following tree.
Subdivisions
Two genera are currently accepted, '' Cleome'' with 199 species, and '' Cleomella'' with 22 species. Plants of the World Online
Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew.
History
Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
considers '' Oxystylis'', '' Peritoma'', and '' Wislizenia'' synonyms of ''Cleomella''.[
]
Description
Cleomaceae are mostly annual, sometimes perennial
In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
herbaceous plant
Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials.
Definitions of "herb" and "herbaceous"
The fourth edition o ...
s, seldom shrubs or small trees, and some are liana
A liana is a long-Plant stem, stemmed Woody plant, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the Canopy (biology), canopy in search of direct sunlight. T ...
s. Most of them have glands
A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...
, a character setting them apart from the related Capparaceae
The Capparaceae (or Capparidaceae), commonly known as the caper family, are a family of plants in the order Brassicales. As currently circumscribed, the family contains 15 genera and about 430 species. The largest genera are ''Capparis'' (about 1 ...
, and contain resin
A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Commo ...
s, and may have an aromatic or foetid smell. The alternately set leaves, have stalks and never form a sheath around the stem. The leaves are always palmately compound, mostly with three to seven leaflets, rarely with just one leaflet. At the base of the leafstalk sit scaly, leafy or spiny stipule
In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole (botany), petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part ...
s, a difference with its sister group
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and ...
Brassicaceae
Brassicaceae () or (the older but equally valid) Cruciferae () is a medium-sized and economically important Family (biology), family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous pla ...
that consistently lack stipules, although a pair of glands can sometimes be present at the base of the leaf stalk. Hairs and papillae may be present.
The flowers of Cleomaceae are somewhat to strongly bilateral symmetric. There are mostly four, sometimes two or six, free or merged sepal
A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106
Etymology
The term ''sepalum'' ...
s, that may be shed quickly or remain, are overlapping in the bud or are already separate. The corolla consists of mostly four, sometimes two or six petal
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 ''corol ...
s that have distinctive claws and limbs, free or with merged limbs. The 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 ...
may be placed on the level of insertion of the perianth
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 tepal ...
or on a stalk, raising the bases of the stamens above the perianth. This occurs for instance in '' Gynandropsis gynandra''. There are four simple stamens or branched in four groups of up to 12 stamens each that have a common base, and are free from the petals. When there are groups of stamens, the anthers
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 ...
mature from the middle outwards. The base of the stamens may or may not be fused with the female parts in the centre of the flower. The filaments may be of equal length or strongly differ, and all may carry anthers, but infertile staminode
In botany, a staminode is an often rudimentary, sterile or abortive stamen, which means that it does not produce pollen.Jackson, Benjamin, Daydon; ''A Glossary of Botanic Terms with their Derivation and Accent''; Published by Gerald Duckworth & Co ...
s may also occur. The anther releases 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 ...
through slits stretching along its length. The ovary is positioned above the perianth and the 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 ...
(a so-called superior ovary), consists of two fully merged carpel
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 of a flower; it consists of (one or more ...
s, and contains one cavity that may be divided in two by a partition (or replum). Between ten and fifty ovule
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the sporangium, megasporangium), ...
s (future seeds) are set along the margins of each of the carpels. As the fruit ripens, a stalk (or gynophore) develops. The dry fruit opens with two valves. When the fruit has a replum, it has the same basic morphology as the Brassicaceae (called silicula or siliqua), and this sets it apart from the Capparaceae, that have a capsule.
References
{{Taxonbar, from=Q132347
Brassicales families