Chloranthaceae ( ) is a

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

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 (angiosperms), the only family in the order Chloranthales. It is not closely related to any other family of flowering plants, and is among the early-diverging lineages in the angiosperms. They are woody or weakly woody plants occurring in Southeast Asia, the Pacific, Madagascar, Central and South America, and the West Indies. The family consists of four extant genera, totalling about 77 known species according to Christenhusz and Byng in 2016. Some species are used in traditional medicine. The type genus is '' Chloranthus''. The fossil record of the family, mostly represented by pollen such as ''Clavatipollenites,'' extends back to the dawn of the history of flowering plants in the

Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...

, and has been found on all continents.

Description

Chloranthaceae are

fragrant An aroma compound, also known as an odorant, aroma, fragrance, flavoring or flavor, is a chemical compound that has a smell or odor. For an individual chemical or class of chemical compounds to impart a smell or fragrance, it must be sufficie ...

shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...

s or

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, that only produce new side branches on the new growth. The stems are mostly cylindrical, with solid internodes, thickened nodes in many species, that carry evergreen leaves arranged in pairs on opposite sides of the stem, with

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 that have merged with that of the opposing leaf. The small

flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...

s are seated directly on the axis of the

inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...

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 are absent in this family, and sometimes so are

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. The flowers can be either hermaphrodite or of separate sexes. The fruit is a

drupe In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...

berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit although many pips or seeds may be present. Common examples of berries in the cul ...

, consisting of one

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 ...

Differences between the genera

The four genera assigned to this family can be distinguished from each other by the following characters: '' Sarcandra'' species are shrubs whose wood lacks vessels. They have bisexual flowers, with only one, club-shaped,

stamen 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 ...

, in which the connective tissue (between the

anther 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 ...

lobes which carry the pollen) is wide, and with a smooth and moist stigma. This genus has four species, which occur in Malaysia, China, Indochina, Japan, India, and Sri Lanka. '' Chloranthus'' species are dwarf shrubs or

s, with

xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...

that contains vessels. Flowers are bisexual, each of which bears three stamens on straight filaments with three anther lobes and a wide connective, and with a smooth and moist stigma. The 20 species occur in southern and eastern Asia. '' Ascarina'' has separate male and female flowers. The male flowers are subtended by two

bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...

s and have between one and five stamens, in which the connective is not widened. The female flower is without bracts, the stigma is dry and covered in papillae. The fruit is a

-like

. There are 12 species, which occur on islands in the Pacific and insular South-East Asia, from New Zealand and the Marquesas to

Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...

, and on

Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...

. '' Hedyosmum'' has separate male and female flowers. The male flowers are without bracts and have one stamen, in which the connective is not widened. The female flower is without bracts; the stigma is dry and covered in papillae. Female flowers have a 3-lobed calyx. The fruit, a drupe, has a kernel with a hard and woody shell. There are 43 species found in Latin America, including the

Antilles The Antilles is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mexico to the northwest, and the Atlantic Ocean to the north and east. The Antillean islands are divided into two smaller groupings: the Greater An ...

, as well as one species found in Southeast Asia.

Taxonomy

The Chloranthaceae have been recognised as a family in most classifications but without clear relatives. Molecular systematics studies have shown that it is not closely related to any other family and is among the early-diverging lineages in the angiosperms. In particular, it is neither a eudicot nor a

monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are flowering plants whose seeds contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but ...

. Fossils assigned to Chloranthaceae, or closely related to the family, are among the oldest angiosperms known. The APG II system (2003) left the family unplaced as to order, but the APG III system (2009) accepted Chloranthales, containing only this family. The cladogram below, from the APG IV system (2016), shows the Chloranthales in a trichotomy with the magnoliids and the monocot-Ceratophyllales-dicot

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 ...

. Earlier, the order was grouped with magnoliids, but studies in 2014 did not support this placement, leaving its phylogeny unclear. A 2021 study sequenced the ''Chloranthus'' genome and found Chloranthales as sister to magnoliids. A 2004 study based on comparisons of homologous DNA fragments indicated that both the family Chloranthaceae and its extant genera ''Ascarina'', ''Chloranthus'', ''Hedyosmum'' and ''Sarcandra'' are probably

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 ...

, with ''Hedyosmum'' being the first to diverge from the rest, and ''Ascarina'' being the

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 ...

of the clade consisting of ''Sarcandra'' and ''Chloranthus''. , these four extant genera are recognized, and insights into their relationships are expressed in the

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...

below: The extinct genus '' Chloranthistemon'' also belongs to this family.

Historical classifications

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 ...

(1981) assigned the family : to the order Piperales :: in subclass Magnoliidae ::: in class Magnoliopsida dicotyledons">dicotyledons.html" ;"title="dicotyledons">dicotyledons :::: of division Magnoliophyta angiosperms The Thorne system (1992) placed it : in the order Magnoliales, which was assigned :: to superorder Magnolianae ::: in subclass Magnoliideae dicotyledons :::: in class Magnoliopsida angiosperms The Dahlgren system raised the family to be : its own order Chloranthales, which was assigned :: to superorder Magnolianae ::: in subclass Magnoliideae dicotyledons :::: in class Magnoliopsida angiosperms

Description

Differences between the genera

Taxonomy

Historical classifications

References

External links