Chloranthaceae is a

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of ...

s (angiosperms), the only family in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

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 (geochronological name) or the Lower Cretaceous ( chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma. Geology Pr ...

, and have been found on all continents.

Description

Chloranthaceae are fragrant shrubs 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 ...

s, that only produce 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). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...

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

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...

s are seated directly on the axis of the

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. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed ...

Petal Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corolla''. Petals are usuall ...

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 The term ''sepalum'' was coined ...

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

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

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 are strawberries, rasp ...

, consisting of one

carpel Gynoecium (; ) 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) '' pistils' ...

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 with wood without vessels, that have bisexual flowers, with only one, club-shaped stamen in which the connective tissue (between the lobes which carry the

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...

) 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 in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived fr ...

that does contain vessels, that has bisexual flowers, each of which containing three stamens on straight, filaments with three lobes and a wide connective, and with a smooth and moist stigma. The twenty species occur in southern and eastern Asia. ''Ascarina'' has separate male and female flowers. The male flowers are subtended by two bracts 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 Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...

. The fruit is a drupe-like berry. Twelve species can be found on islands in the Pacific and insular South-East Asia, from New Zealand and the

Marquesas The Marquesas Islands (; french: Îles Marquises or ' or '; Marquesan: ' (North Marquesan) and ' (South Marquesan), both meaning "the land of men") are a group of volcanic islands in French Polynesia, an overseas collectivity of France in ...

Borneo Borneo (; id, Kalimantan) is the List of islands by area, third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java Isl ...

, and on

Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...

. ''Hedyosmum'' has separate male and female flowers. The male flowers are without bracts and has one stamen in which the connective is not widened. The female flower is without bracts, the stigma is dry and covered in

. Female flowers have a trilobed

calyx Calyx or calyce (plural "calyces"), from the Latin ''calix'' which itself comes from the Ancient Greek ''κάλυξ'' (''kálux'') meaning "husk" or "pod", may refer to: Biology * Calyx (anatomy), collective name for several cup-like structures ...

. The fruit has a kernel with a hard and woody shell (a

). Forty three species are known from Latin America, including the

Antilles The Antilles (; gcf, label=Antillean Creole, Antiy; es, Antillas; french: Antilles; nl, Antillen; ht, Antiy; pap, Antias; Jamaican Patois: ''Antiliiz'') is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mex ...

, and one species lives in Southeast Asia.

Taxonomy

Chloranthaceae have been recognised as a family in most classifications but without clear relatives. Molecular systematic studies have shown that it is not closely related to any other family and is among the early-diverging lineages in the

angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of br ...

. In particular, it is neither a

eudicot 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. Traditionally they were called tricolpates or non- magnoliid dic ...

nor a

monocot Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute on ...

. Fossils assigned to Chloranthaceae, or closely related to the family, are among the oldest angiosperms known. 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-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Gr ...

(2003) left the family unplaced as to order, but 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 fu ...

(2009) accepted

Chloranthales Chloranthaceae is a family of flowering plants (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 ...

, containing only this family. The cladogram below, from the

APG IV system The APG IV system of flowering plant classification is the fourth version of a modern, mostly molecular-based, system of plant taxonomy for flowering plants (angiosperms) being developed by the Angiosperm Phylogeny Group (APG). It was publish ...

(2016), shows the Chloranthales in a

trichotomy A trichotomy can refer to: * Law of trichotomy, a mathematical law that every real number is either positive, negative, or zero ** Trichotomy theorem, in finite group theory * Trichotomy (jazz trio), Australian jazz band, collaborators with Dan ...

with the

magnoliids Magnoliids (or Magnoliidae or Magnolianae) are a clade of flowering plants. With more than 10,000 species, including magnolias, nutmeg, bay laurel, cinnamon, avocado, black pepper, tulip tree and many others, it is the third-largest group of ang ...

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

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...

, 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''. , four extant genera are recognized, and insights in their relationships are expressed in the following tree. The extinct genus '' Chloranthistemon'' also belongs to this family.

Historical classifications

The

Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) ...

(1981) assigned the family : to the order

Piperales Piperales is an order of flowering plants (4,170 recognized species). It necessarily includes the family Piperaceae but other taxa have been included or disincluded variously over time. Well-known plants which may be included in this order inclu ...

:: in subclass Magnoliidae ::: in class Magnoliopsida dicotyledons">dicotyledons.html" ;"title="dicotyledons">dicotyledons :::: of division Magnoliophyta

The

Thorne system (1992) A system of plant taxonomy, the Thorne system of plant classification was devised by the American botanist Robert F. Thorne (1920–2015) in 1968, and he continued to issue revisions over many years (1968–2007). Some versions of the system are ...

placed it : in the order Magnoliales, which was assigned :: to superorder Magnolianae ::: in subclass Magnoliideae

dicotyledons The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...

:::: in class Magnoliopsida

The

Dahlgren system One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 1 ...

raised the family to be : its own order

, which was assigned :: to superorder Magnolianae ::: in subclass Magnoliideae

:::: in class Magnoliopsida

References

External links

Chloranthaceae
in L. Watson and M.J. Dallwitz (1992 onwards)
The families of flowering plants:
descriptions, illustrations, identification, information retrieval''. Version: 3 May 2006. https://web.archive.org/web/20070103200438/http://delta-intkey.com/.
''Andes Trees''

NCBI Taxonomy Browser

Flavon's art gallery: Chloranthaceae
Image of ''Chloranthus oldhamii''

Images of ''Sarcandra glabra''
{{Taxonbar, from=Q133208 Angiosperm families