Myristicaceae
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The Myristicaceae are 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 ...
of
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 native to Africa, Asia, Pacific islands, and the Americas and has been recognized by most taxonomists. It is sometimes called the "nutmeg family", after its most famous member, '' Myristica fragrans'', the source of the spices
nutmeg Nutmeg is the seed or ground spice of several species of the genus '' Myristica''. '' Myristica fragrans'' (fragrant nutmeg or true nutmeg) is a dark-leaved evergreen tree cultivated for two spices derived from its fruit: nutmeg, from its seed, ...
and mace. The best known genera are '' Myristica'' in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an ...
and ''
Virola ''Virola'' is a genus of medium-sized trees native to the South American rainforest and closely related to other Myristicaceae, such as nutmeg. Species are known commonly as ''epená'', ''patricá'', or ''cumala''. They have glossy, dark green ...
'' in the
Neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bi ...
. The family consists of about 21 genera with about 520 species of trees, shrubs and rarely lianas ('' Pycnanthus'') found in tropical forests around the world. Most of the species are large trees that are valued in the timber industry.


Description

They are typically trees with reddish sap and distinctive pagoda-like growth (known as myristicaceous branching) in which horizontal branching only occurs at certain nodes along the main axis of the trunk, each node separated by a large gap where no branching occurs. All genera are
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproducti ...
, except '' Endocomia'' and some '' Iryanthera''. The inner bark is usually pink to reddish or light colored then oxidizing as such. When cut, the tree trunk exudes a red or orange resin; stems and young twigs often will exude clear sap (not colored) that may smell spicy. The foliage is generally spicy-aromatic and the leaves are glossy, dark green, simple, entire, 2-ranked, undersides often whitish or tomentose, with dark brown punctations or not, usually with complex caducous hairs colored golden yellow to red. The flowers are usually small, highly reduced, fragrant, with 3-5 tepals, inner perianth whitish-green, yellow, orange, reddish-pink to rusty-brown, arranged in axillary paniculate inflorescences or unbranched wart-like structures (like '' Knema''). The female flowers are without staminodes, with stigmas often lobed. The male flowers with fused stamens arranged in a synandrium. Pollen is monosulcate, often boat-shaped. The fruit is a leathery dehiscent capsule, with rusty indument or not, containing a single seed that is arillate or not; when present, the aril variously laciniate or entire. In most genera, the aril is colored red but also can be orange or white and translucent. The single seed has ruminate endosperm and is uniform in color or rarely with black blotches ('' Compsoneura''). Many species within the family exhibit highly complex phytochemistry with numerous compounds having been described from the leaves, bark, fruits, arils, and seeds of many species.


Taxonomy

In 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 ...
of 2016, Myristaceae are placed in the order Magnoliales in 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 ...
clade.


List of genera

, the
Angiosperm Phylogeny Website The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis. ...
accepted the following genera: *''
Bicuiba ''Bicuiba'' is a monotypic genus of flowering plants in the nutmeg family, Myristicaceae. The only species is ''Bicuiba oleifera'', which is endemic to southeastern Brazil. It grows in the forests of the Atlantic The Atlantic Ocean is the ...
'' de Wilde *'' Brochoneura'' Warburg *'' Cephalosphaera'' Warburg *''
Coelocaryon ''Coelocaryon'' is a genus of flowering plants in the family Myristicaceae. It is native to tropical Africa. Species According to Kew's Plants of the World Online Plants of the World Online (POWO) is an online database published by the Ro ...
'' Warburg *'' Compsoneura'' Warburg *'' Doyleanthus'' Sauquet *'' Endocomia'' de Wilde *'' Gymnacranthera'' Warburg *'' Haematodendron'' Capuron *'' Horsfieldia'' Willdenow *'' Iryanthera'' Warburg *'' Knema'' Loureiro *'' Mauloutchia'' (Baillon) Warburg *'' Myristica'' Gronovius *'' Osteophloeum'' Warburg *''
Otoba ''Otoba'' is a genus of trees in family Myristicaceae that ranges from Nicaragua to Bolivia. Selected species According to Tropicos and Plants of the World Online Plants of the World Online (POWO) is an online database published by the ...
'' (A. de Candolle) H. Karsten *''
Paramyristica ''Paramyristica'' is a monotypic genus of flowering plants belonging to the family Myristicaceae. The only species is ''Paramyristica sepicana''. Its native range is New Guinea. References {{Taxonbar, from=Q16739307 Myristicaceae Myrist ...
'' de Wilde *'' Pycnanthus'' Warburg *''
Scyphocephalium ''Scyphocephalium'' is a genus of plants in family Myristicaceae native to west Africa, containing two species. One species contains chemicals with significant in vitro antibacterial activity. Species '' Scyphocephalium mannii'' (Benth. & Hook ...
'' Warburg *''
Staudtia ''Staudtia'' is a genus of plant in family Myristicaceae. It contains the following species (but this list may be incomplete): *'' Staudtia pterocarpa'' (Warb.) Warb. *''Staudtia stipitata Staudtia stipitata is a species of plant in the fam ...
'' Warburg *''
Virola ''Virola'' is a genus of medium-sized trees native to the South American rainforest and closely related to other Myristicaceae, such as nutmeg. Species are known commonly as ''epená'', ''patricá'', or ''cumala''. They have glossy, dark green ...
'' Aublet


Ecology

Myristicaceae are found in humid lowland forests, swamp forests, submontane forests, and
cloud forests A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level clo ...
at elevations up to 2100 m. Some of the anatomical characters presented by this family suggest that in the past they could live in xeric (dry) environments, but now their species are linked to tropical rainforests. The species present anthesis at night, and
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
is usually carried out by small beetles from the Anthicidae family that resemble ants and consume pollen (e.g., ''Myristica fragrans'' is probably pollinated by the beetle '' Formicomus braminus''). The strong floral scent that attracts beetles emerges from the ends of the connectives of the
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s. However, ''Myristica'' is probably pollinated by true ants, a case of myrmecophily. A few
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torr ...
''Myristica'' species have evolved hollow stem swellings in which ants reside. This facilitates a mutualistic relationship known as myrmecophily, and is similar to that of ''
Cecropia ''Cecropia'' is a Neotropical genus consisting of 61 recognized species with a highly distinctive lineage of dioecious trees. The genus consists of pioneer trees in the more or less humid parts of the Neotropics, with the majority of the speci ...
''.


Uses

The most important products of the family by far are the
nutmeg Nutmeg is the seed or ground spice of several species of the genus '' Myristica''. '' Myristica fragrans'' (fragrant nutmeg or true nutmeg) is a dark-leaved evergreen tree cultivated for two spices derived from its fruit: nutmeg, from its seed, ...
and
mace Mace may refer to: Spices * Mace (spice), a spice derived from the aril of nutmeg * '' Achillea ageratum'', known as English mace, a flowering plant once used as a herb Weapons * Mace (bludgeon), a weapon with a heavy head on a solid shaft used ...
spices, both derived from respectively the seed and aril of '' Myristica fragrans'', a tree native to the
Moluccas The Maluku Islands (; Indonesian: ''Kepulauan Maluku'') or the Moluccas () are an archipelago in the east of Indonesia. Tectonically they are located on the Halmahera Plate within the Molucca Sea Collision Zone. Geographically they are located ...
. A hallucinogenic snuff (containing a derivative of
tryptamine Tryptamine is an indolamine metabolite of the essential amino acid, tryptophan. The chemical structure is defined by an indole ─ a fused benzene and pyrrole ring, and a 2-aminoethyl group at the second carbon (third aromatic atom, with the f ...
) that is used by certain Amazonian tribes is obtained from the bark of ''
Virola elongata ''Virola elongata'' (syn. ''Virola theiodora '' ) is a species of tree in the family Myristicaceae. The tree is native to Panama, Guyana, Brazil (Acre, Amazonas, Mato Grosso, Pará, Rondônia and Roraima), Bolivia, Colombia, Ecuador and Peru. ...
'' and other closely related species. The wood of some Asian and American species has local commercial use, as is the case of ''
Otoba parvifolia ''Otoba'' is a genus of trees in family Myristicaceae that ranges from Nicaragua to Bolivia. Selected species According to Tropicos and Plants of the World Online Plants of the World Online (POWO) is an online database published by the ...
'' in South America.


Toxicity

Essential oils of Myristicaceae have
antifungal An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as cry ...
action and antimicrobial activity against '' Streptococcus mutans''. The dark-red
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on n ...
of the tree bark in some genera, such as ''
Virola ''Virola'' is a genus of medium-sized trees native to the South American rainforest and closely related to other Myristicaceae, such as nutmeg. Species are known commonly as ''epená'', ''patricá'', or ''cumala''. They have glossy, dark green ...
'', contains several
hallucinogenic Hallucinogens are a large, diverse class of psychoactive drugs that can produce altered states of consciousness characterized by major alterations in thought, mood, and perception as well as other changes. Most hallucinogens can be categorize ...
alkaloid Alkaloids are a class of basic BASIC (Beginners' All-purpose Symbolic Instruction Code) is a family of general-purpose, high-level programming languages designed for ease of use. The original version was created by John G. Kemeny and Th ...
s.
Myristicin Myristicin is a naturally occurring compound found in common herbs and spices, the most well known being nutmeg. It is an insecticide, and has been shown to enhance the effectiveness of other insecticides in combination. Myristicin is also a pre ...
poisoning can induce
convulsions A convulsion is a medical condition where the body muscles contract and relax rapidly and repeatedly, resulting in uncontrolled shaking. Because epileptic seizures typically include convulsions, the term ''convulsion'' is sometimes used as a ...
,
palpitations Palpitations are perceived abnormalities of the heartbeat characterized by awareness of cardiac muscle contractions in the chest, which is further characterized by the hard, fast and/or irregular beatings of the heart. Symptoms include a rapi ...
, nausea, eventual
dehydration In physiology, dehydration is a lack of total body water, with an accompanying disruption of metabolic processes. It occurs when free water loss exceeds free water intake, usually due to exercise, disease, or high environmental temperature. Mi ...
, and generalized body pain. It is also reputed to be a strong
deliriant Deliriants are a subclass of hallucinogen. The term was coined in the early 1980s to distinguish these drugs from psychedelics and dissociatives such as LSD and ketamine, respectively, due to their primary effect of causing delirium, as opposed ...
, and some fatal myristicin poisonings in humans have occurred.


Fossil record

A fossil seed of †''Myristicacarpum chandlerae'' from the early
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
London Clay The London Clay Formation is a marine geological formation of Ypresian (early Eocene Epoch, c. 56–49 million years ago) age which crops out in the southeast of England. The London Clay is well known for its fossil content. The fossils from t ...
flora of southern
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe ...
is the earliest record of Myristicaceae.


References


External links


e-floras
* * {{Taxonbar, from=Q161214 Magnoliid families Dioecious plants