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The Myristicaceae are 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 ...
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 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 the ground spice derived from the seed, of several tree species of the genus '' Myristica''; fragrant nutmeg or true nutmeg ('' M. fragrans'') is a dark-leaved evergreen tree cultivated for two spices derived from its fru ...
and mace. The best known genera are '' Myristica'' in
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
and '' Virola'' 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 biogeog ...
. 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 certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, 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 The Magnoliales are an order of flowering plants. Well-known members of Magnoliales include: magnolias, Liriodendron, tulip trees, Annona squamosa, custard apples, Asimina triloba, American pawpaw, cherimoyas, Cananga odorata, ylang-ylang, Sours ...
in the
magnoliids Magnoliids, 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 angiospe ...
clade. The genera placed in the family are related as shown in the following cladogram:


List of genera

,
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 ...
accepted the following genera:Myristiceae R.Br.
''
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 ...
''. Retrieved 18 November 2023.
*'' Bicuiba'' J.J.de Wilde *'' Brochoneura'' Warb. *'' Coelocaryon'' Warb. *'' Compsoneura'' (DC.) Warb. *'' Doyleanthus'' Sauquet *'' Endocomia'' W.J.de Wilde *'' Gymnacranthera'' Warb. *'' Haematodendron'' Capuron *'' Horsfieldia'' Willd. *'' Iryanthera'' (A.DC.) Warb. *'' Knema'' Lour. *'' Mauloutchia'' Warb. *'' Myristica'' Gronov. *'' Neobrochoneura'' *'' Osteophloeum'' Warb. *'' Otoba'' (DC.) H.Karst. *'' Paramyristica'' W.J.de Wilde *'' Pycnanthus'' Warb. *'' Scyphocephalium'' Warb. *'' Staudtia'' Warb. *'' Virola'' Aubl.


Ecology

Myristicaceae are found in humid lowland forests, swamp forests, submontane forests, and
cloud forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest, is a generally tropical or subtropical, evergreen, Montane forest, montane, Tropical and subtropical moist broadleaf forests, moist forest characteri ...
s 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 Anthesis is the period during which a flower is fully open and functional. It may also refer to the onset of that period. The onset of anthesis is spectacular in some species. In ''Banksia'' species, for example, anthesis involves the extension ...
at night, and
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
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 (: 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 ...
s. However, ''Myristica'' is probably pollinated by true ants, a case of myrmecophily. A few
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
''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''.


Uses

The most important products of the family by far are the
nutmeg Nutmeg is the seed, or the ground spice derived from the seed, of several tree species of the genus '' Myristica''; fragrant nutmeg or true nutmeg ('' M. fragrans'') is a dark-leaved evergreen tree cultivated for two spices derived from its fru ...
and mace spices, both derived from respectively the seed and aril of '' Myristica fragrans'', a tree native to the
Moluccas The Maluku Islands ( ; , ) or the Moluccas ( ; ) are an archipelago in the eastern part of Indonesia. Tectonically they are located on the Halmahera Plate within the Molucca Sea Collision Zone. Geographically they are located in West Melanesi ...
. 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 firs ...
) that is used by certain Amazonian tribes is obtained from the bark of '' Virola elongata'' and other closely related species. The wood of some Asian and American species has local commercial use, as is the case of '' Otoba parvifolia'' 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 ...
action and antimicrobial activity against ''
Streptococcus mutans ''Streptococcus mutans'' is a Facultative anaerobic organism, facultatively anaerobic, gram-positive coccus (round bacteria, bacterium) commonly found in the human oral cavity and is a significant contributor to dental caries, tooth decay. The m ...
''. The dark-red
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 ...
of the tree bark in some genera, such as '' Virola'', contains several hallucinogenic
alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...
s. Myristicin 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 often used as a synony ...
,
palpitations Palpitations occur when a person becomes aware of their heartbeat. The heartbeat may feel hard, fast, or uneven in their chest. Symptoms include a very fast or irregular heartbeat. Palpitations are a sensory symptom. They are often described as ...
, nausea, eventual
dehydration In physiology, dehydration is a lack of total body water that disrupts metabolic processes. It occurs when free water loss exceeds intake, often resulting from excessive sweating, health conditions, or inadequate consumption of water. Mild deh ...
, 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 such as LSD and dissociatives such as ketamine, due to their primary effect of causing delirium, as opposed to th ...
, and some fatal myristicin poisonings in humans have occurred.


Fossil record

A fossil seed of †''Myristicacarpum chandlerae'' from the early
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
London Clay The London Clay Formation is a Sediment#Shores and shallow seas, marine formation (geology), geological formation of Ypresian (early Eocene Epoch, c. 54-50 million years ago) age which outcrop, crops out in the southeast of England. The London C ...
flora of southern
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
is the earliest record of Myristicaceae.


References


External links


e-floras
* * {{Authority control Magnoliid families Dioecious plants