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. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
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 ''Myristica fragrans'' is an evergreen tree indigenous to the Maluku Islands of Indonesia. It is important as the main source of the spices nutmeg and mace. It is widely grown across the tropics including Guangdong and Yunnan in China, Taiwan, ...
'', 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, an ...
and mace. The best known genera are ''
Myristica ''Myristica'' is a genus of trees in the family Myristicaceae. There are over 150 species, distributed in Asia and the western Pacific. The type species of the genus, and the most economically important member, is ''Myristica fragrans'' (the nu ...
'' 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 are ...
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 le ...
'' 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 bioge ...
. 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, except ''
Endocomia ''Endocomia'' is a genus of flowering trees in the family Myristicaceae with a distribution ranging from Indochina to New Guinea. It is distinguished from the other Asian genera by its monoecious inflorescences which are unique in the mostly di ...
'' and some ''
Iryanthera ''Iryanthera'' is a flowering plant genus in the family Myristicaceae. Species include: *'' Iryanthera campinae'' W.A.Rodrigues *''Iryanthera coriacea'' Ducke *''Iryanthera crassifolia'' A.C. Sm. *''Iryanthera dialyandra'' Ducke *''Iryanthera el ...
''. 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 ''Knema'' is a genus of plant in family Myristicaceae, mostly consisting of small-medium trees found in lowland tropical forests from Asia to New Guinea. The highest diversity of species is in Borneo Borneo (; id, Kalimantan) is the thir ...
''). 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 ''Compsoneura'' is a genus comprising 23 species of trees found in tropical lowland forests of the New World.Tropicos, botanical information system at the Missouri Botanical Garden - http://www.tropicos.org/NameSearch.aspx?name=compsoneura It can ...
''). 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 published ...
of 2016, Myristaceae are placed in the order
Magnoliales The Magnoliales are an order of flowering plants. Classification The Magnoliales include six families: * Annonaceae (custard apple family, over 2000 species of trees, shrubs, and lianas; mostly tropical but some temperate) * Degeneriaceae (two ...
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 a ...
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'' de Wilde *'' Brochoneura'' Warburg *'' Cephalosphaera'' Warburg *'' Coelocaryon'' Warburg *''
Compsoneura ''Compsoneura'' is a genus comprising 23 species of trees found in tropical lowland forests of the New World.Tropicos, botanical information system at the Missouri Botanical Garden - http://www.tropicos.org/NameSearch.aspx?name=compsoneura It can ...
'' Warburg *''
Doyleanthus ''Doyleanthus'' is a monotypic genus of flowering plants belonging to the family Myristicaceae. The only species is ''Doyleanthus arillata''. Its native range is Madagascar. Description Similar in form to '' Mauloutchia'' in all vegetative and i ...
'' Sauquet *''
Endocomia ''Endocomia'' is a genus of flowering trees in the family Myristicaceae with a distribution ranging from Indochina to New Guinea. It is distinguished from the other Asian genera by its monoecious inflorescences which are unique in the mostly di ...
'' de Wilde *''
Gymnacranthera ''Gymnacranthera'' is a genus of flowering plants in the family Myristicaceae found from Peninsular Malaysia and Sumatra to New Guinea and the Philippines. Species Species include: #'' Gymnacranthera bancana'' (Miq. Friedrich Anton Wilhelm M ...
'' Warburg *''
Haematodendron ''Haematodendron'' is a monotypic genus of trees in the family Myristicaceae containing a single species, ''Haematodendron glabrum'', endemic to forests of east Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Mad ...
'' Capuron *''
Horsfieldia ''Horsfieldia'' is a genus of evergreen trees. The genus consists of about 100 species and is distributed across South Asia, from India to the Philippines and Papua New Guinea. Some species are used for timber. Species in the genus sometimes con ...
'' Willdenow *''
Iryanthera ''Iryanthera'' is a flowering plant genus in the family Myristicaceae. Species include: *'' Iryanthera campinae'' W.A.Rodrigues *''Iryanthera coriacea'' Ducke *''Iryanthera crassifolia'' A.C. Sm. *''Iryanthera dialyandra'' Ducke *''Iryanthera el ...
'' Warburg *''
Knema ''Knema'' is a genus of plant in family Myristicaceae, mostly consisting of small-medium trees found in lowland tropical forests from Asia to New Guinea. The highest diversity of species is in Borneo Borneo (; id, Kalimantan) is the thir ...
'' Loureiro *''
Mauloutchia ''Mauloutchia'' is a genus of trees endemic to the lowland eastern and northern rain forests of Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: Rép ...
'' (Baillon) Warburg *''
Myristica ''Myristica'' is a genus of trees in the family Myristicaceae. There are over 150 species, distributed in Asia and the western Pacific. The type species of the genus, and the most economically important member, is ''Myristica fragrans'' (the nu ...
'' Gronovius *''
Osteophloeum ''Osteophloeum'' is a genus of plant in family Myristicaceae. It contains a single tree species, ''Osteophloeum platyspermum'' native to Panama and South America South America is a continent entirely in the Western Hemisphere and mostl ...
'' Warburg *'' Otoba'' (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 New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , histo ...
'' de Wilde *'' Pycnanthus'' Warburg *'' Scyphocephalium'' Warburg *'' Staudtia'' 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 le ...
'' Aublet


Ecology

Myristicaceae are found in humid lowland forests, swamp forests, submontane forests, and cloud forests 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 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, a ...
is usually carried out by small beetles from the
Anthicidae The Anthicidae are a family of beetles that resemble ants. They are sometimes called ant-like flower beetles or ant-like beetles. The family comprises over 3,500 species in about 100 genera. Description Their heads constrict just in front of ...
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 stamens. However, ''Myristica'' is probably pollinated by true ants, a case of
myrmecophily Myrmecophily ( , ) is the term applied to positive interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to mutualistic associations with ants, though in its m ...
. 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 Myrmecophily ( , ) is the term applied to positive interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to mutualistic associations with ants, though in its m ...
, 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, an ...
and mace spices, both derived from respectively the seed and aril of ''
Myristica fragrans ''Myristica fragrans'' is an evergreen tree indigenous to the Maluku Islands of Indonesia. It is important as the main source of the spices nutmeg and mace. It is widely grown across the tropics including Guangdong and Yunnan in China, Taiwan, ...
'', 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'' 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 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 le ...
'', 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 categorized ...
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
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 sometimes used as a s ...
, palpitations, 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. Mil ...
, 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 oppose ...
, 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 b ...
is the earliest record of Myristicaceae.


References


External links


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