Moreae
Encyclopedia
Moreae is a tribe
Tribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...

 within the plant family Moraceae
Moraceae
Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 40 genera and over 1000 species. Most are widespread in tropical and subtropical regions, less so in temperate climates...

. It includes 6–10 genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 and 70–80 species including Morus, the genus the includes the mulberries, and Maclura
Maclura
Maclura is a genus of flowering plants in the mulberry family, Moraceae. It includes the inedible Osage-orange, which is used as mosquito repellent and grown throughout the United States as a hedging plant.- Species :* Maclura africana...

, the genus that includes the Osage orange.

Recent work suggests that the tribe is polyphyletic.

Description

The Moreae are a tribe of trees, shrubs, climbers and herbs that are usually dioecious
Dioecious
Dioecy is the property of a group of biological organisms that have males and females, but not members that have organs of both sexes at the same time. I.e., those whose individual members can usually produce only one type of gamete; each individual organism is thus distinctly female or male...

. Their inflorescence
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. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

s are simpler than most other Moraceae. Their flowers are adapted for wind pollination
Anemophily
Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Anemophilous plants may be either gymnosperms or angiosperms ....

.

Members of the tribe are characterised by having inflexed or "urticaceous
Urticaceae
Urticaceae, or the nettle family, is a family of flowering plants. The family name comes from the genus Urtica . Urticaceae includes a number of well-known and useful plants, including the aforementioned nettles, Ramie , māmaki , and ajlai .The family includes approximately 2600 species, grouped...

" stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...

s. The way in which the anther filaments elongate in the developing flower bud causes the anthers to "spring back explosively" when the flower opens. This releases pollen into the air, facilitating wind-pollination. These characters are considered plesiomorphic—traits that were present in the ancestors of the Moraceae
Moraceae
Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 40 genera and over 1000 species. Most are widespread in tropical and subtropical regions, less so in temperate climates...

 which have been retained in the Moreae.

Taxonomy

The tribe is based on Morus, the genus that includes the mulberries. The name Morus was first published by Carl Linnaeus in 1753 in Species Plantarum
Species Plantarum
Species Plantarum was first published in 1753, as a two-volume work by Carl Linnaeus. Its prime importance is perhaps that it is the primary starting point of plant nomenclature as it exists today. This means that the first names to be considered validly published in botany are those that appear...

.

Cornelis Berg included eight genera in the tribe Moreae—Bleekrodea, Broussonetia
Broussonetia
Broussonetia is a genus of four species of trees in the family Moraceae, native to eastern Asia.-References:*...

, Fatoua, Maclura
Maclura
Maclura is a genus of flowering plants in the mulberry family, Moraceae. It includes the inedible Osage-orange, which is used as mosquito repellent and grown throughout the United States as a hedging plant.- Species :* Maclura africana...

, Milicia
Milicia
Milicia may refer to:* the Milicia is the Sicilian river that has its mouth at Palermo.* Milicia is a genus in the plant family Moraceae.* an incorrect spelling of militia...

, Morus
Morus
Morus may refer to:* Morus , a genus of trees* Morus , a genus of seabird in the family Sulidae* Thomas Morus, alias Thomas More, a philosopher...

, Streblus
Streblus
Streblus is a genus of flowering plant in the mulberry family, Moraceae.*Streblus asper Lour. - Siamese rough bush*Streblus banksii*Streblus brunonianus*Streblus elongatus*Streblus heterophyllus...

and Trophis
Trophis
Trophis is a genus in the plant family Moraceae which includes about nine species, six of which are Neotropical and three which are Palaeotropical.Recent work suggests that the genus is polyphyletic....

—which included 73 species.

About 40 generic names have been coined for these species, resulting in a situation with many genera and few species. In 1962 British botanist E. J. H. Corner
E. J. H. Corner
Edred John Henry Corner FRS was a botanist who occupied the posts of assistant director at the Singapore Botanic Gardens and Professor of Tropical Botany at the University of Cambridge...

 reworked this arrangement, merging many genera. This resulting in an expansion of several genera, most notably Maclura, Streblus and Trophis. These were further reorganised by Berg in the 1980s. He found that while it was easy to define the tribe (based on the presence of urticaceous stamens), the definitions of the genera were more challenging.

In an attempt to sort out the evolutionary history of the Moraceae
Moraceae
Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 40 genera and over 1000 species. Most are widespread in tropical and subtropical regions, less so in temperate climates...

 Shannon Datwyler and George Weiblen use the chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...

 ndhF gene to build a phylogeny of the family. They determined that the tribe Moreae was polyphyletic and consisted of two groups—a core "Moreae sensu stricto" (Moreae s.s.), which formed a monophyletic group within the tribe, and a broader "Moreae sensu lato (Moreae s.l.). Streblus and Trophis were found to be polyphyletic, with some species in each genus being placed in Moreae s.s while others were only included in Moreae s.l. They also transferred Bagassa
Bagassa
Bagassa guianensis is a tree in the plant family Moraceae which is native to the Guianas and Brazil. It is valued as a timber tree and as a food tree for wildlife...

and Sorocea
Sorocea
Sorocea is a genus of plant in family Moraceae.Species include:* Sorocea guilleminiana, Gaudichaud* Sorocea sarcocarpa, Lanj. & Wess. Boer...

from the Artocarpeae
Artocarpeae
Artocarpeae is a tribe within the plant family Moraceae. It includes 7 to 12 genera and 70 to 87 species including Artocarpus altilis, the breadfruit.-Description:...

 to the Moreae.

Based on chloroplast ndhF gene sequences, Moreae s.s. is a sister taxon to the Artocarpeae if Bagassa and Sorocea are included in the Moreae. So defined, Moreae s.s. includes Bagassa, Morus, Milicia and Sorocea, together with some (but not all) of the species currently placed in Streblus and Trophis. Moreae s.l. included all of these genera, together with Bleekrodea, Broussonetia, Fatoua, Maclura and the remaining species in Streblus and Trophis. The latter group was found to be a sister to the tribe Dorstenieae
Dorstenieae
Dorstenieae is a tribe within the plant family Moraceae. The tribe includes eight genera and about 120 species.-Genera:*Brosimum - Breadnut*Bosqueiopsis*Dorstenia*Helianthostylis*Scyphosyce*Trilepisium*Trymatococcus...

.

Evolutionary history

Few fossils can be definitively assigned to the Moreae. Fruits from the upper Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...

, Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...

 and Tertiary
Tertiary
The Tertiary is a deprecated term for a geologic period 65 million to 2.6 million years ago. The Tertiary covered the time span between the superseded Secondary period and the Quaternary...

 have been assigned to Broussonetia, and fruit matching modern Morus have been found in the early Eocene. Based on rates of molecular evolution in chloroplast and nuclear genes, the Moreae s.s. is estimated to be about 59–79 million years old. Nyree Zerega and colleagues proposed a Laurasia
Laurasia
In paleogeography, Laurasia was the northernmost of two supercontinents that formed part of the Pangaea supercontinent from approximately...

n origin for the Moreae s.s., with three separate colonisations of South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

 (by Sorocea, Bagassa and Trophis) and a later colonisation of Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 by Milicia.

Distribution

Most genera in the tribe are predominantly tropical, sometimes with a few temperate species (like Maclura
Maclura
Maclura is a genus of flowering plants in the mulberry family, Moraceae. It includes the inedible Osage-orange, which is used as mosquito repellent and grown throughout the United States as a hedging plant.- Species :* Maclura africana...

). Two genera, Broussonetia
Broussonetia
Broussonetia is a genus of four species of trees in the family Moraceae, native to eastern Asia.-References:*...

and Morus, are predominantly temperate.

Genera

  • Antiaris
    Antiaris
    Antiaris is a tropical genus of latex producing evergreen trees in the Moraceae or fig family.Some species include:*Antiaris africana Engl., an economic hardwood of Africa, an African species which grows in the drier forests of the tropics....

  • Antiaropsis
  • Castilla
    Castilla (plant)
    Castilla is a tree genus belonging to the family Moraceae found native in Central America. It is named after Juan Diego del Castillo Castilla (sometimes incorrectly spelled Castilloa) is a tree genus belonging to the family Moraceae found native in Central America. It is named after Juan Diego...

  • Helicostylis
    Helicostylis
    Helicostylis is a genus of plant in family Moraceae.Species include:* Helicostylis heterotricha, Ducke* Helicostylis tomentosa, Rusby...

  • Maquira
  • Mesogyne
    Mesogyne
    Mesogyne is a genus of plant in family Moraceae . The genus is found in east Africa.Species include:* Mesogyne henriquesii * Mesogyne insignis, Engl....

  • Naucleopsis
    Naucleopsis
    Naucleopsis is a genus of plant in family Moraceae.Species include:* Naucleopsis chiguila, Benoist* Naucleopsis oblongifolia, Carauta...

  • Perebia
  • Poulsenia
    Poulsenia
    Poulsenia is a monotypic genus of plants in the family Moraceae, native to rainforests from Mexico south through Venezuela to Bolivia. The only species is P. armata. Unlike any other Neotropical Moraceae, the leaves and twigs of Poulsenia are covered in small prickles...

  • Pseudolmedia
    Pseudolmedia
    Pseudolmedia is a flowering plant genus in the mulberry family . They are known in Latin America as lechechiva and used for timber, construction wood, and sometimes in folk medicine.Species include:...

  • Sparattosyce
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