Moreae is a

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...

within the plant family

Moraceae The Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 38 genera and over 1100 species. Most are widespread in tropical and subtropical regions, less so in temperate climates; however ...

. It includes 6–10

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

and 70–80 species, including '' Morus'', the genus that includes the mulberries, and ''

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. It is dioecious, with male and fe ...

'', the genus that includes the

Osage orange ''Maclura pomifera'', commonly known as the Osage orange ( ), is a small deciduous tree or large shrub, native to the south-central United States. It typically grows about tall. The distinctive fruit, a multiple fruit, is roughly spherical, b ...

. Recent work suggests that the tribe is

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...

Description

The Moreae are a tribe of trees, shrubs, climbers and herbs that are usually

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 reproductio ...

. Their

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 o ...

s are simpler than most other Moraceae. Their flowers are adapted for

wind pollination Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous plan ...

. Members of the tribe are characterised by having inflexed or " urticaceous"

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 filame ...

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 In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

—traits that were present in the ancestors of the Moraceae 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 Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...

in 1753 in ''

Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...

''. Cornelis Berg included eight genera in the tribe Moreae—'' Bleekrodea'', ''

Broussonetia ''Broussonetia'' is a genus of four species (including one hybrid species) of trees in the family Moraceae, native to eastern Asia. These four species have high-quality fiber which consist of more than 90% of cellulose. They are traditionally ap ...

'', ''

Fatoua ''Fatoua'' is a genus of 3 species of herbs in the mulberry family. Species *'' Fatoua villosa'' *'' Fatoua pilosa'' *'' Fatoua madagascariensis'' Bibliography {{Taxonbar, from=Q147612 Moraceae genera Moraceae ...

'', ''

'', '' Milicia'', '' Morus'', ''

Streblus ''Streblus'' is a genus of flowering plants in the mulberry family, Moraceae. The genus is found in the Pacific across Southeast Asia, Eastern Australia, New Zealand and the Pacific Islands. Species include: *''Streblus asper'' Lour. – Siame ...

'' and ''

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. It is dioecious, with male and female flowers borne on separate plants. Recent work suggests ...

''—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 Edred John Henry Corner FRS (12 January 1906 – 14 September 1996) was an English mycologist and botanist who occupied the posts of assistant director at the Singapore Botanic Gardens (1929–1946) and Professor of Tropical Botany at the Univ ...

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

Shannon Datwyler and George Weiblen use the

chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...

''ndhF'' gene to build a

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

of the family. They determined that the tribe Moreae was

and consisted of two groups—a core "Moreae ''

sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

''" (Moreae s.s.), which formed a

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 gro ...

group within the tribe, and a broader "Moreae ''

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'' (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 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. The juvenile leaves are distinctly different in appearance from the mature l ...

'' and ''

Sorocea ''Sorocea'' is a Neotropical genus of woody plants in the family Moraceae. Its distribution ranges from Chiapas to southern Brazil. It is placed within the tribe Moreae, and is closely related to the Monotypic taxon, monotypic ''Bagassa''. Spec ...

'' from the

Artocarpeae Artocarpeae is a tribe (biology), tribe within the plant family Moraceae. It includes 7 to 12 genus, genera and 70 to 87 species including ''Artocarpus altilis'', the breadfruit. Description Species in the Artocarpeae are tropical trees or shru ...

to the Moreae. Based on chloroplast ''ndhF''

gene sequence In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...

s, Moreae s.s. is a

sister taxon 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 t ...

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.

Evolutionary history

Few fossils can be definitively assigned to the Moreae. Fruits from the upper

Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

and

Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...

have been assigned to ''Broussonetia'', and fruit matching modern ''Morus'' have been found in the early Eocene. Based on rates of

molecular evolution Molecular evolution is the process of change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genetics ...

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 () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...

n origin for the Moreae s.s., with three separate colonisations of

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

(by ''Sorocea'', ''Bagassa'' and ''Trophis'') and a later colonisation of

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

by ''Milicia''.

Distribution

Most genera in the tribe are predominantly tropical, sometimes with a few temperate species (like ''

''). Two genera, ''

'' and ''Morus'', are predominantly temperate.

Genera

References

{{Taxonbar, from=Q6911196 Moraceae Rosales tribes