Annonaceae

Annonaceae
File:Annona squamosa.jpg
Annona squamosa fruit
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Magnoliids
Order:	Magnoliales
Family:	Annonaceae Juss.
Genera
See text
Synonyms
Hornschuchiaceae J. Agardh Monodoraceae J. Agardh

The Annonaceae are a family, the custard apple family, of flowering plants consisting of trees, shrubs, or rarely lianas. With 105 accepted genera and ca 2500 known species (Christenhusz & Byng 2016 ) it is the largest family in Magnoliales. Several genera produce edible fruit, most notably Annona, Anonidium, Asimina, Rollinia, and Uvaria. Its type genus is Annona. The family is concentrated in the tropics, with few species found in temperate regions. About 900 species are Neotropical, 450 are Afrotropical, and the other species Indomalayan.

Compared to the species from the Neotropics, very little is known about many species from Indomalaya. Only a few attempts have been made for the phylogeny-based reclassification of the family, and those have been hampered by the Neotropic bias in the available information, with the most of the work having been done on genera and tribes.

Description

Annona muricata

Mostly tropical, some mid-latitude, deciduous or evergreen trees and shrubs, some lianas, with aromatic bark, leaves, and flowers.

Stems, stalks and leaves

Bark is fibrous and aromatic. Pith septate (fine tangential bands divided by partitions) to diaphragmed (divided by thin partitions with openings in them).

Branching distichous (arranged in two rows/on one plane) or spiral.

Leaves are alternate, two-ranked, simple, pinnately veined, and have leaf stalks. Stipules absent.

Flowers

Flower stalks are axillary to (on the opposite side of shoot from) leaf scars on old wood and sometimes from leaves on new shoots. The flowers are usually trimerous; borne singly or in compound inflorescences; bisexual and rarely unisexual. The receptacle might become enlarged, elevated or flat. The outer whorls are inserted below the ovaries, and have valvate (overlapping) or imbricate (nonoverlapping) segments. Usually two to four persistent sepals that are distinct or connate (fused) at the base. Six petals in two unequal whorls of three with larger outer whorls and fleshier inner whorls that might share the same nectar glands, or six to fifteen petals, with impressed veins on their inner face. Ten to twenty (or many more) stamens inserted below the ovary, spirally arranged and forming a ball or flat-topped mass with short and stout filaments and linear to oblong anthers which face outward and open longitudinally. Each flower can have from one to many pistils, distinct to connate, with stigmas distinct. Marginal placentation, each pistil bearing one locule, with one to many ovules. Style short and thick, with terminal stigma.

Fruits and seeds

Fruits are single berries or coalesce from several pistils (into aggregate fruit, syncarps). Seeds are one to many per pistil; have a fleshy and usually brightly colored cover, have ruminate endosperm (nutritive tissue surrounding the embryo) and are oily.

Systematics

Monophyly and inter-familial systematics have been well supported for Annonaceae by a combination of morphological and molecular evidence. The APG II system places Annonaceae as most closely related to the small Magnoliid family Eupomatiaceae.

File:Uvaria ovata II.JPG

Uvaria ovata is part of the Family, Annonaceae.

Magnoliidae Canellales Piperales Magnoliales Myristicaceae Magnoliaceae Degeneriaceae Himantandraceae Eupomatiaceae Annonaceae Laurales

Combined plastid DNA marker and morphological character analysis of Anaxagorea and other Annonaceae and basal angiosperm genera suggest that Anaxagorea is an ancestral clade, sister to the rest of the family. This may confirm the hypothesis that morphological traits shared with other Magnoliales species (such as 2-ranked phyllotaxis, monosulcate pollen, and laminate stamens) represent ancestral characters, while derived characters observed in other genera have evolved independently multiple times. These analyses also support a biogeographic history in which Annonaceae, and perhaps other basal angiosperm clades, originated in South America during the Cretaceous, radiating due to migration events throughout Africa to Asia and Central to North America.

Guatteria, with its approximately 265 species, is the largest genus of Annonaceae, and might be broken into three small genera based on molecular characters from multiple locations.

Tribe Saccopetaleae is another group of Annonaceae that is awaiting review as recent molecular studies suggest that this tribe is not monophyletic at all.

Uses

Asimina triloba fruit.

Food

The large, edible, pulpy fruits of some members typically called "anona" by Spanish and Portuguese speaking people of the family's Neotropical range, include species of Annona: custard apple (A. reticulata), cherimoya (A. cherimola), soursop/guanábana/graviola (A. muricata), sweetsop (A. squamosa), ilama (A. diversifolia), soncoya (A. purpurea), atemoya (a cross between A. cherimola and A. squamosa); and biriba (Rollinia deliciosa, which may require reclassification under Annona.). The names of many of those fruits are sometimes used interchangeably. Recently, consumption of the neotropical annonaceous plant Annona muricata (soursop, graviola, guanabana) has been strongly associated as a causal agent in "atypical Parkinsonism". The causative agent, annonacin, is present in many of the Annonaceae. It is thought to be responsible for up to 70% of Parkinsonian conditions in Guadeloupe. Exposure is typically through traditional food and natural medicines.

Asimina triloba (American pawpaw, prairie banana) has an Eastern U.S. distribution, and is currently under agricultural investigation as a commercial crop.

Folk medicine

The bark, leaves, and roots of some species are used in folk medicines.

Toxicology

The compound annonacin in the seeds and leaves of many Annonaceae including Annona muricata (soursop), is a neurotoxin and it seems to be the cause of a neurodegenerative disease. The disorder is a so-called tauopathy associated with a pathologic accumulation of tau protein in the brain. Experimental results demonstrate that the plant neurotoxin annonacin is responsible for this accumulation.

Other Uses

Lancewood

Lancewood (Oxandra lanceolata) is a tough, elastic, and heavy wood obtained from the West Indies and The Guianas. It was often used for carriage shafts.

It is brought into commerce in the form of taper poles of about 6 metres in length and from 15 to 20 cm in breadth at the butt. The black lancewood or carisiri of The Guianas is of remarkably slender form.

The Yellow lancewood tree Calycophyllum candididissimum, common names lemonwood or degame, is from a different family (Rubiaceae). It is used as an alternative to lancewood and is found in tolerable abundance throughout The Guianas, and used by the Amerinds for arrow-points, as well as for spars, beams, etc. Some bowyers use this wood for making longbows.

Other

File:Ylang-Eden.jpg

Cananga odorata flowers.

• Some species of the family, such as Cananga odorata (ylang-ylang) also have aromatic oil and are used for perfumes or spices.
• The strong bark is used for carrying burdens in the Amazon Rainforest and for wooden implements, such as tool handles and pegs. The wood is valued as firewood.
• Yellow and brown dyes.
• Some species are also grown as ornamental plants, especially the Indian species Polyalthia longifolia pendula.
• The fruit and leaves of Uvariopsis tripetala (pepperfruit) is used as a spice for meats in some parts of Nigeria, due to its "hot" peppery flavor.

Chemical constituents

A large number of chemical compounds, including flavonoids, alkaloids, and acetogenins, have been extracted from the seeds and many other parts of these plants. Flavonoids and alkaloids contained in the leaves and bark of several species of the family have shown insecticidal properties.

Genera

• pl
• Alphonsea
• Ambavia
• Anaxagorea
• Ancana
• Annickia
• Annona
• Anomianthus
• †Anonaspermum
• Anonidium
• Artabotrys—trail-grape
• Asimina
• Asteranthe
• Balonga
• Bocagea
• Bocageopsis
• Boutiquea
• Cananga—Ylang-ylang
• Cardiopetalum
• Cleistochlamys
• Cleistopholis
• Craibella
• Cremastosperma
• Cyathocalyx
• Cyathostemma
• Cymbopetalum
• Dasoclema
• Dasymaschalon
• Deeringothamnus—false pawpaw
• Dendrokingstonia
• Dennettia
• Desmopsis
• Desmos
• Diclinanona
• Dielsiothamnus
• Disepalum
• Duckeanthus
• Duguetia
• Ellipeia
• Ellipeiopsis
• Enicosanthum
• Ephedranthus
• Exellia
• Fissistigma
• Fitzalania
• Friesodielsia
• Froesiodendron
• Fusaea
• Gilbertiella
• Goniothalamus
• Greenwayodendron
• Guamia
• Guatteria—haya minga, haya blanca
• Guatteriella
• Guatteriopsis
• Haplostichanthus
• Heteropetalum
• Hexalobus
• Hornschuchia
• Isolona
• Letestudoxa
• Lettowianthus
• Malmea
• Marsypopetalum
• Meiocarpidium
• Meiogyne
• Melodorum
• Mezzettia
• Mezzettiopsis
• Miliusa
• Mischogyne
• Mitrella
• Mitrephora
• Mkilua
• Monanthotaxis
• Monocarpia
• Monocyclanthus
• Monodora
• Mosannona
• Neostenanthera
• Neo-uvaria
• Onychopetalum
• Ophrypetalum
• Oreomitra
• Orophea
• Oxandra—blacklancewood, vernacular name: haya
• Pachypodanthium
• Papualthia
• Petalolophus
• Phaeanthus
• Phoenicanthus
• Piptostigma
• Platymitra
• Polyalthia
• Polyceratocarpus
• Popowia
• Porcelia
• Pseudartabotrys
• Pseudephedranthus
• Pseudoxandra
• Pseuduvaria
• Pyramidanthe
• Raimondia
• Reedrollinsia
• Richella
• Rollinia—wild sugar-apple
• Ruizodendron
• Sageraea
• Sanrafaelia
• Sapranthus
• Schefferomitra
• Sirdavidia
• es
• Stelechocarpus—kepel
• Stenanona
• Tetrameranthus
• Toussaintia
• es
• Trigynaea
• Trivalvaria
• Unonopsis
• Uvaria
• Uvariastrum
• Uvariodendron
• Uvariopsis
• es
• Xylopia

References

•  This article incorporates text from a publication now in the public domain: 

External links

• List of Annonaceae genera of the GRIN Database
• AnnonBase—online database for Annonaceae

• Informationsdienst Wissenschaft: Tauopathie durch pflanzliches Nervengift, 4. Mai 2007