Annonaceae - WikiHQ

The Annonaceae are a family of flowering plants consisting of trees, shrubs, or rarely lianas commonly known as the custard apple family it is the largest family in the Magnoliales. Several genera produce edible fruit, most notably Annona, Anonidium, Asimina, Rollinia, and Uvaria.<!-- divided by partitions) to diaphragmed (divided by thin partitions with openings in them). simple, pinnately veined, and have leaf stalks. Stipules absent. The APG II system places Annonaceae as most closely related to the small Magnoliid family Eupomatiaceae.

thumb|[[Uvaria ovata]]

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In a phylogeny-based reclassification of the family Annona, and Xylopia belong to Annonoideae. Together, Annonoideae and Malmeoideae comprise the majority of the species and each are further subdivided into a number of tribes. The subfamilial and tribal classification is followed in World Annonaceae which presents an overview of all Annonaceae genera and taxonomic, distribution and photographic information for a large number of species. Keys for the identification of Annonaceae genera (separately for Neotropical, African/Madagascan, and Asian/Australian taxa) are presented in: For a concise bibliographic overview of the taxonomic literature (1900 to 2012) see:

Both plastid DNA markers and morphological characters provide evidence that Anaxagorea is the sister clade to the rest of the family. This may confirm the hypothesis that morphological traits shared between Anaxagorea and 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. The oldest fossil evidence of Annonaceae is described as the genus Futabanthus, from the Late Cretaceous (Coniacian) of Japan, which represents a minimum age of c. 89 million years ago for the most recent common ancestor (crown group) of the family. The ages of Annonaceae clades inferred using fossil evidence and molecular clock-based dating techniques suggests that the pantropical distribution of the family originated subsequent to the break-up of the Gondwanan supercontinent, as the result of a combination of geodispersal tracking the expansion of the boreotropical flora during the Eocene and more recent long-distance dispersal events.

Taxonomic revisions within the subfamily Malmeoideae

The reclassification and establishment of Huberantha

The genus Huberantha (synonym Hubera) was resolved to be sister to Miliusa, with certain species previously under Polyalthia being additionally reclassified. This reclassification was highly supported by maximum parsimony, Bayesian analysis, and morphological characters. Hubera is characterized by reticulate tertiary venation, axillary inflorescences, 1 ovule per ovary, seeds with flat to slightly raised raphes, and other characters. Huberantha<nowiki/>'s phylogenetic distance and morphological difference from Monoon and Polyalthia, distinguish Huberantha on the generic level. Morphologically, Huberantha has a finely and densely granular infratectum whereas Monoon and Polyalthia have columellate or densely granular infratecta. However this is no longer accepted.

Reclassification of Annickia as tribe Annickieae

The genus Annickia was previously included within the tribe Piptostigmateae. However, it is highly supported to being sister to the rest of the Malmeoideae tribes, and weakly supported to being sister to the rest of the Piptostigmateae genera. For these reasons, Annickia is now classified within its own tribe in the Malmeoideae, the Annickieae.

Subfamilies

The taxonomy of the Annonaceae is based on the Angiosperm Phylogeny Website, which recognises four subfamilies and the extinct genus Anonaspermum

Anaxagoreoideae

Auth.: Chatrou et al. 2012 (monotypic)

Anaxagorea St.-Hilaire 1825

Ambavioideae

Auth.: Chatrou et al. 2012

Ambavia Le Thomas
Cananga (DC.) Hook.f. & Thomson (Ylang-ylang)
Cleistopholis Pierre ex Engl.
Cyathocalyx Champ. ex Hook.f. & Thomson
Drepananthus Maingay ex Hook.f.
Lettowianthus Diels
Meiocarpidium Engl. & Diels
Mezzettia Becc.
Tetrameranthus R.E.Fr.

Annonoideae

Auth. Rafinesque, 1815

Tribe Annoneae Endlicher 1839
Annona L. (synonym Rollinia A. St.-Hil.)
Anonidium Engl. & Diels
Asimina Adans.
Diclinanona Diels
Disepalum Hook. f.
Goniothalamus (Blume) Hook.f. & Thomson
Neostenanthera Exell (syn. Boutiquea Le Thomas)
Tribe Bocageeae Endlicher 1839
Bocagea A.St.-Hil.
Cardiopetalum Schltdl.
Cymbopetalum Benth.
Froesiodendron R.E.Fr.
Hornschuchia Nees
Mkilua Verdc.
Porcelia Ruiz & Pav.
Trigynaea Schltdl.
Tribe Duguetieae Chatrou & Saunders 2012
Duckeanthus R.E.Fr.
Duguetia A.St.-Hil.
Fusaea (Baill.) Saff.
Letestudoxa Pellegr.
Pseudartabotrys Pellegr.
Tribe Guatterieae Hooker & Thomson 1855
Guatteria Ruiz & Pav.
Tribe Monodoreae Baill. 1868
Asteranthe Engl. & Diels
Dennettia
Ophrypetalum Diels
Sanrafaelia Verdc.
Tribe Uvarieae Hooker & Thomson 1855
Afroguatteria
Cleistochlamys Oliv.
Dasymaschalon (Hook.f. & Thomson) Dalla Torre & Harms
Desmos Lour.
Dielsiothamnus R.E.Fr.
Fissistigma Griff.
Friesodielsia Steenis
Monanthotaxis Baill.
Pyramidanthe Miq. (synonym Mitrella )
Sphaerocoryne (Boerl.) Ridl.
Toussaintia
Uvaria L. (synonyms Balonga Le Thomas and Melodorum Lour.)
Tribe Xylopieae Endlicher 1839
Artabotrys R.Br.
Xylopia L.

Malmeoideae

Tribe Annickieae Couvreur et al., 2019
Tribe Miliuseae Hooker & Thomson 1855
Alphonsea Hook.f. & Thomson
Desmopsis Saff.
Huberantha (synonym Hubera) Chaowasku
Marsypopetalum Scheff.
Meiogyne Miq. (synonyms: Ancana F. Mueller; Guamia Merrill; Fitzalania F. Muell.; Oncodostigma Diels, Polyaulax Backer)
Miliusa Lesch. ex A.DC. (Saccopetalum Bennett)
Mitrephora (Blume) Hook.f. & Thomson
Monoon Miq. (Cleistopetalum H. Okada, Enicosanthum Becc., Woodiellantha Rauschert)
Neo-uvaria Airy Shaw
Orophea Blume
Phaeanthus Hook.f. & Thomson
Platymitra Boerl.
Polyalthia Blume
Polyalthiopsis Chaowasku
Popowia Endl.
Pseuduvaria Miq.
Sageraea Dalzell
Sapranthus Seem.
Stelechocarpus Hook.f. & Thompson
Stenanona Standl.
Tridimeris Baill.
Trivalvaria (Miq.) Miq.
Wangia X.Guo & R.M.K.Saunders
Wuodendron B.Xue, Y.H.Tan & Chaowasku

Uses

thumb|200px|[[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. macroprophyllata), soncoya (A. purpurea), atemoya (a cross between A. cherimola and A. squamosa); and biriba (A. mucosa). The names of many of those fruits are sometimes used interchangeably.

The American pawpaw (Asimina triloba) has an Eastern United States distribution and has been investigated as a commercial agricultural crop.

Flower petals from sacred earflower (Cymbopetalum penduliflorum) and from related species C. costaricense were traditionally used to flavor chocolate before the arrival of cinnamon and the other Old World spices. The dried petals are still used to flavor atoles, pinoles, and coffee.

Toxicity

Consumption of the neotropical annonaceous plant Annona muricata (soursop, graviola, guanabana) has been strongly associated as a causal agent in "atypical Parkinsonism". The causative agents are acetogenins, especially the most abundant annonacin, found in all parts of the plant. It is thought to be responsible for up to 70% of Parkinsonian conditions in Guadeloupe. Exposure is typically through consumption of the fruits or its juice, traditional food and natural medicines.

The acetogenins, present in significative amounts in Annonaceae, cause atypical Parkinsonism by cumulatively inhibiting NADH dehydrogenase and disrupting mitochondrial function. Because of the cumulative effect, consumption of one fruit, or its juice, everyday for a year has the same effect as the consumption of one fruit every ten days for ten years. Annona muricata, Annona squamata, Annona reticulata, and Asimina triloba were shown to have toxic levels of acetogenins.

Asimina triloba's toxicity has been studied for use as a pesticide, an insecticide, and for cancer treatment. The unripe fruits were the parts that yield the highest levels of toxins.

Folk medicine

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

Toxicology

The acetogenin compounds, which occur in the fruit, seeds, and leaves of many Annonaceae, including soursop (Annona muricata), are neurotoxins and seem 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 indicate that acetogenins are responsible for this accumulation.

Other uses

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 m 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). The wood is valued as firewood.