Annonaceae

'' The species are mostly tropical, some are mid-latitude, deciduous or evergreen trees and shrubs, with some lianas, with aromatic bark, leaves, and flowers. divided by partitions) to diaphragmed (divided by thin partitions with openings in them). 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. '' }} 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'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==

'' 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. 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 the seeds and leaves of many of the Annonaceae, though not in any significant quantity in the fruit flesh. It is thought to be responsible for up to 70% of Parkinsonian conditions in Guadeloupe. Exposure is typically through traditional food and natural medicines. 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. 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. • Yellow and brown natural dyes • Some species are also grown as ornamental plants, especially the Indian species Polyalthia longifolia pendula. • The fruit and leaves of Uvariopsis tripetala (pepperfruit) are 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. ==References==