|Annona squamosa fruit|
Hornschuchiaceae J. Agardh
The Annonaceae are a family, the custard apple family, of flowering plants consisting of trees, shrubs, or rarely lianas. With 2106 accepted species and more than 130 genera, 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.
- 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.
- 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.
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.
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. 
Tribe Saccopetaleae is another group of Annonaceae that is awaiting review as recent molecular studies suggest that this tribe is not monophyletic at all.
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 Parkinsonsism". 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.
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.
- 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.
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.
- Deeringothamnus—false pawpaw
- Guatteria—haya minga, haya blanca
- Oxandra—blacklancewood, vernacular name: haya
- Rollinia—wild sugar-apple
The compound annonacin contained in the seeds of some members of Annonaceae such as 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.
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- Chatrou, Dr. L.W. (2005-07-29). "Molecular Systematics of Annonaceae". Annonaceae research projects (Nationaal Herbarium Nederland). Retrieved 2008-04-20.
- Johnson, D.M. (July–September 2003). "Phylogenetic significance of spiral and distichous architecture in the Annonaceae.". Systematic Botany 28 (3): 503–511.
- Doyle, J.A., H. Sauquet and T. Scharaschkin (2004). "Phylogeny, molecular and fossil dating, and biogeographic history of Annonaceae and Myristicaceae (Magnoliales)". International Journal of Plant Sciences 165 (4): 55–67. doi:10.1086/421068.
- Scharaschkin, T. and J.A. Doyle (2005). "Phylogeny and historical biogeography of Anaxagorea (Annonaceae) using morphology and non-coding chloroplast sequence data.". Systematic Botany 30 (5): 712–735. doi:10.1600/036364405775097888.
- Scharaschkin, T. and J.A. Doyle (2006). "Character evolution in Anaxagorea (Annonaceae)". American Journal of Botany 93 (1): 36–54. doi:10.3732/ajb.93.1.36.
- Erkens, R.H.J.; Chatrou, L.W.; Koek-Noorman, J.; Maas, P.J.M. (2005-07-29). "Phylogeny and evolution of Guatteria". Annonaceae research projects (Nationaal Herbarium Nederland). Retrieved 2008-04-20.
- Mols, J.B.; Chatrou, L.W.; Koek-Noorman, J.; Maas, P.J.M. (2005-07-29). "Phylogeny of Miliusa and allied Asian Annonaceae using cpDNA and morphology". Annonaceae research projects (Nationaal Herbarium Nederland). Retrieved 2008-04-20.
- Lannuzel, A et al. (2003-10-06). "The mitochondrial complex i inhibitor annonacin is toxic to mesencephalic dopaminergic neurons by impairment of energy metabolism". Neuroscience (International Brain Research Organization) 121 (2): 287–296. doi:10.1016/S0306-4522(03)00441-X. PMID 14521988.
- Champy, Pierre et al. (2005-08-02). "Quantification of acetogenins in Annona muricata linked to atypical parkinsonism in guadeloupe". Movement Disorders 20 (12): 1629–1633. doi:10.1002/mds.20632. PMID 16078200.
- Lannuzel A, Höglinger GU, Champy P, Michel PP, Hirsch EC, Ruberg M. (2006). "Is atypical parkinsonism in the Caribbean caused by the consumption of Annonacae?". J Neural Transm Suppl. Journal of Neural Transmission. Supplementa 70 (70): 153–7. doi:10.1007/978-3-211-45295-0_24. ISBN 978-3-211-28927-3. PMID 17017523.
- Caparros-Lefebvre D, Elbaz A. (1999-07-24). "Possible relation of atypical parkinsonism in the French West Indies with consumption of tropical plants: a case-control study". Lancet 354 (9175): 281–6. doi:10.1016/S0140-6736(98)10166-6. PMID 10440304.
- Pomper, K.W. et al. (July 2008). "Flowering and fruiting characteristics of eight pawpaw [Asimina triloba (L.)] Dunal selections in Kentucky". Journal American Pomological Society 62(3):89-97.
- Lincoln, William A (1986). World Woods in Colour. Hertford UK: Stobard Davies Ltd. ISBN 0-85442-028-2.
- University of Southampton (March 2002). "Factsheet No. 5. Annona" (PDF). Fruits for the Future. Department for International Development, International Centre for Underutilised Crops. Retrieved 2008-04-20.
- Manchester, S.R. (1994). "Fruits and Seeds of the Middle Eocene Nut Beds Flora, Clarno Formation, Oregon". Palaeontographica Americana 58: 30–31.
- Informationsdienst Wissenschaft: Tauopathie durch pflanzliches Nervengift, 4. Mai 2007
- This article incorporates text from a publication now in the public domain: Chisholm, Hugh, ed. (1911). Encyclopædia Britannica (11th ed.). Cambridge University Press.
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