The species are mostly tropical, some are mid-latitude,deciduous orevergreen trees and shrubs, with some lianas, with aromatic bark, leaves, and flowers.[3]
Stems, stalks and leaves
Bark is fibrous and aromatic.Pith septate (fine tangential bands[6] divided by partitions) to diaphragmed (divided by thin partitions with openings in them).[3] Branching distichous (arranged in two rows/on one plane) or spiral.[7] Leaves arealternate, two-ranked,[7] simple,pinnately veined, and have leaf stalks.Stipules absent.[3]
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 usuallytrimerous; borne singly or in compoundinflorescences;bisexual and rarelyunisexual. Thereceptacle might become enlarged, elevated or flat. Theouter whorls areinserted below the ovaries, and have valvate (overlapping) or imbricate (nonoverlapping) segments. Usually two to four persistentsepals 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 stoutfilaments 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 onelocule, with one to many ovules. Style short and thick, with terminal stigma.[3]
Fruits and seeds
Fruits are single berries or coalesce from several pistils (intoaggregate 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.[3]
In aphylogeny-based reclassification of the family[5] four subfamilies are recognised:Anaxagoreoideae (including justAnaxagorea),Ambavioideae,Annonoideae, andMalmeoideae. A number of the larger genera, includingGuatteria, with its 177 species,[9]Annona, andXylopia 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 inWorld 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:[10] For a concise bibliographic overview of the taxonomic literature (1900 to 2012) see:[11]
Bothplastid DNA markers and morphological characters provide evidence thatAnaxagorea is the sisterclade to the rest of the family. This may confirm the hypothesis that morphological traits shared betweenAnaxagorea and other Magnoliales species (such as 2-rankedphyllotaxis, monosulcatepollen, and laminatestamens) represent ancestral characters, while derived characters observed in other genera have evolved independently multiple times.[12][13][14] The oldest fossil evidence of Annonaceae is described as the genusFutabanthus, from the Late Cretaceous (Coniacian) of Japan,[15] which represents a minimum age of c. 89 million years ago for themost recent common ancestor (crown group) of the family.[16] The ages of Annonaceae clades inferred using fossil evidence andmolecular clock-based dating techniques suggests that the pantropical distribution of the family originated subsequent to the break-up of theGondwanan supercontinent, as the result of a combination of geodispersal tracking the expansion of theboreotropical flora during theEocene and more recent long-distance dispersal events.[17][18]
Taxonomic revisions within the subfamily Malmeoideae
The genusHuberantha (synonymHubera) was resolved to be sister toMiliusa, with certain species previously underPolyalthia being additionally reclassified.[19] 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 fromMonoon andPolyalthia, distinguishHuberantha on the generic level. Morphologically,Huberantha has a finely and densely granular infratectum whereasMonoon andPolyalthia have columellate or densely granular infratecta.[19]
It was proposed that the genusStelechocarpus, which includesS. burahol andS. cauliflorus be reclassified under a new genusWinitia, which is characterized by mixed flowers, multicolumellar stigmas, and columellate/coarsely granular infratectum. This genus was created after phylogenetic analysis that highly supported an unclassified species from Thailand being sister toS. cauliflorus as a monophyletic group.[20] However this is no longer accepted.[21]
The genusAnnickia 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.[22]
The large, edible, pulpy fruits of some members, typically calledanona by Spanish- and Portuguese-speaking people of the family's Neotropical range, include species ofAnnona: custard apple (A. reticulata), cherimoya (A. cherimola), soursop/guanábana/graviola (A. muricata), sweetsop (A. squamosa), ilama (A. macroprophyllata), soncoya (A. purpurea),atemoya (a cross betweenA. cherimola andA. squamosa); and biriba (A. mucosa).[29] The names of many of those fruits are sometimes used interchangeably.
Consumption of the neotropical annonaceous plantAnnona muricata (soursop,graviola, guanabana) has been strongly associated as a causal agent in "atypicalParkinsonism". 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.[30][31][32][33]
The American pawpaw (Asimina triloba) has an Eastern United States distribution and has been investigated as a commercial agricultural crop.[34]
Theacetogenin compounds, which occur in the fruit, seeds, and leaves of many Annonaceae, including soursop (Annona muricata), areneurotoxins and seem to be the cause of aneurodegenerative disease. The disorder is a so-calledtauopathy associated with a pathologic accumulation oftau protein in the brain. Experimental results indicate that acetogenins are responsible for this accumulation.[39]
Lancewood (Oxandra lanceolata)[40] is a tough, elastic, and heavy wood obtained from theWest Indies andThe Guianas. It was often used forcarriage 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 treeCalycophyllum candididissimum, common nameslemonwood ordegame, is from a different family (Rubiaceae).[40] 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. Somebowyers use this wood for makinglongbows.
Some species of the family, such asCananga odorata (ylang-ylang) also havearomatic oil and are used for perfumes or spices.[6]
The strong bark is used for carrying burdens in theAmazon Rainforest[6] and for wooden implements, such as tool handles and pegs.[41] The wood is valued as firewood.[6]
A large number of chemical compounds, includingflavonoids,alkaloids, andacetogenins, 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 showninsecticidal properties.[41]
^Couvreur, Thomas L.P.; Maas, Paul J.M.; Meinke, Svenja; Johnson, David M.; Keßler, Paul J.A. (2012). "Keys to the genera of Annonaceae".Botanical Journal of the Linnean Society.169 (1):74–83.doi:10.1111/j.1095-8339.2012.01230.x.ISSN0024-4074.
^Scharaschkin, T.; Doyle, J.A. (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.S2CID86018723.
^Couvreur, Thomas L.P.; Pirie, Michael D.; Chatrou, Lars W.; Saunders, Richard M.K.; Su, Yvonne C.F.; Richardson, James E.; Erkens, Roy H.J. (2011). "Early evolutionary history of the flowering plant family Annonaceae: steady diversification and boreotropical geodispersal".Journal of Biogeography.38 (4):664–680.doi:10.1111/j.1365-2699.2010.02434.x.ISSN0305-0270.S2CID85568637.
^Chaowasku, Tanawat; Van Der Ham, Raymond W.J.M. (June 2013). "Integrative systematics supports the establishment of Winitia, a new genus of Annonaceae (Malmeoideae, Miliuseae) allied to Stelechocarpus and Sageraea".Systematics and Biodiversity.11 (2):195–207.doi:10.1080/14772000.2013.806370.ISSN1477-2000.S2CID86528302.
^POWO:Winitia Chaowasku is a synonym ofStelechocarpus Hook.f. & Thomson (retrieved 19 July 2022)
^Manchester, S.R. (1994). "Fruits and Seeds of the Middle Eocene Nut Beds Flora, Clarno Formation, Oregon".Palaeontographica Americana.58:30–31.
^abcDagallier LMJ, Mbago FM, Couderc M, Gaudeul M, Grall A, Loup C, Wieringa JJ, Sonké B, Couvreur TLP. Phylogenomic inference of the African tribe Monodoreae (Annonaceae) and taxonomic revision of Dennettia, Uvariodendron and Uvariopsis. PhytoKeys. 2023 Sep 22;233:1-200. doi: 10.3897/phytokeys.233.103096. PMID 37811332; PMCID: PMC10552675.
^Bangkomnate R., Damthongdee A., Baka A., Aongyong K. & Chaowasku T. 2021:Pyramidanthe andMitrella (Annonaceae, Uvarieae) unified: molecular phylogenetic and morphological congruence, with new combinations in Pyramidanthe. –Willdenowia 51: 383–394.
^Chaowasku, T., A. Damthongdee, H. Jongsook, D.T. Ngo, H.T. Le, D.M. Tran & S. Suddee (2018). Enlarging the monotypic Monocarpieae (Annonaceae, Malmeoideae): recognition of a second genus from Vietnam informed by morphology and molecular phylogenetics.Candollea 73: 261–275. DOI:http://dx.doi.org/10.15553/c2018v732a11
^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.121 (2). International Brain Research Organization:287–296.doi:10.1016/S0306-4522(03)00441-X.PMID14521988.S2CID37873631.
^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.PMID16078200.S2CID31508365.
^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?".Parkinson's Disease and Related Disorders. Journal of Neural Transmission. Supplementa. Vol. 70. pp. 153–157.doi:10.1007/978-3-211-45295-0_24.ISBN978-3-211-28927-3.PMID17017523.{{cite book}}:|journal= ignored (help)
^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.
^Seidemann, Johannes (27 December 2005)."C".World Spice Plants: Economic Usage, Botany, Taxonomy. Springer Science & Business Media. p. 129.ISBN9783540279082. Retrieved2 July 2015.Flavoring of drinking chocolate
^Murray, Nancy A. (6 December 1993).Revision ofCymbopetalum andPorcelia (Annonaceae). Systematic botany monographs. Vol. 40. Ann Arbor, Mich: American Society of Plant Taxonomists.ISBN9780912861401.OCLC29527548.
This article incorporates text from a publication now in thepublic domain: Chisholm, Hugh, ed. (1911). "Custard Apple".Encyclopædia Britannica. Vol. 7 (11th ed.). Cambridge University Press. p. 667.
