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Annonaceae

Article Discussion

Template:Short description Template:Automatic taxobox

The Annonaceae are a family of flowering plants consisting of trees, shrubs, or rarely lianas<ref name="FNA">Template:EFloras</ref> commonly known as the custard apple family<ref name="ITIS">{{#if:18092 | {{#invoke:template wrapper|wrap|_template=cite web|_exclude=id,ID,taxon 

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| Template:Citation error }}</ref><ref name="FNA" /> or soursop family. With 108 accepted genera and about 2400 known species,<ref name="Chatrou et al. 2012">Template:Cite journal</ref> it is the largest family in the 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 remaining are Indomalayan.

DescriptionEdit

File:Annona muricata Blanco1.196.png
Annona muricata

The species are mostly tropical, some are mid-latitude, deciduous or evergreen trees and shrubs, with some lianas, with aromatic bark, leaves, and flowers.<ref name="FNA"/>

Stems, stalks and leaves
Bark is fibrous and aromatic. Pith septate (fine tangential bands<ref name="nh1">Template:Cite journal</ref> divided by partitions) to diaphragmed (divided by thin partitions with openings in them).<ref name="FNA"/> Branching distichous (arranged in two rows/on one plane) or spiral.<ref name="Johnson 2003"/> Leaves are alternate, two-ranked,<ref name="Johnson 2003">Template:Cite journal</ref> simple, pinnately veined, and have leaf stalks. Stipules absent.<ref name="FNA"/>
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.<ref name="FNA"/>
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.<ref name="FNA"/>

SystematicsEdit

Monophyly and inter-familial systematics have been well supported for Annonaceae by a combination of morphological and molecular evidence.<ref name="Doyle 2004">Template:Cite journal</ref> The APG II system places Annonaceae as most closely related to the small Magnoliid family Eupomatiaceae.

File:Uvaria ovata II.JPG
Uvaria ovata
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In a phylogeny-based reclassification of the family<ref name="Chatrou et al. 2012"/> four subfamilies are recognised: Anaxagoreoideae (including just Anaxagorea), Ambavioideae, Annonoideae, and Malmeoideae. A number of the larger genera, including Guatteria, with its 177 species,<ref name="MaasWestra2015">Template:Cite journal</ref> 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:<ref name="CouvreurMaas2012">Template:Cite journal</ref> For a concise bibliographic overview of the taxonomic literature (1900 to 2012) see:<ref name="ErkensMennega2012">Template:Cite journal</ref>

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.<ref name="Scharaschkin and Doyle 2005">Template:Cite journal</ref><ref name="Scharaschkin and Doyle 2006">Template:Cite journal</ref><ref name="DoyleLe Thomas2012">Template:Cite journal</ref> The oldest fossil evidence of Annonaceae is described as the genus Futabanthus, from the Late Cretaceous (Coniacian) of Japan,<ref name="TakahashiFriis2008">Template:Cite journal</ref> which represents a minimum age of c. 89 million years ago for the most recent common ancestor (crown group) of the family.<ref name="PirieDoyle2012">Template:Cite journal</ref> 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.<ref name="RichardsonChatrou2004">Template:Cite journal</ref><ref name="CouvreurPirie2011">Template:Cite journal</ref>

Taxonomic revisions within the subfamily MalmeoideaeEdit

The reclassification and establishment of HuberanthaEdit

The genus Huberantha (synonym Hubera) was resolved to be sister to Miliusa, with certain species previously under Polyalthia being additionally reclassified.<ref name=":0">Template:Cite journal</ref> 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.<ref name=":0" />

Controversy over Stelechocarpus/WinitiaEdit

It was proposed that the genus Stelechocarpus, which includes S. burahol and S. cauliflorus be reclassified under a new genus Winitia, 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 to S. cauliflorus as a monophyletic group.<ref>Template:Cite journal</ref> However this is no longer accepted.<ref>Template:Cite POWO</ref>

Reclassification of Annickia as tribe AnnickieaeEdit

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.<ref name=":1">Template:Cite journal</ref>

SubfamiliesEdit

The taxonomy of the Annonaceae is based on the Angiosperm Phylogeny Website, which recognises four subfamilies<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>Template:Cite journal</ref> and the extinct genus Template:ExtinctAnonaspermum<ref name="Manchester1994">Template:Cite journal</ref>

AnaxagoreoideaeEdit

Auth.: Chatrou et al. 2012 (monotypic)

AmbavioideaeEdit

Auth.: Chatrou et al. 2012

AnnonoideaeEdit

Auth. Rafinesque, 1815

MalmeoideaeEdit

UsesEdit

File:Asimina triloba red fern farm.jpg
Asimina triloba fruit

FoodEdit

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).<ref>Template:Cite iucn</ref> 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.<ref name="health_parkinsons1">Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

The American pawpaw (Asimina triloba) has an Eastern United States distribution and has been investigated as a commercial agricultural crop.<ref name="Pomper">Template:Cite journal</ref>

Flower petals from sacred earflower (Cymbopetalum penduliflorum) and from related species C. costaricense<ref name=Spice>Template:Cite book</ref> were traditionally used to flavor chocolate<ref name="uphof">Template:Cite book</ref> before the arrival of cinnamon and the other Old World spices.<ref name=inventory>Template:Cite book</ref> The dried petals are still used to flavor atoles, pinoles, and coffee.<ref name=names>Template:Cite book</ref>

Folk medicineEdit

The bark, leaves, and roots of some species are used in folk medicines.Template:Citation needed

ToxicologyEdit

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.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Other usesEdit

Lancewood (Oxandra lanceolata)<ref name="Lincoln">Template:Cite book</ref> 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.<ref name="EB1911">{{#if: | 

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The yellow lancewood tree Calycophyllum candididissimum, common names lemonwood or degame, is from a different family (Rubiaceae).<ref name="Lincoln" /> 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.<ref name="EB1911"/> Some bowyers use this wood for making longbows.

OtherEdit

File:Ylang-Eden.jpg
Ylang-ylang (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.<ref name="nh1"/>
  • The strong bark is used for carrying burdens in the Amazon Rainforest<ref name="nh1"/> and for wooden implements, such as tool handles and pegs.<ref name="icuc">{{#invoke:citation/CS1|citation

|CitationClass=web }}</ref> The wood is valued as firewood.<ref name="nh1"/>

  • Yellow and brown natural dyes<ref name="icuc"/>
  • 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 constituentsEdit

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.<ref name="icuc"/>

ReferencesEdit

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External linksEdit

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| {{#invoke:template wrapper|wrap|_template=cite web|_exclude=id,ID,taxon 

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 | title = Annonaceae
 | publisher = Integrated Taxonomic Information System
 }}

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