Annonaceae

The Anonaceae (Annonaceae) are a family of angiosperms of the order Magnoliales. It consists of 130 genera with about 2,300 species that are distributed throughout the tropics of the New and Old World, as far north as Australia and the Pacific islands. They are part of the Ranalian complex, which makes them have a very old evolutionary history.

Description

Fruits and leaves Asimina triloba.

• Trees (sometimes with tabular roots or pneumatophores), bushes or seldom subarbustos or climbers, sometimes with xilopodium, aromatic, with indumento of simple, starry or scaly hairs.
• Simple, whole, alternating, distic, seldom spiral leaves, with craspedódroma nerve to brochydodroma, without stypules. Paracytic stomas. Idioblasts present, oily or mucillaginous. Astrosclereides and osteosclereides present.
• Talles with characteristic tin that presents continuous concentric tangential bands of parenchyma, and small vessels, homogeneous radios to weakly heterogeneous, unseries or more frequently multiseriated. Trilacunar nodes.
• Dial plants, seldom mononic (some species of Uvariopsisor androdioic (enenen) Pseuduvaria, Greenwayodendron and Diclinano).
• Lonely flowers, paired or in fascicles, terminals, axillary or supraaxillary, appearing on trunks or branches, seldom in subterranean, perfect or unisexual serpollos, actinomorfas, hypoogins, frequently with a single adaxial bract. Flat, hemispheric or conical reception. Sepals (2-)3(-4), in a verticil, fenced or imbricated, free or soldiers at least at the base, peristants or caducos. Pétalos (0-)3-6(-12), usually in 2 verticils (2-)3(-4) or in a verticil, fenced or imbricated, free or soldiers at the base, usually alternisépalos, carnosos or coriáceos, rarely membranosos. Numerous and spiral stamens, or 3-15 and verticilados; staminods present at times; short and free filaments, seldom long and monadelfos, linear anterees, with extrorous or rarely latrorceous dehiscence, sometimes transversely septated, connective usually with a truncated, conical or pyramidal extension. Numerous carpals to 1, apocárpicos to syncárpicos or paracárpicos, free styles or soldiers, usually short, cylindrical, stigmas capitados, oblongos or in horseshoe; ovulos numerous to 1 per carpelo, anthropos, basal or marginal placentation in the ventral suture in 1-2 rows.
• Simple fruit, each fleshy or woody element, dehiscent or indehiscent, sessile or stipit, or composed (sincarpo) with numerous unseminated carpelos, or unilocular and pluriseminated. There is a huge variety of forms and types.
• Seeds usually large and with or without arilo, with a peripheral groin in the fruits in dehiscent follicle, abundant endosperm, ruminated, usually in the form of irregular foils (laminiform ruminations), hard, oily, small, straight embryo.
• Very delicate, navicular to triangular, globose or disciforme, monoaperturate distally or inapertured, frequently in dayds, triads or tetrahed, granular exine or columelar, sometimes a lamelada basal layer, reticulated surface to be connected.
• chromosomal number: x = 7, 8, 9; 2n = 16, 24, 32, 48, 64; there is polyploidy in several genres.

Ecology

Flowers are predominantly protogynous. Pollination of many species is carried out by small beetles (Nitidulidae, Curculionidae) attracted by floral odors that imitate rotten fruit and that come to use the flowers as a laying ground, while larger beetles destroy floral parts when feeding on them. To defend themselves against the latter, some species prevent access to the carpels by closing the internal petals, producing self-pollination. Cymbopetalum macropodum, with large flowers on long pendulous stalks, is probably chiropterophilous, while Monodora myristica is dipterophilous (fly-pollinated). The dispersal of fruits and seeds is carried out mainly by animals, such as monkeys, birds, bats, but also iguanas, turtles or alligators. Some of the species with dehiscent follicles are capable of expelling the seeds a considerable distance.

Anonaceae generally develop in a tropical climate, only some species of the genera Asimina and Deeringothamnus appear in temperate zones of North America. In general, there is a stark contrast between the Old World forms, which tend to be climbing or widespread forms in low-lying rainforests, up to 1,500 m, rarely 2,000 m, while in the New World Mundo tend to be trees or shrubs and grow in savannah and cerrado zones, where some species are highly specialized as pyrophytes, e.g. For example, certain species of Annona and Duguetia, which is not an obstacle for some species, such as Raimondia quinduensis to reach 2600 m in the Colombian mountains.

Some New World species are deciduous trees that inhabit lowland deciduous forests.

Phytochemistry

Alkaloids derived from benzylisoquinoline (aporphins and oxoaporphins), berberines with structures similar to morphines, as well as C-methylated alkaloids. Characteristic C-methylated and C-benzylated flavonoids, and nitrophenylethane present. Diterpenes (kauranes and clerodanes), benzylbenzoates, styrylpyrones and polyketides present, as well as tetra- and pentacyclic triterpenoids, and furofuranic lignans (such as (+)-epimembrine and (+)-epieudesmin), but no steroids, the acetogenins of anonaceae have long aliphatic chains with or without tetrahydrofuran or tetrahydropyran rings, in addition, a β-lactone ring is present at one end. These compounds are restricted to the family and due to their toxicity with them, studies are carried out in different parts of the world to find and explain the powerful biological activity that they present.

Uses

Flower of Asimina triloba.

The main economic interest of the family is centered on the edible fruits of some species: the "cherimoya" (Annona cherimolia) whose fruit is the "cherimoya", the "soursop" (Annona muricata) whose fruit is the "soursop" or "graviola", the "anon" (Annona squamosa) whose fruit is the "anón" or the "anona", the "anona colorada" (Annona reticulata), the "Florida custard apple" (Asimina triloba) and the "anonillo", "biribá" or "Amazon apple" (Rollinia mucosa) and the "aratikú" (Anonna emarginata, Anonna rugulosa and Anonna neosalicifolia); all being American species; equally. The flowers of the "ylang-ylang" (Cananga odorata) from Asia are used for the extraction of essence for use in perfumery; its African equivalent is Artabotrys hexapetalus. The "xochinacaztli", sacred flower of the Aztecs, is the "orejuela" or "ear flower" (Cymbopetalum penduliflorum) used as a culinary condiment and for flavoring chocolate in Mexico and Central America; as a substitute for nutmeg in Africa, the seeds of Monodora myristica and those of Xylopia aethiopica are used instead of pepper ("guinea pepper"). Some woods are of special interest, e.g. eg, the "yaya" (Oxandra lanceolata) in the manufacture of handles and billiard cues; others are for local use in construction and for poles. Many species have local uses in pharmacopoeia due to their chemical components. In some places they are used as living fences.

Taxonomy

The family has been divided into 4 subfamilies:

• Anaxagoreoideae Chatrou, Pirie, Erkens, Couvreur " al., 2012
• Ambavioideae Chatrou, Pirie, Erkens, Couvreur " al., 2012
• Malmeoideae Chatrou, Pirie, Erkens & Couvreur " al., 2012
• Annoidae Rafinesque, 1815

Systematic position

The Annonaceae form a very well-defined and natural family, easy to distinguish, always considered as a primitive group of angiosperms, and always included in a basal Order Magnoliales or Annonales, according to different authors. The APW (Angiosperm Phylogeny Website) considers it to be, however, an advanced family of the Order Magnoliales and sister group to the Eupomatiaceae.

Taxa included

Seeds Asimina triloba.

The internal division of the family usually distinguished between two subfamilies, Annonoideae and Monodoroideae, the first having an apocarpic gynoecium that, at most, produced a multilocular syncarp, while the second had a unilocular ovary with parietal placentation; the first subfamily would be divided into 3 tribes: Uvarieae, Miliuseae and Unoneae. This classical classification, which goes back to Hutchinson (1964, see reference), has long been dismissed as artificial (Kessler, 1993, see reference). This author divided the family into informal groups. Recent advances in the methodology of morphological and, above all, molecular phylogeny have made it possible to ensure that there are 3 main branches in the evolutionary tree of the family: 1) a basal branch, formed by the genus Anaxagorea, which is the sister group of the other two; 2) a branch formed by the so-called "Ambavia group" (Ambavia, Cananga, Cleistopholis, Mezzettia, Tetrameranthus, and 3) one terminal branch that encompasses the rest of the genera, in which two main clades can be distinguished: one with little molecular divergence and genera with few species (including Malmea, Piptostigma, Miliusa and Polyalthia), and another clade with more molecular divergence and more specious genera.

See: Complete list of genres