Annonaceae

Cladus: Eukaryota
Regnum: Plantae
Divisio: Magnoliophyta
Classis: Magnoliopsida
Ordo: Magnoliales
Familia: Annonaceae
Genera: Afroguatteria - Alphonsea - Ambavia - Anaxagorea - Ancana - Annickia - Annona - Anomianthus - Anonidium - Artabotrys - Asimina - Asteranthe - Balonga - Bocagea - Bocageopsis - Boutiquea - Cananga - Cardiopetalum - Chieniodendron - Cleistochlamys - Cleistopholis - Cremastosperma - Cyathocalyx - Cyathostemma - Cymbopetalum - Dasoclema - Dasymaschalon - Deeringothamnus - Dendrokingstonia - Dennettia - Desmopsis - Desmos - Diclinanona - Dielsiothamnus - Disepalum - Duckeanthus - Duguetia - Ellipeia - Ellipeiopsis - Enicosanthum - Ephedranthus - Exellia - Fissistigma - Fitzalania - Friesodielsia - Froesiodendron - Fusaea - Gilbertiella - Goniothalamus - Greenwayodendron - Guamia - Guatteria - Guatteriella - Guatteriopsis - Haplostichanthus - Heteropetalum - Hexalobus - Hornschuchia - Isolona - Letestudoxa - Lettowianthus - Malmea - Marsypopetalum - Meiocarpidium - Meiogyne - Melodorum - Mezzettia - Mezzettiopsis - Miliusa - Mischogyne - Mitrella - Mitrephora - Mkilua - Monanthotaxis - Monocarpia - Monocyclanthus - Monodora - Mosannona - Neo-Uvaria - Neostenanthera - Oncodostigma - Onychopetalum - Ophrypetalum - Oreomitra - Orophea - Oxandra - Pachypodanthium - Papualthia - Petalolophus - Phaeanthus - Phoenicanthus - Piptostigma - Platymitra - Polyalthia - Polyaulax - Polyceratocarpus - Popowia - Porcelia - Pseudartabotrys - Pseudephedranthus - Pseudoxandra - Pseuduvaria - Pyramidanthe - Raimondia - Richella - Rollinia - Ruizodendron - Sageraea - Sapranthus - Schefferomitra - Sphaerocoryne - Stelechocarpus - Stenanona - Tetrameranthus - Tetrapetalum - Toussaintia - Tridimeris - Trigynaea - Trivalvaria - Unonopsis - Uvaria - Uvariastrum - Uvariodendron - Uvariopsis - Woodiellantha - Xylopia

Name

Annonaceae Adans.

References

* Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 6, May 2005. [1]
* Royal Botanic Gardens, Kew: Vascular Plant Families and Genera[2]

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Annonaceae, also called the custard apple family[2][3] is a family of flowering plants consisting of trees, shrubs or rarely lianas.[3] With about 2300 to 2500 species and more than 130 genera,[4] it is the largest family in Magnoliales. Only five genera, Annona, Rollinia, Uvaria, Melodorum and Asimina produce edible fruits.[4] 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.[5]

Description
Annona muricata

Mostly tropical, some mid-latitude, deciduous or evergreen trees and shrubs, some lianas, with aromatic bark, leaves, and flowers.[3]

Stems, stalks and leaves
Bark is fibrous and aromatic. Pith septate (fine tangential bands[5] 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.[6]
Leaves are alternate, two-ranked [6], 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 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.[3]

Fruits and seeds
Fruits are distinct (berries) or coalesce (into 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]

Systematics

Monophyly and inter-familial systematics have been well supported for Annonaceae by a combination of morphological and molecular evidence.[7] 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.[8] [9]

Guatteria, with its approximately 265 species, is the largest genus of Annonaceae, and might be broken into three small genera based on molecular characters from multiple locations.[10]

Tribe Saccopetaleae is another group of Annonaceae that is awaiting review as recent molecular studies suggest that this tribe is not monophyletic at all.[11]

Uses

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. diversifolia), soncoya (A. purpurea), atemoya (a cross between A. cherimola and A. squamosa); and biriba (Rollinia deliciosa, which may require reclassification under Annona.[1]). 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, annonancin, 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.[12][13][14][15]

Asimina triloba (pawpaw, prairie banana) has an Eastern U.S. distribution, and is currently under agricultural investigation as a commercial crop.[16]

Medical

The bark, leaves and roots of some species are used in folk medicines. Pharmaceutical research has found antifungal, bacteriostatic, antimalarial, and especially cytostatic capability of some chemical constituents of the leaves and bark. 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 have shown antibacterial properties, and have been used for treatment of medical conditions, such as skin disease, intestinal worms and inflammation of the eye. Pharmaceutical products are currently in animal and human cell-line trials.[17][18] Acetogenins are thought to have anti-HIV and anti-cancer effects. A wide variety of products have been developed and are available for cancer treatment.[19]
Cananga odorata flowers.

Insecticides

Flavonoids and alkaloids contained in the leafs and bark of several species of the family have shown insecticidal properties.[19]

Other

* Some species of the family, such as Cananga odorata (ylang-ylang) also have aromatic oil and are used for perfumes or spices.[5]
* The strong bark is used for carrying burdens in the Amazon Rainforest[5] and for wooden implements, such as tool handles and pegs.[19] The wood is valued as firewood.[5]
* Yellow and brown dyes.[19]
* Some species are also grown as ornamental plants, especially the Indian species Polyalthia longifolia pendula.

Genera

* Afroguatteria
* Alphonsea
* Ambavia
* Anaxagorea
* Ancana
* Annickia
* Annona
* Anomianthus
* Anonidium
* Artabotrys - trail-grape
* Asimina
* Asteranthe
* Balonga
* Bocagea
* Bocageopsis
* Boutiquea
* Cananga - Ylang-ylang
* Cardiopetalum
* Cleistochlamys
* Cleistopholis
* Craibella
* Cremastosperma
* Cyathocalyx
* Cyathostemma
* Cymbopetalum
* Dasoclema
* Dasymaschalon
* Deeringothamnus - false pawpaw
* Dendrokingstonia
* Dennettia
* Desmopsis
* Desmos
* Diclinanona
* Dielsiothamnus
* Disepalum
* Duckeanthus
* Duguetia
* Ellipeia
* Ellipeiopsis
* Enicosanthum
* Ephedranthus
* Exellia
* Fissistigma
* Fitzalania
* Friesodielsia
* Froesiodendron
* Fusaea
* Gilbertiella
* Goniothalamus
* Greenwayodendron
* Guamia
* Guatteria - haya minga, haya blanca
* Guatteriella
* Guatteriopsis
* Haplostichanthus
* Heteropetalum
* Hexalobus
* Hornschuchia
* Isolona
* Letestudoxa
* Lettowianthus
* Malmea
* Marsypopetalum
* Meiocarpidium
* Meiogyne
* Melodorum
* Mezzettia
* Mezzettiopsis
* Miliusa
* Mischogyne
* Mitrella
* Mitrephora
* Mkilua
* Monanthotaxis
* Monocarpia
* Monocyclanthus
* Monodora
* Neostenanthera
* Neo-uvaria
* Onychopetalum
* Ophrypetalum
* Oreomitra
* Orophea
* Oxandra - blacklancewood, haya
* Pachypodanthium
* Papualthia
* Petalolophus
* Phaeanthus
* Phoenicanthus
* Piptostigma
* Platymitra
* Polyalthia
* Polyceratocarpus
* Popowia
* Porcelia
* Pseudartabotrys
* Pseudephedranthus
* Pseudoxandra
* Pseuduvaria
* Pyramidanthe
* Raimondia
* Reedrollinsia
* Richella
* Rollinia - wild sugar-apple
* Ruizodendron
* Sageraea
* Sanrafaelia
* Sapranthus
* Schefferomitra
* Sphaerocoryne
* Stelechocarpus - kepel
* Stenanona
* Tetrameranthus
* Toussaintia
* Tridimeris
* Trigynaea
* Trivalvaria
* Unonopsis
* Uvaria
* Uvariastrum
* Uvariodendron
* Uvariopsis
* Woodiellantha
* Xylopia

References

1. ^ a b Germplasm Resources Information Network (GRIN) (2007-05-12). "Family: Annonaceae Juss., nom. cons.". Taxonomy for Plants. USDA, ARS, National Genetic Resources Program, National Germplasm Resources Laboratory, Beltsville, Maryland. http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?66. Retrieved 2008-04-18.
2. ^ "Annonaceae". Integrated Taxonomic Information System. http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=18092. Retrieved 18 March 2008.
3. ^ a b c d e f g Flora of North America. 2. Annonaceae Jussieu. 3. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10047. Retrieved 2008-04-20.
4. ^ a b Bridg, Hannia (2001-05-03). "Micropropagation and Determination of the in vitro Stability of Annona cherimola Mill. and Annona muricata L.". Zertifizierter Dokumentenserver der Humboldt-Universität zu Berlin. http://edoc.hu-berlin.de/dissertationen/bridg-hannia-2000-03-24/HTML/brigd-ch1.html. Retrieved 2008-04-20.
5. ^ a b c d e Chatrou, Dr. L.W. (2005-07-29). "Molecular Systematics of Annonaceae". Annonaceae research projects (Nationaal Herbarium Nederland). http://www.nationaalherbarium.nl/taskforcemolecular/annonaceae.htm#Molecular%20Systematics%20of%20Annonaceae. Retrieved 2008-04-20.
6. ^ a b Johnson, D.M. (July/September 2003). "Phylogenetic significance of spiral and distichous architecture in the Annonaceae.". Systematic Botany 28 (3): 503–511.
7. ^ 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.
8. ^ 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.
9. ^ Scharaschkin, T. and J.A. Doyle (2006). "Character evolution in Anaxagorea (Annonaceae)". Systematic Botany 93 (1): 36–54.
10. ^ Erkens, R.H.J.; Dr L.W. Chatrou, Dr J. Koek-Noorman and Prof. P.J.M. Maas (2005-07-29). "Phylogeny and evolution of Guatteria". Annonaceae research projects (Nationaal Herbarium Nederland). http://www.nationaalherbarium.nl/taskforcemolecular/annonaceae.htm#Phylogeny%20and%20evolution%20of%20Guatteria. Retrieved 2008-04-20.
11. ^ Mols, J.B.; Dr L.W. Chatrou, Dr J. Koek-Noorman and Prof. P.J.M. Maas (2005-07-29). "Phylogeny of Miliusa and allied Asian Annonaceae using cpDNA and morphology". Annonaceae research projects (Nationaal Herbarium Nederland). Retrieved 2008-04-20.
12. ^ 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.
13. ^ 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.
14. ^ 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. 70 (70): 153–7. doi:10.1007/978-3-211-45295-0_24. PMID 17017523.
15. ^ 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.
16. ^ 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.
17. ^ Cuendet, M.; et al. (2008). "Dietary administration of Asimina triloba (paw paw) extract increases tumor latency in N-methyl-N-nitrosourea-treated rats". Pharmaceutical Biology 46:1-2..
18. ^ Suedee, A.; et al. (2007). "Constituents of Polyalthia jucunda and their cytotoxic effect on human cancer cell lines.". Pharmaceutical Biology 45(7):575-579..
19. ^ a b c d University of Southampton (March 2002). "Factsheet No. 5. Annona" (PDF). Fruits for the Future. Department for International Development, International Centre for Underutilised Crops. http://www.icuc-iwmi.org/files/News/Resources/Factsheets/annona.pdf. Retrieved 2008-04-20.