Susan K. Pell
Brooklyn Botanic Garden, New York, USA.
Trees, shrubs, rarely subshrubs or lianas, frequently with contact dermatitis-causing exudate. Vertical resin canals present in bark and in phloem of petioles and large veins of leaves, also widely present in fruits, flowers, and other tissues. Leaves simple or pinnately compound; alternate, rarely opposite or whorled; sessile or petiolate; leaflets opposite, subopposite, or alternate, entire, serrate, dentate, or crenate; estipulate. Inflorescences terminal and/or axillary, thyrsoid, paniculate, racemose, or spicate; rarely flowers solitary. Flowers actinomorphic, unisexual or bisexual; plants dioecious, monoecious, andromonoecious, polygamous, or hermaphrodite; pedicels often articulate; hypanthium sometimes present; perianth usually 2-whorled, rarely 1-whorled or absent, imbricate or valvate; sepals (3-)4-5, usually basally fused, rarely fully connate and cup-shaped; petals (3-)4-5(-8), rarely absent; stamens (1-)5-10, in 1 or 2 (rarely more) whorls, in some genera only 1 or 2 stamens fertile, filaments free, rarely basally connate, anthers dorsi- or basifixed, usually longitudinally dehiscent, introrse, rarely extrorse; disc intra-staminal, rarely extra-staminal or absent; ovary superior, rarely inferior, solitary or of 2-5 connate carpels, ovule solitary, apotropous, basal, apical, or lateral, style apical or lateral, erect or recurved, rarely sigmoid, stigma capitate, discoid, lobate, or spathulate, rarely punctiform. Fruits drupes or samaras (rarely syncarps, utricles, nut-like, or baccates), fleshy or dry, occasionally subtended by a fleshy hypocarp or an accrescent, chartaceous or fleshy calyx; mesocarp sometimes with prominent black resin canals. Seeds 1-5; endosperm scant or absent; embryo curved or straight (rarely horseshoe-shaped or pyramidal), cotyledons usually planoconvex or flat and free, usually equal in size, rarely fused or ruminate, sometimes bilobed.
Notes on delimitation
- Pell and Mitchell split the family into two subfamilies, Anacardioideae and Spondioideae (Mitchell et al. 2007). However, recent phylogenetic evidence (Pell unpublished data) suggests that subfamily Spondioideae may be polyphyletic.
Distribution in the Neotropics
- Actinocheita F.A.Barkley - endemic to south-central Mexico.
- Amphipterygium Schiede ex Standl. - western Mexico to northwestern Costa Rica.
- Anacardium L. - Honduras to Paraguay, Brazil and Bolivia.
- Antrocaryon Pierre - Amazonian Brazil (also found in Africa).
- Apterokarpos Rizzini - endemic to the Caatinga of northeastern Brazil.
- Astronium Jacq. - Mexico to Paraguay and northern Argentina.
- Bonetiella Rzed. - northern to central Mexico.
- Campnosperma Thwaites - Honduras to Ecuador and Amazonia (also found in the paleotropics).
- Cardenasiodendron F.A.Barkley - endemic to Bolivia.
- Comocladia P.Browne - central Mexico to Guatemala, Belize and the Greater and Lesser Antilles.
- Cyrtocarpa Kunth - southern Baja California, Mexico, and Colombia east to Guyana, Venezuela and northern Brazil.
- Haplorhus Engl. - endemic to the dry inter-Andean valleys of Peru to northern Chile.
- Lithrea Hook. - Brazil, Bolivia, Paraguay, Argentina, Uruguay, and Chile.
- Loxopterygium Hook.f. - Venezuela to Argentina, absent from Amazonia.
- Malosma Engl. - southern California, USA to central Baja California, Mexico.
- Mauria Kunth - El Salvador to eastern Venezuela and northern Argentina.
- Metopium P.Browne - West Indies, southern Florida, USA, Mexico and northern Central America.
- Mosquitoxylum Krug & Urb. - Jamaica, and southern Mexico to northwestern Ecuador.
- Myracrodruon Allem. - sub -Amazonian Brazil, Bolivia, Paraguay, and northern Argentina.
- Ochoterenaea F.A.Barkley - Panama, Andean Venezuela and Colombia to Bolivia.
- Orthopterygium Hemsl. - endemic to western Peru.
- Pachycormus Coville - endemic to central Baja California, Mexico.
- Pistacia L. - Texas, USA, to Nicaragua.
- Pseudosmodingium Engl. - endemic to central and southern Mexico.
- Rhus L. - southern Canada to Panama and Cuba (also found in Africa, Asia, and Europe).
- Schinopsis Engl. - northern Peru, sub -Amazonian and eastern Brazil south to central Argentina.
- Schinus L. - Ecuador to Patagonia, absent from Amazonia.
- Spondias L. - Mexico to Brazil and Bolivia (also found in Asia and the Pacific).
- Tapirira Aubl. - southern Mexico to Brazil, Bolivia and Paraguay.
- Thyrsodium Salzm. ex Benth. - Colombia, Peru, Bolivia, Venezuela, the Guianas, and Amazonian and eastern Brazil, absent from the Andes.
- Toxicodendron Mill. - southern Canada to Bolivia (also found in Asia).
Distinguishing characters (always present)
Other important characters
- Intrastaminal nectariferous disc.
- Diplostemonous or haplostemonous.
- Some genera cause contact dermatitis.
- If compound, usually imparipinnate; or if paripinnate, determinate.
- Fruit never dehiscent.
Key differences from similar families
- Anacardiaceae can be separated from Burseraceae by the latter virtually always having opposite leaflets, whereas Anacardiaceae leaflets are usually alternate or subopposite.
Number of genera
- There are 81 (see list above) genera of Anacardiaceae and ca. 800 species worldwide, of which 31 are native to the Neotropics. Attilaea E.Martνnez & Ramos, a recently published genus endemic to the Yucatan peninsula of Mexico, is not included in this list due to a lack of accessible material. Naturalized Mangifera L. is not listed because it is not native to the Neotropics.
Useful tips for generic identification
Key to genera of Neotropical Anacardiaceae
1. Exudate often present and sometimes causing contact dermatitis; leaves simple, unifoliolate, or multifoliolate; carpels 1-3; exocarp usually thin; opercula absent; fruit sometimes wind dispersed 2 (Anacardioideae)1. Exudate often present and not causing contact dermatitis (very rarely causing dermatitis in Spondias); leaves multifoliolate; carpels (3-) 5; exocarp thick; opercula often present; fruit never wind dispersed 40 (Spondioideae)
3. Unicarpellate 43. Bicarpellate or tricarpellate 5
4. Domatia often present in abaxial secondary vein axils; glandular ridges absent from petals; drupe usually subtended by fleshy hypocarp; mesocarp woody...Anacardium 4. Domatia absent; glandular ridges on petals; hypocarp absent; mesocarp fleshy or resinous Mangifera (naturalised)
9. Calyx deeply lobed; stylodia usually 3; exocarp generally separating from mesocarp at maturity, endocarp bony Schinus p. p.9. Calyx shallowly lobed; stylodia always 3; exocarp generally separating from mesocarp at maturity, endocarp chartaceous Mauria p. p.
11. Fruit with elongated wing-like structure 1211. Fruits without elongated wing-like structure 13
12. Exudate turning black with exposure to air, causes contact dermatitis; leaves entire; fruit a laterally winged samara; South America Schinopsis p. p. 12. Exudate milky, nontoxic; leaves serrate; fruit a syncarp subtended by a winged peduncle; Mexico to Costa Rica Amphipterygium p. p.
13. Leaves entire with marginal secondary vein; exocarp readily separating from mesocarp; mesocarp waxy and white with black resin canals; South America Lithrea p. p.13. Leaves entire to deeply lobed (both on same plant), marginal secondary vein absent; exocarp and mesocarp not as above; Mexico Bonetiella
15. Male flowers in pendent thyrses, female flowers tightly arranged & subtended by an involucre; fruit winged Amphipterygium p. p.15. Female and male inflorescences of the same type or if different, not as above; fruit not winged 16
16. Exudate turning black with exposure to air; exocarp white or grey to yellowish and readily separating from mesocarp, mesocarp waxy with black resin canals 1716. Exudate absent; exocarp reddish with glandular trichomes and remaining attached to mesocarp, mesocarp not as above Rhus subgenus Lobadium p. p.
17. Marginal secondary vein present; hairy tuft domatia absent; androecium diplostemonous; South America Lithrea p. p.17. Marginal secondary vein absent; hairy tuft domatia sometimes present in secondary vein axils abaxially; androecium haplostemonous; Mexico to Guatemala Toxicodendron p. p.
18. Fruit likely wind dispersed (samaras, dry drupes with enlarged trichomes, utricles, or drupes subtended by stiffened enlarged sepals) 1918. Fruit likely animal dispersed (fleshy drupes or dry but not as above) 30
20. Fruit subtended by stiffened enlarged sepals 2120. Fruit not as above 22
23. Male inflorescence pendant and large; sepals 5-8, longer than stamens; Mexico to Costa Rica Amphipterygium p. p.23. Male inflorescence erect and small; sepals 3-8, shorter than stamens; Peru and Colombia Orthopterygium
25. Leaflets entire; fruit a samara, laterally compressed with trichomes occurring prominently on the margins; inflorescence corymbose; Venezuela and Colombia south to Bolivia Ochoterenaea 25. Leaflets broadly pinnately lobed (leaf fern-like); fruit a globose drupe covered with trichomes; inflorescence not corymbose; Mexico Actinocheita
28. Plants polygamodioecious; leaves usually deciduous; wing chartaceous with conspicuous venation, consisting of exocarp tissue Loxopterygium 28. Plants dioecious or monoecious; leaves usually evergreen; wing stiffened and thick, lacking conspicuous venation, consisting of exocarp and mesocarp tissue Schinopsis p. p.
29. Flowers sessile; stylodia 3; ovule basal; drupe with two unequal, narrow lateral wings; Bolivia Cardenasiodendron 29. Flowers pedicellate; style 3-branched; ovule apical; drupe with two equal, broad lateral wings; Mexico Pseudosmodingium
30. Flowers apetalous; stylodia plumose Pistacia 30. Flowers with petals; stylodia not plumose 31
32. Hypanthium present Thyrsodium 32. Hypanthium absent 33
34. Exudate usually turning black with exposure to air; leaves often with black spots where injured; exudate toxic 3534. Exudate not turning black with exposure to air; leaves without conspicuous black spots; exudate not toxic (possibly toxic in Apterokarpos) 36
35. Plants polygamodioecious; leaves usually deciduous (rarely evergreen); stylodia 3; exocarp white to grey or yellow, easily separating from mesocarp; mesocarp white or gray with black resin canals Toxicodendron p. p.35. Plants dioecious; leaves evergreen; style 1; exocarp orange to brown, not easily separating from mesocarp; mesocarp resin canals not conspicuous Metopium
37. Erect or scandent shrubs or small trees (up to 7 m); fruit covered with glandular trichomes; seed fills most of the locule Rhus p. p.37. Medium to large trees (up to 20 m); fruit glabrous; seed fills only a small portion of the locule Mosquitoxylum
39. Leaflets petiolulate; corolla valvate or subvalvate; exocarp not easily separating from mesocarp at maturity Mauria p. p.39. Leaflets sessile to subsessile; corolla imbricate; exocarp easily separating from mesocarp at maturity Schinus p. p.
42. Plants polygamodioecious; leaves deciduous; drupe radially symmetric (pedicel scar on stone usually centered); fruit maturing yellow or light orange, endocarp surface pitted Antrocaryon 42. Plants hermaphrodite; leaves evergreen; drupe oblique (pedicel scar on stone markedly excentric); fruit maturing black or brown, endocarp surface relatively smooth Dracontomelon
43. Female flowers usually solitary; stamens 10-16(23); pistillode absent Sclerocarya 43. Female flowers, if present, not solitary; stamens usually 10; pistillode present, or absent and flowers hermaphroditic 44
44. Leaves evergreen; anther connective eglandular; pistil(ode) sparsely to densely pubescent; endocarp cartilaginous to crustaceous (rarely bony, e.g. Tapirira mexicana) .Tapirira 44. Leaves deciduous; anther connective usually glandular; pistil(ode) usually glabrous; endocarp bony Cyrtocarpa
- Mangifera indica L. has naturalized throughout the Neotropics.
- Toxicodendron succedaneum (L.) Kuntze has naturalized in Brazil and possibly elsewhere.
- Several non-native Anacardiaceae are cultivated in the Neotropics for their edible fruits: Bouea macrophylla Griff., Harpephyllum caffrum Bernh. ex Krauss, Mangifera indica, Schinus terebinthifolia Raddi, Sclerocarya birrea Hochst. subspecies caffra (Sond.) Kokwaro, and Spondias dulcis G.Forst..
Mitchell, J. D., and S. A. Mori. 1987. The cashew and its relatives (Anacardium: Anacardiaceae). Memoirs of the New York Botanical Garden 42: 1-76.
Mitchell, J. D. 1990. The poisonous Anacardiaceae genera of the world. Advances in Economic Botany 8: 103-129.
Mitchell, J. D. 2004. Anacardiaceae. In Smith, N., S. A. Mori, A. Henderson, D. W. Stevenson, and S. V. Heald (eds.) Flowering Plants of the Neotropics. pgs. 14-16, Princeton University Press, Princeton, NJ, USA.
Pell, S. K., J. D. Mitchell, T. Lobova, and A. J. Miller. In press. Anacardiaceae. In K. Kubitzki (Ed.), The Families and Genera of Vascular Plants. Springer.
Ribeiro, J.E.L.S., Hopkins, M.J.G., Vicentini, A., Sothers, C.A., Costa, M.A.S., Brito, J.M., Souza, M.A.D., Martins, L.H.P., Lohmann, L.G., Assunηγo, P.A.C.L., Pereira, E.C., Silva, C.F., Mesquita, M.R. & Procσpio, L.C. 1999. Flora da Reserva Ducke: Guia de identificaηγo das plantas vasculares de uma floresta de terra firme na Amazτnia Central. INPA, Manaus. 816 pp.
Wannan, B. S., and C. Quinn. 1990. Pericarp structure and generic affinities in the Anacardiaceae. Botanical Journal of the Linnean Society 102: 225-252.
Wannan, B. S., and C. Quinn. 1991. Floral structure and evolution in the Anacardiaceae. Botanical Journal of the Linnean Society 107: 349-85.
Various Neotropical floras are important as well. These include, among others: Flora de Nicaragua, Guide to the Vascular Plants of Central French Guiana, Flora of Baja California, and Flora of the Venezuelan Guayana.
How to cite
Pell, S.K. (2009). Neotropical Anacardiaceae. In: Milliken, W., Klitgεrd, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Anacardiaceae.htm.
Click images to enlarge
Anacardium giganteum © William Milliken, Royal Botanic Gardens, Kew.
Anacardium humile © Taylor/Zappi, RBG Kew.
Cashew nut Anacardium occidentale L. © Peter Gasson, Royal Botanic Gardens, Kew.
Habit Cardenasiodendron brachypterum © John Wood, Darwin Initiative Project 161/11/015.
Lithraea sp. © John Wood, Darwin Initiative Project 161/11/016.
Lithraea sp. © John Wood, Darwin Initiative Project 161/11/017.
Lithraea sp. © John Wood, Darwin Initiative Project 161/11/018.
Lithraea sp. © John Wood, Darwin Initiative Project 161/11/019.
Loxopterygium sp.© John Wood, Darwin Initiative Project 161/11/020.
Loxopterygium sp.© John Wood, Darwin Initiative Project 161/11/021.
Loxopterygium sp.© John Wood, Darwin Initiative Project 161/11/022.
Myracrodruon urundeuva © Nigel Taylor & Daniela Zappi/RBG Kew.
Schinopsis brasiliensis © John Wood, Darwin Initiative Project 161/11/015.
Schinus sp. © John Wood, Darwin Initiative Project 161/11/023.
Schinus longifolia © John Wood, Darwin Initiative Project 161/11/025.
Schinus longifolia © John Wood, Darwin Initiative Project 161/11/026.
Schinus molle © John Wood, Darwin Initiative Project 161/11/027.
Schinus molle © John Wood, Darwin Initiative Project 161/11/028.
Schinus molle © John Wood/Darwin Initiative Project 161/11/030
Schinus sp. © John Wood/Darwin Initiative Project 161/11/023
Schinus sp. © John Wood/Darwin Initiative Project 161/11/024
Schinus sp. © John Wood, Darwin Initiative Project 161/11/024.
Schinus sp. © John Wood, Darwin Initiative Project 161/11/024.
Tapirira obtusa © Denise Sasaki, Programa Flora Cristalino.
Tapirira obtusa © Denise Sasaki, Programa Flora Cristalino.