Sapindaceae Juss.

Sapindaceae, Davies & B. Verdcourt. Flora of Tropical East Africa. 1998

Habit

Trees, shrubs or rarely herbs, sometimes climbing by tendrils

Indumentum

Indumentum commonly of simple hairs, sometimes in tufts in the axils of leaf-veins, rarely fasciculate; glands of several kinds occurring on both vegetative and flowering parts

Leaves

Leaves alternate, sometimes crowded (palm-like) at the apex of the tree, variously simple, ternate, biternate, pinnate or bipinnate, in the latter two kinds most often without a terminal leaflet at maturity; stipules lacking or small

Inflorescences

Inflorescences axillary or cauliflorous, thyrsoidal, paniculate or racemose

Flowers

Flowers regular or slightly zygomorphic, unisexual except in some Dodonaea but with non-functional organs of the other sex usually present

Calyx

Calyx of (3–)4–5(–7) free or partially united imbricate or valvate sepals

Corolla

Petals absent or 4–5, often with a basal claw, usually with a simple or elaborated scale on the inner face

Nectaries

Disk conspicuous, simple, or rarely a double ring, or reduced to a pair of glands

Androecium

Stamens 5–20(–74), often 8, but sometimes variable in number within a species, inserted in most cases inside the disk or occasionally on its surface; filaments free, terete or barely flattened, glabrous or hairy; anthers variously ovoid-sagittate to spherical, sometimes pilose, the connective rarely glandular, dehiscing introrsely by longitudinal slits; pollen grains tricolporate

Gynoecium

Ovary 1–8-locular; ovules usually 1–2 (rarely several) per locule; style apical, except in >i>Allophylus where semi-gynobasic, entire or 2–3-lobed

Fruits

Fruit a capsule, sometimes lobed, or drupe, often composed of one matured mericarp with the aborted remaining carpels visible at its base

Seeds

Seed usually with a hard black or brown testa, which in one case contains stomata, often with a conspicuous fleshy aril or sarcotesta, without endosperm

Acevedo-Rodríguez, P. (2009). Neotropical Sapindaceae.

Morphology

Description

Trees, shrubs, lianas, or less often herbaceous vines. Stems of climbing species usually with multiple vascular cylinders, and very often with white, milky latex . Stipules minute to large, present only in climbing plant species (tribe Paullinieae).  Leaves pinnately or ternately compound , rarely simple or unifoliolate, alternate , spirally arranged, rarely opposite; leaf rachis of most arborescent species with a terminal process (rudimentary leaflet ). Inflorescence axillary (usually solitary), terminal , or cauliflorous (usually fascicled), racemose, paniculate, or in spicate thyrses, with lateral dichasial or cincinnal cymes; peduncles angular to terete , bracts and bracteoles usually inconspicuous; pedicels usually articulate near the base.  Flowers actinomorphic or zygomorphic , seemingly bisexual but functionally unisexual, plants dichogamous, monoecious , or dioecious ; sepals 4-5, distinct or connate to various degrees; petals 4-5, rarely wanting, distinct, usually white or yellowish, with an adnate appendage on their adaxial surface, or the petals auriculate ( involute basal margins); petal appendage petaloid , simple , bifurcate, or hood-shaped with a glandular apex , variously pubescent ; disc extrastaminal, annular or unilateral, entire or lobed ; stamens (4-7)8(10), the filaments equal or unequal in length, free or connate at base, the anthers dorsifixed or basifixed, introrsely opening by longitudinal slits; ovary superior , of (2)3(-5) connate carpels, usually with same number of locules , the style usually present, the stigmas elongated or capitate ; placentation axial, the ovules 1 or 2 per locule .  Fruits capsules, schizocarps, or indehiscent and baccate , winged or unwinged, sometimes echinate . Seeds usually one per locule , or less often two, exarillate, arillate, or with a fleshy testa.

Distribution

Distribution in the Neotropics

• From southern United States to northern Argentina and Chile, including the West Indies.
• Widely distributed genera include:  Allophylus L., Cardiospermum L., Cupania L., Dodonaea Mill., Matayba Aubl., Paullinia L., Serjania Mill. and Urvillea Kunth.
• Restricted to South America:  Athyana Radlk., Diplokeleba N.E.Br., Guindilia Gill., Llagunoa Ruiz & Pav., Lophostigma Radlk., Magonia Vell., Pentascyphus Radlk., Porocystis Radlk., Pseudima Radlk., Scyphonychium Radlk., Toulicia Aubl., Tripterodendron Radlk. and Vouarana Aubl.
• Restricted to Caribbean area:  Thouinidium Radkl. and Thouinia Poit.
• Restricted to Mexico:  Ungnadia Engl., Blomia Miranda.
• Restricted to the West Indies:  Hypelate P.Browne.
• Endemic to western Cuba: Euchorium Ekman & Radlk.

Diagnostic

Notable genera and distinguishing features

Serjania: 

• Vines or lianas.
• Samaroid mericarps.
• The most speciose genus of Sapindaceae in the Neotropics with ca. 230 species.
• Abundant in open vegetation such as savannas, shrubby savannas and forest margins.
• Most diverse in the Brazilian central plateau and central Mexico.

Paullinia: 

• Vines or lianas.
• Coriaceous to woody capsules, opening to expose arillate seeds.
• The second largest genus of Sapindaceae with ca. 200 species in the Neotropics.
• Most diverse in the dense lowland to mid-elevation forests; most speciose in Brazil, Peru and Colombia.

Allophylus: 

• Ubiquitous shrubs or small trees.
• Ca. 100 species in humid, lowlands to mid-elevation forests in the Neotropics.

Talisia: 

• Treelets, small to medium-sized trees.
• 59 species most of which occur in the dense, lowlands humid forest of South America.
• Fruits indehiscent.

Cupania: 

• Small trees, ca. 50 species in dense, lowlands, humid Neotropical forests.
• Leaves pinnately compound.
• Leaflets mostly with serrate margins.
• Fruits capsular.
• Seeds arillate at the base.

Matayba: 

• Small trees
• Ca. 50 species in dense, lowlands, humid Neotropical forests.
• Leaves pinnately compound.
• Leaflets mostly with entire margins.
• Fruits capsular.
• Seeds arillate at the base.

Other important characters

• Lianescent species often with milky sap, and polystellar stems showing "anomalous secondary" growth.
• Extra-staminal floral disc.
• Seeds often arillate.

Distinguishing characters (always present)

• Leaf rachis of arborescent species with pinnately compound leaves present a rudimentary leaflet distally on the leaf rachis.
• Leaves biternate in lianescent species.
• Petals usually with an abaxialpetaloidappendage, or with involutebasal elongated marginal lobes.

Key differences from similar families

• Floral disc extra-staminal.
• Petals with adnate petaloidappendage.
• Trees with pinnately compound leaves with a terminal rudimentary leaflet (which does not develop into new leaflets).

Useful tips for generic identification

Key to genera of Neotropical Sapindaceae

1. Lianas or vines; often bearing a pair of coiled tendrils at the base of inflorescence (rarely herbaceous, and lacking tendrils in Cardiospermum ); stems usually producing milky sap; leaves always with a well develop terminal leaflet; stipules present (usually early deciduous) ... 2
1. Shrubs or trees (without tendrils); not producing milky sap; leaves with a rudimentary or a well developed terminal leaflet; stipules absent ... 8

2. Fruit a schizocarp, splitting into 3, indehiscent, winged mericarps (samaras) ... 3
2. Fruit a septicidal or septifragal, marginicidal capsule  ... 6

3. Thyrses with umbel-like units; flowers actinomorphic with an annular disc; mericarps with a basal locule and a distal wing. About 12 species from Central and South America ... Thinouia
3. Thyrse racemiform; flowers zygomorphic with a unilateral discdivided into 2-4 lobes; mericarps with a distal locule and a proximal wing, or the locule central, surrounded by a dorsal wing ... 4

4. Petals conspicuous, much projecting beyond the calyx; pollen grains triangular, triaperturate ... 5
4. Petals inconspicuous, not projecting beyond the calyx; pollen cylindrical ellipsoid, with 4 endoapetures and 2-3 colpi. Two species endemic to Bolivia, Peru and Ecuador ... Lophostigma

5. Mericarps with a distal locule and a proximal wing. Ca. 230 species from the Neotropics ... Serjania
5. Mericarps with a central locule, surrounded by a dorsal wing. Three species from Venezuela, Brazil,    Bolivia, Paraguay, and north eastern Argentina ... Houssayanthus

6. Capsules coriaceous or woody; seeds partially to completely arillate. ca. 200 species from the Neotropics ... Paullinia
6. Capsules membranous; seeds not arillate, black or dark brown, with a contrasting white, reniform or cordiformhilum; plants herbaceous or sub-woody... 7

7. Capsules with completely inflated carpels without a dorsal marginal wing or with a very narrow dorsal wing; stamens of unequal length; plants not producing a white exudate. Fourteen species eleven endemic to the Neotropics, and three cosmopolitan ... Cardiospermum
7. Capsules with partly inflated or flattened carpels with a dorsal marginal wing; stamens of equal length; plants usually with white exudate. Seventeen species from the Neotropics, one of which extends into the Lesser Antilles ... Urvillea

8. Leaves simple or unifoliolate ... 9
8. Leaves compound... 14

9. Flowers apetalous, with 2 ovules per carpel... 10
9. Flowers petaliferous, with a single ovule per carpel... 11

10. Plants not viscous; flowers zygomorphic, solitary or in axillary cymes; floral disc unilateral, semi-annular; stamens 8; stigma subcapitate; fruit a 3-lobed, crustose, loculicidal capsule. Three or four species from the Andean highlands of South America ... Llagunoa
10. Plant viscous; flowers actinomorphic, in axillary or terminal racemose or paniculate thyrses; floral disc wanting or rudimentary; stamens 5-15; stigma grooved or divided; fruit a 2-6-locular, septifragal or septicidalcapsule. Primarily Australian genus with 2 or 3 species in the Neotropics ... Dodonaea

11. Calyx 5-merous... 12
11. Calyx 4-merous... 13

12.  Leaves alternate, lobed or serrate; floral disc semi-annular, 4-lobed; fruit splitting into 3 samaroid, papery mericarps with a short dorsal wing. A single species from Andean Chile ... Bridgesia
12.  Leaves alternate or opposite, entire or tridentate at apex; floral disc 2-lobed; fruit splitting into 1-3 subglobose, crustose mericarps. Three species from southern South America ... Guindilia

13. Fruits schizocarpic, splitting into 3 samaroid mericarps, each bearing a distal wing. About 30 species from Central America and the Greater Antilles ... Thouinia
13. Fruit an indehiscent drupe, 1-2 coccate, with crustose endocarp and fleshy exocarp. Pantropical with about 100 species in the Neotropics ... Allophylus

14. Leaves trifoliolate... 15
14. Leaves pinnately compound; terminal leaflet rudimentary ... 19

15. Leaves opposite. Two species from southern Mexico to northern South America ... Billia
15. Leaves alternate... 16

16. Flowers apetalous. Three or four species from the Andean highlands of South America ... Llagunoa
16. Flowers petaliferous ... 17

17. Fruits schizocarpic, splitting into 3 samaroid mericarps, each bearing a distal wing ... Thouinia
17. Fruit an indehiscent drupe with fleshy exocarp... 18

18. Petals without appendages; disc annular, obsolete 5-lobed; ovary 3-carpellate, with 2 ovules per carpel; stigma capitate; fruit 1-locular, with woody endocarp. A single species endemic to the West Indies and the south eastern United States (Florida) ... Hypelate
18. Petals with a single, 2-lobed appendage or 2 marginal appendages; disc unilateral; ovary 2 (-3)-carpellate, with a single ovule per carpel; style terminal with 2-3 stigmatic branches; fruit 1-2 coccate, with crustose endocarp... Allophylus

19.  Leaves bipinnate or tripinnate... 20
19.  Leaves once pinnate... 21

20. Leaves tripinnate. A single species endemic to east-central Brazil ... Tripterodendron
20. Leaves bipinnate. Three species from tropical continental America ... Dilodendron

21. Petals without appendages ... 22
21. Petals with adaxial or marginal appendages ... 28

22. Ovary with 2-8 ovules per carpel... 23
22. Ovary with uniovulate carpels ... 27

23. Ovules 8 per carpel; floral disc complete, one half consisting of 4 short glands, the other half of two concentric erect, fleshy laminae, the outer one taller; pollen grains in tetrads; fruit a large (8-12 cm long), 3-locular, trigonous, woody loculicidal capsule. One or two species from Brazil, Bolivia and Paraguay ... Magonia
23. Ovules 2 per carpel; floral disc annular; pollen grains in monads; fruit  < 4 cm long ... 24

24. Fruit unknown. Endemic to Western Cuba ... Euchorium
24. Fruits indehiscent, or a tardily loculicidal crustose capsule. Widely distributed ... 25

25. Petals wanting or rudimentary; fruit a loculicidal, crustose capsule. Four species, one in Mexico, the remaining from tropical South America ... Averrhoidium
25. Petals present; fruit indehiscent... 26

26. Inflorescences terminal panicles or racemes; calyx actinomorphic; fruit with leathery pericarp, sub-globose or ellipsoid; seeds 1(2) with edible sarcotesta.  Ten species, nine native to South America, one to Dominican Republic ... Melicoccus
26. Inflorescences of axillary or sub-terminal corymbose thyrses; calyx zygomorphic; fruits globose, baccate; seeds with coriaceous-fleshy testa. Three species distributed throughout the West Indies, Mexico, Guatemala, Colombia, and Ecuador ... Exothea

27. Floral disc 5-lobed; fruit a 2-(3)-coccate, loculicidal, leathery capsule, the cocci equally developed or one of them rudimentary; seeds large, arillate, not winged. One species from the lowlands of tropical continental America ... Pseudima
27. Floral disc cup-shaped, crenulate; fruit a 3-locular, woody loculicidal capsule; seeds winged. Two species from southern South America ... Diplokeleba        

28. Fruit indehiscent or schizocarpic ... 29
28. Fruit a capsule... 36

29. Fruit indehiscent, leathery to woody; young branches usually developing cataphyls or bud scales. Fifty nine species from Central and South America ... Talisia
29. Fruit schizocarpic ... 30

30. Mericarps winged... 31
30. Mericarps globose... 34

31. Sepals 5; ovary 3-carpellate; mericarps 3 ... 32
31. Sepals 4; ovary 2-carpellate; mericarps 2.  Two species from southern South America (Brazil, Bolivia, Paraguay, and Argentina) ... Diatenopteryx

32. Mericarps distally winged, with a coriaceous, non-inflated locule... 33
32. Mericaprs basally winged, with a papery inflated locule. About 12 species from the lowlands of South America ... Toulicia

33. Leaf rachiswinged; petal appendage hood-shaped; disc semi-annular-lobed; stigma capitate. A single species from southern South America (Peru, Bolivia, Argentina) ...  Athyana
33. Leaf rachis un-winged; petal appendage adnate to the petal to form a pocket above the petal's claw; disc cup-shaped; stigma punctiform. Six species from Mexico, Central America and some islands in the Greater Antilles ... Thouinidium

34. Mericarps membranous, wrinkled, inflated. Three species, 2 from Brazil (Amazonas) and 1 from French Guiana ... Porocystis
34. Mericarps coriaceous or woody, non-inflated nor wrinkled ... 35

35. Fruit with 2(1) rudimentary coriaceous cocci, with fleshy pericarp containing much saponin. About ten  species with tropical to sub -temperate distribution, 1 species in the Neotropics ... Sapindus
35. Fruit (1)2-coccate, woody. A single species native to northern and eastern Brazil, and French Guiana ... Scyphonichium

36. Ovary unicarpellate; seeds with thin sarcotesta. A single species distributed in Mexico, Guatemala, and Belize ... Blomia
36. Ovary 2-3-carpellate; seeds naked or arillate at base ... 37

37. Sepals unequal ... 38
37. Sepals of similar size and shape ... 39

38. Ovary 2-carpellate with a single ovule per carpel; floral disc annular, without androgynophore; seed arillate at base Two species from Costa Rica to northern Brazil ... Vouarana
38. Ovary 3-carpellate with two ovules per carpel; floral disc unilateral, undulate with an androgynophore; seeds exarillate, with a large hilum at base. A single species endemic to Mexico and southern United States ... Ungnadia

39. Sepals longer than wide, imbricate... 40
39. Sepals as long as wide or slightly longer than wide, valvate. About 50 species from tropical and subtropical America ... Matayba

40. Petals with a pair of tomentosemarginal appendages; leaflets usually serrate. About 50 species from tropical and subtropical America ... Cupania
40. Petals with a single short, basal appendageconnate to petal's margins; leaflets entire.  A single species occurring in French Guiana, Surinam, and Brazil (Amazonas) ... Pentascyphus

General Description

Status

Exotic genera:

• Blighiasapida Kon. is an important crop in Jamaica; it is cultivated for its edible arils.
• Dimocarpuslongan Lour., Litchichinensis Sonn., and Nepheliumlappaceum L. are cultivated for their edible fruits in small-scale operations throughout tropical America.
• Acer L. spp., Filicium Thwaites spp., Koelreuteria Schreb. spp., and Harpulliaarborea Radlk. are cultivated as ornamental trees throughout the Neotropics.

General notes

• Sapindaceae are the source of numerous products, some of which are economically important. 
• In the Neotropics these include edible fruits such as keneep or genip (Melicoccusbijugatus Jacq.), wild genip (M. oliviformis Kunth) and the pitomba (Talisiaesculenta Radlk.).
• Numerous species of Paullinia have been reported to be useful in the preparation of medicines, caffeine-rich beverages, binding and weaving material, and for fish, human and arrow poisoning.
• The seeds of Paulliniacupana Kunth are the source of the important Brazilian crop guaraná, a source of caffeine and flavoring of soft drinks.
• Almost all Sapindaceae are used around the tropics for fish poisoning.

Number of genera

A family of 141 genera and about 1,900 species worldwide; 38 genera in the Neotropics with about 800 species:

• Allophylus
• Athyana
• Averrhoidium Baill.
• Billia Peyr.
• Blomia
• Bridgesia Bertero ex Cambess.
• Cardiospermum
• Cupania
• Diatenopteryx Radlk.
• Dilodendron Radlk.
• Diplokeleba
• Dodonaea
• Euchorium
• Exothea Macfad.
• Guindilia
• Houssayanthus Hunz.
• Hypelate
• Llagunoa
• Lophostigma
• Magonia
• Matayba
• Melicoccus P.Browne
• Paullinia
• Pentascyphus
• Porocystis
• Pseudima
• Sapindus L.
• Scyphonychium
• Serjania
• Talisia Aubl.
• Thinouia Planch. & Triana
• Thouinia
• Thouinidium
• Toulicia Aubl.
• Tripterodendron
• Ungnadia
• Urvillea
• Vouarana

Notes on delimitation

• Summary of the most current phylogeny of Sapindaceae is presented in the Angiosperm Phylogeny Website by Stevens [http://www.mobot.org/MOBOT/Research/APweb/welcome.html] places Sapindaceae (including Aceraceae and Hippocastanaceae) in its own order along with families Biebersteiniaceae, Nitraceae, Kirkiaceae, Burseraceae, Anacardiaceae, Simaroubaceae, Meliaceae, and Rutaceae.
• Sapindaceae, accordingly, is made of subfamilies Xanthoceroideae, Hippocastanoideae, Dodonaeideae, and Sapindoideae.

Literature

Important literature

Acevedo-Rodríguez, P. 1993. Systematics of Serjania (Sapindaceae). Part I: A revision of Serjania Sect. Platycoccus. Mem. New York Bot. Gard. 67: 1-93.

Acevedo-Rodríguez, P. 1993. A revision of Lophostigma (Sapindaceae). Syst. Bot. 18: 379-388.

Acevedo-Rodríguez P., 2002. Sapindaceae. In: S. Mori, et al., Manual to the Vascular Plants of Central French Guiana. Mem. New York Bot. Gard. 77: (2): 653-667.

Acevedo-Rodríguez P., 2003. Melicocceae (Sapindaceae). Talisia and Melicoccus. Fl. Neotrop. 87: 1-179.

Acevedo-Rodríguez P. & M.S. Ferrucci. 2002. Averrhoidium dalyi (Sapindaceae) a new species from western Amazonia. Brittonia 54: 112-115.

Acevedo-Rodríguez P., 2005. Sapindaceae (excluding Paullinia) In: Steyermark, J.A., P. Berry & B. Holts (eds.) Flora of the Venezuelan Guayana. Timber Press.

Acevedo-Rodríguez P., P. van Welzen, F. Adema, and R.W.J.M. van der Ham. (submitted). Sapindaceae. In: Kubitzki et al. Family and Genera of Flowering Plants, Springer Verlag. Berlin, Heidelberg, New York.

Barkley, F.A. 1957. Sapindaceae of southern South America. Lilloa 28: 112-179.

Beck, H.T. 1990. A survey of the useful species of Paullinia L. (Sapindaceae). Adv. Econ. Bot. 8: 41-56.

Croat, T.B. 1976. Sapindaceae. Flora of Panama. Ann. Missouri Bot. Gard. 63(3): 419-540.

Ferrucci, M.S. 1991. Sapindaceae In: Spichiger & Ramella (Eds.), Flora del Paraguay. Conservatoire et Jardin botaniques de la Ville de Geneve & Missouri Botanical Garden.Ferrucci, M.S. 2000. Revision taxonómica de los géneros Cardiospermum y Urvillea para el neotropico (Sapindaceae). Ph.D. dissertation, Universidad Nacional de Cordoba, Argentina, 262 pp.

Hoehne, F.C. 1925. Sapindaceae Mattogrossenses. Archivos Bot. sao Paulo 1: 134-142.

Klaassen, R. K.W.M. 1999. Wood anatomy of Sapindaceae. IAWA Journal, Suppl. 2: 1-214.

Kramer, K.U. 1972. Studies in the New World Sapindaceae. I. Notes on the genus Talisia Aublet. Act. Bot. Neerl. 21: 671-678.

Kramer, K.U. 1976. Sapindaceae (additions and corrections). In: Lanjouw & A.L. Stoffers, Flora of Surinam. 2(2): 487-511.

Leenhouts, P.W. 1967. A conspectus of the genus Allophylus (Sapindaceae) The problem of the complex species. Blumea 15: 301-358.

Lippold, H. 1978. Die Gattung Dodonaea Miller (Sapindaceae) in Amerika. Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe 6: 79-126.

MacBride, J.F. 1965. Sapindaceae in Flora of Peru. Field Mus. Pub. Bot. 13(3a): 291-391.

Muller, J. and P. W. Leenhouts. 1976. A general survey of pollen types in Sapindaceae in relation to taxonomy. Ppg. 407-445. In: I.K. Ferguson & J. Muller (eds.), The evolutionary significance of the exine. Linnaean Soc. Symposium Ser. 1. Academic Press, London.

Neto, G.G., 1996. Ocorrencia e distribuçao da Familia Sapindaceae Jussieu nos estados de Mato Grosso, Goias e Tocantins. Bol. Mus. Para emilio Goeldi, ser. Bot. 12(2): 227-236.

Radlkofer, L. 1890. Ueber die Gliederung der Familie der Sapindaceen. Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 20: 105-379.

Radlkofer, L. 1905. Sapindaceae Costaricenses. Bull. Herb. Boissier (2 Ser.) 5: 319-328.

Radlkofer, L. 1931-34. Sapindaceae. Pages 1-1539. In: A. Engler (ed.), Das Pflanzenreich IV, 165 (Heft 98a-h). Verlag von Wilhelm Engelmann, Leipzig.

Rambo, B. 1952. Sapindaceae Riograndenses. Anais Botanicos NR 4: 162-185.

Rashford, J. 2001. Those that do not smile will kill me: The ethnobotany of the Ackee in Jamaica. Econ. Bot. 55: 190-211.

Reitz, P. 1980. Sapindaceae in: Flora ilustrada Catarinese. 1st part.

Soukup, J. 1969. Las Aceraceae y Sapindaceas del Peru, sus géneros y lista de especies. Biota VIII 61: 53-68.

Uittien, H. 1937. Sapindaceae. In: A. Pulle, Flora of Surinam. 2(1): 345-396.

West, J.G. 1984. A revision of Dodonaea Miller (Sapindaceae) in Australia. Brunonia 7: 1 194.

Sapindaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Habit

Trees, shrubs or climbers

Leaves

Leaves alternate or rarely opposite, simple, or 1-or 3-foliolate, or pinnate or bipinnate; stipules rarely present

Flowers

Flowers actinomorphic or zygomorphic, sometimes very small, mostly unisexual, variously arranged

Calyx

Sepals imbricate or rarely valvate

Corolla

Petals 1–5, sometimes absent, imbricate

Nectaries

Disk usually present, sometimes unilateral

Androecium

Stamens hypogynous, often 8, inserted within the disk or unilateral; filaments free, often hairy; anthers 2-celled

Gynoecium

Ovary superior, entire or vertically lobed to the base, 1–8-celled; style terminal or gynobasic; ovules 1–2, rarely many in each cell, axile

Fruits

Fruit capsular or indehiscent, rarely winged

Seeds

Seeds without endosperm, often conspicuously arillate; embryo with usually plicate or twisted cotyledons