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The families of flowering plants

L. Watson and M.J. Dallwitz

Sapindaceae Juss.

Including Dodonaeaceae Link, Dyssapindaceae Radlk., Koelreuteriaceae J.G. Agardh, Saponaceae Vent.; excluding Hippocastanceae, Ptaeroxylaceae

Habit and leaf form. Trees, or ‘arborescent’ (a few being palm- or tree-fern like in habit), or shrubs, or lianas, or herbs; laticiferous (often), or non-laticiferous, without coloured juice; not resinous. Plants green and photosynthesizing. Self supporting, or climbing; the climbers tendril climbers (the tendrils representing modified infloresences), or scrambling. Mesophytic. Leaves alternate; spiral; petiolate; non-sheathing; gland-dotted (often), or not gland-dotted; without marked odour; compound (usually), or simple (but it remains to be ascertained how often records of this condition could reasonably be interpreted as ‘unifoliolate’); often pulvinate; usually ternate, or pinnate, or bipinnate, or multiply compound (sometimes biternate). Lamina when simple, dissected to entire; when more or less dissected, pinnatifid; pinnately veined; cross-venulate. Leaves stipulate (in some climbers), or exstipulate. Domatia occurring in the family (recorded in 17 genera and numerous species); manifested as pits, or pockets, or hair tufts.

General anatomy. Plants with ‘crystal sand’ (occasionally), or without ‘crystal sand’.

Leaf anatomy. The leaf lamina dorsiventral (usually), or bifacial to centric. Extra-floral nectaries absent. Abaxial epidermis papillose (often), or not papillose. Mucilaginous epidermis present (very commonly), or absent. Stomata anomocytic (usually), or paracytic. Hairs of numerous kinds present (in the family); eglandular, or glandular (often, on young leaves); unicellular, or multicellular. Unicellular hairs branched (often 2-armed), or simple. Complex hairs present (often), or absent; when present, peltate, or stellate (or tufted). Adaxial hypodermis present (often), or absent. Lamina without secretory cavities. The mesophyll with spherical etherial oil cells, or without etherial oil cells (?); containing mucilage cells, or not containing mucilage cells (?); with sclerenchymatous idioblasts, or without sclerenchymatous idioblasts; usually containing crystals. The crystals druses, or solitary-prismatic. Minor leaf veins without phloem transfer cells (Cardiospermum, Dodonaea, Melicocca).

Axial (stem, wood) anatomy. Secretory cavities absent (i.e., no resin canals). Cork cambium present; initially superficial (usually), or initially deep-seated (e.g. Dodonaea). Nodes tri-lacunar. Primary vascular tissues in a cylinder, without separate bundles, or comprising two or more rings of bundles (only in lianes Serjania and Paulinia); collateral. Internal phloem absent. Secondary thickening developing from a conventional cambial ring, or anomalous (in lianes: see illustration). The anomalous secondary thickening of lianes via concentric cambia, or from a single cambial ring (sometimes involving development of separate xylem masses, and in Serjania and Paulinia very peculiar, where ridged stems result from the activity of fascicular cambia in the several rings of permanently separate vascular bundles). Primary medullary rays in normal forms with a xylem cylinder, narrow (commonly uniseriate).

The wood ring porous to diffuse porous. The vessels typically small, or medium; typically solitary and in radial multiples. The vessel end-walls simple. The vessels without vestured pits; with spiral thickening, or without spiral thickening. The axial xylem without tracheids; with libriform fibres; nearly always including septate fibres. The fibres without spiral thickening. The parenchyma paratracheal, or apotracheal and paratracheal. ‘Included’ phloem present (rarely), or absent. The wood storied (e.g. Diplokeleba), or partially storied (VPI, Sapindus), or not storied.

Reproductive type, pollination. Unisexual flowers present. Plants monoecious, or polygamomonoecious, or polygamodioecious (but seemingly with much scope for confusion re. appearance versus functional fertility of floral components). Female flowers with staminodes (at least, often with ostensibly well developed but non-functional stamens), or without staminodes (?).

Inflorescence, floral, fruit and seed morphology. Flowers solitary (rarely), or aggregated in ‘inflorescences’; in cymes. The ultimate inflorescence units cymose. Inflorescences usually cymes, thyrses or cincinni. Flowers usually bracteate; bracteolate (usually), or ebracteolate; small; regular to very irregular (often obliquely zygomorphic). The floral irregularity involving the perianth and involving the androecium. Flowers often 4 merous, or 5 merous; cyclic; tetracyclic (usually), or tricyclic. Floral receptacle developing an androphore to developing a gynophore (e.g. sometimes in Dodonaea, by elongation of the disk of female and bisexual flowers), or with neither androphore nor gynophore. Free hypanthium absent. Hypogynous disk present (usually), or absent (or at least indistinct, e.g in Dodonaea); extrastaminal (the stamens usually inserted within it, occasionally upon it); annular or one-sided.

Perianth with distinct calyx and corolla (usually), or sepaline (the corolla occasionally lacking); (3–)4–5(–7) (rarely), or 8, or 9, or 10; 1 whorled (rarely), or 2 whorled (usually); isomerous, or anisomerous (e.g. when the abaxial petal is lacking). Calyx (3–)4, or 5(–7); 1 whorled; polysepalous (usually), or partially gamosepalous (sometimes ostensibly 4-merous), or gamosepalous (sometimes basally connate). Sometimes 2 of the members joined. Calyx unequal but not bilabiate, or regular; imbricate (usually), or valvate; when K5, with the median member posterior. Corolla 4, or 5; 1 whorled; appendiculate (often, the appendages basal, scalelike), or not appendiculate; gamopetalous (usually), or polypetalous (e.g. Koelreuteria); imbricate; unequal but not bilabiate, or regular.

Androecium 8, or 10 (usually), or 4–5(–6), or 11–100 (i.e., sometimes ‘many’). Androecial members free of the perianth; free of one another; 2 whorled (rather theoretically 5+5), or 1 whorled, or 2–5 whorled (sometimes polystemonous in several indistinct series, in Distichostemon). Androecium of male-fertile flowers exclusively of fertile stamens. Stamens 4–5(–6) (rarely), or 8, or 10, or 11–100 (rarely ‘many’); isomerous with the perianth (rarely, e.g. in Ganophyllum), or diplostemonous to polystemonous (usually); filantherous (the filaments often hairy). Anthers dorsifixed, or basifixed (to slightly ventrifixed); more or less versatile; dehiscing via longitudinal slits; introrse; tetrasporangiate; appendaged (usually), or unappendaged. The anther appendages apical (by connective extension). Endothecium developing fibrous thickenings. Anther epidermis persistent, or degenerating. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer, or initially with more than one middle layer (1–2 layers). Tapetum glandular. Pollen grains aperturate; (2–)3(–4) aperturate; colporate (usually tricolporate, sometimes bicolporate, sometimes syncolporate), or porate (rarely 3–4 porate); 2-celled (in Koelreuteria, Litchi and Xanthoceras).

Gynoecium (2–)3(–8) carpelled. Carpels reduced in number relative to the perianth, or isomerous with the perianth. The pistil (2–)3(–8) celled. Gynoecium syncarpous; synovarious to eu-syncarpous; superior. Ovary (2–)3(–8) locular. Gynoecium stylate. Styles 1 (usually), or 2–4; free, or partially joined; attenuate from the ovary, or from a depression at the top of the ovary (rarely, from between the ovary lobes); apical. Stigmas wet type; non-papillate; Group IV type. Placentation axile to basal. Ovules 1(–2) per locule, or 2–5 per locule (Dodonaeoideae); funicled, or sessile (then attached to placental protuberances); pendulous, or horizontal, or ascending; apotropous (Engler); with ventral raphe, or with dorsal raphe; arillate (usually), or non-arillate; hemianatropous, or anatropous, or campylotropous, or amphitropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle (usually), or not contributing to the micropyle (rarely). Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (forming up to 14 cells), or not proliferating; ephemeral (usually), or persistent (rarely). Synergids hooked. Hypostase present. Endosperm formation nuclear. Embryogeny asterad.

Fruit fleshy, or non-fleshy; dehiscent, or indehiscent, or a schizocarp. Mericarps when schizocarpic, (2–)3(–8) (?). Fruit when non-schizocarpic a capsule, or a berry, or a drupe, or a nut, or a samara. Capsules loculicidal, or splitting irregularly, or circumscissile. Seeds non-endospermic; with amyloid, or without amyloid. Cotyledons 2; flat (?), or folded, or twisted (and sometimes confluent). Embryo chlorophyllous (4/6), or achlorophyllous (4/4); curved, or bent, or coiled. The radicle dorsal.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3 and CAM. C3 physiology recorded directly in Cardiospermum. CAM recorded directly in Dodonaea (? - a perhaps dubious record involving a non-succulent species. Anatomy non-C4 type (Nephelium). Sugars transported as sucrose (in 9 genera). Cyanogenic, or not cyanogenic. Cynogenic constituents leucine-derived (and CN lipids). Alkaloids present, or absent. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins present; cyanidin, or cyanidin and delphinidin. Flavonols present; quercetin, or kaempferol and quercetin, or quercetin and myricetin, or kaempferol, quercetin, and myricetin. Ellagic acid absent (6 species, 5 genera). Aluminium accumulation not found.

Geography, cytology. Temperate to tropical. Pantropical and subtropical, also Japan, and widespread in Australasia. X = 10–16.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Rutiflorae; Sapindales. Cronquist’s Subclass Rosidae; Sapindales. APG III core angiosperms; core eudicot; Superorder Rosanae; malvid. APG IV Order Sapindales.

Species about 2000. Genera about 140; Alectryon, Allophylus, Allosanthus, Amesiodendron, Aporrhiza, Arfeuillea, Arytera, Atalaya, Athyana, Averrhoidium, Beguea, Bizonula, Blighia, Blighiopsis, Blomia, Boniodendron, Bridgesia, Camptolepis, Cardiospermum, Castanospora, Chonopetalum, Chouxia, Chytranthus, Conchopetalum, Cossinia, Cubilia, Cupania, Cupaniopsis, Deinbollia, Delavaya, Diatenopteryx, Dictyoneura, Dilodendron, Dimocarpus, Diploglottis, Diplokelepa, Diplopeltis, Distichostemon, Dodonaea, Doratoxylon, Elattostachys, Eriocoelum, Erythrophysa, Euchorium, Euphorianthus, Eurycorymbus, Exothea, Filicium, Ganophyllum, Glenniea, Gloeocarpus, Gongrodiscus, Gongrospermum, Guindilia, Guioa, Handeliodendron, Haplocoelum, Harpullia, Hippobromus, Hornea, Houssayanthus, Hypelate, Hypseloderma, Jagera, Koelreuteria, Laccodiscus, Lecaniodiscus, Lepiderema, Lepidopetalum, Lepisanthes, Litchi, Llagunoa, Lophostigma, Loxodiscus, Lychnodiscus, Macphersonia, Magonia, Majidea, Matayba, Melicoccus, Mischocarpus, Molinaea, Neotina, Nephelium, Otonephelium, Pancovia, Pappea, Paranephelium, Paullinia, Pavieasia, Pentascyphus, Phyllotrichum, Placodiscus, Plagioscyphus, Podonephelium, Pometia, Porocystis, Pseudima, Pseudopancovia, Pseudopteris, Radlkofera, Rhysotoechia, Sapindus, Sarcopteryx, Sarcotoechia, Schleichera, Scyphonychium, Serjania, Sinoradlkofera, Sisyrolepis, Smelophyllum, Stadmania, Stocksia, Storthocalyx, Synima, Talisia, Thinouia, Thouinia, Thouinidium, Tina, Tinopsis, Toechima, Toulicia, Trigonachras, Tripterodendron, Tristira, Tristiropsis, Tsingya, Ungnadia, Urvillea, Vouarana, Xanthoceras, Xeropspermum, Zanha, Zollingeria.

Economic uses, etc. Edible fruits: ‘Spanish lime’ (Melicoccus), litchi and longan (Litchi spp.), pulusan and rambutan (Nephelium spp.); Blighea with edible arils (‘akee’).

Illustrations. • Le Maout and Decaisne: Alectryon, Koelreuteria paniculata, Stadmannia. • Deinbollia pycnophylla: Thonner. • Sapindus senegalensis: Lindley. • Boniodendron minus, as Koelreuteria: Hook. Ic. Pl. 27 (1900). • Cardiospermum halicacabum: R. Wight, Ic. Pl. Indiae Orientalis 2 (1843). • Urvillea ferruginea: Bot. Reg. 1077, 1827. • Dodonaea viscosa: Bot. Reg. 1051, 1827. • Diplopeltis huegelii: Bot. Reg. 1839, 69. • Diplopeltis eriocarpa: Hook. Ic. Pl. 28 (1902). • Euphoria longana: as E. longan, Bot. Reg. 1729, 1835. • Euphoria verticillata, probably = Lepisanthes senegalensis: Bot. Reg. 1059, 1827. • Filicium decipiens (as Rhus): R. Wight (1840). • Magonia campestris, as Phaeocarpus: Martius, Nova Gen. et Spec. Pl. Brasiliensium (1824). • Majidea zanguebarica: Hook. Ic. Pl. 11 (1867–71). • Pappea capensis: Hook. Ic. Pl. 4 (1841). • Tristiropsis canarioides: Hook. Ic. Pl. 29 (1906). • Serjania, Thinouia and Urvillea: transverse sections of stems with anomalous vascular anatomy (Solereder, 1908).

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 15th April 2018.’.