| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Staphyleaceae (DC.) Lindl.

Including Ochranthaceae (Lindl.) Endl., Simabaceae Horan. (p.p.); excluding Tapisciaceae (Pax) Takhtajan

Habit and leaf form. Small trees, or shrubs. Leaves opposite; petiolate; without marked odour; ostensibly simple (at least sometimes, in Turpinia), or compound (generally, at least basically); pulvinate; when recognisably compound, unifoliolate (in some Turpinia species?), or ternate (often), or pinnate; imparipinnate. Lamina pinnately veined; cross-venulate. Leaves stipulate. Stipules caducous. Lamina margins usually serrate, or dentate.

Leaf anatomy. The leaf lamina dorsiventral. Mucilaginous epidermis present, or absent. Stomata present; mainly confined to one surface (abaxial); anisocytic. Hairs present; eglandular. The mesophyll containing crystals. The crystals druses. Minor leaf veins without phloem transfer cells (Staphylea).

Axial (stem, wood) anatomy. Pith with diaphragms, or without diaphragms. Secretory cavities absent. Cork cambium present; initially superficial (sometimes in the epidermis itself). Nodes tri-lacunar. Primary vascular tissues in a cylinder, without separate bundles, or comprising a ring of bundles (at a very early stage); collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays narrow.

The wood diffuse porous, or semi-ring porous to diffuse porous. The vessels very to moderately small; solitary (typically exclusively so), or radially paired, or in radial multiples (with radial multiples in some species of Turpinia). The vessel end-walls oblique; scalariform. The vessels with spiral thickening, or without spiral thickening. The axial xylem with tracheids, or without tracheids; often with vasicentric tracheids; with fibre tracheids; including septate fibres (rarely), or without septate fibres. The fibres without spiral thickening. The parenchyma scanty paratracheal; wood not storied.

Reproductive type, pollination. Unisexual flowers present, or absent. Plants hermaphrodite, or monoecious, or dioecious, or polygamomonoecious.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in racemes, or in panicles. Inflorescences terminal, or axillary; drooping panicles or racemes. Flowers bracteate; small; regular; 5 merous; cyclic. Free hypanthium absent. Hypogynous disk present; intrastaminal.

Perianth with distinct calyx and corolla; 10; 2 whorled; isomerous; sepaloid and petaloid, or petaloid. Calyx 5 (often petaloid); 1 whorled; more or less polysepalous (never forming a tube); regular; imbricate. Corolla 5; 1 whorled; polypetalous; imbricate; regular.

Androecium 5. Androecial members free of the perianth; all equal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens 5; isomerous with the perianth; oppositisepalous; alternating with the corolla members; filantherous (the filaments sometimes very flattened). Anthers dorsifixed; versatile; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with one middle layer. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; (2–)3 aperturate, or 4–12 aperturate; colporate, or rugate; 2-celled (Staphylea), or 3-celled (Turpinia).

Gynoecium 2–3(–4) carpelled. Carpels reduced in number relative to the perianth. The pistil when syncarpous (i.e. usually), 2–3(–4) celled. Gynoecium apocarpous (Euscaphis), or apocarpous to syncarpous, or syncarpous; eu-apocarpous (Euscaphis only), or semicarpous to eu-syncarpous (the carpels variously partially separate or separating); superior to partly inferior (the base often embedded in the disk). Carpel of the apocarpous Euscaphis stylate; apically stigmatic; (1–)6–12 ovuled. Ovary usually syncarpous and 2–3(–4) locular. Gynoecium stylate. Styles 1, or 2–3(–4); free, or partially joined (sometimes joined distally); attenuate from the ovary, or from a depression at the top of the ovary; apical. Stigmas wet type; papillate; Group IV type. Placentation axile, or basal to axile. Ovules (2–)6–12 per locule; horizontal, or ascending (commonly); biseriate (commonly with two rows in each locule), or superposed; arillate (Euscaphis), or non-arillate; anatropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked (with filiform apparatus). Endosperm formation nuclear.

Fruit fleshy, or non-fleshy; in Euscaphis, an aggregate (and sometimes the carpels separating in other genera), or not an aggregate. The fruiting carpel of Euscaphis dehiscent; a follicle. Fruit when syncarpous dehiscent, or a schizocarp (the carpels inflating and opening along their inner sutures, sometimes separating completely, in Staphylea), or indehiscent (in Turpinia). Mericarps when schizocarpic, 2, or 3; comprising follicles (becoming inflated). Fruit when syncarpous a capsule, or a berry, or a drupe (inflated and apically dehiscent in Staphylea, indehiscent and dry to softly fleshy in Turpinia). Seeds copiously endospermic. Endosperm oily. Embryo well differentiated. Cotyledons 2; fleshy, flat or planoconvex. Embryo chlorophyllous (1/1); straight.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. Sugars transported as sucrose, or as sugar alcohols + oligosaccharides + sucrose (in different species of Staphylea). Not cyanogenic. Alkaloids present (mostly), or absent. Arbutin absent. Iridoids not detected. Saponins/sapogenins absent. Proanthocyanidins present; cyanidin. Flavonols present; kaempferol and quercetin. Ellagic acid absent (Staphylea). Aluminium accumulation not found.

Geography, cytology. Holarctic, Paleotropical, and Neotropical. Temperate to tropical. Southwest Europe, Eastern Asia, temperate U.S.A., Central America and Northwest South America. X = 13.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Rutiflorae (seemingly misplaced: Gadek et al 1996); Geraniales (? — pending reassignment). Cronquist’s Subclass Rosidae; Sapindales. APG III core angiosperms; core eudicot; Superorder Rosanae; malvid. APG IV Order Crossosomatales.

Species 60. Genera 5; Euscaphis, Staphylea and Turpinia.

Illustrations. • Le Maout and Decaisne: Staphylea pinnata. • Staphylea bumalda: Lindley. • Staphylea colchica: Bot. Mag. 120 (1894). • Staphylea pinnata: Eng. Bot. 322, 1864. • Turpinea arguta: as Ochranthe, Bot. Reg. 1819, 1836. • Turpinia insignis (as Lacepedea): Bot. Mag. 75 (1849).

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: 8th June 2026. delta-intkey.com’.

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