Staphyleaceae (DC.) Lindl.
Including Ochranthaceae (Lindl.) Endl., Simabaceae Horan. (p.p.), Tapisciaceae (Pax) Takhtajan
Habit and leaf form. Small trees, or shrubs; non-laticiferous and without coloured juice. Leaves opposite, or alternate (Tapiscioideae); petiolate; non-sheathing; not gland-dotted; 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. Lamina pinnately veined; cross-venulate. Leaves stipulate, or exstipulate (some Tapisciodeae). Stipules caducous. Lamina margins usually serrate, or dentate.
Leaf anatomy. Mucilaginous epidermis present, or absent. Stomata present; anisocytic.
The mesophyll containing calcium oxalate crystals. The mesophyll crystals druses. Minor leaf veins without phloem transfer cells (Staphylea).
Stem anatomy. Cork cambium present; initially superficial. Nodes tri-lacunar. Secondary thickening developing from a conventional cambial ring. Xylem with tracheids, or without tracheids; with fibre tracheids; with vessels. Vessel end-walls oblique; scalariform. Wood parenchyma scanty paratracheal.
Reproductive type, pollination. 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, or absent (Tapiscioideae); when present, intrastaminal.
Perianth with distinct calyx and corolla; 10; 2 whorled; isomerous; sepaloid and petaloid, or petaloid. Calyx 5 (often petaloid); 1 whorled; polysepalous (Staphyleoideae), or gamosepalous (connate to varying extents in Tapiscioideae); 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. 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 (polyrugate in Huertia); 2-celled, or 3-celled.
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; eu-apocarpous (Euscaphis only), or synstylous, or semicarpous to synstylovarious (the carpels variously completely joined, partially separate or separating, the styles variously free, connate only distally, or completely joined into one); superior to partly inferior (the base often embedded in the disk). Carpel of Euscaphis (apocarpous) 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 1–2 per locule (Tapiscioideae), or (1–)6–12 per locule (Staphyleoideae); horizontal, or ascending (commonly); biseriate (commonly with two rows in each locule), or superposed; 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 legume. Fruit when syncarpous dehiscent, or indehiscent, or a schizocarp. Mericarps when schizocarpic, 2–3(–4); comprising follicles, or comprising drupelets. Fruit when non-schizocarpic a capsule (inflated, apically dehiscent), or a berry, or a drupe. 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, biochemistry. Not cyanogenic. Alkaloids present (mostly), or absent. Iridoids not detected. Proanthocyanidins present; cyanidin. Flavonols present; kaempferol and quercetin. Ellagic acid absent (Staphylea). Arbutin absent. Saponins/sapogenins absent. Aluminium accumulation not found. Sugars transported as sucrose, or as sugar alcohols + oligosaccharides + sucrose (in different species of Staphylea).
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 (1998) Eudicot; core Eudicot; Rosid; Eurosid II; unassigned at ordinal level. Species 60. Genera 5; Euscaphis, Staphylea and Turpinia (Staphyleoideae), Huertea, Tapiscia (Tapiscioideae).
The subfamilies probably merit family rank. Their affinities seem to be with Rosiflorae rather than Rutiflorae.
Illustrations. • Technical details (Staphylea). • Technical details (Staphylea).
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
