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The families of flowering plants

L. Watson and M.J. Dallwitz

Goodeniaceae R. Br. corr. Dum.

Including Goodenovieae (Goodenoviaceae) R. Br., Scaevoleae (Scaevolaceae) Lindl.; excluding Brunoniaceae.

Habit and leaf form. Herbs, or shrubs (or shrublets), or trees (a few). ‘Normal’ plants, or switch-plants; sometimes with the principal photosynthesizing function transferred to stems (sometimes with flattened stems, rarely spinescent). Leaves well developed, or much reduced. Annual, or perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves. Mesophytic, or xerophytic. Leaves alternate (nearly always), or opposite to whorled (rarely); nearly always spiral; ‘herbaceous’, or leathery; petiolate to sessile; non-sheathing; simple; epulvinate. Lamina pinnately veined. Leaves exstipulate. Lamina margins entire, or serrate, or dentate. Leaf development not ‘graminaceous’.

General anatomy. Plants without laticifers.

Leaf anatomy. The leaf lamina dorsiventral, or bifacial, or centric (sometimes varying within a genus, sometimes fleshy, rush-like in Anthotium). Stomata present; mainly confined to one surface (abaxial), or on both surfaces; anomocytic (sometimes sunken). Hairs of numerous kinds present; eglandular and glandular. Complex hairs present, or absent; when present, often stellate. Lamina without secretory cavities. The mesophyll commonly with sclerenchymatous idioblasts; containing crystals (but usually relatively few). The crystals druses and solitary-prismatic. Main veins seldom accompamied by sclerenchyma. Minor leaf veins without phloem transfer cells (Goodenia, Lechenaultia).

Axial (stem, wood) anatomy. Secretory cavities absent. Cork cambium present; initially deep-seated, or initially superficial. Nodes unilacunar, or tri-lacunar, or penta-lacunar. Primary vascular tissues in a cylinder, without separate bundles (in the woodier forms), or comprising a ring of bundles (most herbaceous species, especially of Goodenia, having poorly developed vascular bundles, depending for support on a cylinder of thick-walled sclerenchyma); collateral. Internal phloem absent. Medullary bundles present (sometimes, representing leaf traces), or absent. Secondary thickening absent (in some herbs?), or developing from a conventional cambial ring, or anomalous. The anomalous secondary thickening from a single cambial ring (producing secodary medullary bundles, see illustration). The axial xylem without vessels (reported for Scaevola spinescens), or with vessels (usually).

The vessel end-walls simple. The vessels without vestured pits. The axial xylem with tracheids; with fibre tracheids. The parenchyma in Scaevola frutescens, apotracheal and paratracheal. ‘Included’ phloem absent.

Reproductive type, pollination. Plants hermaphrodite. Pollination entomophilous; mechanism conspicuously specialized (involving a cupular or bifid stylar modification for active pollen presentation).

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; when aggregated, in cymes, in spikes, in heads, and in racemes. The ultimate inflorescence units cymose, or racemose. Inflorescences scapiflorous, or not scapiflorous; terminal, or axillary; with involucral bracts, or without involucral bracts. Flowers small to medium-sized; very irregular; zygomorphic. The floral irregularity involving the perianth and involving the androecium. Flowers 5 merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk present, or absent.

Perianth with distinct calyx and corolla; (8–)10; 2 whorled; isomerous, or anisomerous. Calyx (3–)5 (usually small); 1 whorled; gamosepalous. Corolla 5; 1 whorled; appendiculate (commonly with the two posterior members auriculate), or not appendiculate (but the margins of the lobes nearly always conspicuously winged); gamopetalous. Corolla tube adaxially deeply split (nearly always, often to the base), or not noticeably adaxially split (rarely). Corolla lobes valvate; unequal but not bilabiate (then adaxially split to the base), or bilabiate (the upper lip bilobed, the lower trilobed); white, or yellow, or blue, or orange, or pink (or brownish); spurred, or not spurred.

Androecium 5. Androecial members free of the perianth, or adnate (to the corolla tube); all equal; free of one another (then anthers often connivent), or coherent (the anthers connate); 1 whorled. Androecium exclusively of fertile stamens. Stamens 5; inserted near the base of the corolla tube; isomerous with the perianth; oppositisepalous; alternating with the corolla members. Anthers separate from one another (e.g. Velleia, Scaevola), or connivent to cohering (usually encircling the style, which presents to insects by growing up through the the aznthers and carrying pollen in a ‘cup’); dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with one middle layer. Tapetum glandular. Pollen shed in aggregates (sometimes), or shed as single grains; that of Lechenaultia, in tetrads. Pollen grains aperturate; 3 aperturate (usually), or 4–8 aperturate (Lechenaultia); porate (Lechenaultia), or colporate; 2-celled (in 3 genera).

Gynoecium 2 carpelled. The pistil (1–)2 celled, or 4 celled. Gynoecium syncarpous; synovarious to synstylovarious; superior to inferior. Ovary (1–)2 locular (effectively unilocular, with the septum incomplete above in Verreauxia), or 4 locular (rarely). Locules without ‘false septa’ (usually), or secondarily divided by ‘false septa’ (rarely?). Styles 1 (with a ‘pollen cup’ close beneath the stigma); bearing an ‘indusium’ beneath the stigma; apical. Stigmas 1–3; dry type; papillate; Group II type. Placentation when more or less unilocular, basal (or on the basal septum); axile. Ovules in the single cavity 1; 1–50 per locule (to ‘many’); ascending; non-arillate (usually), or arillate (Coopernookia?); anatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; fairly persistent. Endosperm formation cellular. Embryogeny solanad.

Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule, or a drupe, or a nut. Seeds copiously endospermic. Endosperm oily. Seeds usually flat, winged, or wingless. Cotyledons 2. Embryo straight.

Seedling. Germination phanerocotylar.

Physiology, phytochemistry. Inulin recorded (very widespread). Cyanogenic, or not cyanogenic. Alkaloids present (commonly), or absent. Iridoids detected; ‘Route I’ type (normal and seco). Proanthocyanidins absent. Flavonols absent. Ellagic acid absent (2 genera). Ursolic acid present. Aluminium accumulation not found.

Geography, cytology. Temperate to tropical. Mainly Australia and Malaysia, but a few species in coastal South America and Africa, West Indies, New Zealand and Southeast Asia. X = 7–9.

Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Gentianiflorae; Goodeniales. Cronquist’s Subclass Asteridae; Campanulales. APG III core angiosperms; core eudicot; Superorder Asteranae; campanulid. APG IV Order Asterales.

Species 300. Genera 12; Anthotium, Catosperma, Coopernookia, Dampiera, Diaspasis, Goodenia, Lechenaultia (Lechenaultia), Pentaptilon, Scaevola, Selliera, Velleia, Verreauxia.

Illustrations. • Le Maout and Decaisne: Goodenia. • Lechenaultia splendens and L. biloba: Lindley. • Lechenaultia formosa (photo). • Lechenaultia formosa (close-up photo). • Lechenaultia biloba: Bot. Reg. 2, 1842. • Scaevola gracilis: Hutchinson. • Scaevola nitida: close-up photo. • Scaevola crassifolia: close-up photo. • Scaevola crassifolia: habitat. • Scaevola albida: Bot. Mag. 287 (1796). • Scaevola albida: Bot. Mag. 287 (1796), text. • Scaevola hookeri: Hooker, Fl. Tasmaniae (1860). • Catosperma muelleri: Hook. Ic. Pl. 11 (1867–71). • Dampiera trigona: Hook. Ic. Pl. 11 (1867–71). • Dampiera alata: Hook. Ic. Pl. 11 (1867–71). • Goodenia humilis and Velleia montana: Hooker, Fl. Tasmaniae (1860). • Velleia lyrata: Bot. Reg. 551, 1821. • Velleia macrophylla: as Euthales, Bot. Reg. 3, 1841. • Velleia paradoxa: Bot. Reg. 971, 1826. • Verreauxia dyeri: Hook. Ic. Pl. 28 (1905). • Dampiera: leaf hairs (Solereder, 1908). • TS Goodenia ovata stem, with anomalous secondary thickening via interfascicular medullary bundles: 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.’.