Family Goodeniaceae

Goodeniaceae R. Br. corr. Dum.

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

Habit and leaf form. Herbs, or shrubs, 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 with neither basal nor terminal 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; not gland-dotted; simple; epulvinate. Lamina pinnately veined. Leaves exstipulate. Lamina margins entire, or serrate, or dentate. Leaves without a persistent basal meristem.

General anatomy. Plants without laticifers.

Leaf anatomy. Stomata present; anomocytic. Hairs present. Complex hairs present, or absent; when present, often stellate.

Lamina dorsiventral, or isobilateral, or centric. The mesophyll commonly with sclerencymatous idioblasts. Minor leaf veins without phloem transfer cells (Goodenia, Lechenaultia).

Stem anatomy. Cork cambium present; initially deep-seated, or superficial. Nodes unilacunar, or tri-lacunar, or penta-lacunar. Medullary bundles present, or absent. Internal phloem absent. Secondary thickening absent, or developing from a conventional cambial ring. Xylem with tracheids; with vessels. Vessel end-walls simple. Wood parenchyma apotracheal and paratracheal.

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’; in cymes, in spikes, in heads, and in racemes. The ultimate inflorescence unit 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 them, 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, biochemistry. 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. Inulin recorded (very widespread).


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 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Asterales.

Species 300. Genera 11; Anthotium, Coopernookia, Dampiera, Diaspasis, Goodenia, Lechenaultia (Lechenaultia), Pentaptilon, Scaevola, Selliera, Velleia, Verreauxia.


Illustrations.
• Technical details: Goodenia.
• Technical details: Lechenaultia (Lindley).
• Lechenaultia formosa (photo).
• Lechenaultia formosa (close-up photo).
• Lechenaultia biloba: Bot. Reg. 2, 1842.
• Scaevola nitida (close-up photo).
• Scaevola albida: Bot. Mag. 287 (1796).
• Scaevola albida: Bot. Mag. 287 (1796), text.
• Velleia lyrata: Bot. Reg. 551, 1821.
• Velleia macrophylla: as Euthales, Bot. Reg. 3, 1841.
• Velleia paradoxa: Bot. Reg. 971, 1826.

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