The Families of Angiosperms

L. Watson and M.J.Dallwitz

Goodeniaceae R. Br. corr. Dum.

IncludingGoodenovieae (Goodenoviaceae) R. Br.,Scaevoleae (Scaevolaceae) Lindl.; excludingBrunoniaceae.

Habit and leaf form.Herbs, orshrubs (or shrublets), or trees (a few). ‘Normal’ plants, orswitch-plants; sometimes with the principal photosynthesizing functiontransferred to stems (sometimes with flattened stems, rarely spinescent). Leaveswell developed, or much reduced. Annual, or perennial; with a basal aggregationof leaves, or without conspicuous aggregations of leaves. Mesophytic, orxerophytic. 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 withoutlaticifers.

Leaf anatomy.The leaflamina dorsiventral, or bifacial, or centric (sometimes varying within agenus, sometimes fleshy, rush-like inAnthotium). Stomata present; mainlyconfined to one surface (abaxial), or on both surfaces; anomocytic (sometimessunken). Hairs of numerous kinds present; eglandular and glandular. Complexhairs present, or absent; when present, often stellate. Lamina without secretorycavities.The mesophyllcommonlywith sclerenchymatousidioblasts; containing crystals (but usually relatively few). The crystalsdruses and solitary-prismatic. Main veins seldom accompamied by sclerenchyma.Minor leaf veins without phloem transfer cells (Goodenia,Leschenaultia).

Axial (stem, wood) anatomy.Secretory cavities absent. Cork cambium present; initiallydeep-seated, or initially superficial. Nodes unilacunar, or tri-lacunar, orpenta-lacunar. Primary vascular tissues in a cylinder, without separate bundles(in the woodier forms), or comprising a ring of bundles (most herbaceousspecies, especially ofGoodenia, having poorly developed vascularbundles, 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 someherbs?), or developing from a conventional cambial ring, or anomalous. Theanomalous secondary thickening from a single cambial ring (producing secodarymedullary bundles, see illustration). The axial xylem without vessels (reportedforScaevola spinescens), or with vessels (usually).

The vessel end-walls simple. The vessels without vestured pits. The axialxylem with tracheids; with fibre tracheids. The parenchyma inScaevolafrutescens, apotracheal and paratracheal. ‘Included’ phloemabsent.

Reproductive type, pollination.Unisexual flowers absent. Plants hermaphrodite. Pollinationentomophilous; mechanism conspicuously specialized (involving a cupular or bifidstylar modification for active pollen presentation).

Inflorescence, floral, fruit and seedmorphology.Flowers solitary, or aggregated in‘inflorescences’; when aggregated, in cymes, in spikes, in heads, andin racemes. The ultimate inflorescence units cymose, or racemose. Inflorescencesscapiflorous, or not scapiflorous; terminal, or axillary; with involucralbracts, or without involucral bracts.Flowers small to medium-sized;very irregular; zygomorphic. The floral irregularity involving theperianth 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 twoposterior members auriculate), or not appendiculate (but the margins of thelobes nearly always conspicuously winged); gamopetalous. Corolla tube adaxiallydeeply split (nearly always, often to the base), or not noticeably adaxiallysplit (rarely). Corolla lobes valvate; unequal but not bilabiate (then adaxiallysplit 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 notspurred.

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

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

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

Seedling.Germinationphanerocotylar.

Physiology, phytochemistry.Inulinrecorded (very widespread). Cyanogenic, or notcyanogenic. Alkaloids present (commonly), or absent. Iridoids detected;‘Route I’ type (normal and seco). Proanthocyanidins absent. Flavonolsabsent. Ellagic acid absent (2 genera). Ursolic acid present. Aluminiumaccumulation not found.

Geography, cytology.Temperate totropical. Mainly Australia and Malaysia, but a few species in coastal SouthAmerica 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,Leschenaultia,Pentaptilon,Scaevola,Selliera,Velleia,Verreauxia.

Illustrations.• Le Maout and Decaisne: Goodenia.• Catosperma muelleri: Hook. Ic. Pl.11 (1867–71). • Dampieratrigona: Hook. Ic. Pl. 11 (1867–71). • Dampiera alata: Hook. Ic. Pl. 11(1867–71). • Goodeniamacmillanii: Mueller, Fragm. Phytog. Austral. 1 (1858). • Goodenia vilmorinae: Mueller, Fragm. Phytog.Austral. 3 (1862). • Goodeniaramelii: Mueller, Fragm. Phytog. Austral. 3 (1862). • Goodenia humilis and Velleia montana: Hooker, Fl.Tasmaniae (1860). • Leschenaultiasplendens and L. biloba: Lindley. • Leschenaultia formosa (photo). • Leschenaultia formosa (close-up photo).• Leschenaultia biloba: Bot. Reg. 2,1842. • Leschenaultia laricina(as L. splendens): Bot. Mag. 72 (1846). • Leschenaultia linarioides (as L. arcuata): Bot.Mag. 72 (1846). • Scaevola albida:Bot. Mag. 287 (1796). • Scaevolaalbida: Bot. Mag. 287 (1796), text. • Scaevola crassifolia: close-up photo.• Scaevola crassifolia: habitat.• Scaevola gracilis: Hutchinson.• Scaevola hainanensis: Hook. Ic. Pl.20 (1891). • Scaevola hookeri:Hooker, Fl. Tasmaniae (1860). • Scaevola nitida (as S. attenuata): Bot. Mag. 71(1845). • Scaevola nitida:close-up photo. • Velleiacycnompotamica: Mueller, Fragm. Phytog. Austral. 6 (1867). • Velleia lyrata: Bot. Reg. 551, 1821.• Velleia macrophylla: as Euthales,Bot. Reg. 3, 1841.Velleia macrophylla. 1, the base of the corolla,with stamens and pistil. 2, vertical section of the ovary. 3, a stamen.• Velleia paradoxa: Bot. Reg. 971,1826. • Verreauxia dyeri: Hook.Ic. Pl. 28 (1905). • Dampiera:leaf hairs (Solereder, 1908). • TS Goodenia ovata stem, with anomalous secondarythickening via interfascicular medullary bundles: Solereder, 1908.

We advise against extracting comparative informationfrom the descriptions. This is much more easily achieved using theDELTA data files or theinteractive key, which allows access to the characterlist, illustrations, full and partial descriptions, diagnostic descriptions,differences and similarities between taxa, lists of taxa exhibiting or lackingspecified 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 alsoGuidelines for using data taken from Web publications.