Contributor

Address

Description

Contributor

D.J. Nicholas Hind

Address

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK

Description

Annual or perennial herbs, subshrubs or shrubs, climbers, lianes or ramblers, small or, rarely, large trees, epiphytes, very rarely true aquatics, glabrous or with simple, glandualr, malpighiaceous or stellate hairs, often glabrescent; plants usually monoecious, rarely dioecious, odourless or rarely with distinctive odour. Rootstocks fibrous or fleshy, sometimes with distinct woody perennating rootstocks (xylopodia). Stems usually unarmed, rarely spiny. Leaves alternate or opposite, sometimes rosulate, rarely whorled, estipulate, lamina simple, rarely compound, variously shaped, pinnately or palmately veined, usually herbaceous, sometimes fleshy, coriaceous, or chartaceous, sessile or petiolate, sometimes with an expanded, sheathing or auriculate base but never stipulate, margins lobed or variously toothed, rarely spiny and rarely ending in a tendril, and rarely reduced to scales and falling rapidly. Primary inflorescence a capitulum, usually chasmogamous, very rarely cleistogamous, usually of many small individual flowers (florets), sometimes reduced to 1, surrounded by an involucre of protective bracts (phyllaries) in one or more series, cylindrical, to globular or urceolate; phyllaries imbricate, subimbricate or distant, usually gradate or sometimes subequal, rarely outer longer than inner, usually homomorphic, very rarely heteromorphic, often chartaceous or herbaceous, sometimes with characteristic apical appendages, rarely with an outer series of subinvolucral bracts forming a calyculus; receptacle solid or hollow, concave, flat or convex, rarely conical or almost cylindrical, surface plane, variously ornamented and areolate, alveolate or fimbriate around achene attachment points, commonly naked but sometimes with chaffy bracts (paleae), scales, bristles or hairs between florets. Inflorescence of capitula usually cymosely arranged within flowering structure but also in corymbose panicles, or rarely spicate, sometimes scapiform, rarely secondarily aggregated into spikes or glomerules, or very rarely arranged in compound structures or synflorescences (syncalathia) possessing a secondary receptacle, rarely also with a secondary involucre, sometimes each individual capitulum within compound structure reduced to single-flowered condition. Capitula homogamous and all florets hermaphrodite or more rarely all male (staminate) or all female (pistillate), very rarely unisexual capitula borne on same plant or on different plants, capitula discoid (with all florets tubular and actinomorphic), ligulate (with all florets ligulate), or bilabiate (with all florets bilabiate), and all florets hermaphrodite, all male, or all female, or capitula heterogamous and radiate with outer marginal florets female (pistillate), or sterile or neuter, and radiate in one or more whorls, and inner disc florets usually hermaphrodite with tubular corollas, or heterogamous and disciform (possessing at least 2 types of eradiate florets, usually with filiform outer florets and central tubular hermaphrodite florets), or heterogamous and radiant (possessing inner hermaphrodite florets and outer enlarged, usually sterile or neuter marginal florets). Florets few, very rarely 1, to many, arranged racemosely or indeterminately within capitula, outer opening first; corollas variously coloured, actinomorphic and typically in disk florets in radiate, or throughout discoid, capitula, or with zygomorphic corollas and bilabiate (with a 3-toothed outer lip and 2-lobed inner lip), pseudobilabiate (with a 4-toothed outer lip and a 1-lobed inner lip), rayed (lacking adaxial lobes but with a limb with 1 to 4 apical teeth), or ligulate (with a flat, strap-shaped limb with 5 apical teeth), glabrous or variously pubescent (eglandular or glandular), corolla lobes short or long, glabrous or variously pubescent, often with thickened apical margin; anthers typically connate (exceedingly rarely anthers free), usually 5 (rarely 4) forming a tube around the style, and dehiscing introrsely, dorsifixed or basifixed, conspicuously exserted from, or included within corolla throat, apical anther appendages (diagnostic in many genera in Eupatorieae) acuminate, apiculate, acute, obtuse, usually persistent, basal anther appendages calcarate and caudate, rarely ecalcarate, tails long or short, entire or variously laciniate, rarely branched or pilose; filaments inserted basally inside corolla tube, higher up in tube or just beneath sinuses of corolla-lobes, usually glabrous, rarely papillose or even hairy, very rarely filaments fused together, filaments often with a conspicuous (and diagnostic) anther or filament collar just below insertion point on anther; styles often well exserted from corolla throat and anther cylinder, usually divided into 2 style arms (rarely connate) each with stigmatic surface on inner surface, style hairy, hairs acute or obtuse to rounded, papillose or glabrous, sometimes with distinctive (and diagnostic) basal node, with or without a nectary, glabrous or pubescent; style arms sometimes with characteristic sterile apical appendages, stigmatic papillae often conspicuous and arranged along style arms margins or inner surfaces or inconspicuous and either marginal or all over inner surfaces. Fruit single-seeded, indehiscent, lacking endosperm, developing from an inferior ovary (a cypsela, although commonly termed an achene), obovoid or oblong, fusiform or distinctly beaked, terete, angular, rounded, variously compressed or curved, ribbed, angled, variously ornamented or winged, glabrous or variously pubescent (frequently with diagnostic eglandular duplex, or twin, hairs) or glandular, rarely myxogenic, with or without phytomelanin in achene body walls; carpopodium (basal attachment area to receptacle) usually present (diagnostic in some genera) and of several layers of variously enlarged, sometimes ornamented cells, usually delimited from body of achene in form or colour, usually symmetrical, rarely eccentric, variously annular, cylindrical, or stopper-shaped, sometimes procurrent on base of achene, usually glabrous, rarely replaced by an elaiosome; apically usually possessing a pappus (considered a modified calyx) of uniseriate to few- or multi-seriate smooth, barbellate or plumose fine hairs, bristles, scales, or awns (sometimes more or less fused together), usually separate, rarely basally connate, caducous, deciduous or persistent, rarely a laciniate crown, sometimes completely lacking.