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Published In: Genera Plantarum 196. 1789. (4 Aug 1789) (Gen. Pl.) Name publication detailView in BotanicusView in Biodiversity Heritage Library

Project Name Data (Last Modified On 6/2/2011)
Acceptance : Accepted
Project Data     (Last Modified On 6/3/2011)
Contributor Institution: Department of Botany, University of Karachi, Karachi-75270
General/Distribution: A large family of c. 500 genera and c. 6000 species. Predominantly woody and distributed in the tropics and sub-tropics. Represented in Pakistan by 33 genera and c. 87 species, about half cultivated or introduced for ornamental purposes. Cultivated taxa have been marked with an asterisk (*). Native genera are mostly herbaceous in our area.
Comment/Acknowledgements: Acknowledgements: We are grateful to the Directors/Curators of the following herbaria for the loan of herbarium material: ISL, K. PESH, PPFI-B. Thanks are also due to Dr. B. Verdcourt, Mrs. D. M. Bridson of Royal Botanic Gardens, Kew, Mr. I.C. Hedge of Royal Botanic Garden, Edinburgh, Prof. F. Ehrendorfer and Dr. E. Schonbeck-Temesy of Botanisches Institut and Botanischer Garten der Universitat, Wien, Austria for going through the manuscript and offering very valuable suggestions. We are also indebted to Dr. H.W. Lack and Dr. M. Hakki, Botanischer Garten and Botanisches Museum, Berlin for providing photocopies of relevant literature. The financial assistance provided by the United States Department of Agriculture under P.L. 480 with the coordination of Pakistan Agricultural Research Council, Islamabad is gratefully acknowledged.


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Trees, shrubs, climbers, erect, prostrate or procumbent annual-perennial herbs. Stems cylindrical or quadrangular, unarmed or rarely spiny-prickly. Raphides often present. Leaves opposite or less often verticillate, decussate, simple, mostly entire very rarely toothed. Stipules inter- or intmpetiolar, mostly persistent, distinct, free, or connate and sheathing, sometimes leaf-like, and forming whorls of 4-10 foliar stipules which am sometimes smaller than true leaves, scale-like or filiform, forming glandular setae. Inflorescence various, mainly different forms of dichasial cymes, thyrsoid, spike or head, corymbose, paniculate or solitary. Bracts present or absent. Flowers usually bisexual, rarely unisexual, usually actinomorphic, very rarely zygomorphic or ± bilabiate, 4-6-merous. Calyx 4-5 (or more)-lobed; tube mostly adnate to the ovary, limb (3-) 4-5 (-8)-lobed or toothed, sometimes only minutely so, with valvate, imbricate or contorted aestivation becoming enlarged after anthesis and persistent in fruit. Corolla 4-5 (-10)-lobed, gamopetalous, infundibuliform, campanulate or rotate, lobes twisted, valvate-induplicate or rarely imbricate in bud. Homostylous or heterostylous with 2 or rarely 3 forms. Stamens generally equalling and alternating with the corolla lobes, epipetalous; anthers 2-celled, introrse, usually distinct, dehiscing longitudinally. Disc mostly present, annular or cushion shaped. Ovary inferior, rarely sub-inferior-superior, generally 2-loculed, rarely 3-5 or more loculed. Ovules solitary-numerous in each locule; placentation exile or apparently basal (rarely parietal in Gardenia). Style simple or branched, stigmas linear, filiform or capitate or bilobed. Fruit a loculicidal or septicidal capsule, berry or a drupe, or of 2 dehiscent or indehiscent cocci or mericarps; rarely syncarps (Nauclea & Morinda), 1-many seeded. Seeds rarely winged, the endosperm entire or ruminate, usually copious and fleshy or cartilaginous, absent in Guettardeae.

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1.Plants armed with thorns or spines
2.Flowers 4-merous [cultivated]
2.Flowers 5-merous [wild]
3.Calyx lobes foliaceous, persistent. Fruit 25-38 mm in diameter
3.Calyx lobes neither foliaceous nor persistent. Emit 6-8 mm in diameter
1.Plants unarmed
4.Inflorescence a globose head
5.Annual or perennial herbs. Flowers tetramerous. Fruit crowned with persistent calyx
5.Shrubs or trees, rarely climbers. Flowers pentamerous. Fruit not crowned with persistent calyx
6.Corolla lobes imbricate, throat glabrous. Fruit an aggregate of capsules, few to many seeded. Seeds embedded in fleshy receptacle
6.Corolla lobes valvate, throat hairy. Fruit not as above
7.Shrubs or small trees. Calyx lobes nearly connate. Fruit a compound succulent berry. Seeds not winged
7.Trees. Calyx lobes free. Fruit not as above. Seeds winged
4.Inflorescence not a globose head
8.Nodes with whorl of 4-8 (-10) leaves and leaf like stipules. Herbs and herbaceous climbers. Fruits globose. Plants often adhesive due to prickles or harsh hairs
9.Flowers in dense spikes
9.Flowers in panicles, cymes or capitula
10.Most flowers enclosed by a large membranous white bract
10.Flowers not enclosed by the membranous bracts
11.Fruit fleshy, berry like
11.Fruit dry
12.Ultimate inflorescence branches with bracts and often bracteoles. Corolla with a very conspicuous tube
12.Ultimate inflorescence branches without bracts or bracteoles. Corolla with a very short tube
8.Nodes with opposite leaves, rarely in whorls of 3-5 but plants either shrubs or trees
13.Some calyx lobes in each inflorescence dilated into a large white or coloured lamina in fruit
13.No calyx lobe dilated into large lamina in fruit
14.Each pair of leaves with one large and one small leaf (anisophyllous). Anthers connivent
14.Leaves not anisophyllous. Anthem not connivent
15.Fruit fleshy, drupaceous or a berry
16.Ovary 5-loculed. Placentation axile
16.Ovary 1-2-loculed. Placentation basal or parietal
17.Ovules numerous. Pollen in tetrads
17.Ovules 1-2. Pollen not in tetrads
18.Plants foetid. Corolla tube funnel-shaped, lobes valvale in bud
18.Plants not foetid. Corolla tube salver-shaped, lobes twisted in bud
19.Bracts coriaceous, not sheathing. Style not twice as long as corolla tube; stigmatic lobes divergent. Leaves without bacterial nodules [cultivated]
19.Bracts membranous, lower bracts sheathing. Style twice as long as corolla tube; stigma fusiform. Leaves with bacterial nodules [wild]
15.Emit dry, capsule or mericarp
20.Ovary 5-loculed; style ending in 5 stigmatic branches
21.Capsule 5-valved at the base. Bracteoles united forming a tubular involucre
21.Capsule 5-valved at the apex. Bracteoles free
20.Ovary 2-3 loculed; style either simple or 2-lobed (rarely 3-lobed in Rondeletia) or ending in 2 stigmatic branches
22.Ovules many in each locule. Fruit many-seeded
23.Large shrubs or trees
24.Flowers usually di-or trimorphic. Stipitate glands usually present between each pair of calyx lobes. Capsule beaked. Seeds without wings
24.Flowers monomorphic. Glands absent. Capsule not beaked. Seeds winged
25.Flowers small. Corolla lobes valvate in bud. Stigma clavate to capilate
25.Flowers often large. Corolla lobes imbricate in bud. Stigma not so
26.Flowers small, 0.5 cm long. Anthers exserted [wild]
26.Flowers 3-5 cm long. Anthers included [cultivated]
23.Herbs or shrubs
27.Leaf blades usually broad with obvious lateral and tertiary venation. Flowers large, pentamerous, di-or timorphic. Stipitate glands usually present between each pair of calyx lobes [cultivated]
27.Leaf blades usually narrow and uninerved or lateral nerves obscure. Flowers small, usually tetramerous, monomorphic (only in flora area; elsewhere many species dimorphic). Stipitate glands absent between each pair of calyx lobes [wild]
28.Calyx teeth contiguous on the capsule. Capsule splitting longitudinally
28.Calyx teeth distant on the capsule. Capsule splitting at the apex
29.Corolla tube cylindrical. Anthers and stigma included; stigmas always below the anthers
29.Corolla tube cylindrical or infundibuliform, if cylindrical than either anthers and stigmas exserted or anthers below the stigma
22.Ovules 1-2 in each locule. Fruit 1-few-seeded
30.Erect or prostrate annual or perennial herbs Corolla 4-lobed. Stamens 4
31.Calyx tube ovoid, globose, limb usually absent or inconspicuous. Fruit didymous not crowned
31.Calyx tube ovate, limb 2-4-lobed, well developed. Fruit not didymous, crowned by persistent calyx
30.Subshrub to shrubs. Corolla 5-lobed. Stamens 5
32.Stipules free and scarious. Fruit didymous, not encircled by persistent calyx teeth
32.Stipules connate with petiole forming a sheath with 2-many setae. Fruit not didymous
33.Calyx limb accrescent after anthesis
34.Calyx limb salverform, enlarged in fruit to form large orbicular and horizontal 5-lobed membranous reticulately veined wings. Bracts similar to stipules
34.Calyx limb bifid, enlarged in fruit to form 2 erect wings. Floral bracts forming an obconical 6-8-toothed membranous sheath
33.Calyx limb not accrescent after anthesis
35.Bracts heteromorphic, forming denticulate sixpartite involucre or sheath
35.Bracts homomorphic not forming a sheath or involucre
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