The World Flora Online

SELECTED REFERENCES Aagesen, D. et al. 2005. Phylogeny of the tribe Colletieae (Rhamnaceae)—A sensitivity analysis of the plastid region trnL-trnF combined with morphology. Pl. Syst. Evol. 250: 197–214. Brizicky, G. K. 1964b. The genera of Rhamnaceae in the southeastern United States. J. Arnold Arbor. 45: 439–463. Medan, D. and C. Schirarend. 2004. Rhamnaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 10+ Vols. Berlin etc. Vol. 6, pp. 320–338. Richardson, J. E. et al. 2000. A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences. Amer. J. Bot. 87: 1309–1324. Richardson, J. E. et al. 2000b. A revision of the tribal classification of Rhamnaceae. Kew Bull. 55: 311–340.

Shrubs, trees, or woody vines, [herbs, annual or perennial], evergreen or deciduous, synoecious, dioecious, or polygamous [monoecious], sometimes armed with thorns or stipular spines. Leaves alternate, fascicled, clustered, or opposite, simple; stipules present; petiole present (absent in Condalia); blade margins entire, serrate, serrulate, crenate, crenulate, dentate, or denticulate, sometimes spinose or spinulose; venation pinnate (sometimes obscurely, appearing 1-veined) or 3[–5]-veined from base (acrodromous). Inflorescences bisexual or unisexual, axillary or terminal, fascicles, umbels, panicles, cymes, or thyrses (these sometimes spikelike, racemelike, or paniclelike), or flowers solitary. Flowers bisexual or unisexual; perianth and androecium epigynous or perigynous [hypogynous]; hypanthium free or adnate to ovary proximally and free distally [absent]; sepals [3–]4–5(–8), distinct, valvate; petals 0 or [3–]4–5(–8), distinct; nectary present, intrastaminal, sometimes lining hypanthium; stamens [3–]4–5(–8), opposite petals, distinct, adnate to petal bases; anthers dehiscing by longitudinal slits; pistil 1, 2–4-carpellate, ovary superior to inferior, (1–)2–4-locular, placentation basal; ovules 1 per locule (2 per locule in Karwinskia), anatropous; styles 1–4, connate proximally to completely; stigmas 1–4. Fruits capsules, dehiscence loculicidal, schizocarps, samaras, or drupes. Seeds 1 (sometimes 2 in Karwinskia) per locule.

Arbres, arbustes ou lianes, ordinairement inermes au Gabon. Des vrilles chez Gouania. Stipules ± caduques. Feuilles simples, opposées, subopposées ou alternes. Limbe souvent denté ou denticulé . Nervures pennées, parfois plusieurs basilaires palmées. Nervilles souvent nombreuses et apparentes. Inflorescences pluri- ou multiflores en cymes, glomérules ou grappes spiciformes de glomérules. Fleurs hermaphrodites, pentamères, petites, blanches, jaunâtres ou vertes. Réceptacle conique ou cupuliforme. Calice à 5 sépales valvaires ± carénés sur la face interne. 5 pétales entièrement cachés à la préfloraison , alternisépales, onguiculés ± cucullés et protégeant chacun une étamine en la coiffant au moins dans le bouton; les 5 étamines sont insérées à la base des pétales , au bord ou sous le bord du disque. Anthère introrse à 2 loges. Pollen simple tricolporé, généralement triangulaire angulaperturé. Disque variable remplissant plus ou moins, ou tapissant le réceptacle, entourant ou submergeant l'ovaire. Ce dernier ± adhérent, infère, semi-infère, ou libre et supère, à 1-2-3 loges presque toujours uniovulées. Ovules basilaires ascendants. Style central à stigmate ± lobé, bi- ou trifide, plus rarement pelté (Maesopsis). Fruit tantôt charnu et drupacé (Maesopsis), tantôt sec ± déhiscent et alors ailé samaroïde (1 aile et l graine chez Ventilago; 3 ailes et 3 graines chez Gouania) ou aptère et capsulaire tricoque à 3 graines (Lasiodiscus).

Arbres , arbustes ou arbrisseaux, parfois sarmenteux, parfois munis d'épines ou de vrilles, très rarement plantes herbacées.'Feuilles'simples, alternes, parfois opposées, stipulées, entières ou diversement dentées, parfois munies de glandes.'Inflorescences'le plus souvent en cymes ou fascicules axillaires, ces derniers parfois réunis en racèmes ou panicules.'Fleurs'petites, actinomorphes, ☿, rarement ♂♀, (4)5-mères; réceptacle ± développé : turbiné, disciforme ou obconique; sépales valvaires, triangulaires, souvent munis d'une carène membraneuse sur la face interne; pétales souvent plus petits que les sépales, ± longuement onguiculés, souvent en capuchon ou en cuiller, parfois spatulés, rarement absents; étamines oppositipétales, souvent embrassées par le pétale, à filet court; anthères à 2 loges à déhiscence longitudinale; disque presque toujours présent et souvent bien développé, périgyne ou parfois épigyne, tapissant parfois le réceptacle, circulaire ou 4-5-lobé, toujours intrastaminal; ovaire supère ou ± semi-infère à infère, (1-2)3(4-5)-loculaire; placentation basilaire, 1 ovule par loge, dressé, anatrope, à 2 téguments, rarement 2 ovules par loge, très rarement 1 seul ovule dans l'ovaire.'Fruits'variables, secs, souvent déhiscents en coques, ou charnus, parfois ailés, souvent munis de la cicatrice du disque persistant.'Graines parfois arillées, à albumen en général présent mais peu abondant, parfois absent; embryon grand, à larges cotylédons convexes; radicule petite.\n\t\t\t\n\t\t\t\t Plantules à germination hypo-ou épigée.\n\t\t\tFamille des régions tropicales et tempérées, groupant 58 genres et ± 900 espèces; 8 genres, 21 espèces et 1 variété au Congo belge. Plusieurs espèces de'Ziziphus'sont cultivées pour leurs fruits comestibles;'Colubrina ferruginosa Brongn.1, espèce ornementale, a été introduite au Congo belge.

Seeds mostly with copious endosperm and large straight embryo Fruit various, often drupaceous Ovary sessile, superior or subinferior, 2–4-celled; ovules solitary or rarely paired, erect from the base, anatropous Disk mostly present, perigynous Stamens 4–5, opposite the petals; anthers 2-celled, opening lengthwise Petals 4 or 5, small, or absent Flowers often cymose or fasciculate, small, hermaphrodite or polygamous Calyx-lobes valvate Trees or shrubs, sometimes climbing Leaves simple, alternate or opposite; stipules mostly present Fruit various, often drupaceous Ovary sessile, superior or subinferior, 2–4-celled; ovules solitary or rarely paired, erect from the base, anatropous Disk mostly present, perigynous Stamens 4–5, opposite the petals; anthers 2-celled, opening lengthwise Petals 4 or 5, small, or absent Flowers often cymose or fasciculate, small, hermaphrodite or polygamous Calyx-lobes valvate Trees or shrubs, sometimes climbing Leaves simple, alternate or opposite; stipules mostly present

Flowers often in axillary cymes or umbels (rarely solitary), or in racemes arranged in terminal panicles or thyrses, bisexual (rarely unisexual), actinomorphic Trees, shrubs, shrublets or lianes, glabrous or with simple hairs; branches rarely with coiled tendrils; leaves alternate or rarely opposite, simple, entire to toothed, petiolate, penninerved or 3–5-nerved from the base; stipules present, rarely interpetiolar, sometimes spinescent Fruit a drupe or septicidal capsule or schizocarp, (1)2–3(4)-locular, sometimeswinged Disk usually present and well developed, intrastaminal, perigynous, very variable in shape, large, filling the receptacle or cup-shaped with free margins, or lining the receptacle; ovary syncarpous, sessile, free or immersed in the disk, superior, subinferior or inferior, 2–4-locular; style entire or 2–4-lobed; ovules solitary in each loculus, erect, anatropous Seeds 1 in each loculus; embryo large, straight; endosperm usually copious Sepals (4) 5, valvate Receptacle flattish to obconic or hemispherical Stamens (4) 5, antipetalous; filaments free; anthers 2-thecous (rarely 1-thecous), introrse, dehiscing longitudinally Petals (4) 5 or absent, usually smaller than the sepals and unguiculate, cucullate, closely surrounding the stamens Trees, shrubs, shrublets or lianes, glabrous or with simple hairs; branches rarely with coiled tendrils; leaves alternate or rarely opposite, simple, entire to toothed, petiolate, penninerved or 3–5-nerved from the base; stipules present, rarely interpetiolar, sometimes spinescent Fruit a drupe or septicidal capsule or schizocarp, (1)2–3(4)-locular, sometimeswinged Disk usually present and well developed, intrastaminal, perigynous, very variable in shape, large, filling the receptacle or cup-shaped with free margins, or lining the receptacle; ovary syncarpous, sessile, free or immersed in the disk, superior, subinferior or inferior, 2–4-locular; style entire or 2–4-lobed; ovules solitary in each loculus, erect, anatropous Seeds 1 in each loculus; embryo large, straight; endosperm usually copious Sepals (4) 5, valvate Receptacle flattish to obconic or hemispherical Stamens (4) 5, antipetalous; filaments free; anthers 2-thecous (rarely 1-thecous), introrse, dehiscing longitudinally Petals (4) 5 or absent, usually smaller than the sepals and unguiculate, cucullate, closely surrounding the stamens

Fls regular, 4- or 5-merous, perfect or sometimes unisexual, rather shortly perigynous; pet distinct, often clawed, rarely wanting; stamens as many as and alternate with the sep, opposite and often enfolded by the concave or hooded pet; ovary sessile on the intrastaminal nectary-disk or ± immersed in it (thus sometimes seemingly inferior), 2–3(–5)-locular, with a single erect, basal ovule per locule; style terminal, undivided to often deeply cleft; fr a drupe or sometimes a capsule or schizocarp; embryo large, with 2 cotyledons; endosperm usually scanty or wanting; woody plants with simple lvs and small fls. 55/900, cosmop.

Deciduous or evergreen, often thorny trees, shrubs, woody climbers, or lianas, rarely herbs. Leaves simple, petiolate, alternate or opposite, pinnately veined or 3-5-veined, entire to serrate, sometimes much reduced; stipules small, caducous or persistent, sometimes transformed into spines. Flowers yellowish to greenish, rarely brightly colored, small, bisexual or unisexual, rarely polygamous, (4 or)5-merous, hypogynous to epigynous, in mostly axillary, sessile or pedunculate cymes, or reduced to few in fascicles. Calyx tube patelliform or hemispherical to tubular, sometimes absent, at rim with calyx, corolla, and stamens; sepals 4 or 5, valvate in bud, triangular, erect or ± recurved during anthesis, adaxially often distinctly keeled, alternate with petals. Petals 4 or 5, rarely absent, usually smaller than sepals, concave or hooded, rarely nearly flat, often shortly clawed. Stamens 4 or 5, antepetalous and often ± enclosed by petals; filaments thin, adnate to bases of petals; anthers minute, versatile or not, 2(or 4)-celled, dehiscing by longitudinal slits, usually introrse. Disk intrastaminal, nectariferous, thin to ± fleshy, entire or lobed, glabrous or rarely pubescent, free from ovary or tightly surrounding it, or adnate to calyx tube. Ovary superior to inferior, (1 or)2-4-loculed, with 1(or 2) ovules per locule; ovules anatropous, basal and erect; styles simple or ± deeply 3-lobed or 3-cleft. Fruit either an indehiscent, rarely explosively dehiscent, sometimes winged, schizocarpic capsule, or a ± fleshy drupe with 1-4 indehiscent, rarely dehiscent, pyrenes (stones). Seeds with thin, oily albumen, sometimes exalbuminous; embryo large, oily, straight or rarely bent.

Ovary syncarpous, with 2 or 3 (rarely 4 or 1) cells; ovule solitary in the cell, anatropous; style minute, rarely simple, usually with 2, 3 or rarely 4 or even more rarely 10 microscopic stigmatic lobes at apex Fruit often dryish and splitting into 3 1-seeded parts at maturity (as in the first 5 genera treated here), or fleshy and with 2 or 3 free 1-seeded stones (as in >i>Rhamnus and >i>Scutia) or fleshy or dryish and with a single 1-, 2- or 3-seeded (or 4-seeded but not in Africa) stone (as in the last 4 genera here); placentation basal Seed with raphe dorsal or lateral; embryo large and straight, the cotyledons usually in planes tangential to the ovary-axis; endosperm in a thick or thin layer, rarely nearly absent but not in Africa, rarely ruminate but not in Africa Sepals, petals and stamens attached at the rim of the cup Flowers minute, regular, bisexual (and often strongly protandrous, or reportedly protogynous in >i>Maesopsis) or less commonly unisexual, peri-gynous or epigynous, 4- or usually 5-merous (6-merous very rarely and not in Africa), in basically cymose arrangements but the cymes often either reduced to fascicles (or even to solitary flowers) or arranged in short or elongate thyrses which in turn are sometimes disposed in leafy to leafless panicles; each flower with a cup lined with a thin intrastaminal nectariferous disk or the disk sometimes thickened near and/or produced beyond the rim of the cup and either free from the ovary or adnate to it Petals absent or usually present, enclosed by calyx in bud, nearly always shorter than the sepals at anthesis, each usually with a narrow base or claw plus an expanded hood-like or concave or amplectant body closely associated with the stamen Sepals triangular, valvate in bud (this being one of the most useful traits to distinguish members of this family from those plants often confused with them) Stamens bowed inward in bud, as many as, opposite to, usually shorter than and usually clasped or hooded by the petals Trees, erect, climbing or scandent shrubs or lianes or subshrubs (or annual herbs but not in East Africa); tendrils present in Heliums and >i>Gouania Stipules mostly present, free or interpetiolar or intra-axillary Leaves simple, alternate or opposite, petiolate (or sessile but not in East Africa); blades penninerved or 3–5-nerved from the base, unlobed, serrate or crenate with each tooth or crenation usually associated with a minute gland, or entire Fruit often dryish and splitting into 3 1-seeded parts at maturity (as in the first 5 genera treated here), or fleshy and with 2 or 3 free 1-seeded stones (as in >i>Rhamnus and >i>Scutia) or fleshy or dryish and with a single 1-, 2- or 3-seeded (or 4-seeded but not in Africa) stone (as in the last 4 genera here); placentation basal Seed with raphe dorsal or lateral; embryo large and straight, the cotyledons usually in planes tangential to the ovary-axis; endosperm in a thick or thin layer, rarely nearly absent but not in Africa, rarely ruminate but not in Africa Sepals, petals and stamens attached at the rim of the cup Flowers minute, regular, bisexual (and often strongly protandrous, or reportedly protogynous in >i>Maesopsis) or less commonly unisexual, peri-gynous or epigynous, 4- or usually 5-merous (6-merous very rarely and not in Africa), in basically cymose arrangements but the cymes often either reduced to fascicles (or even to solitary flowers) or arranged in short or elongate thyrses which in turn are sometimes disposed in leafy to leafless panicles; each flower with a cup lined with a thin intrastaminal nectariferous disk or the disk sometimes thickened near and/or produced beyond the rim of the cup and either free from the ovary or adnate to it Petals absent or usually present, enclosed by calyx in bud, nearly always shorter than the sepals at anthesis, each usually with a narrow base or claw plus an expanded hood-like or concave or amplectant body closely associated with the stamen Sepals triangular, valvate in bud (this being one of the most useful traits to distinguish members of this family from those plants often confused with them) Stamens bowed inward in bud, as many as, opposite to, usually shorter than and usually clasped or hooded by the petals Trees, erect, climbing or scandent shrubs or lianes or subshrubs (or annual herbs but not in East Africa); tendrils present in Heliums and >i>Gouania Stipules mostly present, free or interpetiolar or intra-axillary Leaves simple, alternate or opposite, petiolate (or sessile but not in East Africa); blades penninerved or 3–5-nerved from the base, unlobed, serrate or crenate with each tooth or crenation usually associated with a minute gland, or entire