The World Flora Online

Herbs annual or perennial, shrubs, or subshrubs, sometimes scandent, rarely small trees. Leaves simple, usually alternate, sometimes opposite, with small or leaflike stipules, petiolate, margin entire, serrate, or dissected. Flowers bisexual or unisexual, rarely polygamous, actinomorphic or zygomorphic, solitary or in axillary or terminal, spicate, paniculate, or racemose inflorescences, 2-bracteolate, sometimes cleistogamous. Sepals 5, equal or unequal, imbricate, persistent. Petals 5, imbricate or convolute, unequal, anterior one usually larger than others, saccate, gibbous or spurred at base. Stamens 5; anthers erect, free or connivent or connate, connectives often dilated into membranous appendages; filaments very short or absent, anterior 2 stamens with spurlike nectary at base. Ovary superior, 1-loculed, 3-5-carpelled, syncarpous, with 3-5 parietal placentae each with 1 to many anatropous ovules; style simple; stigmas variously shaped. Fruit a loculicidal capsule, usually with elastic and abaxially carinate valves, rarely baccate. Seeds often carunculate; testa hard, nitid, often with oily bodies, sometimes alate; endosperm copious, fleshy; embryo erect.

Flowers actinomorphic or zygomorphic, from solitary to paniculate Leaves alternate or rarely opposite, stipulate Herbs, shrubs, or small trees Ovary superior, sessile, 1-celled, with usually 3 parietal placentas each with one or more ovules; style usually simple Stamens hypogynous, 5, alternate with the petals; anthers 2-celled, opening lengthwise, with the connective produced beyond the cells Petals 5, nearly equal or the lower larger and often clawed, imbricate Sepals 5, imbricate Fruit an elastic loculicidal capsule, one- or more-seeded Seeds with fleshy endosperm and central mostly straight embryo Leaves alternate or rarely opposite, stipulate Herbs, shrubs, or small trees Ovary superior, sessile, 1-celled, with usually 3 parietal placentas each with one or more ovules; style usually simple Stamens hypogynous, 5, alternate with the petals; anthers 2-celled, opening lengthwise, with the connective produced beyond the cells Petals 5, nearly equal or the lower larger and often clawed, imbricate Sepals 5, imbricate Fruit an elastic loculicidal capsule, one- or more-seeded Seeds with fleshy endosperm and central mostly straight embryo

Herbes, arbustes ou arbres.'Feuilles'alternes, rarement opposées ou ternées, simples, entières ou diversement découpées, glabres ou pubescentes, glanduleuses ou non à la face inférieure; stipules 2, petites ou grandes et foliacées, rapidement caduques chez les espèces ligneuses, persistantes chez les espèces herbacées. Inflorescences axillaires ou/et terminales, en fascicules, racèmes ou panicules, ou fleurs solitaires.'Fleurs'actinomorphes ou zygomorphes, ☿, très rarement ♂ ♀, parfois cléistogames; pédicelles articulés ou non, généralement garnis de 2 bractéoles; sépales 5, libres ou courtement soudés à la base, subégaux à inégaux, imbriqués, concaves ou plans, généralement persistants; pétales 5, libres, hypogynes ou faiblement périgynes, imbriqués ou contortés, alternisépales, égaux ou inégaux et dissemblables, l'antérieur le plus souvent plus grand et éperonné; étamines 5, oppositisépales, semblables ou dissemblables, les 2 antérieures, dans les fleurs zygomorphes, appendiculées et nectarifères; filets presque entièrement libres ou soudés en un anneau court ou en un tube entourant l'ovaire; tube formé par la concrescence du disque extrastaminal et des bases des filets staminaux; anthères sessiles ou à court filet, glabres ou pubescentes, introrses, déhiscentes par fente longitudinale, insérées à l'intérieur du tube ou au sommet, à connectif prolongé dorsalement au-dessus des loges en une lame pétaloïde, membraneuse, décurrente ou non jusqu'à la base de l'anthère; thèques appendiculées ou non; ovaire supère, glabre, pubérulent, pubescent ou hirsute, 1-loculaire; placentas (2) 3 (5), pariétaux, chacun portant 1- ovules; style simple, cylindrique ou claviforme, souvent en forme d'S dans les fleurs zygomorphes; stigmate ponctiforme.'Capsules'ovoïdes-trilobées, loculicides à 3 valves élastiques, papyracées à subligneuses.'Graines à petit funicule, ovoïdes ou anguleuses-subtriquêtres, parfois ailées, rarement arillées.\n\t\t\tFamille cosmopolite comptant 16 genres et environ 800 espèces des régions tropicales, subtropicales et tempérées. Pour la Flore, 3 genres, 45 espèces dont 5 représentées par 10 variétés et 4 imparfaitement connues.

Fls hypogynous, perfect, in ours (tribe Violeae) irregular; sep 5, commonly persistent; pet 5, distinct, the lowest one in ours prolonged behind into a spur; stamens 5, the filaments very short, distinct or ± connate, the anthers commonly connivent around the ovary, often the 2 lower ones (Violeae) or all them (the woody genera) with a gland-like or spur-like nectary on the back; connective often prolonged into a membranous appendage; ovary superior, unilocular, with mostly 3 parietal placentas; style solitary, often distally enlarged or otherwise modified, the stigma simple or lobed, often oblique; ovules (1–) several or many on each placenta; fr a loculicidal capsule (ours) or sometimes a berry; seeds with straight embryo and flat cotyledons embedded in the copious, oily endosperm, often arillate; herbs (ours) or less often woody plants; lvs alternate or seldom opposite or even whorled, stipulate, simple and entire or toothed to sometimes lobed or dissected, but without well defined lfls, the fls axillary, often nodding, or in racemes, heads, or panicles. 16/800, cosmop.

Seeds generally with ample endosperm, occasionally arillate Stamens 5, antisepalous, the lower pair (anterior) in zygomorphic flowers each with an appendage which projects into the spur and which secretes nectar; filaments free or united wholly or partly into a ring around the ovary; anthers introrse, usually with a prominent connective appendage, sometimes with thecal appendages also Ovary sessile, ± ovoid, 1-locular, with (2–)3, 4 or 5 parietal placentas; style solitary, often thickened towards the stigma, which is generally undivided Fruit a loculicidal capsule, generally splitting into 3 wide-spreading contractile valves, rarely a nut or berry Annual or perennial herbs, shrubs or small trees Sepals 5, free or united near the base, usually persistent Petals 5, free, equal or unequal, the anterior one (lowermost in flower) sometimes spurred, imbricate, generally deciduous Leaves spirally arranged, rarely opposite or verticillate, simple, entire or toothed, rarely dissected; stipules present, small or foliaceous, the margin often ciliate or laciniate Flowers actinomorphic or zygomorphic, solitary or in a simple or compound inflorescence, often thyrsoid, terminal or axillary, usually hermaphrodite, sometimes unisexual on separate plants Stamens 5, antisepalous, the lower pair (anterior) in zygomorphic flowers each with an appendage which projects into the spur and which secretes nectar; filaments free or united wholly or partly into a ring around the ovary; anthers introrse, usually with a prominent connective appendage, sometimes with thecal appendages also Ovary sessile, ± ovoid, 1-locular, with (2–)3, 4 or 5 parietal placentas; style solitary, often thickened towards the stigma, which is generally undivided Fruit a loculicidal capsule, generally splitting into 3 wide-spreading contractile valves, rarely a nut or berry Annual or perennial herbs, shrubs or small trees Sepals 5, free or united near the base, usually persistent Petals 5, free, equal or unequal, the anterior one (lowermost in flower) sometimes spurred, imbricate, generally deciduous Leaves spirally arranged, rarely opposite or verticillate, simple, entire or toothed, rarely dissected; stipules present, small or foliaceous, the margin often ciliate or laciniate Flowers actinomorphic or zygomorphic, solitary or in a simple or compound inflorescence, often thyrsoid, terminal or axillary, usually hermaphrodite, sometimes unisexual on separate plants

SELECTED REFERENCES Ballard, H. E., J. de Paula-Souza, and G. A. Wahlert. 2014. Violaceae. In: The Families and Genera of Vascular Plants. Vol. 11, pp. 303–322. Berlin, Heidelberg. Brainerd, E. 1921. Violets of North America. Bull. Vermont Agric. Exp. Sta. 224. Brizicky, G. K. 1961b. The genera of Violaceae in the southeastern United States. J. Arnold Arbor. 42: 321–333. Feng, M. 2005. Floral Morphogenesis and Molecular Systematics of the Family Violaceae. Ph.D. dissertation. Ohio University. Feng, M. and H. E. Ballard. 2005. Molecular systematic, floral developmental and anatomical revelations on generic relationships and evolutionary patterns in the Violaceae. In: International Botanical Congress. 2005. XVII International Botanical Congress, Vienna, Austria, Europe, Austria Center Vienna, 17–23 July 2005. Abstracts. P. 169. Gershoy, A. 1928. Studies in North American violets. I. General considerations. Bull. Vermont Agric. Exp. Sta. 279. Hodges, S. A. et al. 1995. Generic relationships in the Violaceae: Data from morphology, anatomy, chromosome numbers and rbcL sequences. [Abstract.] Amer. J. Bot. 82(6, suppl.): 136. McKinney, L. E. and N. H. Russell. 2002. Violaceae of the southeastern United States. Castanea 4: 369–379. Tokuoka, T. 2008. Molecular phylogenetic analysis of Violaceae (Malpighiales) based on plastid and nuclear DNA sequences. J. Pl. Res. 121: 253–260. Wahlert, G. A. et al. 2014. Phylogeny of the Violaceae (Malpighiales) inferred from plastid DNA sequences: Implications for generic diversity and intrafamilial classification. Syst. Bot. 39: 239–252.

Herbs, annual or perennial, [subshrubs, shrubs, lianas, and trees], glabrous or hairy, hairs simple; taprooted or rhizomatous, sometimes stoloniferous. Stems 0–20, prostrate to erect. Leaves cauline or basal, (attached directly to rhizome, some Viola), alternate (and opposite in Hybanthus [and other genera]), simple or compound, stipulate [estipulate], petiolate or sessile; blade unlobed or lobed. Inflorescences 1(–4)[–5]-flowered, axillary from leaf axils or scapose from rhizomes or stolons (or in racemes of umbels), pedunculate; bracteoles usually present on peduncles, usually alternate. Flowers bisexual [unisexual, plants dioecious], perianth and unequal, imbricate in bud [convolute], lowermost petal often larger with gibbous or elongated spur; stamens 5, alternate with petals, surrounding ovary, connivent or syngenesious; filaments 0–1 mm, filaments of 2 anterior stamens often with nectaries protruding into spur, anther dehiscence by longitudinal slits; pistil 1, [2–]3[–5]-carpellate; ovary superior, 1-locular; placentation parietal; ovules [1–2]8–75, anatropous, bitegmic, crassinucellate; style [0–]1, usually enlarged distally, solid or hollow; stigma 1 [3–5], with or without hairs. Fruits capsular [berry, nut], 3-valved, dehiscence loculicidal. Seeds [1–](3–)6–75, hard, embryo not developed at time of dispersal, spheroid or ovoid [strongly flattened], glabrous [hairy], some arillate, some with elaiosome [seeds winged in some woody vines].

Sepals 5, free or shortly united, quincuncial or open in bud, usually persistent Petals 5, free, equal or ± unequal, the anterior one frequently ± spurred, imbricate, usually deciduous, alternating with the sepals Leaves alternate (rarely opposite or whorled), simple, entire or serrate to dentate (rarely ± dissected), usually with 2 stipules Flowers actinomorphic or, more often, ± zygomorphic, bisexual (rarely polygamous or dioecious) Shrubs or perennial or annual herbs (more rarely trees) Seeds sometimes with a small aril, usually with abundant endosperm Fruit a loculicidal capsule usually with contractile carinate valves, rarely a berry or nut Ovary free, sessile, usually ± ovoid, 1-locular, with (2) 3 (4–5) parietal placentas each bearing 1–? ovules; styles completely united, usually thickened above, often ± S-shaped in zygomorphic flowers, stigma usually undivided Stamens 5, antisepalous, similar or ± dissimilar, the anterior pair in zygomorphic flowers with appendages which project into the spur, filaments free ± united, often forming a cylinder round the ovary; anthers usually introrse, free ± united, usually with a prolongation of the connective Petals 5, free, equal or ± unequal, the anterior one frequently ± spurred, imbricate, usually deciduous, alternating with the sepals Leaves alternate (rarely opposite or whorled), simple, entire or serrate to dentate (rarely ± dissected), usually with 2 stipules Flowers actinomorphic or, more often, ± zygomorphic, bisexual (rarely polygamous or dioecious) Shrubs or perennial or annual herbs (more rarely trees) Seeds sometimes with a small aril, usually with abundant endosperm Fruit a loculicidal capsule usually with contractile carinate valves, rarely a berry or nut Ovary free, sessile, usually ± ovoid, 1-locular, with (2) 3 (4–5) parietal placentas each bearing 1–? ovules; styles completely united, usually thickened above, often ± S-shaped in zygomorphic flowers, stigma usually undivided Stamens 5, antisepalous, similar or ± dissimilar, the anterior pair in zygomorphic flowers with appendages which project into the spur, filaments free ± united, often forming a cylinder round the ovary; anthers usually introrse, free ± united, usually with a prolongation of the connective