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Key to Viola, Key E: Acaulescent violets lacking stolons, with unlobed leaves

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1 At least some petioles prominently winged; lowest sepals linear-lanceolate, acuminate (ovate-triangular, acute to obtuse in V. villosa); auricles prominent, elongating in fruit, erose; cleistogamous capsule unspotted, on declined or erect peduncle
..2 Petioles, lower surface of leaf blades and peduncles moderately to densely hirtellous (rarely glabrous in V. fimbriatula); at least the outer leaves widely spreading or prostrate on the substrate in life; largest leaf blades elliptical or narrowly oblong-ovate in outline; peduncle hirtellous; calyx ciliate; corolla purple, petals broadly rounded, spurred petal not medially compressed; peduncle of cleistogamous capsule declined; seeds 1.4-1.9 × 0.9-1.3 mm, medium to dark brown or gray-brown, unspotted or with small weak darker blotches.
....3 Upper surface of leaf blades concolorous; larger leaf blades all distinctly longer than broad, elliptical or oblong to oblong-ovate (rarely lanceolate), base shallowly cordate or truncate to cuneate, apex acute; petioles shorter than leaf blades (chasmogamous flower) to 2 × as long as blades (cleistogamous fruit); lowest sepals linear-lanceolate, acuminate; upper and lateral petals of chasmogamous flower ≥ 3 mm wide; sepals of cleistogamous fruit ≥ 1/2 as long as capsule; seeds (1.3-) avg. 1.6 (-1.8) × (0.9-) avg. 1.1 (-1.3) mm, medium to dark brown or gray-brown with small weak or prominent darker spots, streaks or blotches
....3 Upper surface of leaf blades commonly bicolorous, with conspicuous darker green or purple veins; at least some larger leaf blades usually scarcely longer than broad to broader than long, narrowly ovate or orbicular to reniform (rarely elliptical), base deeply cordate, apex obtuse to narrowly or broadly rounded; petioles variable, sometimes shorter than leaf blades and sometimes > 2 × as long as blades; lowest sepals lanceolate to ovate-triangular, acute to obtuse or rounded (uncommonly acuminate); upper and lateral petals of chasmogamous flower < 3 mm wide; sepals of cleistogamous fruit < 1/2 as long as capsule; seeds (1.6-) avg. 1.9 (-2.2) × (1.0-) avg. 1.3 (-1.5) mm, brown with minute raised black spots or dark red-brown to black and unspotted
..2 Foliage glabrous or surfaces of leaf blades sparsely appressed-hirsute near margins; leaves erect in life; petioles > 2 × as long as leaf blades; largest leaf blades narrowly ovate-triangular, triangular or deltate-triangular; peduncle glabrous; calyx eciliate; corolla violet, petals commonly emarginate, spurred petal commonly medially compressed; peduncle of cleistogamous capsule erect; seeds 1.4–2.2 × 0.9–1.5 mm, pinkish-gray to dark gray with prominent darker streaks and spots
......4 Petals commonly emarginate; spurred petal commonly medially compressed; [dry sandy soils of open dry woodlands and forest edges on the Piedmont and Atlantic Coastal Plain]
......4 Petals broadly rounded; spurred petal not medially compressed; [dry or wet soils west of the Appalachians].
........5 Outline of largest leaf blades convexly tapering to (broadly) rounded apex; [low wet prairies, s. MO and e. KS south to se. OK]
........5 Outline of largest leaf blades concavely tapering to narrowly acuminate or narrowly rounded apex; [open dry oak and oak-pine woodlands and forest edges, WV & s. OH south to n. AL, LA, and n. AR]
1 Petioles not prominently winged; lowest sepals lanceolate to (more commonly) ovate-lanceolate, oblong or ovate, acuminate to rounded; auricles inconspicuous and remaining short and rounded, or prominent and elongating in fruit and often erose; cleistogamous capsule unspotted or spotted, on prostrate, declined or erect peduncle.
..........6 All petals glabrous within (lateral petals densely bearded within in V. inconspicua); spur > 2× as long as thick, cylindrical or club-shaped; stipules at least half-adnate to petioles.
............ 7 Leaf blades narrowly ovate-triangular to deltate-triangular, broadly cordate with divergent basal lobes with non-overlapping or divergent lobes, the blade margins above the basal lobes straightish to concave; spur 1-3 mm long; upper surface of blades glabrous; lateral petals densely bearded within; [alien, usually in human-maintained habitats]
............ 7 Leaf blades narrowly ovate to ovate, deeply cordate with inner edges of basal lobes commonly overlapping in life, the blade margins above the basal lobes convex; spur 4-7 mm long; upper surface of blades with scattered subappressed hairs; lateral petals glabrous within; [circumboreal, NL to AK, south to RI, PA, WI, n. MN and n. WA, disjunct in SD and CO; Greenland, Eurasia]
..........6 Lateral petals bearded within, spurred petal in some species bearded; spur < 1.5× as long as thick, rounded; leaf blades variously shaped, inner edges of basal lobes not overlapping in life; stipules free.
............ ..8 Leaves spreading to prostrate on the substrate in life; upper leaf blade surface silvery- or gray-green with contrasting dark green or red-purple veins, lower blade surface purple-tinged; foliage glabrous except for conspicuous stiff ascending or spreading hairs uniformly distributed over upper surface of leaf blade; calyx eciliate; lowest sepals oblong to ovate, obtuse to rounded; spurred pet densely bearded; cleistogamous capsule purple-spotted, on initially prostrate peduncle shorter than petioles; seeds 1.5–2.1 × 1.1–1.4 mm, light brown, commonly with slightly darker streaks and spots
............ ..8 Leaves held above the substrate (prostrate in V. villosa, at least outer leaves prostrate especially in fruit in V. fimbriatula); upper surface of leaf blades uniformly green, lower green or purple tinged; foliage glabrous or variously pubescent, if hairs confined to upper surface of leaf blade, then these very small, scattered and (sub)appressed.
............ ....9 Margins of largest leaf blades closely pectinately serrate, surfaces glabrous or sparsely appressed-hirtellous; chasmogamous flowers held above the leaves; lowest sepals linear-lanceolate, acuminate; auricles prominent, elongating to 3 mm in fruit; [sporadic along Atlantic Coastal Plain]
............ ....9 Margins of largest leaf blades merely crenate to serrate, various in indument; chasmogamous flowers held above or among the leaves; lowest sepals and auricles various; [most species absent from or rare on the Atlantic Coastal Plain].
............ ......10 Plant in chasmogamous flower.
............ ........11 Largest leaf blades approx. as broad as long to broader than long (length:width ratio < 1.2), broadly ovate or orbicular to deltate or reniform (one largest leaf blade often longer than broad in V. floridana, V. latiuscula, V. septentrionalis, these are keyed here).
............ ..........12 Spurred petal densely bearded within; lower surface of leaf blades commonly purple-tinged (blades uniformly green in V. retusa).
............ ............ 13 Foliage and peduncles sparsely to densely hirsute, leaf blade margins ciliate, hairs often long; calyx glabrous or appressed-hirtellous, ciliate to apex; auricles prominent, quadrate
............ ............ 13 Foliage strictly glabrous or upper surface of leaf blades with small scattered subappressed hairs, blade margins eciliate; peduncle glabrous; calyx glabrous, eciliate; auricles short and rounded.
............ ............ ..14 Leaf blades orbicular to reniform, broadly obtuse to rounded at apex; lowest sepals oblong to ovate, obtuse to rounded; [dry or wet calcareous open sites throughout ne. N America, Great Plains and Rocky Mtns., sporadic southward along Appalachian Mtns. to w. VA]
............ ............ ..14 Leaf blades deltate-triangular, acute to short-acuminate at apex; lowest sepals acuminate to acutish.
............ ............ ....15 Petals broadly obovate, broadly rounded; lowest sepals oblong-lanceolate to ovate-lanceolate, acute or acutish; lower surface of leaf blades commonly flushed with purple; summit of petioles commonly papillate or granular; [dry sandy woodland soils in NY, PA, VT]
............ ............ ....15 Petals narrowly obovate, upper commonly retuse; lowest sepals linear-lanceolate, acuminate; leaves uniformly green; petioles smooth; [gravelly riverbanks in the Great Plains]
............ ..........12 Spurred petal glabrous within; foliage green (foliage blue-green, petioles, lower surface of leaf blades and peduncles purple-tinged, leaf blades ovate to orbicular in V. sororia var. 2 [“hirsutula-like” variant]).
............ ............ ......16 Lowest sepals oblong to ovate, obtuse to rounded; auricles not prominent, short and rounded or truncate.
............ ............ ........17 Foliage (at least the petioles and lower surface of leaf blades) hirsute; calyx ciliate
............ ............ ........17 Foliage strictly or essentially glabrous; calyx eciliate.
............ ............ ..........18 Foliage, peduncles and calyx uniformly medium green; apex of leaf blades obtuse or abruptly acutish; margins incurved-serrate; sepals oblong to narrowly ovate (uncommonly broadly so), obtuse to rounded at apex; [widely distributed]
............ ............ ..........18 Foliage, peduncles and calyx blue-green, midrib of upper leaf blade surface purple, lower surface of leaf blades, petioles, peduncles and calyx flushed with purple; apex of leaf blades broadly rounded; margins shallowly crenate; sepals ovate, broadly rounded at apex; [Appalachian highlands and Western Allegheny Plateau]
............ ............ ......16 Lowest sepals linear-lanceolate to ovate-triangular, acuminate; auricles prominent, narrowly linear to trapezoidal.
............ ............ ............ 19 Corolla violet with contrasting dark purple eyespot around throat; lowest sepals linear-lanceolate; lateral petal beards very short, exposing throat in life, hairs strongly clavate to reniform
............ ............ ............ 19 Corolla violet to deep purple without contrasting eyespot (white with contrasting purple-gray eyespot around throat in V. communis ['V. priceana']; lowest sepals lance- to ovate-triangular; lateral petal beards long, obscuring throat in life, hairs filiform or slightly clavate.
............ ............ ............ ..20 Corolla purple (white with contrasting purple-gray eyespot around throat in V. communis f. priceana, ined.); petioles ascending in life; leaf blades suborbicular or ovate-deltate, broadly rounded to obtuse at apex, deep green, glossy above; foliage strictly glabrous or upper leaf blade surface with scattered minute appressed hairs requiring magnification; auricles weakly prominent, trapezoidal, commonly erose, 1.5-2 mm; [widespread in floodplains, upland/lowland transitions, thickets, lawns and suburban woodlots, mostly absent from the Coastal Plain]
............ ............ ............ ..20 Corolla whitish (with prominent dark violet near throat on lateral and spurred petals in V. floridana [“peninsular Florida” variant]), pale violet or rose-violet; petioles ascending to widely spreading in life; leaf blades ovate to broadly so, obtuse to subacute at apex, medium to dark green, not glossy above; marginal dentition incurved-serrate; upper surface of leaf blades with easily visible scattered appressed hairs; auricles short and rounded or truncate, < 1 mm; silty; [lowland stream and river terraces and hammocks, FL and Gulf Coastal Plain].
............ ............ ............ ....21 Petioles ascending; flowers held among or slightly overtopping leaves; corolla rose-violet; [local along w. Gulf Coast in LA, MS]
............ ............ ............ ....21 Petioles and leaf blades widely spreading; flowers greatly overtopping leaves; corolla whitish to violet; [e. Gulf Coast region in FL, s. AL and s. GA].
............ ............ ............ ......22 All petals pale violet distally and white in proximal half near throat, spurred petal with a few slender well separated purple nectar-guide-lines on proximal half near throat; lateral petals obovate and broadly rounded, spurred petal broadly obovate and broadly rounded to subtruncate at apex; [throughout range of complex]
............ ............ ............ ......22 Upper and lateral petals whitish distally with conspicuous darker violet flush near throat, spurred petal whitish distally with broad prominent darker violet flush and dense heavily coalescing purple nectar-guides on proximal half near throat; spurred petal obovate or rhombic-obovate and angulate-rounded or trilobate at apex; [central peninsular Florida]
............ ........11 Largest leaf blades distinctly longer than broad (length:width ratio > 1.2).
............ ............ ............ ........23 Spurred petal densely bearded within; foliage glabrous or sparsely to moderately hirtellous [naturally occurring in open sites]
............ ............ ............ ..........24 Largest leaf blades narrowly elliptical, ovate-lanceolate, lance-triangular or sagittate; foliage glabrous or sparsely to moderately hirtellous; flowers overtopping leaves; auricles prominent and quadrate; [dry to moist sand of sand barrens and dry prairies, lakeshores, mostly east of the Great Plains]
............ ............ ............ ..........24 Largest leaf blades narrowly ovate- to deltate-triangular; foliage glabrous or upper surface of leaf blades bearing scattered small subappressed hairs; flowers held among the leaves; auricles short and rounded.
............ ............ ............ ............ 25 Largest leaf blades narrowly ovate-triangular; upper surface of leaf blades bearing scattered small subappressed hairs; upper petals broadly rounded; [sandy and silty soils of floodplains and swamps, absent from the Great Plains region]
............ ............ ............ ............ 25 Largest leaf blades deltate-triangular or an occasional leaf narrowly ovate-triangular; leaves strictly glabrous; upper petals frequently retuse; [gravelly riverbanks in the Great Plains region, KS, OK, and TX westward and northward]
............ ............ ............ ........23 Spurred petal glabrous within; foliage strictly glabrous or upper surface of leaf blades with small to minute scattered subappressed hairs; [naturally occurring in forested sites, V. cucullata also in marshes and meadows].
............ ............ ............ ............ ..26 Calyx ciliolate; lowest sepals oblong to ovate-lanceolate, obtuse to rounded; auricles short and rounded
............ ............ ............ ............ ..26 Calyx eciliate; lowest sepals linear-lanceolate to ovate-triangular, acuminate; auricles prominent to elongate.
............ ............ ............ ............ ....27 Corolla violet with contrasting dark purple "eyespot" around throat; lowest sepals linear-lanceolate; lateral petal beards very short, with strongly clavate to reniform hairs; [widely distributed over e. North America, infrequent on Atlantic Coastal Plain]
............ ............ ............ ............ ....27 Corolla violet lacking contrasting "eyespot" around throat but with flush of dark purple at base of lateral pet; lowest sepals lance- to ovate-triangular; lateral petal beards long, with slightly clavate hairs.
............ ............ ............ ............ ......28 Petioles and leaves widely spreading; auricles prominent but scarcely elongate, < 1.5 mm long; [Gulf Coastal Plain, e. TX to w. FL panhandle]
............ ............ ............ ............ ......28 Petioles ascending; auricles prominent, elongate, 1.5-4 mm long; [s. Atlantic Coastal Plain, se. VA to se. GA]
............ ......10 Plant in cleistogamous fruit.
............ ............ ............ ............ ........29 Largest leaf blades substantially longer than broad (length:width ratio > 1.2), lance-triangular to sagittate
............ ............ ............ ............ ........29 Largest leaf blades about as long as broad to broader than long (length:width ratio < 1.2), ovate or deltate-triangular to reniform.
............ ............ ............ ............ ..........30 Foliage (at least petioles and/or lower surface of leaf blades) hirsute; calyx ciliate.
............ ............ ............ ............ ............ 31 Margins of leaf blades ciliate with spreading hairs (hairs long in Appalachian populations); sepals ciliate to apex (cilia long in Appalachian populations); auricles prominent, elongate to 2 mm; seeds 1.2–2.3 × 0.8–1.2 mm, brown to dark brown, unspotted (rarely with small weak darker spots); [inhabiting limestone and dolomite-associated soils in boreal region and s. at higher elevations to s. Appalachian Mtns]
............ ............ ............ ............ ............ 31 Margins of leaf blades ciliate with short ascending hairs; sepals ciliate in basal ½ (2/3) with short hairs; auricles short and rounded; seeds 1.5–2.6 × 1.0–1.6 mm, dark gray to grayish-brown with faint to prominent rectangular darker blotches; [inhabiting various substrates in upland and lowland forests in much of e. N America]
............ ............ ............ ............ ..........30 Foliage strictly glabrous or upper surface of leaf blades with small to minute scattered subappressed hairs.
............ ............ ............ ............ ............ ......34 Petioles and leaves widely spreading; leaf blades deltate-triangular, broadly acute or right-angled at apex; lowest sepals lanceolate, acuminate; auricles prominent, elongating to 2 mm; seeds yellow-brown; [silt or sand in floodplains of streams and rivers in Gulf Coast region and w. panhandle of FL]
............ ............ ............ ............ ............ ......34 Petioles ascending; leaf blades reniform, broadly obtuse to rounded at apex; lowest sepals oblong to ovate, obtuse to rounded; auricles short and rounded; seeds dark brown; [in dry or wet calcareous substrates of open sites in northeastern states and sporadic along Appalachian highlands, widely distributed across the Cordillera]
............ ............ ............ ............ ............ ....33 Peduncle erect, or initially prostrate and coiled then elongating to become sinuous-erect.
............ ............ ............ ............ ............ ........35 Largest leaf blades suborbicular to deltate-reniform, apex broadly obtuse to rounded, upper surface glossy in life; peduncle initially prostrate and coiled but eventually becoming sinuous-erect; lowest sepals ovate-triangular, < ½ as long as capsule; auricles weakly elongate to 2 mm; seeds olive-brown or brownish-black to black, often with minute raised black spots; [in moist clay or loam soils of thickets, grassy banks and meadows, slope-floodplain transitions and lawns]
............ ............ ............ ............ ............ ........35 Largest leaf blades ovate, apex acute, upper surface not glossy in life; peduncle erect; lowest sepals linear-lanceolate to lanceolate, > ½ as long as the capsule; auricles weakly or strongly elongate; seeds medium to dark brown or reddish-brown, unspotted or spotted; [intermittently or permanently wet soils of wetlands, swamps and riverbanks].
............ ............ ............ ............ ............ ..........36 Auricles strongly elongate, 3–8 mm; seeds 1.4–1.9 × 0.8–1.3 mm, dark brown to dark reddish-brown, unspotted; [inhabiting open or forested wet sites including marshes, sedge meadows, seeps, bog forests and swamps throughout most of e. N Am. e. of the Great Plains]
............ ............ ............ ............ ............ ..........36 Auricles weakly elongate to 1.5 mm; seeds 1.7–2.2 × 1.0–1.3 mm, medium brown with minute raised black spots; [gravel riverbanks in the Great Plains]
............ ............ ............ ............ ............ ..32 Capsule finely spotted, or heavily purple-spotted or blotched.
............ ............ ............ ............ ............ ............ 37 Lowest sepals oblong- to ovate-lanceolate or ovate, obtuse to rounded at apex.
............ ............ ............ ............ ............ ............ ..38 Calyx ciliolate; seeds 1.5–2.2 × 1.0–1.3 mm, medium orange-brown, unspotted or often with minute dark brown to blackish spots; [sandy soils of bottomland forests, w. Midwest, Great Plains and Lower Midwest]
............ ............ ............ ............ ............ ............ ..38 Calyx eciliate; seeds 1.5–2.6 × 1.0–1.6 mm, light gray to medium grayish-brown with weak to dark rectangular blotches; [inhabiting various substrates in upland and lowland forests in much of e. North America].
............ ............ ............ ............ ............ ............ ....39 Foliage, peduncles and calyx uniformly medium green, often turning yellow green in cleistogamous fruit; apex of leaf blades obtuse or abruptly acutish; margins incurved-serrate; sepals oblong to narrowly ovate (uncommonly broadly so), obtuse to rounded at apex; [widely distributed across e. N America to e. Great Plains]
............ ............ ............ ............ ............ ............ ....39 Foliage, peduncles and calyx blue-green, midrib of upper leaf blade surface often purple, lower surface of leaf blades, petioles, peduncles and calyx flushed with purple; apex of leaf blades broadly rounded; margins shallowly crenate; sepals ovate, broadly rounded at apex; [Appalachian highlands and Western Allegheny Plateau]
............ ............ ............ ............ ............ ............ 37 Lowest sepals linear-lanceolate to ovate-triangular, acuminate from middle or base (narrowly rounded to acute in V. latiuscula).
............ ............ ............ ............ ............ ............ ......40 Peduncle declined, or initially prostrate and coiled but becoming erect-sinuous; auricles prominent, weakly to strongly elongate, 1.5–4 mm; seeds olive-brown or brownish-black to black, unspotted or with minute black spots.
............ ............ ............ ............ ............ ............ ........41 Largest leaf blades broadly obtuse to rounded at apex, upper surface glossy in life; peduncle stout, initially prostrate and coiled, elongating to become erect-sinuous prior to dehiscence; calyx green; auricles triangular-trapezoidal, weakly elongate to 1.5(2) mm; capsule finely spotted to heavily blotched with dark purple; seeds 1.5–2.5 × 0.9–1.5 mm, olive-brown, brownish-black to black, often with minute raised spots; [in moist soils of thickets, grassy banks and meadows, forest edges, and lawns across much of e. N. Am. e. of the Great Plains, rare on the Atlantic Coastal Plain]
............ ............ ............ ............ ............ ............ ........41 Largest leaf blades (abruptly) acute at apex, upper surface not glossy in life; peduncle slender, declined or ascending from the beginning, gently curved; calyx purple-pigmented (rarely green); auricles narrowly linear, strongly elongate, (1.5)2–4 mm; capsule finely purple-spotted at least at base; seeds 1.3–1.8 × 0.9–1.2 mm, dark brown with white raphe, unspotted; [in frequently flooded sandy soils along streams and rivers on the s. Atlantic Coastal Plain]
............ ............ ............ ............ ............ ............ ......40 Peduncle initially prostrate, arching upward prior to capsule dehiscence; auricles short and rounded or truncate; seeds yellow- to orange-brown or purple-black and unspotted, or light red-brown with dark brown or red-brown blotches.
............ ............ ............ ............ ............ ............ ..........42 Largest leaf blades ovate-triangular, deltate-triangular or deltate-reniform, shallowly cordate to subcordate at base, abruptly acute to right-angled at apex; seeds 1.4-2.0 mm long, yellow- to orange-brown or purple-black, unspotted; [absent on the Gulf Coastal Plain].
............ ............ ............ ............ ............ ............ ............ 43 Largest leaf blades approx. as broad as long, ovate- to deltate-triangular, 37–69 mm wide, broadly acute at apex, shallowly to moderately cordate at base; petioles smooth throughout; lowest sepals linear-lanceolate or lanceolate, acuminate, > 1/2 as long as mature or dehisced cleistogamous capsule; cleistogamous capsules frequently hirtellous; seeds 1.4–2.0 mm, light to medium yellow- or orange-brown (rarely red-brown) [moist to wet soils of swamps, floodplains and riverside meadows over much of e. N. Am. e. of the Great Plains, mostly absent from the c. and s. Atlantic Coastal Plain, not reaching Gulf Coastal Plain]
............ ............ ............ ............ ............ ............ ............ 43 Largest leaf blades much broader than long, broadly deltate-reniform, 34–145 mm wide, abruptly obtuse to acute at apex, truncate to subcordate at base; summit of petioles granular or papillate-puberulent; lowest sepals broadly acute, mostly < 1/2 as long as mature or dehisced cleistogamous capsule; cleistogamous capsules glabrous; seeds purple-black; [dry sandy soils in upland forests of NY, PA, VT]
............ ............ ............ ............ ............ ............ ..........42 Largest leaf blades ovate to suborbicular, deeply cordate at base, broadly obtuse to rounded at apex; seeds 1.7-2.3 mm long, light red-brown with prominent dark brown or red-brown streaks or blotches; [Gulf Coastal Plain].
............ ............ ............ ............ ............ ............ ............ ..44 Petioles and leaf blades widely spreading; seeds broadly obovoid; [e. Gulf Coast region in FL, s. AL and s. GA].