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Key to Ilex

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1 Leaves coriaceous, evergreen.
..2 Leaves with a well-developed apical spine (and usually also marginal spines) 2-6 mm long.
....3 Flowers in 1-few-flowered axillary cymes, on growth of the same year; upper leaf surfaces somewhat shiny to matte; marginal leaf spines (when present) in the plane of the leaf or mostly declined < 30 degrees from that plane; [native trees of a wide variety of habitats]; [section Cassinoides]
......4 Leaves somewhat yellowish green above, 1.5-5.0 cm long, 1.0-2.5 cm wide, strongly revolute; fruits orange to red; [of dry sandy habitats of FL]
......4 Leaves dark green above, 3-12 cm long, 2.0-5.5 cm wide, flat to slightly revolute; fruits red (rarely yellow); [of moist to fairly dry habitats, widespread in our area]
....3 Flowers in axillary clusters, on growth of the previous year; upper leaf surfaces very shiny; marginal leaf spines (when present) often strongly oriented below or above the plane of the leaf; [exotic shrubs or small trees usually in suburban or urban areas]; [section Ilex]
........5 Lateral leaf margin spines > 5 per leaf side (rarely 0, in some cultivars)
........5 Lateral leaf margin spines usually 0, 2, or 4 per leaf side
..2 Leaves with margins either entire, crenate, serrate, or with marginal spinose prickles < 2 mm long (the apex sometimes mucronate, but not stiff and spinose).
..........6 Leaves crenate from base to apex, 0.5-4.5 cm long; calyx and corolla 4-lobed.
............ 7 Fruits black; leaf undersurface with punctate glands; leaf apex obtuse to broadly acute, tipped with a small but obvious sharp tooth (best seen at 10× or greater); [exotic shrub, rarely naturalized, especially in suburban areas]; [section Paltoria]
............ 7 Fruits red or yellow; leaf undersurface lacking punctate glands; leaf apex notched (retuse), with a very small mucro in the notch (best seen at 10× or greater); [native shrub of the Coastal Plain, sometimes planted and naturalized elsewhere]; [section Repandae]
..........6 Leaves entire, crenate (if so, only beyond the midpoint), serrate, or with marginal spinose prickles, 2-10 cm long; calyx and corolla 4-lobed or 5-9-lobed.
............ ..8 Fruits black; calyx and corolla 5-9-lobed; leaves crenate near the tip or with a few marginal spinose prickles, or entire, with dark punctate dots beneath; [section Glabra].
............ ....9 Leaves 1.5-3× as long as wide, generally about 2-3 cm wide, broadly elliptic and widest near the midpoint of the blade; leaf margins either entire or with a few, irregularly spaced, marginal spinose prickles that diverge from the margin
............ ....9 Leaves 3-4× as long as wide, generally about 1 cm wide (almost never > 2 cm wide), obovate and widest beyond the midpoint of the blade; leaf margins crenate in the apical 1/2 to 1/3 (almost never entire)
............ ..8 Fruits red, yellow, or black; calyx and corolla 4-lobed; leaves entire (or with a few spinose serrations), lacking dark punctate dots beneath.
............ ......10 Fruits black or purple when ripe; peduncle absent or very short (< 3 mm long), the flowers (later fruits) thus clustered in the leaf axils; [of s. FL only]; [section undetermined]
............ ......10 Fruits red or yellow when ripe; peduncle elongate; [more widespread in our area]; [section Cassinoides].
............ ........11 Leaves oblanceolate, oblong, or elliptic, 3-12 cm long, (8-) 15-40 mm wide, 2-4× as long as wide; petioles (3-) 5-15 mm long; leaf apex acute, obtuse, or rounded; branchlets strongly ascending, most of them forming an angle of < 45 degrees to the branch
............ ........11 Leaves lanceolate to narrowly oblong, 2-4 cm long, 3-8 mm wide, 3-7× as long as wide; petioles 1-3 (-5) mm long; leaf apex acute to acuminate; branchlets ascending to spreading, most of them forming angles greater than 45 degrees to the branch, and often at right angles
............ ..........12 Leaves entire, or nearly so; [of moist to wet sites, from WV northward]; ["clade IV"]
............ ..........12 Leaves toothed; [collectively widespread in our area].
............ ............ 13 Leaves oblanceolate or obovate, broadest above the middle (to near the middle), 8-30 (-45) mm wide, narrowly cuneate basally, mostly 2-3× as long as wide; [section Prinoides].
............ ............ ..14 Pedicels of fruits and pistillate flowers 2-6 mm long; pedicels of staminate flowers (2-) 4-8 (-16) mm long; leaves mostly gray green, often revolute, especially toward the base; pubescence of the lower leaf surface tomentose, primarily on or near the midrib; leaf margins rarely ciliate.
............ ............ ....15 Leaves 2-4.8 cm long, 0.6-1.5 cm wide; fruits 4-5 mm in diameter; sepals usually ciliate; [apparently endemic to the Suwanee River drainage of sc. GA and e. Panhandle FL]
............ ............ ....15 Leaves 4.5-8.5 (-10) cm long, 1.5-3 cm wide; fruits (4-) 5-8 (-9) mm in diameter; sepals not ciliate; [widespread in our area, in the Coastal Plain, Piedmont, and rarely Mountains of our area]
............ ............ ..14 Pedicels of fruits and pistillate flowers (5.5-) 10-30 mm long; pedicels of staminate flowers (10-) 15-25 mm long; leaves rarely revolute; pubescence of the lower leaf surface strigose, distributed on the surface; leaf margins often ciliate.
............ ............ ......16 Upper leaf surface with trichomes throughout; sepals ciliate; leaf blades entire to shallowly crenate
............ ............ ......16 Upper leaf surface glabrous, or with trichomes confined to the veins or their vicinity; sepals eciliate; leaf blades crenate to distinctly serrate
............ ............ 13 Leaves elliptic or ovate, broadest near the middle, (10-) 20-55 mm wide, rounded to broadly cuneate basally, mostly 1-2.5× as long as wide.
............ ............ ........17 Tertiary veins on undersurface of leaf blades reticulate, defining areoles between the raised primary and secondary veins; fruit surface dull; fruiting pedicels 6-14 mm long (averaging about 10 mm); [of blackwater floodplains and clay-based Carolina bays of the Coastal Plain]; ["clade IV"]
............ ............ ........17 Tertiary veins on undersurface of leaf blades obscure, not defining obvious areoles between the raised primary and secondary veins; fruit surface shiny; fruiting pedicels either (8-) 10-30 mm long or 2-9 mm long (averaging either < 6 mm or > 15 mm long); [collectively of various habitats, widespread in our area].
............ ............ ..........18 Fruiting pedicels (8-) 10-30 mm long; fruit (7-) 8-12 mm in diameter, bright cherry-red; petiole with a deeply U- to V-shaped channel on its upper side (made by the decurrent leaf edges), with dark ascending trichomes in the channel; bark of 2-3 year old twigs usually light tan; [of bogs, seepages, and very moist forests of the Mountains]; [section Prinoides]
............ ............ ..........18 Fruiting pedicels 2-9 mm long; fruit 5-9 (-12) mm in diameter, duller red to orange; petiole with U-shaped channel on its upper side, with white appressed trichomes in the channel, or the petiole nearly terete; bark of 2-3 year old twigs usually brown, gray, or purplish; [collectively of various habitats, widespread in our area].
............ ............ ............ 19 Nutlets (5-) 6-8 per fruit, smooth on the (curved) back; staminate flower clusters on peduncles 2-6 mm long; pistillate flowers with entire corolla lobes; flowers mostly in axils of leaves on normal shoots; petiole nearly terete in cross-section (or very shallowly channeled on the upper surface); [section Prinos].
............ ............ ............ ..20 Sepals glabrous (in flower or fruit), acute; leaves lighter green, slightly if at all rugose
............ ............ ............ ..20 Sepals ciliate (in flower or fruit), obtuse; leaves darker green, often moderately rugose
............ ............ ............ 19 Nutlets 4-5 per fruit, with striate ridges on the (curved) back; staminate flower clusters sessile or very short-peduncled (0-2 mm long); pistillate flowers with ciliate corolla lobes; flowers mostly in axils of leaves on lateral short-shoots; petiole with U-shaped channel on its upper side, with white appressed trichomes in the channel; [section Prinoides].
............ ............ ............ ....21 Leaves 2-9 (-10.5) cm long, elliptic to broadly ovate, often nearly round, the base usually rounded, the apex abruptly to gradually acuminate, the marginal teeth usually inconspicuous; petioles of mature leaves usually < 1 cm long; fruits 5-9 mm in diameter; plant a shrub to 6 m tall; [of the Coastal Plain, Piedmont, and Mountains]
............ ............ ............ ....21 Leaves 6-16 cm long (the largest, at least, > 8 cm long), narrowly to broadly ovate, the base usually cuneate, the apex long acuminate to attenuate, the marginal teeth rather coarse; petioles of mature leaves usually > 1 cm long; fruits 9-12 mm in diameter; plant a shrub or small tree to 10 m tall; [of the Mountains and upper Piedmont]

Key G4: shrubs and subshrubs with alternate, simple, unlobed, entire leaves

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1 Leaves evergreen. {add to 1a: Scaevola in GOODENIACEAE, Morella (inodora) in MYRICACEAE, Ternstroemia in PENTAPHYLACACEAE, Myrsine in PRIMULACEAE, Pyracantha in ROSACEAE, Dodonaea in SAPINDACEAE, Cestrum in SOLANACEAE, Thymelaea in THYMELAEACEAE, Conocarpus in COMBRETACEAE}
..2 Leaves 1-7 mm long, either acicular and spreading or ovate and appressed to the stems
..2 Leaves > 10 mm long.
....3 Leaves linear, > 15× as long as wide; [Monocots]
....3 Leaves broader, < 15× as long as wide; [Eudicots, Basal Angiosperms, or Monocots].
......4 Plant a creeping subshrub, < 1 dm tall
......4 Plant not creeping, > 3 dm tall.
........5 Inflorescence solitary (Illicium in ILLICIACEAE) or variously branched, spicate, racemose, or fascicled, not an involucrate head.
..........6 Carpels separate; fruit an aggregate; fresh foliage strongly fragrant; [Basal Angiosperms]
............ 7 Fruit an aggregate of woody follicles arranged in a whorl
............ 7 Fruit an aggregate of red to blackish berries, 4-6 mm long, each on a long stipe, giving the aggregate almost the appearance of an umbel
..........6 Carpels fused; fruit a berry, drupe, acorn (nut), capsule, or legume; fresh foliage not strongly fragrant; [Eudicots, Monocots, and Basal Angiosperms].
............ ..8 Ovary with 3 carpels; fruit a berry; “leaves” actually cladodes; [Monocots]
............ ..8 Ovary with 1, 2, 3, 4, or 5 carpels; fruit a berry, drupe, capsule, legume, or nut; leaves actually leaves; [Eudicots and Basal Angiosperms].
............ ....9 Leaves largely covered with silver and/or bronze lepidote scales and/or dense stellate hairs below (visible at 10× or higher magnification), giving the lower leaf surface a slightly shiny to almost metallic appearance. {add Lyonia ferruginea and L. fruticosa in ERICACEAE; add Loropetalum in HAMAMELIDACEAE}
............ ......10 Petals present, conspicuous, connate, white, the corolla rotate; fruit a berry with several seeds; fresh foliage with a strong, tar-like odor
............ ......10 Petals absent or inconspicuous, greenish and separate if present (note that the calyx is petaloid and white or yellowish in Elaeagnus of ELAEAGNACEAE); fruit a dry capsule with 3 seeds, or a drupe with a single seed; fresh foliage lacking a strong odor.
............ ........11 Perianth 4-merous; petals absent; petaloid sepals white to cream, fused and salverform; carpel 1; fruit a fleshy, red drupe, with a single seed
............ ........11 Perianth 5-merous; petals green and separate, or absent; sepals greenish, separate; carpels 3; fruit a 3-valved capsule with 3 seeds
............ ....9 Leaves with various vestiture (or glabrous), but not as above.
............ ..........12 Flowers in spikes, these solitary opposite leaves or in axillary umbels
............ ..........12 Flowers in other types of inflorescences, not spikes.
............ ............ 13 Leaves 1-foliolate on the upper stems, sometimes 3-foliolate below, or all reduced to phyllodial spines; flowers papilionaceous, bright yellow; fruit a legume; stems bright green
............ ............ 13 Leaves simple throughout; flowers either small, inconspicuous, tannish, borne in catkins (Quercus), or larger and urceolate, or with almost separate and spreading petals, white to pink or reddish-orange, in various terminal or axillary, branched inflorescences; fruit either a nut in a cupule (an acorn), or a (3-) 5-valved capsule, or a spherical berry or drupe; stems generally brown or tan (sometimes green).
............ ............ ..14 Flowers small, inconspicuous, tannish, borne in catkins; fruit a nut in a cupule (an acorn)
............ ............ ..14 Flowers white to pink or reddish-orange, either urceolate or tubular or with separate and spreading petals, in various terminal or axillary inflorescences, or solitary; fruit either a (3-) 5-valved capsule, or a spherical berry with 10+ seeds, or a 1-8 seeded dry or fleshy drupe.
............ ............ ....15 Flowers white to pink or reddish-orange, rotate or urceolate (the petals united at least basally), in various terminal or axillary inflorescences or solitary; fruit either a 2-5 valved capsule or a spherical berry with 10+ seeds.
............ ............ ......16 Flowers reddish-orange; corollas bearing post-staminal hairs (appearing as tufts of hair opposite each anther)
............ ............ ......16 Flowers white to pink; corollas without obvious tufts of hair opposite each anther.
............ ............ ........17 Leaves 1 per node or also paired (on one side of the stem) at some nodes (the leaves then uneven in size); inflorescences leaf-opposed; fruit a berry.
............ ............ ........17 Leaves 1 per node; inflorescences terminal or axillary, never leaf-opposed; fruit a valved capsule.
............ ............ ..........18 Seeds without fleshy aril or attachment; ovaries 5-10 carpellate; flowers urceolate; leaves serrate or entire
............ ............ ..........18 Seeds with fleshy aril or attachment; ovaries 2-4 carpellate; flowers rotate; leaves entire or very remotely serrulate.
............ ............ ............ 19 Flowers few, not showy, green to greenish-white, inflorescence not densely arranged (flowers also sometimes solitary); capsules not beaked; leaves not revolute or undulate
............ ............ ............ 19 Flowers numerous, showy, white; inflorescence densely arranged; capsules with short beak; leaf margins revolute or slightly undulate
............ ............ ....15 Flowers white, petals spreading, separate even at the base, in axillary fascicles or racemes; fruit either a fleshy drupe with 4-8 pyrenes, or a dry single-seeded drupe.
............ ............ ............ ..20 Shrub not rhizomatous; fruit either a fleshy or dry drupe, < 1 cm long.
1 Leaves deciduous. {add: Ditrysinia in EUPHORBIACEAE, Glochidion in PHYLLANTHACEAE, Phyllanthopsis in PHYLLANTHACEAE, Nierembergia in SOLANACEAE, Edgeworthia in THYMELAEACEAE, Ipomoea (I. carnea) in CONVOLVULACEAE; Swida (S. alternifolia) in CORNACEAE}
............ ............ ............ ......22 Inflorescence branched, spicate, a catkin, or consisting of a solitary flower or axillary clusters or whorls, not an involucrate head.
............ ............ ............ ............ ..26 Fruit a globose, spinose capsule bearing a longitudinal ridge across 1 or both faces, thus essentially resembling a spikey ball; perianth (4-)5-merous, consisting of 3 upper connate petaloid claws and two lower sessile petals; fresh plants without a strange musky odor; [Eudicots]
............ ............ ............ ............ ..26 Fruit an oblong berry, greenish-yellow when ripe, not bearing small spines; perianth 3-4-merous, without connate petaloid claws; fresh plants fragrant with a strange, musky odor; [Basal Angiosperms]
............ ............ ............ ............ 25 Inflorescence of 2 or more flowers; perianth 3-5-merous; fresh plants not musky-fragrant; fruits various, not as above.
............ ............ ............ ............ ....27 Flowers 3-merous; fruit fleshy, red or greenish-yellow at maturity; ovary superior; [Basal Angiosperms or Eudicots].
............ ............ ............ ............ ......28 Leaves elliptic or narrowly elliptic, broadest near the middle; fresh plants strongly fragrant with a citrus-like aroma; stems unarmed; fruit a drupe, with a single seed
............ ............ ............ ............ ......28 Leaves obovate or oblanceolate, broadest near the apex; stems armed with nodal spines; fresh plants not fragrant; fruit a berry, with several seeds.
............ ............ ............ ............ ....27 Flowers 4-5-merous; fruit fleshy or dry, black, blue, brown, tan, or red at maturity; ovary superior or inferior; [Eudicots].
............ ............ ............ ............ ........29 Fruit either a drupe or berry (indehiscent, and variously fleshy or dry) or a dry 3-valved capsule with 1 seed; inflorescence axillary (solitary, clusters, fascicles, or racemes), or in a terminal raceme (Pyrularia in SANTALACEAE).
............ ............ ............ ............ ..........30 Leaves largely covered with silver and/or bronze shiny lepidote scales below, giving the lower leaf surface an almost metallic appearance
............ ............ ............ ............ ............ ..32 Fruit an elongate drupe (definitely longer than thick), with 1 seed.
............ ............ ............ ............ ............ ........35 Fruit a red or orange berry, 8-20 mm long; leaves usually on spur-shoots; [salty coastal areas, or exotics of disturbed situations]
............ ............ ............ ............ ............ ........35 Fruit a yellowish-green drupe, 12-15 mm long; leaves on main stems; [rich forests, mainly inland]
............ ............ ............ ............ ............ ..........36 Inflorescence a narrowly cylindrical raceme, clustered several to many at the tip of the previous year’s wood and below the current season’s growth; fruit < 3 mm in diameter
............ ............ ............ ............ ............ ............ ..38 Fruit dark red to black, the pedicel < 10 mm long; leaf venation very neatly pinnate, with the secondary veins nearly straight and parallel to one another
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