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Key to Scrophulariaceae

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1 Plant a shrub; inflorescence a terminal thyrse
1 Plant an herb; inflorescence either a raceme, a spike, or a diffuse panicle (Scrophularia and Verbascum), or the flowers solitary from basal axils (Limosella).
..2 Plants aquatic, < 6 cm tall; flowers solitary on pedicels from basal axils
..2 Plants terrestrial, > 60 cm tall; inflorescence a raceme, a spike, or a diffuse panicle.
....3 Corolla cylindric, purplish; fertile stamens 4
....3 Corolla rotate, yellowish; fertile stamens 5

Key C4: rooted aquatics with basal and simple, linear leaves

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1 Leaves thread-like or quill-like, about as thick as wide.
..2 Plants bulbous at base, and with the leaf bases expanded and containing sporangia; plant tufted or with very short rhizomes; [Lycophytes]
..2 Plants either somewhat bulbous or not at the base, the leaf-bases not containing sporangia; plant rhizomatous; [Pteridophytes, Eudicots, Monocots].
....3 New leaves unfurling with circinate vernation (a fiddlehead); plants reproducing by spores, from sporocarps on short stalks from the rhizome; [Pteridophytes]
....3 New leaves lacking circinate vernation; plants reproducing by flowers and seeds.
......4 Perianth differentiated, with either 3 sepals and 3 petals or 5 sepals and 5 petals; stamens either 7-many or stamens 4.
........5 Sepals 3; petals 3; stamens 7-many; [Monocots]
........5 Sepals 5; petals 5; stamens 4; [Eudicots]
......4 Perianth undifferentiated, with 0, 3, or 6 tepals; stamens 1, 2, or 3; [Monocots].
..........6 Gynoecium of 2 or more pistils, each pistil with 1 carpel and with 1 stigma
..........6 Gynoecium of 1 pistil, each pistil with (2-) 3 carpels and (2-) 3 stigmas.
............ 7 Fruit an achene; perianth absent
............ 7 Fruit a capsule; perianth of 6 tepals
1 Leaves ribbon-like or strap-like, distinctly flattened (sometimes only near the tip of the leaf).
............ ..8 Subterranean portions of plant bearing bladder-traps; flowers yellow or purple, bilaterally symmetrical
............ ..8 Subterranean portions of plant lacking bladder traps; flowers white, green, gray, radially symmetrical (except bilaterally symmetrical in Glossostigma in PHRYMACEAE).
............ ....9 Leaves broadened towards the tip; [Monocots, Eudicots].
............ ......10 Flowers 3-merous; [Monocots]
............ ......10 Flowers 4- or 5-merous; [Eudicots].
............ ........11 Leaves phyllodial, with obvious cross-partitions (septa); leaves alternate
............ ........11 Leaves not phyllodial; leaves opposite
............ ....9 Leaves parallel-margined or tapering towards the apex over much of their length; [Monocots].
............ ..........12 Leaves tapering towards the apex over much of their length; plant either tufted and not rhizomatous, or short rhizomatous;.
............ ............ 13 Plant tufted, not rhizomatous; leaves spreading radially; inflorescence a tightly button-like head of very numerous small flowers, white, gray, tan, yellowish, or blackish; roots thickened, septate (not requiring magnification), unbranched
............ ............ 13 Planted short-rhizomatous; leaves distichous, equitant; inflorescence either a subglobular, ovoid, or cylindrical head, of spirally imbricate scales, or a diffuse corymb; roots not thickened, not septate, branched.
............ ............ ..14 Inflorescence a diffuse corymb; rhizomes and roots bright red
............ ............ ..14 Inflorescence a subglobular, ovoid, or cylindrical head of spirally imbricate scales subtending individual flowers; rhizomes and roots not bright red
............ ..........12 Leaves parallel-margined; plant usually rhizomatous.
............ ............ ....15 Plants of marine habitats, growing submersed in salt water; [FL and the Gulf Coast of AL, MS, and LA]
............ ............ ....15 Plants of freshwater or slightly to somewhat brackish habitats; [collectively widespread].
............ ............ ......16 Leaves lacking any midvein; flowers and fruits in globose heads
............ ............ ......16 Leaves with a midvein; flowers and fruits solitary or in diffuse inflorescences.
............ ............ ........17 Leaves with a distinct, broad lacunar band along the midvein
............ ............ ........17 Leaves lacking a distinct lacunar band along the midvein.

Key N2: herbaceous dicots with mainly basal, simple leaves

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1 Inflorescence an involucrate head subtended by phyllaries, the heads solitary or many and variously arrayed in secondary inflorescences, the ovary inferior, the corolla connate and tubular at least basally, the calyx absent, the stamens 5, the fruit a cypsela
1 Inflorescence, flower, and fruit structure various, but not with the combination of features as above (sometimes the flowers in a head subtended by bracts, e.g. Eryngium in APIACEAE, but then with other features differing, such as stamens 4, or green calyx present, or petals separate, or fruit a schizocarp of mericarps, etc.).
..2 Basal leaves 2-lobed, pinnately lobed, or palmately lobed (not considering cordate, hastate, or auriculate leaf bases as “lobed”).
....3 Basal leaves 2-lobed, hinged between the lobes, each lobe with stiff, marginal, eyelash-like bristles; [Coastal Plain of NC and SC, rarely planted and weakly naturalized elsewhere]
....3 Basal leaves 3-many-lobed, palmately or pinnately; [collectively widespread].
......4 Leaf lobing pinnate.
........5 Gynoecium of separate pistils (each with a single carpel); fruit an aggregate
........5 Gynoecium of a single pistil (with 2, rarely more, carpels); fruit simple.
..........6 Stamens many; sepals 2, petals 4; fresh plants with yellow, orange, or white milky juice
..........6 Stamens 4, 5, or 6; sepals 4 or 5; petals 4 or 5.
............ 7 Petals 4, distinct; stamens 6
............ 7 Petals 5, fused; stamens 2, 4, or 5.
............ ..8 Corolla radially symmetrical; stamens 5
............ ..8 Corolla 2-lipped, bilaterally symmetrical or asymmetrical; stamens 2 or 4.
............ ....9 Corolla lobes not twisted, the flower bilaterally symmetrical; stamens 2
............ ....9 Corolla lobes twisted so as to make the flower asymmetrical; stamens 4
......4 Leaf lobing palmate.
............ ......10 Petiole attachment peltate.
............ ........11 Leaves < 10 cm in diameter
............ ........11 Leaves > 15 cm in diameter
............ ......10 Petiole attachment marginal.
............ ..........12 Ovary inferior.
............ ............ 13 Petals 4; stamens 8; fruit a capsule
............ ............ 13 Petals 5; stamens 5; fruit a schizocarp of 2 mericarps.
............ ............ ..14 Fruit tuberculate; leaves 3-lobed
............ ............ ..14 Fruit smooth; leaves with 5 or more lobes
............ ..........12 Ovary superior, or half-inferior by fusion of a hypanthium a part of the way up the ovary wall.
............ ............ ....15 Gynoecium of separate pistils (each with a single carpel); fruit an aggregate.
............ ............ ......16 Perianth of 5 green sepals and 5 colored petals.
............ ............ ........17 Carpels partly fused, arrayed in a ring of 10-20
............ ............ ........17 Carpels separate, spiral, many
............ ............ ......16 Perianth of a single whorl of 3-12 petaloid sepals (the petals absent or small and rudimentary).
............ ............ ..........18 Leaves 2, the single flower terminal and associated with the upper leaf; fruit an aggregate of berries
............ ............ ..........18 Leaves normally > 2, flowers not as above; fruit an aggregate of achenes, utricles, or follicles
............ ............ ....15 Gynoecium of a single pistil (with 1-5 carpels); fruit simple.
............ ............ ............ 19 Hypanthium present, partially fused or not fused to the pistil; ovary partially inferior to superior
............ ............ ............ 19 Hypanthium absent; ovary superior.
............ ............ ............ ..20 Petals connate at base; leaves sharply toothed
............ ............ ............ ..20 Petals distinct; leaves with rounded lobes or teeth.
............ ............ ............ ....21 Corolla radially symmetrical; petals 8-12; capsule fusiform, narrowed to both ends, > 5× as long as wide
............ ............ ............ ....21 Corolla bilaterally symmetrical; petals 5; capsule ovoid, < 2× as long as wide
..2 Basal leaves not lobed, at most serrate or crenate (and sometimes also cordate, hastate, auriculate, or peltate at the base).
............ ............ ............ ......22 Basal leaves petiolate, the blade with a cordate, hastate, auriculate, or peltate base.
............ ............ ............ ..........24 Inflorescence a terminal and/or axillary raceme, panicle, or cyme of many small flowers; fruit an achene; perianth uniseriate, of 0, 4-5, or 6 tepals.
............ ............ ............ ............ 25 Flowers unisexual; staminate flowers 4-5 tepals, pistillate flowers lacking a perianth
............ ............ ............ ..........24 Inflorescence either a terminal spike, or a 1-7-flowered terminal cyme, or of a solitary axillary or terminal flower; fruit various; perianth biseriate (of differentiated sepals and petals (except uniseriate, of 3 fused sepals in ARISTOLOCHIACEAE).
............ ............ ............ ............ ..26 Flowers bilaterally symmetrical; inflorescence a terminal spike (with > 20 flowers); petals 4, usually scarious, transparent; sepals 4, green; stamens 4
............ ............ ............ ............ ..26 Flowers radially symmetrical; inflorescence either of a solitary flower or of a 1-7-flowered terminal cyme; petals 5, 8-12, or 0; sepals 5 (green), 3 (brown), or 5-9 (yellow); stamens 5, 12, or many.
............ ............ ............ ............ ....27 Gynoecium of separate pistils (each with a single carpel); fruit an aggregate of achenes or follicles; flowers bright yellow, either of 5-9 distinct petaloid sepals, or of 8-12 distinct petals subtended by 3-4 green distinct sepals
............ ............ ............ ............ ....27 Gynoecium either of a single pistil with 6 carpels or of a single pistil with 4 carpels or of 2 nearly separate carpels; fruit a simple capsule (or deeply 2-lobed); flowers white, brown, or greenish, either of 5 fused or distinct white petals and 5 fused or distinct green sepals, or of 3 fused brown or greenish petaloid sepals.
............ ............ ............ ............ ......28 Flowers brown or green, of 3 fused brown or greenish petaloid sepals (and 0 petals); carpels 6; stamens 12; leaves 4-10 cm wide
............ ............ ............ ............ ......28 Flowers white, of 5 white or whitish petals and 5 green sepals; carpels 2; stamens 5; leaves 1-12 (-15) cm wide
............ ............ ............ ............ ........29 Petals separate; sepals separate; plant glabrous
............ ............ ............ ............ ........29 Petals fused; sepals fused; carpels 2; plant pubescent.
............ ............ ............ ........23 Leaf margins crenate, serrate, or incised.
............ ............ ............ ............ ............ 31 Gynoecium of separate pistils (each with a single carpel); fruit an aggregate; perianth of 5 green sepals and 5 colored petals.
............ ............ ............ ............ ............ ..32 Carpels 10-20, partly fused, arrayed in a ring; petals white, pink, red, or purplish
............ ............ ............ ............ ............ ..32 Carpels many, separate, spiral; petals yellow or white.
............ ............ ............ ............ ............ ....33 Flowers lacking a hypanthium; fruit an aggregate of achenes or aggregate of follicles
............ ............ ............ ............ ............ ....33 Flowers with a hypanthium; fruit an aggregate of drupelets or aggregate of achenes
............ ............ ............ ............ ............ 31 Gynoecium of a single pistil (with 1-5 carpels); fruit simple.
............ ............ ............ ............ ............ ......34 Flowers bilaterally symmetrical; inflorescence of a solitary flower; fruit a 3-locular capsule
............ ............ ............ ............ ............ ......34 Flowers radially symmetrical; inflorescence an umbel (or composite of umbelliform units, or a terminal panicle.
............ ............ ............ ............ ............ ........35 Ovary superior; inflorescence a terminal panicle or terminal raceme.
............ ............ ............ ............ ............ ..........36 Inflorescence a terminal raceme; perianth of 4 green sepals and 4 white petals; fruit a silique/silicle; fresh foliage in spring and summer with a strong garlic odor; larger leaves < 10 cm in diameter
............ ............ ............ ............ ............ ..........36 Inflorescence a terminal panicle; perianth of 6 cream-colored petaloid sepals; fruit a winged achene; fresh foliage lacking a garlic odor; larger leaves typically > 30 cm in diameter
............ ............ ............ ............ ............ ........35 Ovary inferior; inflorescence an umbel (or a composite of umbellate units); fruit a schizocarp of mericarps.
............ ............ ............ ............ ............ ............ 37 Petiole attachment marginal (the blade cleft to the petiole.
............ ............ ............ ............ ............ ............ ..38 Leaf blades longer than wide, sharply V-cleft at the base and otherwise shallowly denticulate
............ ............ ............ ............ ............ ............ ..38 Leaf blades as broad or broader than long, cleft at the base and also irregularly serrate or crenate around the margin
............ ............ ............ ......22 Basal leaves petiolate or not, with a truncate, rounded, or cuneate leaf base.
............ ............ ............ ............ ............ ............ ....39 Leaves tubular, with a sutured ventral flange, erect or reclining, adapted as a pitfall for insects (flat, phyllodial leaves sometimes present as well, common in the winter in some species, such as S. oreophila)
............ ............ ............ ............ ............ ............ ......40 Stem leaves opposite; perianth 5-merous, at least the corolla bilaterally symmetrical (barely so in VALERIANACEAE), or the parts curved so as to be asymmetrical (Pedicularis in OROBANCHACEAE); stamens 2, 3, or 4.
............ ............ ............ ............ ............ ............ ..........42 Corolla narrowly tubular, the five lobes flaring at nearly 90 degrees and nearly radially symmetrical
............ ............ ............ ............ ............ ............ ..........42 Corolla distinctly 2-lipped (with prominently large upper and lower corolla lobes) or hooded (the upper lip hood-like), distinctly bilaterally symmetrical, or the lobes twisted so as to make the corolla asymmetrical.
............ ............ ............ ............ ............ ............ ............ 43 Corolla yellow, the upper lip often slightly to strongly maroon, hooded but the corolla lobes twisted so as to make the flower asymmetrical
............ ............ ............ ............ ............ ............ ............ 43 Corolla white, lavender, or blue, 2-lipped and bilaterally symmetrical.
............ ............ ............ ............ ............ ............ ......40 Stem leaves alternate; perianth radially symmetrical (less commonly bilaterally symmetrical); stamens 5, 6-8, 9, 10 (rarely 4).
............ ............ ............ ............ ............ ............ ............ ..........48 Pistils many, each with a single carpel; fruit an aggregate of achenes
............ ............ ............ ............ ............ ............ ............ ..........48 Pistil 1, with 1-5 carpels; fruit simple (a legume, silique/silicle, capsule, utricle, or schizocarp of 4 nutlets).
............ ............ ............ ............ ............ ............ ............ ............ 49 Corolla bilaterally symmetrical (barely so in Limosella in SCROPHULARIACEAE); stamens 2, 4, 6, 8, or 10.
............ ............ ............ ............ ............ ............ ............ ............ ............ ..56 Perianth of green sepals and more brightly colored petals; stamens 5 or 10; carpels 2, 3, 4, or 5.
............ ............ ............ ............ ............ ............ ............ ............ ............ ....57 Leaves covered with sticky, gland-tipped hairs (often red), as flypaper traps for insects
............ ............ ............ ............ ............ ............ ............ ............ ............ ............ ......64 Inflorescence a somewhat to very diffuse panicle, with 3 or more orders of branching, not giving at all the impression that the overall inflorescence is made of racemose units.
............ ............ ............ ............ ............ ............ ............ ............ ............ ............ ........65 Leaves serrate or crenate; stamens 10; [plants of various habitats, especially rock outcrops and bottomland forests and streambanks, never in tidal marshes]
............ ............ ............ ............ ............ ............ ............ ............ ............ ............ ......64 Inflorescence either a single terminal raceme (sometimes spike-like), or of 1 to several terminal and axillary racemes (these sometimes combined into a diffuse panicle, but one whose structure is clearly made up of many racemes).
............ ............ ............ ............ ............ ............ ............ ............ ............ ............ ..........66 Inflorescence of 1-several terminal and axillary racemes, the plant typically well-branched, especially from the base; stamens 5
............ ............ ............ ............ ............ ............ ............ ............ ............ ............ ..........66 Inflorescence of a single, terminal raceme, the plant unbranched; stamens 10 (or 5, with 5 staminodes)
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