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euphrasia, eyebright

broom-rape family

Habit Herbs, annual [perennial]; hemiparasitic. Herbs, rarely subshrubs or shrubs, annual, biennial, or perennial, sometimes fleshy, hemiparasitic or holoparasitic (without chlorophyll) [autotrophic].
Stems

erect, not fleshy, retrorsely hairy.

subterranean or aerial;

aerial stems prostrate to decumbent, ascending, or erect [viny].

Leaves

cauline, opposite;

petiole absent or nearly so;

blade sometimes fleshy, not leathery, margins crenate, serrate, dentate, or incised.

deciduous, cauline or basal and cauline, rarely basal only or absent, sometimes scales, opposite, alternate, whorled, or spiral, simple;

stipules absent;

petiole present or absent;

blade usually not fleshy or leathery, rarely fleshy, leathery, or chartaceous, margins entire, toothed, or lobed.

Inflorescences

terminal, subcapitate to diffuse spikes, flowers solitary in each axil;

bracts present, subopposite or irregularly alternate.

terminal and/or axillary, racemes, panicles, spikes, corymbs, or flowers 1 or 2.

Pedicels

present or absent;

bracteoles absent.

Flowers

sepals 4, calyx bilaterally symmetric, not flattened laterally, tubular, not accrescent in fruit, lobes triangular;

petals 5, corolla white or cream to purple, lilac, violet, brownish purple, or yellow with violet veins (often without violet veins in E. subarctica) and yellow spot in throat and on abaxial lip, adaxial lip sometimes lilac, purple, or yellow, contrasting with rest of corolla, bilabiate, funnelform, abaxial lobes 3, emarginate, adaxial 2, adaxial lip cucullate;

stamens 4, didynamous, filaments glabrous, anther mucros unequal;

staminode 0;

ovary 2-locular, placentation axile;

stigma capitate.

bisexual, perianth and androecium hypogynous;

sepals (0 or)2–5(–8), connate, calyx radially or bilaterally symmetric;

petals [4 or]5, connate, corolla bilaterally symmetric, bilabiate or strongly bilabiate, tubular, funnelform, campanulate, salverform, or club-shaped, sometimes cylindric, subrotate, or curved;

stamens (2 or)4, adnate to corolla tube, didynamous, subequal, or equal, staminodes 0 or 2;

pistil 1, 2[or 3]-carpellate, ovary superior, 1- or 2-locular, placentation axile, sometimes parietal;

ovules anatropous or campylotropous-like (Rhinanthus), unitegmic, tenuinucellate;

style 1;

stigma 1.

Fruits

capsules, dehiscence loculicidal and/or septicidal or indehiscent (Conopholis).

Capsules

dehiscence septicidal, opening in distal 1/2, margins ciliate, sometimes glabrous or short-ciliate (E. salisburgensis).

Seeds

10–18, grayish, fusiform, wings absent.

1–2500(–5000), brown or black, sometimes tan, white, yellow, amber, or gray, ovoid to ellipsoid, reniform, globular, oblong, or angled;

embryo straight, endosperm present.

x

= 11.

Euphrasia

Orobanchaceae

Distribution
from USDA
North America; South America; Eurasia; Africa (Morocco); Atlantic Islands (Azores, Iceland); Pacific Islands (Oceania); Australia
[BONAP county map]
nearly worldwide; especially in warm temperate regions
[BONAP county map]
Discussion

Species ca. 350 (18 in the flora).

Euphrasia is distributed throughout temperate regions of the northern and southern hemispheres, with one transtropical connection across the high mountains of Oceania. All Euphrasia species in the flora area belong to sect. Euphrasia, the largest of 15 sections in the genus (G. Gussarova et al. 2008). The section is noted for its taxonomic complexity. Major factors explaining the complex patterns of variation in sect. Euphrasia include interspecific hybridization, recent radiations, parasitism, polyploidization, and breeding system transitions (T. Karlsson 1974, 1986; P. F. Yeo 1978; G. C. French et al. 2008; R. A. Ennos et al. 2012). All North American species are tetraploids except, probably, E. disjuncta, E. farlowii, E. oakesii, E. randii, and E. subarctica. Of 18 species in the flora, nine are considered endemic. To facilitate identification of the species, the aggregate species concept is sometimes used to show relationships among the so-called more conventional, discrete taxa (Gussarova 2005). Following that concept, four aggregate species are represented in the flora area and their associated species treated here are: E. borealis (F. Townsend) Wettstein agg. (species 3 and 4), E. nemorosa agg. (species 6–10), E. oakesii agg. (species 13–15), and E. minima Jacquin ex de Candolle agg. (species 16 and 17).

The key makes no provision for hybrids, which often have intermediate features or a mixture of character states typical of their putative parental species. For a reliable identification, multiple representative plants should be collected and examined from a population to account for individual variation in key characters. The bracts referred to in the key are proximal floral leaves, which are most representative of the variation along the stem. The teeth of the bracts become progressively more acute up the stem as well as on any one leaf from apex to base. To characterize teeth shape, the key uses the most basal ones. Sessile glands are often present on the abaxial surface of bracts; they have no taxonomic value. Glandular (on short versus long stalks) and eglandular hairs are used in characterizing indumentum of different species.

Two infraspecific levels are used in this treatment. This is consistent with the usage in other Euphrasia treatments (for example, P. F. Yeo 1978). Subspecific rank is used for geographic or ecological integrity; varietal rank is used to indicate transitions between the extreme forms without any geographic or ecological distinction.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Genera ca. 100, species ca. 2000 (27 genera, 292 species in the flora).

Orobanchaceae are now defined to include both the holoparasitic members traditionally included in the family (A. Cronquist 1981) and the hemiparasitic genera formerly included in Scrophulariaceae. Although multiple research groups focus on members of the Orobanchaceae, a widely accepted infrafamilial classification of the family in the sense of Angiosperm Phylogeny Group (2016) has not yet appeared.

The classification by J. R. McNeal et al. (2013), who found that Orobanchaceae comprise six clades, is followed herein (their named clades are roughly equivalent to tribes). The autotrophic Lindenbergia Lehmann (12 species in the Old World) corresponds to the basal clade sister to the rest of the clades. Species in our region are distributed among the remaining five clades: Cymbarieae D. Don (genus 1), Orobancheae Lamarck & de Candolle (genera 2–6), Rhinantheae Lamarck & de Candolle (genera 7–12), Buchnereae Bentham (genera 13 and 14), and Pedicularideae Duby (genera 15–27). Within the family, genera are arranged alphabetically within tribes, or within Pedicularideae, in subgroups within the tribe.

Parasitic plants attach to their hosts via haustoria (L. J. Irving and D. D. Cameron 2009). Haustoria are produced by both hemiparasitic and holoparasitic Orobanchaceae (E. Fischer 2004). In hemiparasitic taxa, haustoria usually tap their host’s xylem, mostly taking up water, mineral nutrients, and nitrogen from their host, and sometimes also carbon. Holoparasitic taxa derive all of their growth requirements predominantly from the host’s phloem (Irving and Cameron).

Parasitism has evolved once in the family (N. D. Young et al. 1999; J. R. McNeal et al. 2013); holoparasitism has arisen independently three times from the hemiparasitic condition (J. R. Bennett and S. Mathews 2006; McNeal et al.).

Some Orobanchaceae are serious pests, primarily on legume and grain crops in warmer and drier areas, especially in sub-Saharan Africa. Striga is a particularly serious pest that parasitizes mostly monocots; S. gesnerioides attacks eudicots (K. I. Mohamed et al. 2006). Orobanche parasitizes eudicot crops primarily in temperate parts of the world (E. S. Teryokhin 1997). All Striga species and non-native species of Orobanche in the flora area are listed on the Federal Noxious Weed List (http://www.aphis.usda.gov/plant_health/plant_pest_info/weeds/downloads/weedlist.pdf) in the United States.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Bracts oblong to lanceolate, lengths 2.5+ times widths; capsule margins eciliate or short-ciliate.
E. salisburgensis
1. Bracts ovate, obovate, deltate, suborbiculate, elliptic, oval, oblong-ovate, or oblong, lengths not more than 2 times widths; capsule margins conspicuously long-ciliate.
→ 2
2. Bract surfaces hairy, hairs glandular (with stalks sometimes flexuous, 3–6-celled, 0.2–0.6 mm).
→ 3
3. Stems simple, sometimes with 1 or 2 pairs of branches; corollas 3–4 mm, white to yellow; Alberta, British Columbia, Manitoba, Northwest Territories, Saskatchewan, Yukon, Alaska, Montana.
E. subarctica
3. Stems simple or with 1–6 pairs of branches; corollas 4–5.5 mm, white; Newfoundland and Labrador, Nova Scotia, Nunavut, Quebec, Maine.
E. disjuncta
2. Bract surfaces glabrous or hairy, hairs eglandular or glandular (with stalks 1- or 2-celled, 0.1–0.2 mm).
→ 4
4. Bract surfaces hairy, hairs glandular.
→ 5
5. Inflorescences beginning at nodes (5–)7–9(–11); corollas 8–10 mm; capsules narrowly oblong, apices truncate to retuse; usually pastures, scrub, marshy places.
E. stricta
5. Inflorescences beginning at nodes 2–4; corollas 5–8(–10) mm; capsules oblong to elliptic or obovate, apices truncate, retuse, or emarginate; usually coastal.
→ 6
6. Bract teeth much longer than wide, apices acute to acuminate; corollas 6–8(–13) mm.
E. arctica
6. Bract teeth as long as or slightly longer than wide, apices subacute to acute, rarely aristate; corollas 5–8 mm.
E. frigida
4. Bract surfaces glabrous or hairy, hairs eglandular.
→ 7
7. Corollas yellow.
→ 8
8. Corollas 3–4 mm; leaf blades oblanceolate to broadly ovate; bract surfaces sparsely hirsute.
E. subarctica
8. Corollas 4–5.5 mm; leaf blades ovate to orbiculate; bract surfaces densely hirsute.
E. mollis
7. Corollas white, cream, lilac, violet, purple, or brownish purple.
→ 9
9. Bracts elliptic to ovate, bases strongly cuneate, tooth apices acute, sometimes aristate.
E. hudsoniana
9. Bracts oval, ovate to obovate, suborbiculate, deltate, or oblong-ovate, bases round, truncate, or cuneate, tooth apices obtuse, acute, or subacute, rarely aristate.
→ 10
10. Corollas 6–11(–13) mm.
→ 11
11. Inflorescences beginning at nodes 3–5.
E. arctica
11. Inflorescences beginning at nodes (5–)7–9(–11).
E. stricta
10. Corollas 2.5–7.5(–8.5) mm.
→ 12
12. Sinuses between teeth rounded, bract tooth apices obtuse or acute; corollas purple or lilac with darker lines.
E. oakesii
12. Sinuses between teeth acute, bract tooth apices obtuse, subacute, or acute, rarely aristate; corollas white, cream, lilac, violet, or brownish purple, rarely purple.
→ 13
13. Inflorescences capitate, beginning at nodes 4 or 5; calyx lobes falcate, apices obtuse.
E. oakesii
13. Inflorescences sparsely or densely spicate, beginning at nodes 2–12; calyx lobes straight, apices acute.
→ 14
14. Inflorescences densely spicate, sometimes 4-angled.
E. tetraquetra
14. Inflorescences sparsely spicate, not 4-angled.
→ 15
15. Inflorescences beginning at nodes 2–5; stems with 0–2 pairs of branches; arctic regions, sometimes non-arctic regions.
→ 16
16. Capsules 6–8 mm, elliptic to obovate, equal to or slightly longer than calyces, apices emarginate.
E. wettsteinii
16. Capsules 4–6 mm, oblong to narrowly elliptic, shorter than or equal to calyces, apices retuse.
E. vinacea
15. Inflorescences beginning at nodes 3–12; stems with 0–7 pairs of branches; non-arctic regions or, if arctic, inflorescences usually beginning at nodes 5+.
→ 17
17. Bracts suborbiculate, broadly ovate, or oblong-ovate, surfaces +/- densely hirsute, teeth shorter than or as long as wide, apices obtuse or subacute to acute.
→ 18
18. Stems to 12(–15) cm; bract teeth as long as wide.
E. ostenfeldii
18. Stems to 27(–37) cm; bract teeth shorter than wide.
E. suborbicularis
17. Bracts ovate to obovate, oblong, or oval, surfaces glabrous or hirsute, teeth longer than or as long as wide, apices obtuse to acute, sometimes aristate.
→ 19
19. Corollas 4.5–7.5(–8.5) mm.
→ 20
20. Corollas 5–7.5(–8.5) mm; bracts not glossy, surfaces glabrous or hirsute, tooth apices acute, sometimes aristate.
E. nemorosa
20. Corollas 4.5–6.5 mm; bracts glossy, surfaces glabrous, tooth apices acute.
E. micrantha
19. Corollas 3–5.5 mm.
→ 21
21. Stems branched from middle and/or distal cauline nodes, branches erect; cauline internode lengths (1–)2–5 times subtending leaves; inflorescences: proximal internode lengths 0.8–3(–4) times bracts.
→ 22
22. Stems simple, sometimes branched, branches 1 or 2 pairs; corollas 3–4 mm, white to yellow; Alberta, British Columbia, Manitoba, Northwest Territories, Saskatchewan, Yukon, Alaska, Montana.
E. subarctica
22. Stems simple or branched, branches 1–6 pairs; corollas 4–5.5 mm, white; Newfoundland and Labrador, Nova Scotia, Nunavut, Quebec, Maine.
E. disjuncta
21. Stems branched from basal, middle, and/or distal cauline nodes, branches ascending; cauline internode lengths 1–3(–5) times subtending leaves; inflorescences: proximal internode lengths 0.8–1.5 times bracts.
→ 23
23. Cauline internode lengths 1–3(–5) times subtending leaves; bracts 2–7 mm, abaxial surfaces setulose on veins, adaxial puberulent.
E. randii
23. Cauline internode lengths 1–2 times subtending leaves; bracts 2–4 mm, surfaces coarsely, densely hirsute.
E. farlowii
1. Plants holoparasitic, achlorophyllous.
→ 2
2. Corollas salverform; annuals.
Striga
2. Corollas short-tubular, tubular, or funnelform; perennials or annuals.
→ 3
3. Flowers cleistogamous and chasmogamous; petals 5 (appearing as 4); stems absent.
Epifagus
3. Flowers chasmogamous; petals 5; stems present.
→ 4
4. Capsules indehiscent; calyces divided abaxially, not divided adaxially; stamens exserted.
Conopholis
4. Capsules dehiscent; calyces divided roughly uniformly abaxially and adaxially; stamens included.
→ 5
5. Corollas tinged pink to purple, yellow, or blue, pallid proximally; palatal folds present (longitudinal folds in abaxial side of tube); calyces narrowly campanulate to campanulate; roots short, sometimes coralloid.
Orobanche
5. Corollas dark red or purple, sometimes yellow; palatal folds absent; calyces cup-shaped; roots absent.
→ 6
6. Inflorescences dense spikes; pedicels absent, bracteoles absent; corollas short-tubular.
Boschniakia
6. Inflorescences compact or open racemes; pedicels present, bracteoles present, rarely absent; corollas funnelform.
Kopsiopsis
1. Plants hemiparasitic, chlorophyllous.
→ 7
7. Corollas bilabiate, adaxial lips not galeate, cucullate, or beaked.
→ 8
8. Corollas salverform.
→ 9
9. Corollas purple, blue-purple, blue, violet, rosy, or white; filaments pilose.
Buchnera
9. Corollas red, brownish red, or purple, rarely white or yellow; filaments glabrous.
Striga
8. Corollas tubular, campanulate, or subrotate.
→ 10
10. Leaves whorled.
Brachystigma
10. Leaves alternate, opposite, or subopposite.
→ 11
11. Leaves alternate.
Agalinis
11. Leaves opposite or subopposite.
→ 12
12. Corollas pale pink to rose purple or purple, rarely white; leaf blade margins entire, rarely proximally cleft, pinnatifid, or 2-pinnatifid.
Agalinis
12. Corollas yellow or bright orange; leaf blade margins toothed or irregularly lobed, pinnatifid, or 2-pinnatifid, sometimes entire.
→ 13
13. Corollas bright orange, tubular; stamens equal.
Macranthera
13. Corollas yellow, campanulate; stamens didynamous, subequal, or equal.
→ 14
14. Anthers villous.
Aureolaria
14. Anthers glabrous.
→ 15
15. Calyx lobes ovate to oblong-ovate; stamens didynamous.
Dasistoma
15. Calyx lobes linear to lanceolate; stamens equal to subequal.
Seymeria
7. Corollas strongly bilabiate or bilabiate, adaxial lips galeate, cucullate, or beaked.
→ 16
16. Perennials, caudices woody or fleshy.
→ 17
17. Bracteoles present; sepals 5.
Schwalbea
17. Bracteoles absent; sepals 2, 4, or 5.
→ 18
18. Cauline leaves decussate.
Bartsia
18. Cauline leaves alternate, rarely whorled.
→ 19
19. Pollen sacs equal; corollas: adaxial lips sometimes with an upcurved or coiled beak.
Pedicularis
19. Pollen sacs unequal; corollas: adaxial lips straight, rarely hooked.
Castilleja
16. Annuals, rarely biennials, caudices absent.
→ 20
20. Cauline leaves opposite, sometimes subopposite or alternate.
→ 21
21. Leaf blade margins entire, sometimes margins of distal leaves proximally toothed; calyx lobes subulate; seeds 1–4.
Melampyrum
21. Leaf blade margins toothed; calyx lobes deltate, triangular, or lanceolate; seeds (2–)10–450.
→ 22
22. Calyces ovate to suborbiculate, flattened laterally, accrescent in fruit.
Rhinanthus
22. Calyces tubular to campanulate, not flattened laterally, not accrescent in fruit.
→ 23
23. Anther mucros unequal; capsule dehiscence septicidal.
Euphrasia
23. Anther mucros equal or absent; capsule dehiscence loculicidal.
→ 24
24. Filaments glabrous; inflorescences spikelike racemes.
Bellardia
24. Filaments papillose; inflorescences unilateral racemes.
Odontites
20. Cauline leaves alternate, proximals rarely subopposite to opposite.
→ 25
25. Stamens 2.
→ 26
26. Leaf blades: margins of proximals 3-lobed, margins of distals entire.
Cordylanthus
26. Leaf blades: margins entire or pinnately 5–11-lobed.
→ 27
27. Leaf blade margins entire or pinnately 5- or 7-lobed; pollens sacs approximate, connectives not elongate.
Chloropyron
27. Leaf blade margins pinnately 8–11-lobed; pollens sacs separate, connectives elongate.
Dicranostegia
25. Stamens 4.
→ 28
28. Corollas: adaxial lips ± straight, openings directed forward, rarely beaked, bent, or hooked at tip and openings directed downward; stigmas capitate or 2-lobed.
→ 29
29. Cauline leaves alternate; pollen sacs 2.
Castilleja
29. Cauline leaves: proximals usually subopposite to opposite, distals alternate; pollen sacs 1.
Triphysaria
28. Corollas: adaxial lips rounded at apex, sometimes obscurely so, openings directed downward; stigmas not or slightly expanded.
→ 30
30. Sepals 4, calyces tubular.
Orthocarpus
30. Sepals 2, calyces spathelike.
→ 31
31. Leaf blade margins entire; corollas: middle lobes of abaxial lips not revolute; saline marshes, alkaline flats.
Chloropyron
31. Leaf blade margins entire or 3–7-lobed; corollas: middle lobes of abaxial lips tightly revolute; sagebrush scrub, chaparral, woodlands, forests.
Cordylanthus
Source FNA vol. 17, p. 492. Author: Galina Gussarova. FNA vol. 17, p. 456. Authors: Craig C. Freeman, Richard K. Rabeler, Wayne J. Elisens.
Parent taxa Orobanchaceae
Subordinate taxa
E. arctica, E. disjuncta, E. farlowii, E. frigida, E. hudsoniana, E. micrantha, E. mollis, E. nemorosa, E. oakesii, E. ostenfeldii, E. randii, E. salisburgensis, E. stricta, E. subarctica, E. suborbicularis, E. tetraquetra, E. vinacea, E. wettsteinii
Agalinis, Aureolaria, Bartsia, Bellardia, Boschniakia, Brachystigma, Buchnera, Castilleja, Chloropyron, Conopholis, Cordylanthus, Dasistoma, Dicranostegia, Epifagus, Euphrasia, Kopsiopsis, Macranthera, Melampyrum, Odontites, Orobanche, Orthocarpus, Pedicularis, Rhinanthus, Schwalbea, Seymeria, Striga, Triphysaria
Name authority Linnaeus: Sp. Pl. 2: 604. (1753): Gen. Pl. ed. 5, 263. (1754) Ventenat
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