Flora of North America species comparison

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Euphrasia

Euphrasia subarctica

euphrasia, eyebright

arctic eyebright, subarctic eyebright
Habit Herbs, annual [perennial]; hemiparasitic.
Stems

erect, not fleshy, retrorsely hairy.

simple, sometimes branched, to 30 cm;

branches 1 or 2 pairs, erect, from distal cauline nodes;

cauline internode lengths (1–)2–5 times subtending leaves.
Leaves

cauline, opposite;

petiole absent or nearly so;

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

blade oblanceolate to broadly ovate, 2–6(–11) mm, margins crenate, teeth 1–3(–5) pairs, apices obtuse to subacute.
Inflorescences

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

bracts present, subopposite or irregularly alternate.

sparsely spicate, not 4-angled, beginning at node 3 or 4(–6);

proximal internode lengths 0.8–3(–4) times bracts;

bracts green, often purplish adaxially, broader than leaves, ovate to obovate, length not more than 2 times width, 3–10 mm, base cuneate, surfaces sparsely hirsute and hairs eglandular or +/- densely, sometimes sparsely, glandular-pilose and hairs glandular, stalks sometimes flexuous, 3–6-celled, 0.2–0.6 mm, teeth 2–6 pairs, longer than wide, apices subacute to acute, rarely aristate, sinuses between teeth acute.
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.

calyx lobes straight, apex acute;

corolla white to yellow, seldom with violet veins, 3–4 mm, lips +/- equal.
Capsules

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

elliptic to oblong, (2.5–)5–6.5 mm, apex truncate to retuse.
Seeds

10–18, grayish, fusiform, wings absent.

x

= 11.

Euphrasia

Euphrasia subarctica
Phenology Flowering summer.
Habitat Sandy, gravelly, or damp grassy places, stream banks, shores, thickets, heathlands, tundra.
Elevation 0–2500 m. (0–8200 ft.)
Distribution
from USDA
North America; South America; Eurasia; Africa (Morocco); Atlantic Islands (Azores, Iceland); Pacific Islands (Oceania); Australia
[BONAP county map]
from FNA
AK; MT; AB; BC; MB; NT; SK; YT
[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.)

The holotype (Raup & Abbe 4633, GH) fixes the name Euphrasia subarctica to individuals with long, glandular hairs. Eglandular individuals are sometimes found growing mixed together with typical, glandular E. subarctica in Alaska and northwestern Canada. Similarly, eglandular and long glandular-pubescent individuals are intermixed in populations of otherwise morphologically similar, but not identical, E. disjuncta, including the specimens of its paratype (Fernald & Wiegand 6166, CAN). Due to the clear tendency for the glandular forms to be more frequent in the west, while eglandular ones are more typical in the east, as well as other morphological differences, E. subarctica and E. disjuncta are treated as separate species here. However, it is likely that they represent a single lineage with a disjunct distribution, which could be a result of postglacial colonization from two separate refugia in western and eastern North America. Co-occurrence of glandular and eglandular states is documented in other diploids [for example, E. picta Wimmer in the Austrian Alps (P. F. Yeo 1978)]. Occasional individuals of E. subarctica with yellow corollas may be the result of introgression from E. mollis.

(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
Source FNA vol. 17, p. 492. Author: Galina Gussarova. FNA vol. 17, p. 494.
Parent taxa Orobanchaceae Orobanchaceae > Euphrasia
Sibling 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. suborbicularis, E. tetraquetra, E. vinacea, E. wettsteinii
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
Synonyms E. arctica var. dolosa, E. disjuncta var. dolosa, E. pennellii var. incana
Name authority Linnaeus: Sp. Pl. 2: 604. (1753): Gen. Pl. ed. 5, 263. (1754) Raup: Rhodora 36: 87, plate 278. (1934)
Web links 
Unless otherwise noted, all data comes from the online version of the Flora of North America, who have made most of their content available via a Creative Commons license (CC BY 4.0).



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