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broom-rape family

elephant head, fernflower, lousewort, pedicularis

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

subterranean or aerial;

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

erect, fleshy, glabrous, hispid, villous, or woolly.

Leaves

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.

basal rosette [absent], petiole present, blade not fleshy, not leathery, margins entire, serrate, or crenate;

cauline alternate, rarely whorled [nearly opposite], sometimes absent, petiole present or absent, blade chartaceous, margins entire, serrate, or crenate.

Inflorescences

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

terminal or axillary, racemes;

bracts present.

Pedicels

present;

bracteoles absent.

Flowers

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.

sepals 2 or [4]5, calyx bilaterally symmetric, campanulate or tubular, lobes triangular, spatulate, or filiform;

petals 5, corolla pink, purple, red, yellow, or white, strongly bilabiate, cylindric to funnelform, abaxial lobes 3, adaxial 2, adaxial lip galeate, enclosing anthers and style, beaked or beakless, margins entire medially and distally, with 1 set of teeth medially and entire distally, with 1 set of teeth medially and distally, or entire medially and with 1 set of teeth distally;

stamens 4, didynamous, filaments glabrous or hairy, pollen sacs equal;

staminode 0;

ovary 2-locular, placentation axile;

stigma capitate.

Fruits

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

Capsules

dehiscence loculicidal.

Seeds

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.

5–100, dark gray, brown, or tan, ovoid, wings absent.

x

= 8.

Orobanchaceae

Pedicularis

Distribution
nearly worldwide; especially in warm temperate regions
[BONAP county map]
map from USDA
North America; Mexico; South America; Europe; s Asia; c Asia; Atlantic Islands (Iceland)
[BONAP county map]
Discussion

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.)

Species 400–600 (37 in the flora).

E. Hultén (1968) listed three Pedicularis taxa in his Flora of Alaska and Neighboring Territories that do not occur in the flora area; P. amoena Adams ex Steven, P. kanei Durand subsp. adamsii (Hultén) Hultén, and P. villosa Ledebour ex Sprengel are found on the Chukchi Peninsula of Siberia.

The most common chromosome number for Pedicularis is 2n = 16. Two species are reported to have a diploid number of 2n = 12, and at least five species have a number of 2n = 14. See reviews in Jie C. et al. (2004) and M. I. S. Saggoo and D. K. Srivastava (2009).

Infrageneric classification of Pedicularis is both difficult and incomplete. The author has not attempted to present one for the North American taxa. Inconsistencies result from using either floral morphology or phyllotaxy to construct major subgeneric groups (C. Steven 1823; G. Bentham 1835; C. J. Maximowicz 1888; D. Prain 1890; G. Bonati 1918; Tsoong P. C. 1955); the latter author proposed 130 series to accommodate the worldwide variation in Pedicularis. Recognizing that floral form probably evolved in parallel among groups, Li H. L. (1948) employed phyllotaxy to erect three informal subgeneric taxa (greges) for the Pedicularis of China, including grex Allophyllum, species with alternate leaves; grex Cyclophyllum, species with opposite or verticillate arrangement of leaves; and grex Poecilophyllum, species with alternate to subopposite leaves on the same plant. However, molecular phylogenies (Yang F. S. et al. 2003; R. H. Ree 2005; B. W. Robart et al. 2015; Yu W. B. et al. 2015) indicate inconsistencies even in the treatment by Li, although the study by Yu et al., which included 257 species, did show that grex Cyclophyllum is monophyletic.

Pediculariopsis was described to account for species (for example, Pedicularis verticillata) that exhibit a different base chromosome number (x = 6 versus x = 8). That concept has not been accepted by others, because there is little morphological support.

The use of infraspecific ranks in Pedicularis follows T. F. Stuessy (1990), who argued that the subspecific rank is appropriate when several obvious morphological differences are associated with an allopatric or peripatric geographical pattern, whereas the varietal rank is appropriate when one to few differences are associated with more or less geographic overlap. Unfortunately, Pedicularis specialists have not used these levels consistently. The subspecies rank is used here, except in P. bracteosa, P. centranthera, and P. contorta.

Pedicularis is a common element of arctic and alpine habitats in the Northern Hemisphere, with the greatest concentration of species occurring in Asia.

Etymology: Latin pediculus, louse, alluding to belief that livestock feeding on P. palustris developed lice

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

Parent taxa Orobanchaceae
Subordinate taxa
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
P. angustifolia, P. attollens, P. aurantiaca, P. bracteosa, P. canadensis, P. capitata, P. centranthera, P. chamissonis, P. contorta, P. crenulata, P. cystopteridifolia, P. densiflora, P. dudleyi, P. flammea, P. furbishiae, P. groenlandica, P. hirsuta, P. howellii, P. labradorica, P. lanata, P. lanceolata, P. langsdorffii, P. lapponica, P. oederi, P. ornithorhynchos, P. palustris, P. parryi, P. parviflora, P. pennellii, P. procera, P. pulchella, P. racemosa, P. rainierensis, P. semibarbata, P. sudetica, P. sylvatica, P. verticillata
Key
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
1. Racemes verticillate; cauline leaves whorled.
→ 2
2. Galeas beaked; basal leaf blades 15–40 mm.
P. chamissonis
2. Galeas beakless; basal leaf blades 10–20 mm.
P. verticillata
1. Racemes simple or paniculate; cauline leaves alternate.
→ 3
3. Calyx lobes 2(–4).
→ 4
4. Galeas beaked.
→ 5
5. Beaks sickle-shaped.
P. racemosa
5. Beaks straight.
→ 6
6. Calyx lobe apices distally serrate.
P. lanceolata
6. Calyx lobe apices distally entire.
P. lapponica
4. Galeas beakless.
→ 7
7. Galea margins 1-toothed medially, entire distally.
P. parviflora
7. Galea margins entire or 1-toothed medially, 1-toothed distally.
→ 8
8. Galea margins 1-toothed medially.
→ 9
9. Galeas 6.5–9 mm.
P. palustris
9. Galeas 3–6.5 mm.
P. pennellii
8. Galea margins entire medially.
→ 10
10. Racemes paniculate, or buds present in cauline leaf axils.
→ 11
11. Basal leaves 0; cauline leaf blades undivided.
P. angustifolia
11. Basal leaves 2 or 3; cauline leaf blades undivided or 1- or 2-pinnatifid.
P. labradorica
10. Racemes simple.
→ 12
12. Cauline leaf blades 1-pinnatifid, lobe margins 1- or 2-serrate.
P. canadensis
12. Cauline leaf blades undivided, lobe margins 2-crenate.
P. crenulata
3. Calyx lobes 5.
→ 13
13. Galeas beaked.
→ 14
14. Galea beaks coiled.
→ 15
15. Galea beak apices surrounded by abaxial lips.
P. contorta
15. Galea beak apices not surrounded by abaxial lips.
→ 16
16. Galea beaks abruptly upcurved; beaks 3–6 mm.
P. attollens
16. Galea beaks gradually upcurved; beaks 5–18 mm.
P. groenlandica
14. Galea beaks straight.
→ 17
17. Beaks 0.8–2.5 mm; bracts undivided.
→ 18
18. Corolla tubes and galeas yellow.
P. bracteosa
18. Corolla tubes and galeas white.
P. howellii
17. Beaks 2–8 mm; bracts undivided or +/- lobed.
→ 19
19. Galeas 4–6.5 mm.
P. ornithorhynchos
19. Galeas 7–10 mm.
P. parryi
13. Galeas beakless.
→ 20
20. Galea margins entire medially, 1-toothed distally.
→ 21
21. Calyces glabrous.
→ 22
22. Bracts 2-pinnatifid to midrib.
P. sylvatica
22. Bracts undivided proximally, 1-pinnatifid 1/2 to midrib distally, or undivided with or without long auricles, or 1-pinnatifid.
→ 23
23. Pedicels 2.5–5 mm; corolla tubes 11–13 mm.
P. langsdorffii
23. Pedicels 1–2.5 mm; corolla tubes 9–11 mm.
P. sudetica
21. Calyces +/- tomentose, hispid-glandular, hispid to hirsute, white- or yellowish white-lanate, sparsely pilose, or +/- woolly.
→ 24
24. Corolla tubes yellow to light or greenish yellow.
→ 25
25. Corollas 14–19 mm.
P. furbishiae
25. Corollas 22–30 mm.
P. procera
24. Corolla tubes red, violet-red, lavender, purple, magenta, or pink, sometimes white.
→ 26
26. Basal leaf blades 150–250 mm.
P. procera
26. Basal leaf blades 5–110 mm.
→ 27
27. Basal leaves: margins of adjacent lobes nonoverlapping to extensively overlapping.
→ 28
28. Bracts undivided or 1- or 2-auricled, sometimes 1-pinnatifid.
P. cystopteridifolia
28. Bracts 2-pinnatifid, not auricled.
P. pulchella
27. Basal leaves: margins of adjacent lobes nonoverlapping or slightly overlapping distally.
→ 29
29. Corollas 11–19 mm, galea apices nearly straight to arching slightly over abaxial lips.
P. hirsuta
29. Corollas 16–25 mm, galea apices strongly arching over abaxial lips.
→ 30
30. Pedicels 2.5–5 mm.
P. langsdorffii
30. Pedicels 1–2.5 mm.
P. sudetica
20. Galea margins entire medially and distally.
→ 31
31. Racemes not exceeding basal leaves.
→ 32
32. Galeas 13–15 mm.
P. centranthera
32. Galeas 5–12 mm.
P. semibarbata
31. Racemes exceeding basal leaves.
→ 33
33. Galea apices straight.
→ 34
34. Corolla abaxial lips 3–7 mm.
P. aurantiaca
34. Corolla abaxial lips 8–15 mm.
P. densiflora
33. Galea apices arching over or beyond abaxial lips.
→ 35
35. Calyces glabrous or ciliate.
→ 36
36. Galeas yellow.
P. bracteosa
36. Galeas yellow proximally, dark red to purple distally.
P. flammea
35. Calyces hispid to tomentose, hirsute, or densely woolly.
→ 37
37. Racemes capitate, each 2–8-flowered, galea apices arching beyond abaxial lips.
P. capitata
37. Racemes simple, each 6–100-flowered, galea apices arching over or beyond abaxial lips.
→ 38
38. Galeas bicolored.
P. oederi
38. Galeas concolored.
→ 39
39. Corolla tubes light to dark yellow or dark blood red.
→ 40
40. Pedicels 0.5–1 mm.
P. bracteosa
40. Pedicels 1–3.5 mm.
P. rainierensis
39. Corolla tubes pink or reddish purple, rarely white.
→ 41
41. Bracts undivided.
P. dudleyi
41. Bracts 1-pinnatifid distally.
P. lanata
Synonyms Elephantella, Pediculariopsis
Name authority Ventenat Linnaeus: Sp. Pl. 2: 607. (1753): Gen. Pl. ed. 5, 266. (1754)
Source FNA vol. 17, p. 456. Treatment authors: Craig C. Freeman, Richard K. Rabeler, Wayne J. Elisens. FNA vol. 17, p. 510. Treatment author: Bruce W. Robart.
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