bird's-beak, birdbeak
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broom-rape family
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Herbs, annual; hemiparasitic. |
Herbs, rarely subshrubs or shrubs, annual, biennial, or perennial, sometimes fleshy, hemiparasitic or holoparasitic (without chlorophyll) [autotrophic]. |
erect or ascending, rarely decumbent, not fleshy, hairy or glabrous. |
subterranean or aerial; aerial stems prostrate to decumbent, ascending, or erect [viny]. |
cauline, alternate; petiole absent; blade not fleshy, not leathery, margins entire or 3–7-lobed. |
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. |
terminal, spikes or flowers solitary, often capitate; bracts present. |
terminal and/or axillary, racemes, panicles, spikes, corymbs, or flowers 1 or 2. |
absent; bracteoles absent. |
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sepals 2, calyx bilaterally symmetric, spathelike, lobes triangular; petals 5, corolla yellow, yellow-green, purple, pink, or red, strongly bilabiate, club-shaped, abaxial lobes 3, middle lobe tightly revolute, tip distinctly folded inside-out, adaxial 2, adaxial lip galeate, rounded at apex, opening downward; stamens (2 or)4, didynamous, filaments hairy or glabrous; staminode 0; ovary 2-locular, placentation axile; stigma slightly expanded at apex. |
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. |
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capsules, dehiscence loculicidal and/or septicidal or indehiscent (Conopholis). |
dehiscence loculicidal. |
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4–25, pale brown to dark brown, ovoid to reniform, 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. |
= 6, 7. |
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w United States; nw Mexico |
nearly worldwide; especially in warm temperate regions |
Species 13 (13 in the flora). Cordylanthus is similar to Orthocarpus but differs in having spathelike calyces deeply cut along one side and narrowly lanceolate, entire bracteoles subtending each flower. Also, the abaxial lip of Orthocarpus is spreading; it usually is appressed to the adaxial lip in Cordylanthus. The mature flowers often appear to be buds. Cordylanthus is sometimes defined to include Dicranostegia and Chloropyron as subgenera (T. I. Chuang and L. R. Heckard 1986). D. C. Tank et al. (2009) found that those genera are a sister group of Triphysaria. The monograph by Chuang and Heckard provided the basis for most of the species delineation presented here. (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.) |
1. Stamens 2, filaments glabrous; calyx apices 2-fid, cleft 3–5 mm. | C. capitatus |
1. Stamens 4, filaments hairy; calyx apices entire or 2-fid, cleft 0–3 mm. | → 2 |
2. Bract margins 5–7-lobed, flabelliform; corollas 8–9 mm. | C. pringlei |
2. Bract margins entire, 3-lobed, or (4 or)5–9-lobed, not flabelliform; corollas 10–30 mm. | → 3 |
3. Fertile pollen sacs 1 per filament. | → 4 |
4. Calyx apices 2-fid; bracts green or purple distally; abaxial corolla lips 7–10 mm, slightly spreading. | C. laxiflorus |
4. Calyx apices entire; bracts white to cream distally; abaxial corolla lips 3–5 mm, appressed to adaxial. | C. nevinii |
3. Fertile pollen sacs 2 per filament. | → 5 |
5. Stems decumbent, 5–10(–15) cm; corollas white with purple veins. | C. nidularius |
5. Stems erect or ascending, 10–150 cm; corollas pale or bright yellow, yellow, yellow-green, pale pink, lavender-pink, pink, purple, purple-pink, or purple-red. | → 6 |
6. Bract margins (4 or)5–9-lobed. | → 7 |
7. Calyx tubes 0 mm; fertile pollen sacs unequal. | C. ramosus |
7. Calyx tubes 1–4 mm; fertile pollen sacs equal. | → 8 |
8. Corollas 10–20 mm, throats 4–6 mm diam.; calyx tubes 1–3 mm; capsules 7–10 mm. | C. eremicus |
8. Corollas 15–30 mm, throats 6–8 mm diam.; calyx tubes 3–4 mm; capsules 10–15 mm. | C. wrightii |
6. Bract margins entire or 3-lobed. | → 9 |
9. Calyx tubes 5–8 mm; corolla abaxial lips shorter than and not appressed to adaxial. | C. parviflorus |
9. Calyx tubes 0–2.5 mm; corolla abaxial lips ca. equal to and appressed to adaxial. | → 10 |
10. Calyx apices 2-fid, clefts 2–3 mm; bracteole margins pinnately lobed. | C. kingii |
10. Calyx apices entire or 2-fid, clefts 0.5–1 mm; bracteole margins entire or toothed. | → 11 |
11. Calyx tubes 1–2 mm; inflorescences (2–)5–15-flowered; corollas with abaxial, U-shaped, purple markings. | C. rigidus |
11. Calyx tubes 0–1 mm; inflorescences 2–7-flowered, or flowers solitary; corollas streaked and spotted with maroon or marked with purple. | → 12 |
12. Stems densely puberulent and glandular-puberulent, and pilose; abaxial corolla lips 5–10 mm. | C. pilosus |
12. Stems glabrous or sparsely glandular-puberulent, puberulent, and/or pilose; abaxial corolla lips 4–6 mm. | C. tenuis |
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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 |
| → 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 |
| Brachystigma |
10. Leaves alternate, opposite, or subopposite. | → 11 |
| 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 |
| Aureolaria |
| → 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 |
| → 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 |
| → 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 |
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FNA vol. 17, p. 669. Author: Kerry A. Barringer. |
FNA vol. 17, p. 456. Authors: Craig C. Freeman, Richard K. Rabeler, Wayne J. Elisens. |
Orobanchaceae |
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C. capitatus, C. eremicus, C. kingii, C. laxiflorus, C. nevinii, C. nidularius, C. parviflorus, C. pilosus, C. pringlei, C. ramosus, C. rigidus, C. tenuis, C. wrightii |
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 |
Adenostegia |
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Nuttall ex Bentham: in A. P. de Candolle and A. L. P. P. de Candolle, Prodr. 10: 597. (1846) — name conserved |
Ventenat |
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