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bellardia, glandweed, lineseed, parentucellia

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 short-strigose or glandular-hairy.

subterranean or aerial;

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

Leaves

cauline, opposite, sometimes some subopposite or alternate;

petiole absent;

blade not fleshy, not leathery, margins coarsely crenate-dentate.

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, spikelike racemes;

bracts present.

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

Pedicels

present;

bracteoles absent.

Flowers

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

petals 5, corolla white with purple galea, yellow, or red-purple, strongly bilabiate, tubular-funnelform, abaxial lobes 3, adaxial 2, adaxial lip galeate;

stamens 4, didynamous, filaments glabrous, anther mucros equal or absent;

staminode 0;

ovary 2-locular, placentation axile;

stigma clavate, capitate, or +/- 2-lobed.

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 loculicidal.

Seeds

150–450, white, reddish brown with age, ellipsoid or ellipsoid-oblong, 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

= 12.

Bellardia

Orobanchaceae

Distribution
from USDA
Europe [Introduced in North America; introduced also in s South America, Asia, Africa, Atlantic Islands, Pacific Islands, s Australia]
[BONAP county map]
nearly worldwide; especially in warm temperate regions
[BONAP county map]
Discussion

Species 48 (3 in the flora).

Bellardia trixago has been included in a monospecific section of Bartsia in the broad sense (U. Molau 1990). Morphology and molecular phylogenetic analyses indicate a close relationship among Bellardia, Parentucellia, and New World species of Bartsia (A. D. Wolfe et al. 2005; J. R. Bennett and S. Mathews 2006; S. Uribe-Convers and D. C. Tank, http://2010.botanyconference.org/engine/search/index.php?func=detail&aid=685; A. Scheunert et al. 2012). Molecular phylogenetic analyses suggest that South American Bartsia are nested within a highly supported clade including Bellardia and Parentucellia (Scheunert et al.). An expanded Bellardia includes P. latifolia (Linnaeus) Caruel, P. viscosa (Linnaeus) Caruel, and the South American species of Bartsia. After this treatment had been completed, the author became aware of the alternative view presented by Uribe-Convers and Tank (2016), where all of the South American Bartsia taxa are transferred to the new genus Neobartsia Uribe-Convers & Tank, Parentucellia is recognized, and Bellardia includes only B. trixago and B. viscosa.

(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. Corollas white with purple galea; calyx lobes unequal.
B. trixago
1. Corollas red-purple or yellow; calyx lobes +/- equal.
→ 2
2. Corollas red-purple; leaf blades 4–10(–12) mm.
B. latifolia
2. Corollas yellow; leaf blades (10–)20–46 mm.
B. viscosa
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. 490. Author: Elizabeth H. Zacharias. FNA vol. 17, p. 456. Authors: Craig C. Freeman, Richard K. Rabeler, Wayne J. Elisens.
Parent taxa Orobanchaceae
Subordinate taxa
B. latifolia, B. trixago, B. viscosa
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
Synonyms Bartsia section bellardia, Parentucellia
Name authority Allioni: Fl. Pedem. 1: 61. (1785) Ventenat
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