According to Neotropikey[NTK]
Souza, V.C. (2012). Neotropical Orobanchaceae.
Annual , biennial , perennial herbs or shrubs, sometimes climbers (Velloziella), hemiparasites or holoparasites on roots ; rarely non-parasitic (Lindenbergia), usually producing orobanchin and iridoid compounds, turning plant black after drying. Leaves simple , opposite or less frequently alternate or whorled ; sometimes scale-like; venation pinnate , palmate or one-veined; margin entire , serrate or sometimes deeply divided ; without stipules. Inflorescence most racemose, frequently with bracts similar to normal leaves. Flowers mostly showy (bracts more attractive than flowers in Castilleja), bisexual ; calyx gamosepalous, actinomorphic to zygomorphic , generally 4-5- lobed ; corolla gamopetalous, zygomorphic , usually bilabiate, generally 5- lobed ; stamens 4, rarely with a staminode , generally didynamous; epipetalous, alternating with corolla lobes, anthers dithecal or less frequently monothecal (Buchnera, Cordylanthus), introrse, opening by longitudinal slits or rarely by pores (some Euphrasia); gynoecium syncarpous of 2(-3-5) carpels, locules as many as carpels or 1, style 1, ovules many, placentation axile or parietal . Fruit a septicidal (sometimes also loculicidal) capsule . Seeds numerous, minute, testa often ornamented, usually wind-dispersed.
- General Description
Notes on delimitation
- Orobanchaceae is positioned in Lamiales, as a probable sister to Paulowniaceae, but the evolutionary relationships between families of this order are still poorly known.
- Recent studies (summarized by Olmstead et al. 2001 and Tank et al., 2006) suggest that the traditional circumscription of Orobanchaceae should be reviewed to include some hemiparasitic genera traditionally included in Scrophulariaceae and Lindenbergia, a nonparasitic genus, probably basal in this group.
- Orobanchaceae has a cosmopolitan distribution, including about 100 genera and 2000 species. Of these, 21 genera and 300-330 species are native in the Neotropics and one (Parentucellia Viv.) is naturalised.
- This family include some damaging weeds such as the witchweeds (Striga asiatica (L.) Kuntze, Striga hermonthica (Delile) Benth. and allies) and they present in the agriculture in many countries, particularly in Africa and Asia, affecting very poor areas. The parasitic habit, plus the quick life cycle, and extraordinary capacity for propagation (thousands of small seeds carried by wind can be produced by a single plant), makes the species of Striga some of the worst weeds in the world.
- There is no record of Striga infestation in the Neotropics, but some areas in U.S.A. have been infected for decades, with little perspective of eradication.
- Considering that many places where Striga species occur in Asia and Africa have a climate similar to the Neotropics and also a similarity in crops cultivated, it is realistic to be concerned about these species. Buchnera, which is morphologically similar to Striga and its sister group, has many native Neotropical species, but there is no record of Buchnera species behaving as weeds.
- Most species of Neotropical Orobanchaceae are native; only a few species are naturalised, including Bartsiatrixago L. and Parentucellia viscosa (L.) Caruel.
- Nine genera of Orobanchaceae are endemic to the Neotropics.
- Five genera are also native in other New World regions (Canada and United States).
- Seven genera are also native in the Old World.
Distribution in the Neotropics
Throughout the Neotropics, especially the Andes and the highlands of Brazil.
- Agalinis Raf. (including Anisantherina Pennell, Brachystigma Pennell, Gerardia Benth. and Schizosepala G.M. Barroso): (80-90 species, with c. 40 in the Neotropics) from Canada to northern Argentina, with an Andean and a Central Brazilian centre of diversity.
- Aureolaria Raf.: (8 species) U.S.A. to Mexico.
- Bartsia L. (including Bellardia All.): (50-60 species, with c. 48 in the Neotropics) mainly Colombia, Venezuela and Peru to Chile and NW Andean Argentina, but also in tropical Africa, Europe and North America. Bartsia trixago L.is a cosmopolitan weed, probably with a Mediterranean origin, which occurs in Peru, Brazil (Molau, 1990) and Chile (Marticorena & Quezada, 1985).
- Buchnera L. (130-140 species, with 16 in the Neotropics) Africa (centre of diversity) and southern U.S.A. to Argentina.
- Castilleja Mutis ex L.f. (including Clevelandia Greene, Gentrya Breedlove & Heckard and Ophiocephalus Wiggins) - (200-210 species, with 80-90 in the Neotropics) Pantropical, with centre of diversity in North America.
- Conopholis Wallr. (2 species) eastern Canada through U.S.A., Mexico and Panama.
- Cordylanthus Nutt. ex Benth. (including Chloropyron Behr) (16 species, with 5 in the Neotropics) U.S.A. and Mexico.
- Eremitilla Yatskievych & J.L. Contreras (1 species) endemic to the state of Guerrero, Mexico.
- Escobedia Ruiz & Pav. (10-15 species) Mexico to South Brazil.
- Esterhazya J.C.Mikan (5 species, but many undescribed (Castro Sousa in prep.)) Brazil and Paraguay.
- Euphrasia L. (250-350 species, 15-20 in Latin America) cosmopolitan, mainly Asia and Europe, but also in Chile, Argentina and Bolivia.
- Lamourouxia Kunth (25-30 species) northern Mexico to Andean Colombia, Ecuador and Peru.
- Magdalenaea Brade (1 species) endemic to the Pedra Dubois, Santa Maria Madalena in eastern Brazil.
- Melasma P.J. Bergius (including AlectraThunb.) (40-50 species, 4 in the Neotropics) Mexico to southern Brazil.
- Nothochilus Radlk. (1 species) endemic to the Serra do Caparaó in eastern Brazil.
- Orobanche L. (80-100 species, 3 in South America), Colombia, Chile, Peru and Argentina.
- Parentucellia Viv. (1 species) naturalized in Chile and Brazil, native in Europe.
- Pedicularis L. (400 species with 10 Neotropical species) from Mexico to Ecuador.
- Physocalyx Pohl (4 species) endemic to the Espinhaço Range in Bahia and Minas Gerais, Brazil.
- Seymeria Pursh (25-30, 15-20 in the Neotropics) U.S.A. to Central Mexico.
- Silviella Pennell (2 species) endemic to Mexico.
- Velloziella Baill. (3 species) Venezuela to southern Brazil.
Distinguishing characters (always present)
- Hemiparasitic or holoparasitic (except Lindenbergia).
- Herbs to shrubs.
- Simple leaves.
- Stipules absent.
- Calyx synsepalous.
- Ovules numerous.
- Leaves usually opposite.
- Fruit usually a capsule.
- In a traditional sense, Orobanchaceae could be differentiated from Scrophulariaceae by number of locules in the ovary (1 in Orobanchaceae, 2 in Scrophulariaceae).
- In the current circunscription, the main difference between the two families is the parasitic habit, not always obvious, since some Orobanchaceae, such as Esterhazya, Agalinis and Physocalyx can be robust shrubs, apparently independent of other plants.
- From Verbenaceae, Acanthaceae and Lamiaceae, which Orobanchaceae is sometimes confused, Orobanchaceae can be differentiated by the number of ovules (many in these three families and 1 in Orobanchaceae) in addition to the parasitic habit.
Key to genera of Neotropical Orobanchaceae
The morphological distintion between some genera of Euphrasinae is unclear and the key, in this sense, is provisional, waiting for a comprehensive phylogenetic and morphological work concerning to this group.
1. Plants holoparasitic, without chlorophyll; ovary unilocular — 2
1. Plants hemiparasitic (mostly not evidently so), with chlorophyll; ovary bilocular — 4
2. Ovaries and fruits 5-ribbed, the outer surface muricate — Eremitilla
2. Ovaries and fruits terete or slightly flattened, not ribbed, the outer surface smooth — 3
3. Sepals free or basally connate — Orobanche
3. Sepals connate for most part of their length, forming a tube, sometimes with a longitudinal slit — Conopholis
4. Calyx strongly zygomorphic, sometimes spathe -like — 5
4. Calyx actinomorphic or almost so — 11
5. Climbers or scandent plants; corolla with the upper lip not domed — Velloziella
5. Erect plants; corolla generally with the upper lip domed — 6
6. Anthers 1-thecate — Cordylanthus
6. Anthers 2-thecate — 7
7. Leaves alternate; bracts showy (more than flowers) — Castilleja
7. Leaves generally opposite or verticillate; bracts not showy — 8
8. Calyx spathe -like — Nothochilus
8. Calyx campanulate, not spathe -like — 9
9. Corolla ventricous (inflated) — Lamourouxia
9. Corolla not ventricous — 10
10. Corolla with upper lip laterally compressed and entire or bilobed — Pedicularis
10. Corolla with upper lip not laterally compressed and always entire — Bartsia
11. Corolla salver-shaped — 12
11. Corola mostly campanulate or bilabiate, not salver-shaped — 13
12. Anthers 1-thecate; corolla up to 3 cm long; generally pink, purple or violet — Buchnera
12. Anthers 2-thecate; corolla more than 3 cm long; white or cream — Escobedia
13. Corolla evidently 2-lipped — 14
13. Corolla tubular, campanulate, subrotaceous or subglobose, not 2-lipped — 17
14. Corolla with upper lip laterally compressed — Pedicularis
14. Corolla with upper lip not laterally compressed — 15
15. Sepals 5 — Parentucellia
15. Sepals 4 — 16
16. Corolla inflated, red or pink and over 2.5 cm long — Lamourouxia
16. Corolla not inflated, yellow, white, pink, generally under 2.5 cm long — 17
17. Corolla with upper lip entire; capsule ovoid to subglobose — Bartsia
17. Corolla with upper lip generally bilobed; capsule oblong to lanceolate, flattened — Euphrasia
18. Corolla yellow — 19
18. Corolla red, pink or orange — 23
19. Calyx with multipartite lobes — Agalinis
19. Calyx not with multipartite lobes — 20
20. Corolla lobes longer or with the same size of the tube — Seymeria
20. Corolla lobes smaller than tube — 21
21. Corolla tubular or subglobose — Melasma
21. Corolla campanulate to funelform — 22
22. Stemerect; filaments lanose — Aureolaria
22. Stemprostrate; filaments glabrous — Silviella
23. Leaves with serrate margins — Magdalenaea
23. Leaves with entire margins — 24
24. Stamens exsert, anthers villous — Esterhazya
24. Stamens included; anthers glabrous or almost so — 25
25. Corolla rose to pink; calyx not inflated, green or purple — AgalinisNotable genera and distinguishing features
24. Corolla orange; calyx inflated, also orange — Physo calyx
- Many genera have big and attractive flowers, with ornamental potential. These include Esterhazya, Escobedia, Agalinis and Physocalyx, but cultivation are difficult since they are parasites.
- Nothochilus - Monoespecific genus, endemic to Serra do Caparaó - Brazil.
- Magdalenaea - Monoespecific genus, endemic to Santa Maria Madalena region. Many recent expedictions were made ir order to find this species in field with no results. It is possible to be extincted.
- Eremitilla - Recently described genus, unique in having five carpels.
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This treatment was produced with support from
Bolsista produtividade do CNPq
According to Flora of Tropical East Africa[FTEA]
Orobanchaceae, R. A. Graham. Flora of Tropical East Africa. 1957
- Leafless, scapigerous, parasitic herbs subsisting by attachment to roots of other plants; variously and sometimes beautifully coloured, but without chlorophyll thus never green
- Stems erect, scaly, sometimes fleshy, simple or branched, glabrous to glandular-hairy
- Inflorescence in East African species a cylindrical spike or spiciform raceme
- Flowers hermaphrodite, zygomorphic, subtended by scale-like bracts with or without bracteoles
- Calyx hypogynous, persistent, tubular (long-sided) or campanulate (short-sided), sessile or stalked, 2–5-lobed or -toothed, gamosepalous or split dorsally and ventrally into two free sections
- Corolla hypogynous, gamopetalous, tubular, limbate, sometimes funnel-shaped above, curved throughout or in part
- Stamens didynamous, inserted at or below the middle of the corolla tube; filaments filiform, glabrous to glandular hairy; anther cells in pairs, attached to the filament dorsally, longitudinally dehiscent, glabrous to densely lanate
- Ovary superior, unilocular, bi- (rarely tri-) carpellary; placentas 4, parietal, separate to partially fused; style bilobed, peltate, orbicular or funnel-shaped
- Capsule usually bivalved, ovoid-ellipsoid or ovoid-spheroid, loculicidally dehiscent; seeds small, numerous, obscurely rounded
According to Flora of West Tropical Africa[FWTA]
Orobanchaceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963
- Herbs, parasitic on roots, often covered with scales at the base, never green; stems with alternate, often crowded, scales
- Flowers solitary in the axils of bracts, often crowded, hermaphrodite, zygomorphic
- Calyx 4-5-toothed or lobed or variously split, lobes open or valvate
- Corolla gamopetalous, often curved; limb oblique or 2-lipped, lobes 5, imbricate, the adaxial 2 interior
- Stamens 4, didynamous, inserted below the middle of the corolla-tube, alternate with the lobes, the fifth (adaxial) one reduced to a staminode or absent; anthers often connivent in pairs, opening lengthwise
- Ovary superior, 1-celled, with 4 parietal placentas; style terminal; ovules numerous
- Capsule often enveloped by the calyx, opening by 2 valves
- Seeds very numerous, small, with fleshy endosperm and minute embryo
According to Flora of Tropical East Africa under the synonym Cyclocheilaceae[FTEA]
Cyclocheilaceae, W. Marais. Flora of Tropical East Africa. 1984
- Shrubs with simple and glandular hairs
- Leaves opposite or subopposite, simple, entire, exstipulate
- Flowers axillary, solitary on bracteate pedicels
- Bracteoles lateral, membranous, free or joined, flat or convex and winged, their margins closely adhering and enclosing the bud, accrescent
- Calyx absent or reduced to a disc-like rim
- Corolla gamopetalous, obliquely infundibuliform, 5-lobed
- Stamens didynamous; filaments long pilose; connective short; thecae pendulous, ± divergent, often apiculate at the base
- Ovary compressed, completely or incompletely 2-locular or 1-locular; ovules 2–10, anatropous on long funicles; placentation axile or parietal; style curved, stigma lingulate
- Fruit strongly compressed, a discoid unilocular capsule, or bilocular, breaking up into 1-seeded mericarps
- Seeds without endosperm
According to Flora of Tropical East Africa under the synonym Nesogenaceae[FTEA]
Nesogenaceae, W. Marais. Flora of Tropical East Africa. 1984
- Herbs with both simple and multicellular and usually also glandular hairs
- Leaves opposite, exstipulate, simple, dentate or entire
- Inflorescence axillary of 1 or more superposed cymes, these 1–several-flowered (when several-flowered then clearly bracteate in one species), or flowers in spicate inflorescences
- Calyx (4–)5-lobed, accrescent
- Corolla sympetalous, slightly zygomorphic, unequally 5-lobed, the lobes imbricate
- Stamens didynamous, inserted low down in the tube, included; anthers 2-thecous, the thecae pendulous from a very short connective, apiculate at the base, ± divergent
- Ovary superior, 2-locular; ovules 1 per locule, axile, basal, erect
- Fruit indehiscent, oblong-ovoid, compressed, shallowly grooved between the two halves, pubescent around the thickened persistent base of the style
- Seeds containing endosperm; testa foveolate, the foveoles arranged in longitudinal lines
According to Flora Zambesiaca under the synonym Nesogenaceae[FZ]
Flora Zambesiaca. Vol 8, Pt 7. Avicenniaceae, R. Fernandes. Nesogenaceae, M.A. Diniz. Verbenaceae, R. Fernandes. Lamiaceae, R. Fernandes. 2005.
- Annual or perennial herbs with intermixed simple and multicellular unbranched hairs, and usually also with gland-tipped hairs Annual or perennial herbs with intermixed simple and multicellular unbranched hairs, and usually also with gland-tipped hairs
- Leaves opposite, simple, toothed or entire, exstipulate Leaves opposite, simple, toothed or entire, exstipulate
- Flowers axillary, solitary or clustered cymes (bracteate one species) or bracteate spikes Flowers axillary, solitary or clustered in cymes (bracteate in one species) or in bracteate spikes
- Pedicels short (sometimes absent)
- Stamens didynamous, epipetalous, inserted low down the corolla tube, included; anthers 2-locular, the thecae pendulous, from a very short connective, tailed, ± divergent
- Ovary superior, 2-locular; ovules 1 per locule, axial at the base of the locule, erect
- Style filiform; stigma small, capitate
- Fruit with pericarp crustaceous, indehiscent or at least tardily dehiscent, oblong-ovoid or obovoid, compressed, grooved at the base between the locules, pubescent at the apex around the thickened base of the style Fruit with pericarp crustaceous, indehiscent or at least tardily dehiscent, oblong-ovoid or obovoid, compressed, grooved at the base between the locules, pubescent at the apex around the thickened base of the style
- Seeds oblong, foveolate; endosperm present. Seeds oblong, foveolate; endosperm present
- A monogeneric family, previously included in the Verbenaceae until separated as a family in 1981.
- Pedicels short (sometimes absent)
- Calyx (4)5-lobed, accrescent Calyx (4)5-lobed, accrescent
- Corolla gamopetalous slightly zygomorphic, funnel-shaped, unequally (4)5-lobed; lobes imbricate, either the lower or the two lateral lobes outermost Corolla gamopetalous slightly zygomorphic, funnel-shaped, unequally (4)5-lobed; lobes imbricate, either the lower or the two lateral lobes outermost
- Stamens didynamous, epipetalous, inserted low down in the corolla tube, included; anthers 2-locular, the thecae pendulous, from a very short connective, tailed, ± divergent
- Ovary superior, 2-locular; ovules 1 per locule, axial at the base of the locule, erect Style filiform; stigma small, capitate
- Aeginetia L.
- Agalinis Raf.
- Alectra Thunb.
- Anisantherina Pennell
- Aphyllon Mitch.
- Asepalum Marais
- Aureolaria Raf.
- Bardotia Eb.Fisch., Schäferh. & Kai Müll.
- Bartsia L.
- Baumia Engl. & Gilg
- Bellardia All.
- Boschniakia C.A.Mey.
- Brandisia Hook.f. & Thomson
- Buchnera L.
- Bungea C.A.Mey.
- Buttonia McKen ex Benth.
- Campbellia Wight
- Castilleja Mutis ex L.f.
- Centranthera R.Br.
- Christisonia Gardner
- Cistanche Hoffmanns. & Link
- Conopholis Wallr.
- Cordylanthus Nutt. ex Benth.
- Cyclocheilon Oliv.
- Cycniopsis Engl.
- Cycnium E.Mey. ex Benth.
- Cymbaria L.
- Dasistoma Raf.
- Epifagus Nutt.
- Eremitilla Yatsk. & J.L.Contr.
- Escobedia Ruiz & Pav.
- Esterhazya J.C.Mikan
- Euphrasia L.
- Gerardiina Engl.
- Ghikaea Volkens & Schweinf.
- Gleadovia Gamble & Prain
- Graderia Benth.
- Harveya Hook.
- Hedbergia Molau
- Hiernia S.Moore
- Hyobanche L.
- Kopsiopsis (Beck) Beck
- Lamourouxia Kunth
- Lathraea L.
- Leptorhabdos Schrenk
- Leucosalpa Scott Elliot
- Lindenbergia Lehm.
- Macranthera Nutt. ex Benth.
- Magdalenaea Brade
- Mannagettaea Harry Sm.
- Melampyrum L.
- Melasma P.J.Bergius
- Micrargeria Benth.
- Micrargeriella R.E.Fr.
- Monochasma Maxim. ex Franch. & Sav.
- Neobartsia Uribe-Convers & Tank
- Nesogenes A.DC.
- Nothobartsia Bolliger & Molau
- Nothochilus Radlk.
- Odontitella Rothm.
- Odontites Ludw.
- Omphalotrix Maxim.
- Orobanche L.
- Orthocarpus Nutt.
- Parasopubia H.-P.Hofm. & Eb.Fisch.
- Parentucellia Viv.
- Pedicularis L.
- Phacellanthus Siebold & Zucc.
- Phelypaea L.
- Phtheirospermum Bunge ex Fisch. & C.A.Mey.
- Physocalyx Pohl
- Pseudobartsia D.Y.Hong
- Pseudosopubia Engl.
- Pterygiella Oliv.
- Radamaea Benth.
- Rehmannia Libosch. ex Fisch. & C.A.Mey.
- Rhamphicarpa Benth.
- Rhaphispermum Benth.
- Rhinanthus L.
- Rhynchocorys Griseb.
- Schwalbea L.
- Seymeria Pursh
- Seymeriopsis Tzvelev
- Sieversandreas Eb.Fisch.
- Silviella Pennell
- Siphonostegia Benth.
- Sopubia Buch.-Ham. ex D.Don
- Striga Lour.
- Tetraspidium Baker
- Thunbergianthus Engl.
- Tozzia L.
- Triphysaria Fisch. & C.A.Mey.
- Vellosiella Baill.
- Xylanche Beck
- Xylocalyx Balf.f.
First published in Tabl. Regn. Vég. 2: 292. 1799 [5 May 1799] (as "Orobanchoideae") (1799)
- APG IV (2016) http://dx.doi.org/10.1111/boj.12385
Flora of Tropical East Africa
Flora of Tropical East Africa
Flora of West Tropical Africa
Flora of West Tropical Africa
Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2019. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0
Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.