Orobanchaceae

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Template:Short description Template:More citations needed Template:Automatic taxobox

Orobanchaceae, the broomrapes, is a family of mostly parasitic plants of the order Lamiales, with about 90 genera and more than 2000 species.[1] Many of these genera (e.g., Pedicularis, Rhinanthus, Striga) were formerly included in the family Scrophulariaceae sensu lato.[2][3][4] With its new circumscription, Orobanchaceae forms a distinct, monophyletic family.[4] From a phylogenetic perspective, it is defined as the largest crown clade containing Orobanche major and relatives, but neither Paulownia tomentosa nor Phryma leptostachya nor Mazus japonicus.[5][6]

The Orobanchaceae are annual herbs or perennial herbs or shrubs, and most (all except Lindenbergia, Rehmannia and Triaenophora) are parasitic on the roots of other plants—either holoparasitic or hemiparasitic (fully or partly parasitic). The holoparasitic species lack chlorophyll and therefore cannot perform photosynthesis.

File:Striga bilabiata MS4167.jpg
Striga bilabiata
File:Broomrape (Cistanche tubulosa) Negev.jpg
Cistanche tubulosa
File:Castilleja purpurea.jpg
Castilleja purpurea
File:Pedicularis zeylanica-Silent Valley-2016-08-13-001.jpg
Pedicularis zeylanica

Description

Orobanchaceae is the largest of the 20–28 dicot families that express parasitism.[7] Apart from a few non-parasitic taxa, the family displays all types of plant parasitism: facultative parasite, obligate parasite, hemiparasites, and holoparasites.

Roots and stems

Parasitic plants are attached to their host by means of haustoria, which transfer nutrients from the host to the parasite. Only the hemiparasitic species possess an additional extensive root system referred to as the lateral or side haustoria. In most holoparasitic species there is a swollen mass of short, bulky roots or one big swollen haustorial organ, which may be simple or composite, commonly called the terminal or primary haustorium.[8]

Plants are reduced to short vegetative stems, their alternate leaves are reduced to fleshy, tooth-like scales, and have multicellular hairs interspersed with glandular hairs.[9]

The hemiparasitic species (transferred from Scrophulariaceae) with green leaves are capable of photosynthesis, and may be either facultative or obligate parasites.

Flowers

The hermaphroditic flowers are bilaterally symmetrical and grow either in racemes or spikes or singly at the apex of the slender stem. The tubular calyx is formed by 2–5 united sepals. There are five united, bilabiate petals forming the corolla and they may be yellowish, brownish, purplish, or white. The upper lip is two-lobed, the lower lip is three-lobed. There are two long and two short stamens on slender filaments, inserted below the middle, or at the base of the corolla tube, alternating with the lobes of the tube. A fifth stamen is either sterile or lacking completely. The anthers dehisce via longitudinal slits. The pistil is one-celled. The ovary is superior. The flowers are pollinated by insects or birds (e.g., hummingbirds, as in Castilleja).

Fruits

The fruit is a dehiscent, non-fleshy, 1-locular capsule with many very minute endospermic seeds. Fruits of Orobanchaceae are small and abundant and can produce between 10,000–1,000,000 seeds per plant.[10] These are dispersed by the wind over long distances, which increases their chances of finding a new host.

Taxonomy

The taxon was first described in 1799 by Étienne Pierre Ventenat as Orobanchoideae. The family name Orobanchaceae is a conserved name.[11][12] Despite the similar morphological traits found in both Scrophulariaceae and Orobanchaceae, the latter is now considered a separate monophyletic taxon, on both molecular and mophological grounds. The 2016 APG IV system expanded Orobanchaceae to include genera previously placed in Scrophulariaceae, so that the family absorbed the former Lindenbergiaceae and Rehmanniaceae.[13] These two former families may be treated as tribes.[14] Molecular phylogenetic studies show that they are sisters to the other Orobanchaceae genera:[15][16] Template:Barlabel

Evolution

Development of the haustoria was a significant evolutionary event that allowed for the advancement of parasitic plants. The holoparasitic clade, Orobanche, delineates the first transition from hemiparasitism to holoparasitism within Orobanchaceae.Script error: No such module "Unsubst".

Genomics

The parasitism and its different modes have been suggested to have an impact on genome evolution, with increased DNA substitution rates in parasitic organisms compared to non-parasitic taxa.[17] For example, holoparasite taxa of Orobanchaceae exhibit faster molecular evolutionary rates than confamilial hemiparasites in three plastid genes.[18]

In a study comparing the rates of molecular evolution of parasitic versus non parasitic taxa for 12 pairs of angiosperm families — including Apodanthaceae, Cytinaceae, Rafflesiaceae, Cynomoriaceae, Krameriaceae, Mitrastemonaceae, Boraginaceae, Orobanchaceae, Convolvulaceae, Lauraceae, Hydnoraceae, and Santalaceae/Olacaceae —, parasitic taxa evolve on average faster than their close relatives for mitochondrial, plastid, and nuclear genome sequences.[19] Whereas Orobanchaceae fit to this trend for plastid DNA, they appear to evolve slower than their non parasitic counterpart in comparisons involving nuclear and mitochondrial DNA.[19]

Genera

Template:As of, Plants of the World Online accepted 99 genera.[20] Three further genera are accepted by other sources, and are included in the following list. Template:Div col

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  • Agalinis Raf.Script error: No such module "Check for unknown parameters".
  • Alectra Thunb.Script error: No such module "Check for unknown parameters".
  • Anisantherina PennellScript error: No such module "Check for unknown parameters".
  • Aphyllon Mitch.Script error: No such module "Check for unknown parameters".[21]
  • Asepalum MaraisScript error: No such module "Check for unknown parameters".
  • Aureolaria Raf.Script error: No such module "Check for unknown parameters".
  • Bardotia Eb.Fisch., Schäferh. & Kai Müll.Script error: No such module "Check for unknown parameters".
  • Bartsia L.Script error: No such module "Check for unknown parameters".
  • Baumia Engl. & GilgScript error: No such module "Check for unknown parameters".
  • Bellardia All.Script error: No such module "Check for unknown parameters".
  • Boschniakia C.A.Mey. ex BongardScript error: No such module "Check for unknown parameters".
  • Brachystigma PennellScript error: No such module "Check for unknown parameters".
  • Brandisia Hook.f. & ThomsonScript error: No such module "Check for unknown parameters".
  • Buchnera L.Script error: No such module "Check for unknown parameters".
  • Bungea C.A.Mey.Script error: No such module "Check for unknown parameters".
  • Buttonia McKen ex Benth.Script error: No such module "Check for unknown parameters".
  • Campbellia WightScript error: No such module "Check for unknown parameters".
  • Castilleja Mutis ex L.f.Script error: No such module "Check for unknown parameters".
  • Centranthera R.Br.Script error: No such module "Check for unknown parameters".
  • Chloropyron BehrScript error: No such module "Check for unknown parameters".
  • Christisonia GardnerScript error: No such module "Check for unknown parameters".
  • Cistanche Hoffmanns. & LinkScript error: No such module "Check for unknown parameters".
  • Conopholis Wallr.Script error: No such module "Check for unknown parameters".
  • Cordylanthus Nutt. ex Benth.Script error: No such module "Check for unknown parameters".
  • Cyclocheilon Oliv.Script error: No such module "Check for unknown parameters".
  • Cycniopsis Engl.Script error: No such module "Check for unknown parameters".
  • Cycnium E.Mey. ex Benth.Script error: No such module "Check for unknown parameters".
  • Cymbaria L.Script error: No such module "Check for unknown parameters".
  • Dasistoma Raf.Script error: No such module "Check for unknown parameters".
  • Dicranostegia (A.Gray) PennellScript error: No such module "Check for unknown parameters".
  • Epifagus Nutt.Script error: No such module "Check for unknown parameters".
  • Eremitilla Yatsk. & J.L.Contr.Script error: No such module "Check for unknown parameters".
  • Escobedia Ruiz & Pav.Script error: No such module "Check for unknown parameters".
  • Esterhazya J.C.MikanScript error: No such module "Check for unknown parameters".
  • Euphrasia L.Script error: No such module "Check for unknown parameters".
  • Gerardiina Engl.Script error: No such module "Check for unknown parameters".
  • Ghikaea Volkens & Schweinf.Script error: No such module "Check for unknown parameters".
  • Gleadovia Gamble & PrainScript error: No such module "Check for unknown parameters".
  • Graderia Benth.Script error: No such module "Check for unknown parameters".
  • Harveya Hook.Script error: No such module "Check for unknown parameters".
  • Hedbergia MolauScript error: No such module "Check for unknown parameters".
  • Hiernia S.MooreScript error: No such module "Check for unknown parameters".
  • Hyobanche L.Script error: No such module "Check for unknown parameters".
  • Kopsiopsis (Beck) BeckScript error: No such module "Check for unknown parameters".
  • Lamourouxia KunthScript error: No such module "Check for unknown parameters".
  • Lathraea L.Script error: No such module "Check for unknown parameters".
  • Leptorhabdos Schrenk ex Fisch. & C.A.Mey.Script error: No such module "Check for unknown parameters".
  • Leucosalpa Scott ElliotScript error: No such module "Check for unknown parameters".
  • Lindenbergia Lehm.Script error: No such module "Check for unknown parameters".
  • Macranthera Nutt. ex Benth.Script error: No such module "Check for unknown parameters".
  • Magdalenaea BradeScript error: No such module "Check for unknown parameters".
  • Mannagettaea Harry Sm.Script error: No such module "Check for unknown parameters".
  • Melampyrum L.Script error: No such module "Check for unknown parameters".
  • Melasma P.J.BergiusScript error: No such module "Check for unknown parameters".
  • Micrargeria Benth.Script error: No such module "Check for unknown parameters".
  • Micrargeriella R.E.Fr.Script error: No such module "Check for unknown parameters".
  • Monochasma Maxim. ex Franch. & Sav.Script error: No such module "Check for unknown parameters".
  • Neobartsia Uribe-Convers & TankScript error: No such module "Check for unknown parameters".
  • Nesogenes A.DC.Script error: No such module "Check for unknown parameters".
  • Nothobartsia Bolliger & MolauScript error: No such module "Check for unknown parameters".
  • Nothochilus Radlk.Script error: No such module "Check for unknown parameters".
  • Odontitella Rothm.Script error: No such module "Check for unknown parameters".
  • Odontites Ludw.Script error: No such module "Check for unknown parameters".
  • Omphalotrix Maxim.Script error: No such module "Check for unknown parameters".
  • Orobanche L.Script error: No such module "Check for unknown parameters".
  • Orthocarpus Nutt.Script error: No such module "Check for unknown parameters".
  • Parasopubia H.-P.Hofm. & Eb.Fisch.Script error: No such module "Check for unknown parameters".
  • Parentucellia Viv.Script error: No such module "Check for unknown parameters".
  • Pedicularis L.Script error: No such module "Check for unknown parameters".
  • Phacellanthus Siebold & Zucc.Script error: No such module "Check for unknown parameters".
  • Phelipanche PomelScript error: No such module "Check for unknown parameters".[21]
  • Phelypaea Tourn. ex L.Script error: No such module "Check for unknown parameters".
  • Phtheirospermum Bunge ex Fisch. & C.A.Mey.Script error: No such module "Check for unknown parameters".
  • Physocalyx PohlScript error: No such module "Check for unknown parameters".
  • Pseudobartsia D.Y.HongScript error: No such module "Check for unknown parameters".
  • Pseudomelasma Eb.Fisch.Script error: No such module "Check for unknown parameters".
  • Pseudosopubia Engl.Script error: No such module "Check for unknown parameters".
  • Pseudostriga BonatiScript error: No such module "Check for unknown parameters".
  • Pterygiella Oliv.Script error: No such module "Check for unknown parameters".
  • Radamaea Benth.Script error: No such module "Check for unknown parameters".
  • Rehmannia Libosch. ex Fisch. & C.A.Mey.Script error: No such module "Check for unknown parameters".
  • Rhamphicarpa Benth.Script error: No such module "Check for unknown parameters".
  • Rhaphispermum Benth.Script error: No such module "Check for unknown parameters".
  • Rhinanthus L.Script error: No such module "Check for unknown parameters".
  • Rhynchocorys Griseb.Script error: No such module "Check for unknown parameters".
  • Schwalbea L.Script error: No such module "Check for unknown parameters".
  • Seymeria PurshScript error: No such module "Check for unknown parameters".
  • Seymeriopsis TzvelevScript error: No such module "Check for unknown parameters".
  • Sieversandreas Eb.Fisch.Script error: No such module "Check for unknown parameters".
  • Silviella PennellScript error: No such module "Check for unknown parameters".
  • Siphonostegia Benth.Script error: No such module "Check for unknown parameters".
  • Sopubia Buch.-Ham. ex D.DonScript error: No such module "Check for unknown parameters".
  • Striga Lour.Script error: No such module "Check for unknown parameters".
  • Tetraspidium BakerScript error: No such module "Check for unknown parameters".
  • Thunbergianthus Engl.Script error: No such module "Check for unknown parameters".
  • Tozzia L.Script error: No such module "Check for unknown parameters".
  • Triaenophora Soler.Script error: No such module "Check for unknown parameters".[21]
  • Triphysaria Fisch. & C.A.Mey.Script error: No such module "Check for unknown parameters".
  • Vellosiella Baill.Script error: No such module "Check for unknown parameters".
  • Xylanche BeckScript error: No such module "Check for unknown parameters".
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Genera by life history trait

Orobanchaceae genera listed according to their life history trait.Script error: No such module "Unsubst".

Non-parasitic

Hemiparasitic

Template:Div col

Template:Div col end

Holoparasitic

Template:Div col

Template:Div col end

Distribution

The family Orobanchaceae has a cosmopolitan distribution, found mainly in temperate Eurasia, North America, South America, parts of Australia, New Zealand, and tropical Africa. The only exception to its distribution is Antarctica, though some genera may be found in subarctic regions.[22]

Ecology

This family has tremendous economic importance because of the damage to crops caused by some species in the genera Orobanche and Striga. They often parasitize cereal crops like sugarcane, maize, millet, sorghum, and other major agricultural crops like cowpea, sunflower, hemp, tomatoes, and legumes. Because of the ubiquitous nature of these particular parasites in developing countries, it is estimated to affect the livelihood of over 100 million people, killing 20 to 100 percent of crops depending on infestation.[23]

Some genera, especially Cistanche and Conopholis, are threatened by human activity, including habitat destruction and over-harvesting of both the plants and their hosts.

Research for this plant family can often be difficult due to its permit requirements for collection, travel, and research.

Notes

Template:Notelist

References

Template:Reflist

External links

Template:Sister project

Template:Angiosperm families Template:Taxonbar Template:Authority control

  1. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named McNeal2013
  2. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named dePamphilis1997
  3. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named Young1999
  4. ↑ a b Cite error: Invalid <ref> tag; no text was provided for refs named Olmstead2001
  5. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named Xia2009
  6. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named deQueiroz2020
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  10. ↑ Script error: No such module "citation/CS1".
  11. ↑ Template:IPNI
  12. ↑ Script error: No such module "citation/CS1".
  13. ↑ Script error: No such module "Citation/CS1".
  14. ↑ Stevens, P.F. (2001 onwards). "Orobanchaceae". Angiosperm Phylogeny Website. Retrieved 2025-02-28.
  15. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named LiFengRandSchn19
  16. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named MortBoykBeauTank22
  17. ↑ Script error: No such module "Citation/CS1".
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  19. ↑ a b Script error: No such module "Citation/CS1".
  20. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named POWO_30003446-2
  21. ↑ a b c Stevens, P.F. (2001 onwards). List of Genera in Orobanchaceae. Angiosperm Phylogeny Website. Retrieved 2025-02-28.
  22. ↑ Script error: No such module "Citation/CS1".
  23. ↑ Script error: No such module "Citation/CS1".
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