Research Article |
Corresponding author: Adam C. Schneider ( acschneider@berkeley.edu ) Academic editor: Eberhard Fischer
© 2016 Adam C. Schneider.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Schneider AC (2016) Resurrection of the genus Aphyllon for New World broomrapes (Orobanche s.l., Orobanchaceae). PhytoKeys 75: 107-118. https://doi.org/10.3897/phytokeys.75.10473
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Recent phylogenetic studies support a monophyletic clade of New World broomrapes (Orobanche sects. Gymnocaulis and Nothaphyllon) sister to the Old World genus Phelipanche. I place the New World taxa in the genus Aphyllon, propose 21 new combinations, and provide a list of currently accepted taxa.
Aphyllon , broomrape, Gymnocaulis , Myzorrhiza , Nothaphyllon , nomenclature, Orobanche , Orobanchaceae
Phylogenetic analysis of broomrapes and related holoparasites using nuclear DNA have found that the small eastern Mediterranean genus Diphelypaea Nicolson is nested within Orobanche sensu lato (s.l.) as circumscribed by
Broomrape species native to the New World constitute two well-supported clades that together form a clade sister to Phelipanche (
In order to compare molecular branch lengths of major clades of Orobanche s.l., a maximum likelihood (ML) phylogram of Aphyllon and related holoparasites was inferred from 3 nuclear DNA loci (ITS, phytochrome A, and phytochrome B). All sequences were downloaded from Genbank, aligned, and concatenated into a supermatrix using SUMAC (
Information about type specimens, basionyms, and synonomy of these new combinations was gathered by examining protologues and images of type specimens using major databases, including Tropicos (http://www.tropicos.org), JSTOR Global Plants (http://plants.jstor.org), and the International Plant Names Index (http://www.ipni.org). Types for all North American taxa and O. weberbaueri Mattf. have been designated by previous authors and are presented here. For three of the four South American taxa, typification would require more careful efforts beyond the scope of this article. No repository is given in the protologue for two syntypes of Orobanche tacnaensis Mattf. (Woitschach 71 and F. J. F. Meyen s.n.). The current existence of these specimens could not be verified, although a photograph of the Woitscach 71 (possibly from a specimen at B) is available at F. No specimens are cited by Rodolfo Phillipi in the protologues of the two taxa that he described.
Molecular phylogenetic analyses have consistently supported a sister-group relationship between two strongly supported two American clades, representing O. sect. Gymnocaulis and O. sect. Nothaphyllon (
Maximum likelihood (ML) phylogram of Aphyllon and related holoparasite species inferred from 3 nuclear DNA loci (ITS, phytochrome A and phytochrome B). Bootstrap support values >70% are labeled. Due to space constraints, several clades have been collapsed. For a more detailed and thorough study of phylogenetic relationships within Aphyllon, see
The genus Aphyllon was described by
Due to the biogeographical, morphological, cytological, and phylogenetic affinities of the New World broomrapes, I recommend treating them in a single genus, Aphyllon, composed of sections Aphyllon (= O. sect. Gymnocaulis) and Nothaphyllon (= O. sect. Nothaphyllon). Below, I present a key to sections and a list of recognized taxa in Aphyllon, proposing new combinations as necessary. Combinations are made at the most recently treated rank for the taxon in Orobanche, with the exception of Orobanche uniflora subsp. occidentale Greene, which is recognized at species rank under the available name Aphyllon purpureum (A. Heller) Holub due to its unique hosts, long molecular branch lengths, and recent discovery of sympatric populations of A. purpureum and A. uniflorum in southwestern British Columbia (
1 | Bracteoles subtending the calyx absent; pedicels much longer than flower (2-8× length); stems subterranean or rising to about ground level | Aphyllon sect. Aphyllon (syn.: O. sect. Gymnocaulis) |
1’ | Bracteoles subtending the calyx 2; pedicels equal to or shorter than flower, occasionally 2× length; stems usually rising above ground level | Aphyllon sect. Nothaphyllon (syn.: O. sect. Nothaphyllon) |
Loxanthes Raf. Neogenyton 3. 1825. [Type: Loxanthes fasciculatus (Nutt.) Raf.]
Anoplanthus Endl., nom. superfl., Gen. Pl. [Endlicher] pt. 10: 727. 1839.
Thalesia Raf. ex Britton, nom. superfl., Mem. Torrey Bot. Club 5: 298. 1894.
Aphyllon uniflorum (L.) Torr. & A. Gray, Manual 290. 1848.
Herb, annual or rarely perennial, achlorophyllous, holoparasitic. Stems fleshy. Leaves reduced to scale-like bracts. Inflorescences terminal racemes, spikes, corymbs, or panicles. Calyx 5-toothed. Corolla sympetalous, bilabiate to regular, tubular and often curved. Style long, stigma crateriform and peltate, or bilamellar. Fruit loculicidal capsules.
About 22 species: 18 in North America, 4 in South America.
Orobanche sect. Gymnocaulis Nutt., Gen. N. Amer. Pl. [Nuttall]. 2: 59. 1818.
Stems subterranean or rising to about ground level. Pedicels long and slender, much longer than flower. Bracteoles subtending the calyx absent.
Orobanche fasciculata Nutt., Gen. N. Amer. Pl. 2: 59. 1818.
Phelipaea fasciculata (Nutt.) Spreng., Syst. Veg. [Sprengel] 2: 818. 1825.
Loxanthes fasciculatus (Nutt.) Raf., Neogenyt. 3. 1825.
Anoplon fasciculatum (Nutt.) G. Don., Gen. Hist. 4: 633. 1838.
Anoplanthus fasciculatus (Nutt.) Walp., Repert. Bot. Syst. 3: 480. 1844.
Thalesia fasciculata (Nutt.) Britton, Mem. Torrey Bot. Club 5: 298. 1894.
USA: “Missouri”, ca. 1811, Nuttal s.n., (holotype,
Thalesia purpurea A. Heller, Bull. Torrey Bot. Club 24: 313. 1896.
Orobanche porphyrantha Beck, Pflanzenr. 96[IV,261]: 49. 1930.
Orobanche uniflora var. purpurea (A. Heller) Achey, Bull. Torrey Bot. Club 60: 445. 1933.
USA: Idaho: Nez Perce Co.: near mouth of the Potlatch, 20 May 1896, Heller 3099. (no holotype designated; isotypes,
Orobanche uniflora L., Sp. Pl. 2: 633. 1753.
Anoplanthus uniflorus (L.) Endl., Gen. Pl. [Endlicher] 727. 1839.
Thalesia uniflora (L.) Britton, Mem. Torrey Bot. Club 5: 298. 1894.
USA: Virginia (lectotype, Clayton 387,
Myzorrhiza Phil., Linnea 29: 36. 1858. [Type: Myzorrhiza chilensis Phil.]
Orobanche sect. Myzorrhiza Beck, Bibliotheca Botanica 4(19): 78. 1890.
Orobanche sect. Nothaphyllon (A. Gray) Heckard, Madroño 22: 41. 1973.
Aphyllon californicum (Cham. & Schltdl.) A. Gray, lectotype designated by Heckard, Madroño 22: 41. 1973.
Stems clearly rising above ground. Pedicels equal to or shorter than flower. Bracteoles subtending the calyx 1 or 2.
Orobanche arizonica L.T. Collins, Phytoneuron 2015–48: 16, f. 1, 2, 4, 5, 6A, 7. 2015.
USA: Arizona, Coconino Co.: near Tuba City, 1539 m, 27 September 1935, Kearney & Peebles 12867 (holotype,
Orobanche californica Cham. & Schltdl., Linnea 3: 134–136. 1828.
Phelypaea californica (Cham. & Schltdl.) G. Don, Gen. Hist. 4: 632. 1838.
Myzorrhiza californica (Cham. & Schltdl.) Rydb., Bull. Torrey Bot. Club 36: 696. 1909.
USA: California: Near Port of San Francisco, Aug 1816, Chamisso s.n (holotype,
Orobanche californica subsp. condensa Heckard, Madroño 22: 59–60, f. 1I-L, 5. 1973.
USA: California: San Luis Obispo Co.: Yaro Creek, 25 May 1955, Bacigalupi, Ferris & Robbins 5242 (holotype,
Orobanche grayana var. feudgei Munz, Bull. Torrey Bot. Club 57: 616–617, pl. 38, f. 8. 1930.
Orobanche californica subsp. feudgei (Munz) Heckard, Madroño 22: 62. 1973.
USA: California: San Bernardino Co.: Baldwin Lake, 2 June 1924, Munz 8177 (holotype,
Orobanche californica subsp. grandis Heckard, Madroño 22: 60–62, f. 1P-R, 3A, 4E, 5. 1973.
USA: California: Santa Barbara Co.: dunes at Surf, 22 July 1954, H. M. Pollard (holotype,
Orobanche grayana Beck, Biblioth. Bot. 4: 79. 1890.
Myzorrhiza grayana (Beck) Rydb., Bull. Torrey Bot. Club 36: 695. 1909.
Orobanche californica subsp. grayana (Beck) Heckard, Madroño 22: 54. 1973.
USA: Oregon: banks of the Columbia River, 1825, Douglas s.n. (lectotype, K).
Orobanche grayana var. jepsonii Munz, Bull. Torrey Bot. Club 57: 617, pl. 38, f. 10. 1930.
Orobanche californica subsp. jepsonii (Munz) Heckard, Madroño 22: 57. 1973.
USA: California: Colusa Co.: Princeton, October 1905, H. P. Chandler s.n. (holotype:
Myzorrhiza chilensis Phil., Linnea 29: 36–37. 1857.
Orobanche chilensis (Phil.) Beck, Biblioth. Bot. 4: 82–83. 1890.
Orobanche ludoviciana var. cooperi (A. Gray) Beck, Biblioth. Bot. 4(Heft 19): 81. 1890.
Orobanche cooperi (A. Gray) A. Heller, Cat. N. Amer. Pl. 7. 1898.
Myzorrhiza cooperi (A. Gray) Rydb. Bull. Torrey Bot. Club 36: 695. 1909.
USA: Arizona: Fort Mojave (lectotype designated by Munz, Bull. Torrey Bot. Club 57: 620-21, Cooper s.n. in 1860-61,
Orobanche ludoviciana var. latiloba Munz, Bull. Torrey Bot. Club 57: 621–622, pl. 39, f. 18. 1930.
Orobanche cooperi subsp. latiloba (Munz) L.T. Collins, Phytoneuron 2015–48: 15. 2015.
USA: California: Riverside Co.: Colorado Desert, 22 April 1922, Munz & Keck 4960 (holotype:
Orobanche multicaulis var. palmeri Munz, Bull. Torrey Bot. Club 57: 613, pl. 38, f. 2. 1930.
Orobanche cooperi subsp. palmeri (Munz) L.T. Collins, Phytoneuron 2015–48: 16. 2015.
Mexico, Durango, April-November 1896, Palmer 7 (holotype:
Myzorrhiza corymbosa Rydb., Bull Torrey Bot. Club 36: 696. 1909.
Orobanche corymbosa (Rydb.) Ferris, Contr. Dudley Herb. 5: 99. 1958.
USA: Reynold’s Creek, 2 July 1892, Isabel Mulford s.n. (holotype,
Orobanche corymbosa subsp. mutabilis Heckard, Canad. J. Bot. 56: 187–188. 1978.
USA: Washington: Grant Co.: O’Sullivan Dam, 11 July 1950, S. W. Harris 97 (holotype,
Orobanche dugesii (S. Watson) Munz, Bull. Torrey Bot. Club 57: 613, t. 38, f. 3. 1931.
Mexico: Gueanajatao, Dugès s.n. (holotype,
Orobanche ludoviciana Nutt. Gen. N. Amer. Pl. 2: 58–59. 1818.
Phelypaea ludoviciana (Nutt) Walp. Repert. Bot. Syst. 3: 461. 1844.
Myzorrhiza ludoviciana (Nutt) Rydb. Fl. S.E. U.S 1338. 1903.
USA: Fort Mandan, 1810-1811, Nuttall s.n. (holotype,
Orobanche multiflora Nutt., J. Acad. Nat. Sci. Philadelphia, ser. 2 1: 179. 1848.
USA: Rio Grande, 1845, Gambel s.n. (neotype designated by White & Holmes, Sida 19: 623, USA: Texas: Jim Wells Co., 19 April 1944, Lundell & Lundell 12809,
Orobanche californica var. parishii Jeps. Man. Fl. Pl. Calif. 952. 1925.
Orobanche parishii (Jeps.) Heckard. Madroño 22: 66. 1973.
USA: California: San Bernardino Co.: Bear Valley, 1894, S. B. Parish s.n. (holotype,
Orobanche parishii subsp. brachyloba Heckard, Madroño 22: 68–70, 2J, 3N, 5. 1973.
USA: California: Ventura Co.: Dutch Harbor, San Nicolas Island, 23 April 1966, Raven & Thompson 20794 (holotype,
Orobanche pinorum Geyer ex Hook., Hooker’s J. Bot. Kew Gard. 3:297–298. 1851.
USA: Idaho/Washington border, Geyer 445 (holotype, K).
Orobanche riparia L.T. Collins, J. Bot. Res. Inst. Texas 3: 7–10, f. 1A-B, 2. 2009.
USA: Indiana, Gibson Co.: Griffin, 16 August 1931, Deam 50941 (holotype,
Orobanche robbinsii Heckard ex Colwell & Yatsk., Phytoneuron 2016-58: 2. 2016.
USA: California: San Francisco Co.: Lands End, 13 August 1956, Robbins 3707 (holotype,
Orobanche tacnaensis Mattf., Notizbl. Bot Gart. Berlin-Dahlem 8: 185–186. 1922.
Peru: Tacna, 1890, Woitschach 71 (photograph of type: F); Peru: Tacna, 1833, F. J. F. Meyen s.n.
Orobanche tarapacana Phil., Anales Mus. Nac. Santiago de Chile 1891: 69. 1891.
Phelypaea tuberosa A. Gray, Proc. Amer. Acad. Arts 7: 371. 1868.
Orobanche bulbosa Beck, Biblioth. Bot. 4: 83–84. 1890.
USA: California: Monterey Co: Gavilan Mountains, 1860-1862, Brewer 743 (holotype,
Orobanche valida Jeps., Madroño 1: 255–256. 1929.
Orobanche ludoviciana var. valida (Jeps.) Munz, Bull. Torrey Bot. Club 57: 621. 1930.
USA: California: Rock Creek, San Gabriel Mountains, 2 June 1923, F. W. Peirson 7937 (holotype:
Orobanche valida subsp. howellii Heckard & L.T Collins, Madroño 29: 98–100, f. 1A–E. 1982.
USA: California: Mendocino Co.: Impassable Rock, 14 July 1951, Donald V. Hemphill s.n. (holotype:
Orobanche comosa var. vallicola Jeps., Man. Fl. Pl. Calif. 952. 1925.
Orobanche vallicola (Jeps.) Heckard, Madroño 22: 64. 1973.
USA: California: Santa Clara Co.: Coyote, 14 October 1914, W. L. Jepson 6196 (holotype:
Orobanche weberbaueri Mattf., Notizbl. Bot Gart. Berlin-Dahlem 8: 185. 1922.
Peru: Camaná: Areuipa, Hafen Chala, 26 November 1915, A. Weberbauer 7185 (isotypes:
I thank Bruce Baldwin, Alison Colwell, Gerald Schneeweiss, John Strother, and George Yatskievych for reviewing earlier versions of this article and for nomenclatorial advice.