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Description

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Plants rupestral or terrestrial. Rhizomes compact to long-creeping, ascending or horizontal, scaly. Rhizome scales lanceolate to acicular, concolorous (tan to dark brown) or bicolorous (with dark central stripe and brown margins). Leaf vernation non-circinate to circinate. Petiolescastaneous to black, scaly and/or pubescent, rarely almost glabrous. Rachises terete or flattened or grooved adaxially, with indument similar to that of the petioles. Blades2- to 4-pinnate (rarely pinnate-pinnatifid), lanceolate to ovate-deltate, occasionally linear or pentagonal; adaxial surfaces glabrous or pubescent; abaxial surfaces scaly and/or pubescent or rarely glabrous. Ultimate segments round to oblong-ovate, minute to >1 cm long, the veins obscure and not ending in prominent hydathodes. Segment margins usually recurved, with a poorly differentiated false indusium (strongly differentiated in Myriopteris lendigera and Myriopteris marsupianthes). Sori usually partly to completely covered by the recurved segment margins, the sporangia clustered at vein tips. Sporangia 64-spored (in sexual species) or 32-spored (in apomicts). Spores globose-tetrahedral, tan to brown, cristate to rugulate. Chromosome numbers n = 29, 30, 58, 60 (sexual species); n = 2n = 87, 90 (apomictic triploids); n = 2n = 120 (apomictic tetraploids).
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Amanda Lee Grusz, Michael Dennis Windham
bibliographic citation
Grusz A, Windham M (2013) Toward a monophyletic Cheilanthes: The resurrection and recircumscription of Myriopteris (Pteridaceae) PhytoKeys 32: 49–64
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Amanda Lee Grusz
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Michael Dennis Windham
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Phytokeys (archived)

Distribution

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Species of Myriopteris range from southern Canada through the Caribbean and Central America to southern Chile, with one species (Myriopteris rawsonii) endemic to Namibia and South Africa. Mexico is the center of species diversity for the genus; 34 of the 44 species can be found in Mexico, and seven of these are endemic. 1) Myriopteris aemula (Maxon) Grusz & Windham, comb. nov. Cheilanthes aemula Maxon, Contr. U.S. Natl. Herb. 10: 495. 1908. Type: Mexico. Tamaulipas: Victoria, in river canyon, under overhanging rocks, altitude about 320 meters, February 1 to April 9, 1907, Palmer 187 (holotype: US; isotype: US). urn:lsid:ipni.org:names:77134841-1 2) Myriopteris alabamensis(Buckley) Grusz & Windham, comb. nov. Pteris alabamensis Buckley, Amer. J. Sci. Arts 45: 177. 1843. Cheilanthes alabamensis (Buckley) Kunze, Linnaea 20: 4. 1847. Type: USA. Alabama: Growing in tufts on limestone rocks that form the banks of the Tennessee River, at the foot of Muscle Shoals, Buckley s.n. (holotype: PH; isotypes: MO, NY). urn:lsid:ipni.org:names:77134842-1 3) Myriopteris allosuroides (Mett.) Grusz & Windham, comb. nov. Cheilanthes allosuroides Mett., Abh. Senckenberg. Naturf. Ges. 3: 78. 1859. Pellaea allosuroides (Mett.) Hieron., Hedwigia 62: 18. 1920. Type: Mexico, Schmitz s.n. (holotype: location unknown). urn:lsid:ipni.org:names:77134843-1 4) Myriopteris aurea (Poir.) Grusz & Windham, comb. nov. Pteris aurea Poir. Encyclopédie Méthodique, Botanique 5: 710. 1804. Type: Peru. Elle a été recueillie au Pérou par Joseph de Jussieu s.n. (sheet 1333 in hb. Jussieu; holotype: P). urn:lsid:ipni.org:names:77134844-1 Acrostichum bonariense Willd., Sp. Pl., ed. 4, 5(1): 114. 1810. Notholaena bonariensis (Willd.) C. Chr., Index Filic. 459. 1906. Cheilanthes bonariensis (Willd.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 15. 1953. In Cheilanthes, this has been called Cheilanthes bonariensis (Willd.) Proctor because use of the oldest applicable epithet (based on Pteris aurea Poir.) was blocked by the earlier publication of Cheilanthes aurea Baker (Proctor 1953). With the transfer of this species to Myriopteris we revert to the older epithet and thus avoid the typification difficulties associated with the basionym Acrostichum bonariense Willd. (Ponce and Zimmer 2011). 5) Myriopteris chipinquensis (Knobloch & Lellinger) Grusz & Windham, comb. nov. Cheilanthes chipinquensis Knobloch & Lellinger, Amer. Fern J. 59: 8. 1969. Type: Mexico. Nuevo Leon: Chipinque Mesa, outside Monterey, Knobloch 1996B (holotype: MSC; isotypes: F, GH, MEXU, MICH, UC, US). urn:lsid:ipni.org:names:77134845-1 6) Myriopteris cinnamomea(Baker) Grusz & Windham, comb. nov. Notholaena cinnamomea Baker in Hook. & Baker, Syn. Fil. ed. 2. 515. 1874. Cheilanthes cinnamomea (Baker) Domin., Biblioth. Bot. 20: 133. 1913. hom. illeg. non Cheilanthes cinnamomea D. C. Eaton, Proc. Amer. Acad. Arts 18: 186. 1883. Type: Guatemala. Mo[n]tagua, 1862, Salvin & Goodman s.n. (holotype: K; isotype: BM). urn:lsid:ipni.org:names:77134870-1 Cheilanthes tryonii T. Reeves, Brittonia 32: 504. 1980. In Cheilanthes, this species has been called Cheilanthes tryonii T. Reeves because use of the oldest applicable epithet (based on Notholaena cinnamomea Baker) was blocked by the earlier publication of Cheilanthes cinnamomea D. C. Eaton (Reeves 1980). With the transfer of this species to Myriopteris, we revert to the older epithet. 7) Myriopteris clevelandii (D. C. Eaton) Grusz & Windham, comb. nov. Cheilanthes clevelandii D. C. Eaton, Bull. Torrey Bot. Club 6: 33. 1875. Type: USA. California: Growing on a mountain about forty miles from San Diego at an elevation of about 2500 feet, Cleveland s.n. (holotype: YU; isotypes: GH, P, US). urn:lsid:ipni.org:names:77134846-1 8) Myriopteris cooperae (D. C. Eaton) Grusz & Windham, comb. nov. Cheilanthes cooperae D. C. Eaton, Bull. Torrey Bot. Club 6: 33. 1875. Type: USA. California: near Santa Barbara, Mrs. Ellwood Cooper (syntype: YU); Sierra Valley, Lemmon s.n. (syntype: YU). urn:lsid:ipni.org:names:77134847-1 9) Myriopteris covillei (Maxon) Á. Löve & D. Löve, Taxon 26: 325. 1977. Cheilanthes covillei Maxon, Proc. Biol. Soc. Wash. 31: 147. 1918. Type: USA. California: Surprise Canyon, Panamint Mountains, 13 April 1891, 1550 meters, Coville & Funston 593 (holotype: US). urn:lsid:ipni.org:names:77134848-1 10) Myriopteris cucullans (Fée) Grusz & Windham, comb. nov. Cheilanthes cucullans Fée, Mém. Fam. Foug. 7: 39, t. 25, f. 4. 1857. Type: Mexico, ad vallem Mexicanum, Schaffner 82 [holotype: RB; isotypes: K, US (fragment)]. urn:lsid:ipni.org:names:77134873-1 11) Myriopteris fendleri (Hook.) E. Fourn., Mex. Pl. 1: 125. 1872. Cheilanthes fendleri Hook., Sp. Fil. 2: 103, p. 107b. 1852. Type: USA. New Mexico, 1847, Fendler 1015 [holotype: K; isotypes: GH, MO, NY, US (fragment)]. 12) Myriopteris × fibrillosa(Davenp.) Grusz & Windham, comb. nov. Cheilanthes lanuginosa var. fibrillosa Davenp., Bull. Torrey Bot. Club 12: 21. 1885. Cheilanthes fibrillosa (Davenp.) Davenp., Bull. Torrey Bot. Club 15: 225. 1888. Type: USA. California: San Jacinto Mountains, June 1882, Parish & Parish s.n. (holotype: GH). urn:lsid:ipni.org:names:77134880-1 13) Myriopteris fimbriata (A. R. Sm.) Grusz & Windham, comb. nov. Cheilanthes microphylla (Sw.) Sw. var. fimbriata A. R. Sm., Amer. Fern J. 70: 19, 21., f. 9–10. 1980. Type: Mexico. Chiapas: Munic. Frontera Comalapa, 6–8 km east of Frontera Comalapa, Breedlove 39018 (holotype: DS). urn:lsid:ipni.org:names:77134881-1 Cheilanthes fimbriata (A. R. Sm.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 112. 1988. hom. illeg., non Cheilanthes fimbriata Vis., Fl. Dalmat. 1. 42 t. 1 f. 1. 1842. 14) Myriopteris gracilis Fée, Mém. Fam. Foug. 5: 150, t. 29, f. 6. 1852. Cheilanthes gracilis (Fée) Mett. ex Riehl, Abh. Senckenberg. Naturf. Ges. 80. 1859. hom. illeg., non Cheilanthes gracilis (Michx.) Kaulf., Enum. Filic. 209. 1824. Type: USA. Missouri: Jefferson County, Habitat ad rupes circa Hillsboro, Americâ septentr., Riehl 529 (isotypes: MO, US). Cheilanthes feei T. Moore, Index Fil., 38. 1857. Myriopteris lanuginosa J. Sm. Hist. Fil. 280. 1875. [non M. lanuginosa (Mart. & Gal.) E. Fourn. Mexic. Pl. 1: 125. 1872.] In Cheilanthes, this has been called Cheilanthes feei T. Moore because use of the oldest applicable epithet (based on Myriopteris gracilis Fée) was blocked by the earlier publication of Cheilanthes gracilis (Michx.) Kaulf. With the transfer of this species to Myriopteris, we revert to the original name published by Fée in 1852. 15) Myriopteris gracillima (D. C. Eaton) J. Sm., Hist. Fil. 280. 1875. Cheilanthes gracillima D. C. Eaton, Rep. U.S. Mex. Bound. Botany 2: 234. 1859. Type: USA. Oregon: Cascade Mountains, 7000 feet of altitude, latitude 44°, Bigelow s.n. (lectotype: YU). 16) Myriopteris intertexta (Maxon) Grusz & Windham, comb. nov. Cheilanthes covillei Maxon subsp. intertexta Maxon, Proc. Biol. Soc. Wash. 31: 149. 1918. Cheilanthes intertexta (Maxon) Maxon in Abrams, Ill. Fl. Pacific States 1: 28. 1923. Type: USA. California: Santa Clara County, Santa Cruz Mountains, collected at the top of Black Mountain, 6 July 1903, Dudley s.n. (holotype: DS). urn:lsid:ipni.org:names:77134849-1 17) Myriopteris jamaicensis (Maxon) Grusz & Windham, comb. nov. Cheilanthes jamaicensis Maxon, Contr. U.S. Natl. Herb. 24: 51. 1922. Type: Jamaica. Below Cinchona, 28 February 1919, Harris 12905 (holotype: US; isotypes: GH, MO, NY). urn:lsid:ipni.org:names:77134850-1 18) Myriopteris lanosa (Michx.) Grusz & Windham, comb. nov. Nephrodium lanosum Michx. Fl. Bor.-Amer. 2: 270. 1803. Cheilanthes lanosa (Michx.) D. C. Eaton, Rep. U.S. Mex. Bound., Botany 2: 234. 1859. Type: USA. Tennassee (sic) et Carolinae septentrionalis (non designatus). urn:lsid:ipni.org:names:77134851-1 Myriopteris vestita (Sw.) J. Sm., Cul. Ferns 29. 1857. (fide C. Chr. 1906.) Adiantum vestitum Spreng., Anleit. Kenntn. Gew. 3: 122. 1804. 19) Myriopteris lendigera (Cav.) Fée, Mém. Fam. Foug. 5: 149. 1852 (as Myriopteris lentigera). Pteris lendigera Cav., Descr. Pl. 268. 1801. Cheilanthes lendigera (Cav.) Sw., Syn. Fil. 128, 328. 1806. Type: Mexico. Hidalgo: Ixmiquilpan en la Nueva España, Nee s.n. [syntype: MA, US (fragment)]; Ecuador. Bolivar: junto á Guaranda en el Reyno de Quito, Nee s.n. (syntype: MA). Cheilanthes minor Mart. & Gal. Mém. Act. Brux. 75, pl. 21, f. 1. 1842. Myriopteris minor (Mart. & Gal.) Fée, Mém. Fam. Foug. 5: 150. 1852. Cheilanthes lanuginosa Mart. & Gal. Mém. Act. Brux. 75, pl. 20, f. 2. 1842. Myriopteris lanuginosa (Mart. & Gal.) E. Fourn. Mex. Pl. 1: 125. 1872. Myriopteris villosa Fée, Mém. Fam. Foug. 5: 149. t. 28, f. 1. 1852. Cheilanthes frigida Linden ex T. Moore, Gard. Chr. 772. 1857. Myriopteris frigida (Linden ex T. Moore) J. Sm. Cat. Cult. Ferns 28. 1857. Myriopteris lendigera (Cav.) J. Sm., Cat. Cult. Ferns 28. 1857. hom. illeg. Pomataphytum pocillatum M. E. Jones, Contributions to Western Botany 16: 12. 1930. 20) Myriopteris lindheimeri (Hook.) J. Sm., Bot. Voy. Herald. 340. 1856. Cheilanthes lindheimeri Hook., Sp. Fil. 2: 101, t. 107a. 1852. Type: USA. Western Texas, 1847, Lindheimer 744 [lectotype: K; isolectotypes: GH, P (2 sheets), SD, US, YU]. 21) Myriopteris longipila (Baker) Grusz & Windham, comb. nov. Cheilanthes longipila Baker, Ann. Bot. (Oxford) 5: 211. 1891. Type: Mexico. San Luis Potosí, 22°N Lat., 6000–8000 ft., Parry & Palmer 989 [holotype: K; isotype: US (fragment)]. urn:lsid:ipni.org:names:77134852-1 22) Myriopteris longipila subsp. brevipila (Mickel) Grusz & Windham, comb. nov. Cheilanthes longipila var. brevipila Mickel, Mem. New York Bot. Gard. 88: 198–199, f. 84N–Q, 87J–M. 2004. Type: Mexico. Guerrero: 2 km al SE de Amatitlán, 1600 m, 13 August 1994, Soto 1052 (holotype: NY; isotype: FCME). urn:lsid:ipni.org:names:77134882-1 23) Myriopteris marsupianthes Fée, Mém. Fam. Foug. 5: 149, t. 12A, f. 1. 1852. Cheilanthes marsupianthes (Fée) T. Reeves ex Mickel & A. R. Sm. Mem. New York Bot. Gard. 88: 201, f. 83M–P. 2004.Type: Mexico. Veracruz: Pic d’Orizaba, Martens & Galeotti 6256 (holotype: P; isotype: BR). 24) Myriopteris maxoniana (Mickel) Grusz & Windham, comb. nov. Cheilanthes maxoniana Mickel, Mem. New York Bot. Gard. 88: 201, f. 87A–D. 2004. Type: Mexico. Tamaulipas: San Lucas, Viereck 76 (holotype: US). urn:lsid:ipni.org:names:77134853-1 25) Myriopteris mexicana (Davenp.) Grusz & Windham, comb. nov. Cheilanthes mexicana Davenp., Bull. Torrey Bot. Club 15: 227. 1888. Type: Mexico. Chihuahua: on the verge of a high cliff near the summit of Potrero Peak (Santa Eulalia Mts.), October 1886, 7300 ft., Pringle 827 (holotype: GH; isotypes: MO, BR, DS, NY, P, UC, US, YU). urn:lsid:ipni.org:names:77134854-1 26) Myriopteris mickelii (T. Reeves) Grusz & Windham, comb. nov. Cheilanthes mickelii T. Reeves, Brittonia 32: 502, f. 1–5. 1980. Type: Mexico. Oaxaca: Distr. Yautepec, Mickel 4210 (holotype: NY; isotypes: MO, UC). urn:lsid:ipni.org:names:77134855-1 27) Myriopteris microphylla (Sw.) Grusz & Windham, comb. nov. Adiantum microphyllum Sw., Prodr. 135. 1788. Cheilanthes microphylla (Sw.) Sw., Syn. Fil. 127. 1806. Type: Jamaica, Swartz s.n. (holotype: S). urn:lsid:ipni.org:names:77134856-1 28) Myriopteris moritziana (Kunze) Grusz & Windham, comb. nov. Cheilanthes moritziana Kunze, Linnaea 23: 307. 1850. Type: Venezuela. Caracas: La Guayra, Moritz 263 (lectotype: B; isolectotype: GH). urn:lsid:ipni.org:names:77134857-1 29) Myriopteris myriophylla (Desv.) J. Sm., Bot. Voy. Herald, 340. 1856. Cheilanthes myriophylla Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 328. 1811. Type: South America. Anon. s.n. (holotype: P). Cheilanthes elegans Desv. Ges. Naturf. Freunde Berlin Mag. 5: 328. 1811. Myriopteris elegans (Desv.) J. Sm., Cat. Cult. Ferns 29. 1857. Cheilanthes paleacea Mart. & Gal., Mém. Foug. Mexique 76, pl. 21, f. 2. 1842. Myriopteris paleacea (Mart. & Gal.) Fée, Mém. Fam. Foug. 5: 149, t. 29, f. 6. 1852. Myriopteris intermedia E. Fourn., Bull. Soc. Bot. Fr. 27: 328. 1880. hom. illeg., non Fée, Mém. Fam. Foug. 5: 149. 1852. 30) Myriopteris newberryi(D. C. Eaton) Grusz & Windham, comb. nov. Notholaena newberryi D. C. Eaton, Bull. Torrey Bot. Club 4: 12. 1873. Cheilanthes newberryi (D. C. Eaton) Domin, Biblioth. Bot. 20: 133. 1913. Types: USA. California: San Diego, 9 November 1857, Newberry 1352 (syntype: MO, YU); San Diego, 1866, Wood s.n. (syntype: YU); Southern California: S. W. corner of San Bernardino County, rocks in the Temescal range, 22 January 1861, W. H. Brewer s.n. (syntype: YU). urn:lsid:ipni.org:names:77134858-1 31) Myriopteris notholaenoides (Desv.) Grusz & Windham, comb. nov. Pteris notholaenoides Desv., Mém. Soc. Linn. Paris 6: 299. 1827. Cheilanthes notholaenoides (Desv.) Maxon ex Weath., Contr. Gray Herb. 114: 34. 1936. Type: Hispaniola, Anon. s.n. (holotype: P). urn:lsid:ipni.org:names:77134859-1 32) Myriopteris × parishii (Davenp.) Grusz & Windham, comb. nov. Cheilanthes parishii Davenp., Bull. Torrey Bot. Club 8: 59. 1881. Type: USA. California: San Diego County, W. J. Parish s.n. (holotype: GH; isotypes: GH, YU). urn:lsid:ipni.org:names:77134860-1 33) Myriopteris parryi (D. C. Eaton) Grusz & Windham, comb. nov. Notholaena parryi D. C. Eaton, Amer. Naturalist 9: 351. 1875. Cheilanthes parryi (D. C. Eaton) Domin, Biblioth. 85: 133. 1913. Type: USA. Utah: C. C. Parry 263 (holotype: YU; isotypes: GH, US, YU). urn:lsid:ipni.org:names:77134861-1 34) Myriopteris peninsularis (Maxon) Grusz & Windham, comb. nov. Cheilanthes peninsularis Maxon, Contr. U.S. Natl. Herb. 10: 496. 1908. Type: Mexico. Baja California, T. S. Brandegee s.n. (holotype: US). urn:lsid:ipni.org:names:77134862-1 35) Myriopteris peninsularissubsp. insularis (Weath.) Grusz & Windham, comb. nov. Cheilanthes peninsularis (Maxon) var. insularis Weath., Amer. Fern J.21: 25. 1931. Type: Mexico. Socorro Island, Mason 1616 (holotype: CAS). urn:lsid:ipni.org:names:77134884-1 36) Myriopteris pringlei (Davenp.) Grusz & Windham, comb. nov. Cheilanthes pringlei Davenp., Bull. Torrey Bot. Club 10: 61, t. 34. 1883. Type: USA. Arizona: C. G. Pringle s.n. (holotype: GH; isotypes: DS, MO, NY, US, YU). urn:lsid:ipni.org:names:77134863-1 37) Myriopteris pringlei subsp. moncloviensis (Baker) Grusz & Windham, comb. nov. Cheilanthes moncloviensis Baker, Ann. Bot. (Oxford) 5: 210. 1891. Cheilanthes pringlei var. moncloviensis (Baker) Mickel, Mem. New York Bot. Gard. 88: 207–208, f. 79J–M. 2004. Type: Mexico. Coahuila: Soledad, E. Palmer 1378 (holotype: K; isotypes: MO, NY, US). urn:lsid:ipni.org:names:77134864-1 38) Myriopteris rawsonii(Mett. ex. Kuhn) Grusz & Windham, comb. nov. Cheilanthes rawsonii Mett. ex. Kuhn, Filices Africanae 75. 1868. Type: Africa. Cape Province: Namaqualand, between Specktakel and Komaggas, Whitehead s.n. (holotype: BM; isotype: K). urn:lsid:ipni.org:names:77134878-1 39) Myriopteris rufaFée, Mém. Fam. Foug. 8: 77. 1857. Type. Mexico. Veracruz: Volcan de Orizaba, Schaffner 83 (holotype: P?; isotype: RB?). Cheilanthes eatonii Baker in Hook. & Baker, Syn. Fil. 140. 1867. Cheilanthes castanea Maxon, Proc. Biol. Soc. Wash. 32: 111. 1919. In Cheilanthes, this has been called Cheilanthes eatonii Baker. Examination of putative type specimens of Myriopteris rufa housed at RB (digital image) and P indicates that the latter name very likely represents the same species as broadly defined by recent authors (e.g., Mickel and Smith 2004). Because Myriopteris rufa (published in 1857) has priority over Cheilanthes eatonii (1867), we take up Fée’s original name for this taxon in Myriopteris. 40) Myriopteris scabra (C. Chr.) Grusz & Windham, comb. nov. Pellaea scabra C. Chr., Index Filic. 483. 1906. Type: USA. Texas: crevices of rock on hills, Turkey Creek, 25 June 1849, Wright 824 (holotype: K; isotypes: GH, NY, US). Cheilanthes aspera Hook., Sp. Fil. 2: 111, t. 108A. 1852. hom. illeg., non Cheilanthes aspera Kaulf., Linnaea 6(1): 186. 1831. urn:lsid:ipni.org:names:77134865-1 Cheilanthes horridula Maxon, Amer. Fern J. 8: 94. 1918. In Cheilanthes, this has been called Cheilanthes horridula Maxon because use of the oldest legitimate epithet (based on Pellaea scabra C. Chr.) was blocked by the earlier publication of Cheilanthes scabra H. Karst. (Maxon 1918). With the transfer of this species to Myriopteris, we revert to the older, exceedingly appropriate epithet. 41) Myriopteris tomentosa (Link) Fée, Mém. Fam. Foug. 5: 149. 1852. Cheilanthes tomentosa Link, Hort. Berol. 2: 42. 1833. Type: Mexico. Anon. s.n. [holotype: B; isotypes: PH, US (fragment)]. Cheilanthes bradburii Hook., Sp. Fil. 2: 97, t. 109b. 1852. Myriopteris bradburii (Hook.) J. Sm. Hist. Fil. 280. 1875. 42) Myriopteris viscida(Davenp.) Grusz & Windham, comb. nov. Cheilanthes viscida Davenp., Bull. Torrey Bot. Club 6: 191. 1877. Types: USA. California: Eastern slope of the Sierra Nevada near San Gogorio Pass, April 1876, Parry & Lemmon 427 (syntype: NY); California/Nevada: Downieville Buttes and bluffs of White Water River on the Colorado Desert, April–May, Lemmon s.n. (syntype: NY). urn:lsid:ipni.org:names:77134866-1 43) Myriopteris windhamii Grusz, Amer. Fern J. 103: 113. 2013. Type: USA. Arizona: Huachuca Mountains, Windham 4165 (holotype: DUKE; isotypes: ARIZ, ASC, ASU, GH, MO, NMC, NY, TEX/LL, UNM, US, UT). Cheilanthes villosa Davenp. ex Maxon, Proc. Biol. Soc. Wash. 31: 142. 1918. In Cheilanthes, this has been called Cheilanthes villosa Davenp. ex Maxon. Because transfer of the epithet villosa to Myriopteris is blocked by the earlier publication of Myriopteris villosa Fée (= Myriopteris lendigera fide Reeves 1979), we use the replacement name for this distinctive taxon published by Grusz (2013). 44) Myriopteris wootonii (Maxon) Grusz & Windham, comb. nov. Cheilanthes wootonii Maxon, Proc. Biol. Soc. Wash. 3: 146. 1918. Type: USA. Arizona: Santa Rita Mountains, Wooton s.n. (holotype: US). urn:lsid:ipni.org:names:77134867-1 45) Myriopteris wrightii (Hook.) Grusz & Windham, comb. nov. Cheilanthes wrightii Hook., Sp. Fil. 2: 87, t. 110A. 1858. Type: USA. Texas–New Mexico: Wright 823 (holotype: K; isotypes: GH, NY, US). urn:lsid:ipni.org:names:77134868-1 46) Myriopteris yatskievychiana (Mickel) Grusz & Windham, comb. nov. Cheilanthes yatskievychiana Mickel, Mem. New York Bot. Gard. 88: 212–213, f. 74F–K. 2004. Type: Mexico. Sonora: Sierra del Aliso, A. Búrquez M. 96-302 (holotype: MO). urn:lsid:ipni.org:names:77134869-1 47) Myriopteris yavapensis (T. Reeves ex Windham) Grusz & Windham, comb. nov. Cheilanthes yavapensis T. Reeves ex Windham, Contr. Univ. Michigan Herb. 19: 32. 1993. Type: USA. Arizona: Yavapai County, Windham 202 (holotype: UT; isotypes: ASC, ASU, US). urn:lsid:ipni.org:names:77134879-1 Name of uncertain application Myriopteris cheiloglyphisFée, Mém. Fam. Foug. 8: 77. 1857. Excluded names Myriopteris contracta(Kunze) Fée, Mém. Fam. Foug. 5: 149. 1852. = Cheilanthes contracta (Kunze) Mett. ex Kuhn Myriopteris hirta(Sw.) J. Sm., Ferns Brit. and For. 174. 1866. = Cheilanthes hirta Sw. Myriopteris induta (Kunze) Fée, Mém. Fam. Foug. 5: 149. 1852. = Cheilanthes induta Kunze Myriopteris intermedia(Kunze) Fée, Mém. Fam. Foug. 5: 149. 1852. = Cheilanthes hirta Sw. fide Christensen (1906) Myriopteris macleaniiJ. Sm., Hist. Fil. 280. 1875. = Cheilanthes pilosa Goldm. fide Christensen (1906) Myriopteris scariosa(Sw.) Fée, Mém. Fam. Foug. 5: 149, t. 29, f. 6. 1852. = Cheilanthes scariosa Sw. Myriopteris szovitzii(Fisch. & Meyer) J. Sm., Hist. Fil. 281. 1875. = Cheilanthes persica (Bory) Mett. ex Kuhn fide Christensen (1906)
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Amanda Lee Grusz, Michael Dennis Windham
bibliographic citation
Grusz A, Windham M (2013) Toward a monophyletic Cheilanthes: The resurrection and recircumscription of Myriopteris (Pteridaceae) PhytoKeys 32: 49–64
author
Amanda Lee Grusz
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Michael Dennis Windham
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Phytokeys (archived)

Myriopteris

provided by wikipedia EN

Myriopteris, commonly known as the lip ferns, is a genus of cheilanthoid ferns. Like other cheilanthoids, they are ferns of dry habitats, reproducing both sexually and apogamously. Many species have leaves divided into a large number of small, bead-like segments, the probable inspiration for the generic name. Hairs and/or scales are often present on both the upper and lower surfaces of the leaf, and their presence and appearance are useful in distinguishing between species. The genus is most diverse in Mexico, but species are found from southwestern Canada south to southern Chile, and one species is endemic to southern Africa.

Description

No single morphological character divides Myriopteris, as presently circumscribed, from the other cheilanthoids. Convergent evolution in arid environments is thought to be responsible for widespread homoplasy in the morphological characters traditionally used to classify this group.[1] While small, bead-like ultimate segments are associated with the genus, they only appear in about 40% of its species,[2] and appear in some cheilanthoids outside the genus as well. Cheilanthes sensu stricto bears 32 spores per sporangium in sexual species and 16 in apogamous species; with the exception of a few species of Notholaena, Myriopteris and the other cheilanthoids bear 64 spores per sporangium when sexual and 32 per sporangium when apomictic. Myriopteris can also be separated from Cheilanthes s.s., although less reliably, by spores bearing crests or wrinkles (rather than spines or bumps) and a lack of enlarged vein endings (rather than prominent hydathodes).[1]

The three subgroups of the genus, informally called the alabamensis, covillei, and lanosa clades, likewise lack unique defining features. The small, bead-like ultimate segments are found in the core subclade of the covillei clade and also in M. gracilis, which is deeply nested in the alabamensis clade.[3] Circinate vernation (the unfolding of new leaves as fiddleheads) is found throughout the lanosa clade and also in M. wrightii, the most basal member of the alabamensis clade.[3] Most species have round rachises, although early-diverging members of the alabamensis and lanosa clades have rachises deeply grooved on the upper surface and flattened rachises shallowly grooved near the frond tip, respectively.[2] Leaf indument (hairs and scales) is highly diverse across the genus and a key feature in species identification.[2]

Myriopteris covillei has large, prominent scales beneath the leaf.
Myriopteris gracillima growing on basalt in the lower Columbia River gorge, WA

The base chromosome number for the genus appears to be x=30, except for a portion of the alabamensis clade, where x=29; the latter is typical for other cheilanthoids. Most species are either sexually reproducing diploids or apogamous triploids, with the exception of M. lendigera and some specimens of M. microphylla and M. scabra, which are sexually reproducing tetraploids.[4]

Taxonomy

The genus was first described in 1852 by A.L.A. Fée, who separated it from Cheilanthes proper by the presence of red hairs among the sporangia and a scarious (hardened) indusium formed from the leaf margin. He typified it on Myriopteris marsupianthes. Fée described the division of the leaf into numerous small, beadlike segments, all capable of bearing spores,[5] which may have led him to choose the name Myriopteris; "myrio-" means "very many"[6] and "pteris" means "fern". John Smith recognized Myriopteris in his Cultivated Ferns of 1857, noting the "minute, orbicular or cuneiform, concave" ultimate segments typical of species in the genus.[7] However, most authors until the 21st century preferred to include the genus in Cheilanthes.[8]

The development of molecular phylogenetic methods showed that the traditional broad circumscription of Cheilanthes is polyphyletic. Many of the morphological characters that have traditionally been used to separate the cheilanthoid ferns into genera, including Cheilanthes, are homoplasious; that is, they have appeared independently in unrelated groups, probably as a result of convergent evolution in arid environments.[1]

A molecular phylogeny of 157 cheilanthoid species by Michael D. Windham et al. in 2009 revealed seven well-supported clades within the group. One of these included a number of mostly North American species usually placed in Cheilanthes sensu lato. They informally referred to this group as the myriopterids, Myriopteris being the most senior genus whose type specimen was included in the clade.[9] Further phylogenetic analysis by Eiserhardt et al., in the course of a study on cheilanthoid evolutionary radiation in the Cape Floristic Region, also supported the existence of the myriopterid clade and showed that Cheilanthes rawsonii, an African endemic, was deeply nested in it, its closest relative being C. parryi.[10] After more extensive sampling within the clade, Amanda Grusz and Windham revived Myriopteris in 2013 and provided names in it for all species in the myriopterid clade.[1]

Further molecular studies by Grusz et al. confirmed the monophyly of this group, and showed that its species can be divided among three well-supported clades. These were informally referred to by the epithet of a prominent species in the clade, without formal taxonomic rank; they are the alabamensis clade (A), covillei clade (C), and lanosa clade (L).[11]

The genus Cheilosoria was described by Conde Vittore Trevisan in 1877 to accommodate Cheilanthes allosuroides, now M. allosuroides, and a few other species of Cheilanthes with long sori along the veins and relatively unmodified false indusia.[12] That genus was lectotypified on C. allosuroides by Edwin Copeland in 1947.[13] If the alabamensis clade were to be treated as a separate genus, Cheilosoria would be the senior name for it.[14]

Species

The following species (including two hybrids) are those recognized by Grusz & Windham in 2013, with some additions from the Checklist of Ferns and Lycophytes of the World (version 8.11).[15] Letters in parentheses following the scientific names indicate which of the three clades the species belong to, if known.

Distribution and habitat

Myriopteris parryi growing in a rock outcrop in Death Valley.

The greatest diversity of species occurs in Mexico.[16] Myriopteris species are mostly confined to the Americas, ranging from southwestern Canada[17] to southern Chile. M. rawsonii is endemic to Namibia and South Africa.[16] It is most closely related to M. parryi,[10] a species of the Sonoran and Mojave Deserts.[4]

Like other cheilanthoids, Myriopteris species occupy dry habitats, growing on rocks or soil.[18] The closely-related species M. gracillima, M. intertexta, M. covillei, and M. clevelandii [19] all grow mostly in rock crevices, with western North American ranges that overlap in increasingly hot and dry climates.[20]

Notes and references

References

  1. ^ a b c d Grusz & Windham 2013, pp. 53–54.
  2. ^ a b c Grusz et al. 2014, p. 709.
  3. ^ a b Grusz et al. 2014, p. 705.
  4. ^ a b Grusz et al. 2014, p. 708.
  5. ^ Fée 1852, p. 152.
  6. ^ Short & George 2013, p. 213.
  7. ^ Smith 1857, p. 28.
  8. ^ Grusz & Windham 2013, p. 53.
  9. ^ Windham et al. 2009, pp. 128–131.
  10. ^ a b Eiserhardt et al. 2011, p. 1274.
  11. ^ Grusz et al. 2014, pp. 702, 704.
  12. ^ Trevisan 1877, p. 579.
  13. ^ Copeland 1947.
  14. ^ Grusz & Windham 2013, p. 52.
  15. ^ Hassler, Michael & Schmitt, Bernd (November 2019). "Myriopteris". Checklist of Ferns and Lycophytes of the World. 8.11. Retrieved 2020-01-01.
  16. ^ a b Grusz & Windham 2013, p. 55.
  17. ^ Kartesz 2014.
  18. ^ Grusz & Windham 2013, p. 54.
  19. ^ Grusz et al. 2014.
  20. ^ Jepson Flora Project (eds.) 2022, Jepson eFlora, https://ucjeps.berkeley.edu/eflora/, accessed on September 20, 2022

Works cited

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Myriopteris: Brief Summary

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Myriopteris, commonly known as the lip ferns, is a genus of cheilanthoid ferns. Like other cheilanthoids, they are ferns of dry habitats, reproducing both sexually and apogamously. Many species have leaves divided into a large number of small, bead-like segments, the probable inspiration for the generic name. Hairs and/or scales are often present on both the upper and lower surfaces of the leaf, and their presence and appearance are useful in distinguishing between species. The genus is most diverse in Mexico, but species are found from southwestern Canada south to southern Chile, and one species is endemic to southern Africa.

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