Abstract
Gamochaeta (tribe Gnaphalieae, Asteraceae) is composed of ca. 60 species primarily distributed in tropical and subtropical America. Within the tribe Gnaphalieae, the genus is characterized by capitula arranged in spikes or head-like clusters, few hermaphroditic central florets, truncate style branches with apical sweeping trichomes, pappus bristles connate at the base into a ring falling as a unit, and achenes with globose twin trichomes. Previous molecular phylogenetic studies have suggested the paraphyly of the genus, but have not provided a basis for redefining generic limits due to incomplete taxon sampling. To address this problem, DNA sequences from the plastid (trnL-F) and nuclear (ETS and ITS) genomes were analyzed from a broad taxon sample representing the full range of morphological variation known in the genus. Our results affirm that Gamochaeta is paraphyletic as presently circumscribed. Two clades can be recognized: one clade that includes the majority of the species currently assigned to Gamochaeta and a second clade that includes Gamochaetopsis, Stuckertiella and seven species of Gamochaeta. We present here a new circumscription of Gamochaeta, including two new combinations, Gamochaeta alpina and Gamochaeta peregrina, and the resurrection of Gamochaeta capitata. Our results also show Omalotheca supina, O. norvegica and O. sylvatica, which were placed by some authors in Gamochaeta or in Gnaphalium, form a monophyletic group distantly related to both genera.
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Acknowledgments
We express our thanks to the reviewers for critical reading of the manuscript; to S. Beck (LPB, Bolivia) for his assistance during field work in Bolivia; to F. Zuloaga (SI, Argentina) and A. Tribsch (SZU, Austria) who provided material from their own collections; to L. Aagesen for comments on early drafts of the manuscript; to the curators of CONC, LP, LPB, MNCS, MO, SI, SZU, USMS and W for providing materials; to F. Zuloaga and S. Beck for permission to use their photographs included in Fig. 6; J. Hurst for technical assistance. This work was supported by Consejo Nacional de Investigaciones Científicas y Técnicas for financial support (PIP 112-200801-02196) to S. E. Freire and FONDECYT No. 1150425 to A. Moreira Muñoz.
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Appendix
Appendix
Species used in the molecular studies with voucher information and GenBank accession numbers (ETS, ITS and trnL-F). Accession number in bold indicates data obtained from GenBank. * indicates sequence not obtained for that marker.
Achyrocline tomentosa. KM091358, KM091389, KM091420. Antennaria chilensis var. magellanica. KM091373, KM091395, *. Antennaria dioica. FN645610, FJ639964, FN645790. Belloa chilensis KM091349, KM091388, KM091430. Berroa gnaphalioides. KM091355, KM091386, KM091418. Facelis plumosa. KM091372, KM091394, KM091428. Filago fuscescens. FN645579, FN645846, FN645764. Filago lutescens. FN645596, FN645883, FN645779. Filago pyramidata. FN645589, FN645872, HM364535. Gamochaeta affinis. Argentina. Tierra del Fuego: Ushuaia, Bahía Valentín, 16 Oct 1971, Dudley et al. 303 (MO). KX078016, *, *. Gamochaeta alpina. (under Gamochaetopsis alpina). KM091356, KM091390, KM091417. Gamochaeta ambatensis. Argentina. Jujuy: Valle Grande, de San Francisco a Alto de Calilegua, 21 Feb 2008, Zuloaga et al. 10324 (SI). KX078017, *, KX078049. Gamochaeta americana. KM091382, KM091411, KM091437. Gamochaeta andina. Chile. Maule: Maule, Lueberg and Teiller 2290 (CONC). KX078018, KX077990, KX078050. Gamochaeta antillana. USA. Mississippi: Lamar County, Alford 3892 (USMS). KX078019, KX077991, KX078051. Gamochaeta argyrinea. USA. Nesom WMGT8 (LP). KX078020, *, *. Gamochaeta beckii. Bolivia. La Paz: Nor Yungas, camino a Coroico, entrada del camino viejo hacia la Mina Lourdes, 30 Mar 2010, Urtubey et al. 508 (SI). KX078021, KX077992, KX078052. Gamochaeta calviceps. Argentina. Entre Ríos: Colón, camino al arroyo El Palmar, 51 m a. s. l., 7 Dec 2008, Urtubey and Baztarrica 384 (SI). KX078022, *, *. Gamochaeta camaquaensis. Brasil. Rio Grande do Sul: Cacapava do Sul, 30 Sep 2009, Deble and Deble 11879 (SI). KX078023, KX077993, KX078053. Gamochaeta capitata (under Stuckertiella capitata) KM091369, KM091398, KM091412. Gamochaeta chamissonis. Chile. Metropolitana: El Yeso, 1283 m a. s. l., 26 Jan 2012, Urtubey and Freire 713 (SI). KX078024, KX077994, KX078054. Gamochaeta chionesthes. USA. Giorgia: 15 Apr 2001, Nesom GASC04-26 (LP). KX078025, KX077995, KX078055. Gamochaeta coarctata. Argentina. Neuquén: Minas, de Chos Malal a Andacollo, ca. 40 km junto al arroyo El Manzanito, 1443 m a. s. l., 12 Jan 2009, Urtubey et al. 409 (SI). KX078026, KX077996, KX078056. Gamochaeta depilata. Chile. Biobío: Mihoc et al. 4092 (CONC). KX078027, KX077997, KX078057. Gamochaeta deserticola. Bolivia. La Paz: Murillo, 4100 m a. s. l., 4.8 km al NE del autopista por el camino subiendo el Valle del Río Kaluyo 28 Feb 1987, Solomon 16209 (LPB). KX078028, *, KX0780580. Gamochaeta erecta. Brasil. Rio Grande do Sul: Cacapava do Sul, 28 Oct 2009, Deble and Deble 11874 (SI). KX078029, KX077998, KX078060. Gamochaeta erythractis. Argentina. Jujuy: Humahuaca, camino a la Mina Aguilar, 3940 m a. s. l., 23 Mar 2009, Urtubey and Freire 442 (SI). KX078030, KX077999, KX078061. Gamochaeta falcata. Argentina. Córdoba: Calamuchita, desde Yacanto de Calamuchita al Champaquí, ca. 2100 m a. s. l., 18 Jan 2006, Urtubey and Baztarrica 197 (SI). *, KX078000, *. Uruguay. Colonia: Conchillares, playa, 18 Nov 2010, Urtubey 520 (SI). KX0780310, *, *. Gamochaeta filaginea. Argentina. Buenos Aires: Isla Martín García, 24 Sep 2000, Hurrell 4363 (LP). KX078032, *, *. Gamochaeta grazielae. Brasil. Río de Janeiro: Nov 2006, Deble and Deble 6018 (SI). KX078033, KX078001, KX078062. Gamochaeta humilis Bolivia. La Paz: Nor Yungas, camino a Coroico, bajando de la ruta en la entrada del camino viejo hacia la Mina Lourdes, 3919 m a. s. l., 30 Mar 2010, Urtubey et al. 506 (SI). KX078034, KX078002, KX078059. Gamochaeta longipedicellata. Bolivia. La Paz: Murillo, 24 Mar 2010, Urtubey et al. 473 (SI). KM091381, KM091410, KX078063. Gamochaeta lulioana. Bolivia. La Paz: Murillo, en el borde de bofedal, 4661 m a. s. l., 24 Mar 2010, Urtubey et al. 480 (SI). KX078035, KX078003, KX078064. Gamochaeta nivalis. Argentina. Río Negro: Bariloche, Cerro Catedral, 16 Jan 2009, Urtubey et al. 415 (SI). KX078036, *, KX078065. Gamochaeta pensylvanica. Argentina. Buenos Aires: La Plata, ciudad Vieja, 6 Oct 2012, Urtubey 755 (SI). *, *, KX078066. USA. Mississippi: Lamar County, Alford 4156 (USMS). KX078037, KX078004, *. Gamochaeta procumbens. Argentina. Tierra del Fuego: Rio Grande, Estancia Cullén, 6 Jan 1971, Goodall 3118 (Herb. Goodall). KX078038, KX078005, *. Gamochaeta purpurea. USA, Nesom WMGT13 (LP) KX078039, KX078006, KX078067. Gamochaeta serpyllifolia Chile. Los Ríos: Valdivia, Comuna de Panguipulli, faldeos Volcán Chosuenco, 39º55’/71º59’, 1558 m a. s. l., 5 Feb 2012, Baeza 4357 (CONC). KM091380, KX078010, KX078068. Gamochaeta simplicicaulis. Argentina. Jujuy: Tilcara, camino de Molulo a Huairahuasi, 3100 m a. s. l., 11 Feb 2010, Zuloaga et al. 11702 (SI). KX078040, KX078007, KX078069. Gamochaeta sphacelata. Argentina. Salta: Santa Victoria, Nazareno, 3308 m a. s. l., Adler 4 (MNCS 1665). KX078041, *, KX078070. Gamochaeta spiciformis. Chile. Magallanes: Domínguez 984 (CONC). *, KX078011, KX078071. Argentina. Tierra del Fuego: Ushuaia, Brown Sawmill, Goodall 286 (LP). KX078042, *, *. Gamochaeta stachydifolia. Argentina. Buenos Aires: Tandil, hacia sierras de Las Animas desde el Hotel Elegance, 12 Nov 2005, Urtubey and Baztarrica 179 (SI) KX078043, *, *. Ibid. Frente al hotel Elegance, 1 Oct 2010, Urtubey 513 (SI). *, KX078008, KX078072. Gamochaeta subfalcata. Uruguay. Canelones: La Floresta, 19 Nov 2010, Urtubey 524 (SI). KX078044, KX078009, KX078073. Gamochaeta ustulata USA. Oregon: Lincoln County, Shannon Straub 201 (USMS). KX078045, KX078012, KX078074. Gnaphalium austroafricanum. FN645630, FN645830, FN645756. Gnaphalium declinatum FR821617, *, FR822634. Gnaphalium uliginosum Austria. Upper Austria: Innviertel, Reichersberg am Inn, Hübing, 22 Aug 2010, Pflugbeil 936 (SZU). KX078048, KX078015, KX07807077. Jalcophila boliviensis. KM091370, KM091402, KM091429. Jalcophila ecuadorensis. KM091383*,*. Leontopodium alpinum FM173135, GU943413, AF141821 + AF141733. Leontopodium microphyllum. FJ640015, FJ639947, FJ640047 + FJ639977. Lucilia acutifolia. KM091374, KM091396, KM091432. Omalotheca norvegica (Gunnerus) Sch.-Bip. & F.W.Schultz. Russia. Republic Altai: surroundings of Zeminsky (Zeminskij) pass to area NE of the pass, ca. 105 km S Gorno Altaysk (Altajsk), subalpine Siberian pine forest, 1700-1730 m a. s. l., 11 Aug 2003, Tribsch and Essl 10383 (W). KX078047, KX078013, KX078075. Omalotheca supina FN645558, AY445230, FN645789. Omalotheca sylvatica. Austria. Carinthia: Lavanttal, Koralpe, Hartelsberg SE Wolfsberg, 1345 m a. s. l., Riegler-Hager 921 (W). KX078046, KX078014, KX078076.
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Urtubey, E., López, A., Chemisquy, M.A. et al. New circumscription of the genus Gamochaeta (Asteraceae, Gnaphalieae) inferred from nuclear and plastid DNA sequences. Plant Syst Evol 302, 1047–1066 (2016). https://doi.org/10.1007/s00606-016-1316-4
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DOI: https://doi.org/10.1007/s00606-016-1316-4