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Is morphology supporting a monophyletic Proustia Lag., (Nassauvieae, Asteraceae)?

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Abstract

Proustia is an Andean genus traditionally considered to have three species and ten infraspecific taxa, which has often been placed in tribes other than Nassauvieae mainly because of its distinctive style features. Currently, each of the three species of Proustia is included in its own section denoting a consistent morphological variability within the genus. The heterogeneity among Proustia species mainly relies on habit, spiny structures arrangement, capitula arrangement, and pappus features. On the contrary, the forms or varieties within each species are apparently more homogeneous. The phylogenetic analysis performed here is based on morphological features and includes all the species, and all but two varieties and forms of Proustia. We test the boundaries of Proustia and the hypothesis that morphological discontinuities in the genus are phylogenetically uninformative. A total of 26 taxa and 29 morphological, anatomical, and palynological characters were studied. A maximum parsimony analysis yield six most parsimonious trees that showed almost identical topologies. Our results do not support the current generic concept of Proustia. Independently, each species of Proustia was monophyletic when taxa at sub-specific level were considered. Proustia pyrifolia was retrieved sister to the genus Berylsimpsonia although this clade did not obtain significant support. The spiny structures usually referred to as diagnostic for Proustia are not homologous for the genus. Characteristics of habit, spiny structures, secondary inflorescences and pappus support clades representing each species of Proustia, which may in turn, represent different genera.

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Acknowledgments

We would like to thank Jorge V. Crisci, the editors, and the anonymous reviewers for critical reading and suggestions on the manuscript and Chilebosque.cl project for allowing us to use their photographs for this scientific article. We are also grateful to the curators of the herbaria who provided material for examination. Research for this article was supported by Agencia Nacional de Promoción Científica y Tecnológica (SECYT), and Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), MINCYT, Argentina.

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  1. División Plantas Vasculares, Museo de La Plata, Paseo del Bosque s/n, B1900FWA, La Plata, Argentina

    Gisela Sancho, Liliana Katinas & Anabela Plos

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  1. Gisela Sancho
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  2. Liliana Katinas
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  3. Anabela Plos
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Correspondence to Gisela Sancho.

Appendices

Appendix 1

Morphological characters and character scores used in the phylogenetic analyses of Proustia 1. Habit: Shrubs (0); subshrubs (1); herbs (2). 2. Stem position: Erect (0); scandent (1). 3. Inflorescence axes distally spiny (thorns): absent (0); present (1). 4. Infrapetiolar spines: absent (0); present (1). 5. Brachyblasts: absent (0); present (1). 6. Leaves disposition: sparce (0); clustered at the base (1). 7. Leaf margin spiny: absent (0); present (1). 8. Leaf consistency: coriaceous to sub-coriaceous (0); herbaceous (1). 9. Leaf glandular trichomes: absent (0); present (1). 10. Leaf glandular trichomes: 1-seriate (1); 2-seriate (2); multiseriate (3). 11. Leaf structure: isolateral (0); dorsiventral (1); undifferentiated (2). 12. Number of capitula per capitulescence: few (2-8) (0); solitary (1); numerous (more than 15) (2). 13. Receptacle: pubescent (0); glabrous (1). 14. Capitulum sexuality: homogamous (0); heterogamous (female florets in the margin, bisexual in the center) (1). 15. Corolla morphology in homogamous capitula: all tubular (0); all bilabiate (1). 16. Corolla color: yellow (0); white (1); pink to purple (2). 17. Corolla vestiture: pubescent (0); glabrous or with apical papillae (1). 18. Style branches (according to Crisci, 1974): type 1 (1); type 2 (2); type 3 (3). 19. Style base widened: absent (0); present (1). 20. Cypsela apex: truncate (0); constricted to attenuate (1). 21. Carpopodium: absent (0); present (1). 22. Cypsela vestiture: pubescent (0); glabrous (1). 23. Cypsela type of trichomes: non-glandular (0); glandular (1). 24. Cypsela type of non-glandular hair: barnadesioid (0); 1-seriate (1); 2-seriate (twin hairs) (2). 25. Pappus bristle apex: plumose (0); scabrous or barbelate (1). 26. Pappus bristle body: plumose (0); scabrous or barbelate (1). 27. Pappus bristles color: tawny to pinkish (0); white (1). 28. Pollen exine type (Crisci, 1974; Zhao et al., 2000; Tellería et al., 2003): Chuquiraga type (0); Trixis type (1); Proustia type (2); Oxyphyllum type (3) Mutisia type (4). 29. Capitula arrangement: glomerules or solitary (0); spikes or glomerules in racemes (1); panicles (2).

Appendix 2

List of selected specimens examined for the morphological analysis and voucher data. Abbreviations for herbaria follow Holmgren et al. (1990). For analyzed specimens of Lophopappus see Katinas et al. (2013) Acourtia parryi (A. Gray) Reveal & R.M. King: I. M. Johnston 7370-Mexico, Zacatecas (2 Mar 1938), (LP!). Adenocaulon chilense Less.: A. L. Cabrera & M. M. Job 136-Argentina, Rio Negro, Cerro Otto (Jul 1935), (LP!); J. C. Montiel 205-Argentina, Rio Negro, Puerto Blest (1 Feb 1946), (LP!); M. J. Bonifacino, D. Gutiérrez & P. Simón 248-Argentina, Rio Negro, Cerro López (19 Jan 2000), (LP!). Berylsimpsonia vanillosma (C. Wright) B. L. Turner: E. L. Ekman 3034-Haiti, Port au Prince (without date), (LP!); E. L. Ekman 4306-Haiti, Ile La Tortue (14 Jun 1925), (LP!); J. I. Otero 391-Puerto Rico, Bayamon, Aibonito (19 Dec 1937), (GH!); A. A. Heller 6263 PIO4306-Puerto Rico, along Bucana river east of Ponce (11 Dec 1902), (GH!); E. C. Leonard 7445-Haiti, vicinity of St, Michael de l’ Atalaye (26 Nov 1925), (F!); N. L. Britton & E. G. Britton 9143-Puerto Rico, between Coano and Aibonito (11 Jan 1929), (F!). Calopappus acerosus Meyen: O. Zöllner 1406-Chile, Aconcagua, Los Andes (24 Mar 1967), (LP!); M. Ricardi 3226-Chile, Colchagua, San Fernando, Vegas del Flaco (10 Feb 1955), (CONC!); F. Schegel 2492-Chile, Santiago, Cerro Pirámide (5 Apr 1959), (CONC!); Ricardi 2971-Chile, Aconcagua, Portillo (11 Mar 1954), (CONC!). Chuquiraga avellanedae Lorentz: A. L. Cabrera, S. Botta, C. Ezcurra, R. Kiesling 33197-Argentina, Chubut, Sarmiento (14 Dec 1981), (LP!); A. L. Cabrera, A. Giaotti, R. Kiesling, M. Ronco, E. Zardini, F. O. Zuloaga 23245-Argentina, Chubut, Sarmiento (25 Jan 1973), (LP!); M. Gentili 524-Argentina, Neuquén, Cerro El Marucho (4 Jan 1973), (LP!). Jungia polita Griseb.: A. L. Cabrera 3740-Argentina, Salta, Guachipas (12 Aug 1936), (LP!); A. L. Cabrera 20777-Argentina, Jujuy, Capital, Cuesta de Las Lajitas (28 Aug 1970), (LP!); S. Venturi 9674-Argentina, Jujuy, sierra de Santa Bárbara, (8 Oct 1929), (LP!); J. B. Sotelo 10051-Argentina, Jujuy, Capital, Yala, (9 Jun 1948), (LP!) H. Fabris 4695-Argentina, Jujuy, Capital, alrededores de San Salvador de Jujuy (20 Oct 1963), (LP!); A. L. Cabrera & H. Fabris 15991-Argentina, Jujuy, Ledesma, entrada al camino a Valle Grande (15 Oct 1964), (LP!). Leucheria achillaeifolia Hook. et Arn.: J. J. Neumeyer 407-Argentina, Chubut, Cushamen, Lago Puelo (10 Dec 1940), (LP!); G. Covas 428-Argentina, Mendoza, Malargüe, cerros al norte de Calmuco (16 Feb 1942), (LP!); A. L. Cabrera 11161-Argentina, Neuquén, Minas, Andacollo (9 Dec 1952), (LP!); R. Hauthal s.n.-Argentina, Santa Cruz, Lago Argentino, (Jan 1902), (LP!); A. Soriano 3783-Argentina, Chubut, Futaleufú, Esquel, (10 Nov 1949), (LP!). Macrachaenium gracile Hook. f.: E. Pisano 3081-Chile, Magallanes, Fiordo Parry (21 Feb 1971), (LP!); E. Pisano 2876-Chile, Magallanes, Fiordo Parry (13 Dec 70), (LP!). Nassauvia cumingii Hook. & Arn.: A. Ruíz Leal 61-Argentina, Mendoza, La Cuevas (without date), (LP!); F. Pastore 121-Argentina, Neuquén, Arroyo Memanque (18 Apr 1912), (LP!); A. Ruíz Leal 16786-Argentina, Mendoza, San Rafael (6-7 Feb 1955), (LP!). Proustia cuneifolia f. angustifolia (Wedd.) Fabris: A. Jiménez 39-Bolivia, Cochabamba, Angostura (3 Apr 1955), (LP!); T. Herzog 2021-Bolivia (Apr 1911), (LP); S. Venturi 8297-Argentina, Jujuy, Humahuaca, Sierra del Zenta (27 Feb 1929), (US!, GH!); J. R. I. Wood 8313-Bolivia, Chuquisaca, on the escarpment between Zudanez and Tarabuco (11 Apr 1994), (US!); E. Asplund 4182-Bolivia, Cochabamba, Capinata (11 Jun 1921), (GH!). Proustia cuneifolia f. cuneifolia D. Don: F. Vervoorst 8638-Argentina, Catamarca, Andalgalá, Cuesta de la Chilca, (10 Jan 1973), (US!); S. Venturi 6634-Argentina, Catamarca, Santa María (1 Feb 1925), (US!); A. Bridarolli 1067-Argentina, Córdoba, San Javier (13 Jan 1940), (LP!); T. Meyer, R. Cuezzo & R. Legname 21083-Argentina, Jujuy, Humahuaca (8 Mar 1960), (LP!); S. Venturi 4878-Argentina, Jujuy, Tilcara (8 Feb 1927), (US!, F!, GH!); A. Bohi & D. Liesner 4-Argentina, Jujuy, Yavi (11 Jan 1967), (LP!); T. Meyer 3933-Argentina, La Rioja, Capital (30 Dec 1941), (US!); Rodríguez 1435 (LIL 27912)-Argentina, Salta, Cafayate (6 Apr 1914), (GH!); A.L. Cabrera 9008-Argentina, Salta, Rosario de Lerma (9 Feb 1946), (LP!); R. F. Steinbach 186-Bolivia, Cochabamba, pie de colina San Pedro (11 May 1966), (F!); S. G. Beck 8473-Bolivia, La Paz, Loayza (27 Jun 1983), (US!); C. Grünge 3162-Chile, Biobío, Los Ángeles (29 Jan 1961), (US!); G. Montero O. 755-Chile, Calchagua, San Fernando (24 Feb 1928), (F!, GH!); C. Punge 2447-Chile, Biobío, Antuco (19 Jan 1941), (LP!); R. Ferreyra 13939-Perú, Arequipa, Carevalí (19 Dec 1959), (LP!); H. Fabris & J. V. Crisci 6986-Argentina, Jujuy, Valle Grande, Caspalá (12 Mar 1967), (HUT!, LP!). Proustia cuneifolia f. mendocina (Phil.) Fabris: A. L. Cabrera 1179-Argentina, Catamarca, Belén (5 Mar 1929), (LP!); M. Layaga 2269-Argentina, Catamarca, La Ciénaga (Jan 1956), (LP!); R. Schreiter 10588 (LIL 58106)-Argentina, Catamarca, Las Mansas (Mar 1938), (GH!); J. H. Hunziker 1821-Argentina, La Rioja, Famatina (12 Jan 1947), (LP!); R. Falcone & J. Castellanos 3643-Argentina, La Rioja, General Belgrano (12 Mar 1957), (LP!); A. T. Hunziker, A. E. Cocucci & R. Subils 15866-Argentina, La Rioja, Independencia (3 Feb 1961), (NY!); M. I. H. Scott de Birabén & M. Birabén 997-Argentina, La Rioja, Tinogasta (23 Nov 1939), (LP!); D. O. King 1926-Argentina, Mendoza, Las Heras (30 Oct 1936), (LP!); E. M. García 509-Argentina, Mendoza, Luján (6 Jan 1948), (GH!); L. Serra s.n. (LP 900747)-Argentina, Mendoza, Luján de Cuyo (26 Jan 1952), (LP!); O. Boelcke, N. M. Bacigalupo & M. N. Correa 10415-Argentina, Mendoza, Malargüe (31 Jan 1963), (LP!); A. Burkart, N. S. Troncoso & E. G. Nicora 14362-Argentina, Mendoza, San Carlos (10 Feb 1942), (LP!); O. Boelcke 4196-Argentina, Mendoza, San Rafael (6 Feb 1950), (LP!); A. Ruiz Leal 1099-Argentina, Mendoza, Tunuyán (19 Feb 1933), (LP!); A. L. Cabrera, D. Añon Suarez, M. A. Torres, J. Crisci & N. Tur 18008-Argentina, San Juan, Iglesia, Pismanta (26 Feb 1967), (LP!); T. M. Pedersen 15228-Argentina, San Juan, Ullún (13 Mar 1989), (F!, NY!). Proustia cuneifolia var. mollis Cabrera: A. L. Cabrera & H. A. Fabris 21042-Argentina, Jujuy, Santa Bárbara (21 Nov 1970), (LP!); A. Burkart 13287-Argentina, Salta, Coronel Moldes (14 Nov 1942), (LP!); A. L. Cabrera, J. Frangi, A. M. de Frangi, R. Kiesling & E. M. Zardini 22074-Argentina, Salta, Rosario de Lerma (12 Feb 1972), (LP!); C. Spegazzini s.n. (LPS 1873)-Argentina, Salta, Pampa Grande (Jan 1897), (LP!). Proustia cuneifolia f. oblongifolia (Wedd.) Fabris: R. S. Shepard 172-Bolivia, La Paz, Cañon La Paz River (6 Aug 1920), (GH!); A. Gentry, M. Dillon, P. Berry & J. Aronson 23340-Perú, Apurimac, río Chalhuanca (24 Jun 1978), (F!); A. Weberbauer 5762-Perú, Ayacucho, Tal v. Huacata (1909-1914), (F!). Proustia cuneifolia f. tipia (Phil.) Fabris: M. O. Dillon, D. Dillon, V. Asencio & M. Villarroel 5735-Chile, Antofagasta (23 Oct 1988), (F!); A. Borchers s.n.-Chile, Atacama (1887), (LP!); H. Niemeyer s.n. (F 2065915)-Chile, II Región, Comuna Taltal (Apr 1985), (F!). Proustia ilicifolia f. baccharoides (D. Don) Fabris: E. M. L. Kausel 5150-Chile, Atacama, zona interior, Panamericana (20 Sep 1966), (LP!); F. Ruiz s.n. (LP 071041)-Chile, Coquimbo, Illapel (Feb 1931), (LP!); E. Barros 2369-Chile, Coquimbo, Ovalle (15 Jan 1942), (LP!); F. Claude Joseph 4054-Chile, Río Blanco (Nov 1925), (US!); A. Garaventa Limache 332-Chile, Cerro de la Virgen, 6 Mar 1928), (LP!). Proustia ilicifolia f. ilicifolia Hook. & Arn.: E. Werdermann 423-Chile, Atacama, Copiapó (Sep 1924), (GH!, F!); M. Ricardi 3751-Chile, Atacama, Estancia Manflas (2 Nov 1956), (LP!); M. Ricardi & C. Marticorena 4888 (CONC 25683)-Chile, Atacama, Cachiyuyo (10 Oct 195), (LP!); M. Ricardi 3667-Chile, Atacama, La Puerta (28 Oct 1956), (LP!); M. Ricardi 3851-Chile, Atacama, San Félix (10 Nov 1956), (LP!); I. M. Johnston 6267-Chile, Coquimbo, Elqui (18-19 Jan 1926), (GH!); M. Ricardi 5553 (CONC 37022)-Chile, Coquimbo, La Serena (16 Feb 1963), (LP!); C. Jiles 1256-Chile, Coquimbo, Ovalle (17 Jan 1949), (LP!); C. Jiles 1074-Chile, Coquimbo, Ovalle (10 Oct 1948), (LP!); G. Montero O. 2845-Chile, Coquimbo, Rivadavia (17 Sep 1936), (GH!); J. P. Simón 257-Chile, Coquimbo, Road Punitaqui to Combarbalá (24 Oct 1970), (US!); without colector s.n. (LP s.n.)-Chile, Coquimbo, Paihuano (5 Feb 1883), (LP!); G. T. Hastings 618 (US 580477)-Chile, Santiago, San Cristóbal (Dec 1901), (US!); without colector s.n. (LP s.n.)-Chile, Santiago, San Cristóbal (1881), (LP!). Proustia pyrifolia DC.: C. Jiles 2329-Chile, Aconcagua, Petorca (16 Nov 1952), (LP!); O. Boelcke 3991-Chile, Aconcagua (5 Jan 1949), (LP!); J. Acuña s.n. (CONC 9638)-Chile, Cautín, Villarrica (23 Feb 1950), (LP!); M. Ricardi s.n. (CONC 10027)-Chile, Colchagua, San Fernando (3 Jan 1951), (LP!); H. Riffo 12888-Chile, Concepción (1-4 Mar 1925), (GH!); A. Lourteig 2514-Chile, Concepción, Hualpén (3 Jan 1971), (LP!). H. A. Fabris & J. Crisci 7549-Chile, Concepción, Rocoto (Jan 1969), (LP!); A. L. Cabrera 19661-Chile, Concepción, Rocoto (19 Jan 1969), (LP!); Mertens s.n.-Chile, Concepción, Camarico (without data), (GH!); Y. Mexia 7857-Chile, Curicó (13 Feb 1936), (F!); F. Fredrich s.n. (LP 071038)-Chile, río Tolten cerca de Villarica, Cautín (Mar 1935), (LP!); A. Lourteig 2514-Chile, Concepción, Huelpén (3 Jan 1971), (LP!); E. Werdermann 569-Chile, Curicó, Hacienda Monte Grande (Dec 1924), (F!); P. Germain s.n. (F 1013019)-Chile, Guillota, Colchagua (without date), (F!); M. H. de Looser 5176-Chile, Maule, Constitución (Jan 1946), (GH!); E. M. L. Kausel 4347-Chile, Santiago, San Antonio (Feb 1958), (F!); A. Caldeleugh s.n. (F 1013154)-Chile, Santiago, Colchagua (without date), (F!); L. Moreira s.n.-Chile, Talca, Camarico (Feb 1926), (GH!); A. Hollermayer 86-Chile, Valdivia (4 Mar 1943), (LP!); F. Schlegel 579 (CONC 40338)-Chile, Valparaíso (17 Jan 1955), (F!); C. L. G. Bertero 1263-Chile, Valparaíso (1829), (GH!, F!); J. West 5197-Chile,Valparaíso, Cajón de San Pedro, (19 Jan 1936), (GH!). Perezia pungens (Bonpl.) Less.: I. Sánchez Vega & W. M. Ruíz Vigo 468-Peru, Cajamarca, Challuayaco (4 May 1970), (LP!); A. López 605-Peru, La Libertad, Otuzco (1 Jun 1951), (LP!). Triptilion achilleae DC.: A. Soriano 2454-Argentina, Chubut, Futaleufú, Estancia Pampa Chica (22 Jan 1947), (LP!); A. L. Cabrera 11286-Argentina, Neuquén, Huiliches (17 Dec 1952), (LP!); A. L Cabrera & Job 357-Argentina, Río Negro, Bariloche, Nahuel Huapi (16 Jan 1935), (LP!); A. Burkart 9602-Chile, Malleco, Liucura, (4 Mar 1939), (LP!); A. Ruiz Leal 26763-Argentina, Neuquén, Aluminé, camino Aluminé-Las Coloradas, (6 Jan 1970), (LP!). Trixis lessingii DC.: A. Macêdo 2852-Brazil, Minas Gerais, Ouro Preto, Andorinha (5 Jan 1951), (LP!); A. P. Duarte 2116-Brazil, Serra do Cipó (6 Dec 1948), (LP!); G. Hatschbach 17851-Brazil, Paraná, Campina Grande Sul (15 Nov 1967), (LP!); M. Magalhaes 5393-Brazil, Minas Gerais, Rawcharia, (30 Jan 1948), (LP!); R. Klein 3569-Brazil, Santa Catarina, Matos Costa (8 Mar 1962), (LP!).

Appendix 3

See Table 2

Table 2 Data matrix of morphological characters used in the phylogenetic analysis of Proustia and allied genera
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Sancho, G., Katinas, L. & Plos, A. Is morphology supporting a monophyletic Proustia Lag., (Nassauvieae, Asteraceae)?. Plant Syst Evol 300, 2265–2276 (2014). https://doi.org/10.1007/s00606-014-1052-6

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