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A phylogenetic analysis of the genus Paspalum (Poaceae) based on cpDNA and morphology

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Abstract

With about 350 species, Paspalum is one of the richest genera within the Poaceae. Its species inhabit ecologically diverse areas along the Americas and they are largely responsible for the biodiversity of grassland ecosystems in South America. Despite its size and relevance, no phylogeny of the genus as a whole is currently available and infrageneric relationships remain uncertain. Many Paspalum species consist of sexual-diploid and apomictic-polyploid cytotypes, and several have arisen through hybridization. In this paper we explore the phylogenetic structure of Paspalum using sequence data of four non-coding cpDNA fragments from a wide array of species which were combined with morphological data for a subset of diploid taxa. Our results confirmed the general monophyly of Paspalum if P. inaequivalve is excluded and the small genus Thrasyopsis is included. Only one of the four currently recognized subgenera was monophyletic but nested within the remainder of the genus. Some informal morphological groups were found to be polyphyletic. The placement of known allopolyploid groups is generally congruent with previously stated hypotheses although some species with shared genomic formulae formed paraphyletic arrangements. Other species formed a basal grade including mostly umbrophilous or hygrophilous species. It is hypothesized that the genus may have diversified as a consequence of the expansion of C4 grass-dominated grasslands in South America.

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Acknowledgments

We are indebted to Pamela Soltis for laboratory facilities at the University of Florida, USA, to the Willi Hennig Society for making available a sponsored version of TNT, and to J. Marcos Mirande for kindly making available his unpublished TNT script. The following persons collaborated with us during field collection work and/or made available plant material from their own collections: Lone Aagesen, Alejandro Asenjo, Isabel B. Boccaloni, M. Eugenia Buela, Irene Caponio, Ana M. Carrión, Julio Daviña, Juan R. Fernández, Luis Frank, M. Cecilia Gróttola, Diego H. Hojsgaard, Ana I. Honfi, Graciela Lavia, Regina C. Oliveira, Florencia Perret, Camilo L. Quarin, Yamil Ramírez, Germán G. Roitman, José L. Rosa, G. Mónica Tourn, Nicolás Trillo, José F. M. Valls. This work received financial support through grants PIP 6568 (CONICET, Argentina), UBACyT JG21, and G806 (University of Buenos Aires, Argentina), and to GHR from the “Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)”, Brazil. GHR is a member of the “Carrera del Investigador” of the “Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)”, Argentina.

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Authors and Affiliations

  1. Cátedra de Botánica Agrícola, Facultad de Agronomía, Universidad de Buenos Aires, Avenida San Martín 4453, C1417DSE, Buenos Aires, Argentina

    Gabriel H. Rua

  2. Depto. de Biología Vegetal, Facultad de Agronomía, Universidad de la República, Garzón 780, 12900, Montevideo, Uruguay

    Pablo R. Speranza & Magdalena Vaio

  3. Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St., Box G-W, Providence, RI, 02912, USA

    Mónica Arakaki

Authors
  1. Gabriel H. Rua
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  2. Pablo R. Speranza
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  3. Magdalena Vaio
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  4. Mónica Arakaki
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Corresponding author

Correspondence to Gabriel H. Rua.

Additional information

G. H. Rua and P. R. Speranza contributed equally to this paper.

Appendix

Appendix

Morphological characters used for phylogenetic analysis.

1. Lower glume, presence: lacking [0], present [1]. 2. Lower glume, position: on the medial plane of the spikelet [0], turned to one side of the spikelet [1]. 3. Upper glume, presence: lacking [0], present [1]. 4. Upper glume, length relative to the lower lemma: nearly equal [0], conspicuously shorter [1]. 5. Upper glume, edge: not edge-forming [0], forming an edge around the lower lemma [1]. 6. Upper glume, distal portion: rounded [0], obtuse [1], acute [2], acuminate [3]. 7. Upper glume, apex: rounded [0], obtuse [1], apiculate [2], acute [3], acuminate [4], truncate [5]. 8. Upper glume, dorsum: flat [0], bowed [1]. 9. Upper glume, a tuft of long hairs at the base: lacking [0], present [1]. 10. Upper glume, consistency: hyaline, tiny [0], membranous [1]. 11. Upper glume, a marginal fringe of tuberculate hairs: lacking [0], present [1]. 12. Upper glume, number of lateral veins on each glume half: none [0], one [1], two [2], three [3], four or more [4]. 13. Upper glume, abaxial vein protrusion: not prominent [0], prominent [1]. 14. Upper glume, distal convergence of inner lateral veins: not convergent [0], convergent [1]. 15. Upper glume, midrib: lacking [0], present [1]. 16. Upper glume, vein distribution: equidistant, no veins approximate towards margins [0], only outer lateral veins approximate towards margins [1], all lateral veins approximate towards margins [2]. 17. Upper glume, outer vein position: marginal [0], not marginal [1]. 18. Upper glume, marginal region: folded [0], flat [1]. 19. Upper glume, between-vein indumentum: glabrous to scabrous [0], pubescent at the base [1], fully pubescent [2], pubescent towards the apex [3]. 20. Upper glume, marginal region (beyond outer veins) indumentum: glabrous to scabrous [0], scarcely pubescent at base [1], pubescent [2]. 21. Upper glume, hair base: simple [0], with [tinged] cushions [1], both [2]. 22. Upper glume, apex: navicular or cucullate [0], flat [1]. 23. Upper glume, surface: smooth [0], transversely crumpled or wrinkled [1]. 24. Upper glume, whether flabby or not: tight [0], flabby [1]. 25. Upper glume, symmetry: symmetrical [0], asymmetrical [1]. 26. Lower lemma, distal portion: rounded [0], obtuse [1], acute [2], acuminate [3]. 27. Lower lemma, apex: rounded [0], obtuse [1], apiculate [2], acute [3], acuminate [4], truncate [5], emarginate [6]. 28. Lower lemma, dorsum: flat [0], bowed [1], concave [2], sulcate [3]. 29. Lower lemma, consistency: hyaline, tiny [0], membranous [1], laterally indurate [2]. 30. Lower lemma, a basal tuft of hairs: wanting [0], present [1]. 31. Lower lemma, a marginal fringe of tuberculate hairs: wanting [0], present [1]. 32. Lower lemma, between-vein indumentum: glabrous [0], distally pubescent [1], basally pubescent [2], fully pubescent [3]. 33. Lower lemma, number of lateral veins on each side: none [0], one [1], two [2], three [3]. 34. Lower lemma, midvein: lacking [0], present [1]. 35. Lower lemma, distal convergence of inner lateral veins: not convergent [0], convergent [1]. 36. Lower lemma, marginal region: folded [0], flat [1]. 37. Lower lemma, marginal region (beyond outer veins) indumentum: glabrous [0], pubescent [1]. 38. Lower lemma, apex: navicular to cucullate [0], flat [1], folded [2]. 39. Lower lemma, internerval space: smooth [0], wrinkled [1]. 40. Lower lemma, axillary flower: wanting [0], reduced to a palea [1], well developed [2]. 41. Upper floret, pigmentation: pale to stramineous [0], brown [1], dark brown, shining [2], purplish [3]. 42. Upper floret, induration: not indurate [0], slightly indurate [1], strongly indurate [2]. 43. Upper floret, length relative to the lower lemma: nearly equal [0], conspicuously shorter [1]. 44. Upper floret, shape: elliptical [0], ovate [1], obovate [2], rhomboid [3], orbicular [4]. 45. Upper floret, basal constriction: lacking [0], present [1]. 46. Upper floret, callus: glabrous [0], laterally tufted [1], tufted around [2]. 47. Upper floret, lemma dorsum: more or less flattened [0], bowed towards the base [1], sharply bowed [2], gibbose [3]. 48. Upper floret, palea dorsum: convex [0], flat [1], concave [2]. 49. Upper floret, lemma apex: acute, pointed [0], acute but blunt at the very apex [1], rounded [2], acuminate [3], obtuse [4]. 50. Upper floret, lemma nerves: not prominent [0], sharply prominent [1]. 51. Upper floret, epidermal papillae on the lemma: wanting [0], present [1]. 52. Upper floret, abscission at maturity: none [0], occurring [1]. 53. Upper floret, apex indumentum: wanting [0], present [1]. 54. Upper floret, palea margins: not winged [0], with non-overlapping wings [1], with overlapping wings [2]. 55. Upper floret, palea adaxial surface: smooth [0], papillose [1]. 56. Upper floret, lemma apex: open [0], cucullate [1]. 57. Upper floret, lemma margins: not thickened [0], slightly thickened [1], sharply thickened [2]. 58. Upper floret, number of lateral veins on each lemma half: none [0], one [1], two [2], three or more [3]. 59. Upper floret, midvein: lacking [0], present [1]. 60. Upper floret, lemma [inner]-nerves: distant from margins [0], submarginal [1]. 61. Upper floret, anther pigmentation: yellow [0], purple-tinged [1], deep purple [2]. 62. Pigmentation of stigmas: whitish [0], lila [1], purple [2], yellow [3]. 63. Upper floret, caryopsis hilum: punctiform [0], elliptical [1], linear [2]. 64. Inflorescence, terminal spikelet: wanting [0], present [1]. 65. Inflorescence, arrangement of primary branches: several branches along an axis with conspicuous internodia [0], several branches, the two distal ones conjugate [1], only two conjugate primary branches [2], one branch alone [3]. 66. Inflorescence, arrangement of primary branches II: all alternate [0], verticillate at least in the lower nodes [1]. 67. Maximum number of orthostichies: none [0], one [1], two [2], three [3], four [4]. five [5], six [6]. 68. Inflorescence, main axis cross section: polygonal [0], flattened [1]. 69. Inflorescence, pubescence on pulvinula: glabrous [0], [shortly] pubescent [1]. 70. Inflorescence, long cilia arising from pulvinula: wanting [0], present [1]. 71. Inflorescence, a spikelet ending each primary branch: present [0], lacking, lateral spikelets becoming rudimentary towards apex [1]. 72. Inflorescence, rachis cross section: trigonous, not expanded [0], laterally expanded into wings having chlorenchyma [1], laterally expanded into membranous epidermal wings [2]. 73. Inflorescence, rachis, sclerenchima ring: lacking [0], present [1]. 74. Inflorescence, rachis, medullar lacunae: lacking [0], present [1]. 75. Inflorescence, rachis venation: a midnerve thick and prominent [0], several parallel equal-range nerves [1]. 76. Inflorescence, rachis margin: smooth [0], scabrous [1], with more or less scattered cilia [2], with a dense fringe of tuberculate cilia [3]. 77. Inflorescence, rachis: straight to slightly sinuous [0], sharply sinuous [1]. 78. Inflorescence, rachis surface indumentum: glabrous [0], scabrous [1], pubescent [2]. 79. Inflorescence, arrangement of spikelets: solitary [0], homogeneously paired [1], proximally paniculate, distally paired [2]. 80. Inflorescence, concrescence of sPc branchlets with the rachis: (nearly) free, the spikelets appear pedicellate [0], concrescent, the spikelets appear subsessile [1]. 81. Inflorescence, a crown of hairs at the top of pedicells: lacking [0], present [1]. 82. Inflorescence, pedicells cross section: terete [0], trigonous/tetragonous [1], flattened [2]. 83.

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Rua, G.H., Speranza, P.R., Vaio, M. et al. A phylogenetic analysis of the genus Paspalum (Poaceae) based on cpDNA and morphology. Plant Syst Evol 288, 227–243 (2010). https://doi.org/10.1007/s00606-010-0327-9

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