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Morphometric analysis of the Saccharum complex (Poaceae, Andropogoneae)

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Abstract

Saccharum s.l. comprises 35–40 species, including sugarcane (S. officinarum), and is distributed in the tropics and subtropics. The term “Saccharum complex” refers to four related genera: Saccharum, Erianthus s.l. (including E. sect. Ripidium, or Tripidium), Narenga, and Sclerostachya. Miscanthus and Miscanthidium are also closely related to this group. The delimitation of these genera is contentious and largely unresolved, with most species recognized under Saccharum. This study aimed to assess the morphological variation of the Saccharum complex, especially Saccharum s.s. and Erianthus s.l., and contribute to clarifying their generic circumscriptions. To accomplish this purpose, 31 exomorphological characters (17 qualitative and 14 quantitative) were analyzed using cluster (UPGMA) and discriminant analyses. The existence of significant differences among groups was analyzed by a one-way MANOVA. The morphological evidence clearly indicated three groups: (1) Saccharum s.s. from the Old World, (2) Tripidium + some species of Saccharum s.l. sometimes recognized as Narenga, Miscanthus, or Miscanthidium (all taxa from the Old World), and (3) Erianthus s.s. from the New World. The results support that the native species from the New World (Erianthus s.s.) are morphologically different from the species of Saccharum s.s. often cultivated, suggesting that Erianthus should be accepted as a genus distinct from Saccharum. For the species of Tripidium, we provide the morphological traits that distinguish them from Saccharum s.s. and Erianthus s.s. and revalidate the recent differential recognition postulated by molecular works.

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Acknowledgements

We thank the directors and curators of the CTES, ICN, K, MO, and SI herbaria for the loan of specimens that made this study possible, Liliana Gómez for the illustration, Dr. Jinshun Zhong (Max Planck Institute for Plant Breeding Research) for the translation of the Chinese labels of some specimens, and Dr. Maria S. Vorontsova (Royal Botanic Gardens, Kew) for her help in understanding some specimen labels of material from K herbarium. Steven P. Sylvester and an anonymous reviewer are thanked for constructive comments and edits on an earlier version of the manuscript. Funding of this research was provided by grants from the Secretaría General de Ciencia y Técnica-Universidad Nacional del Nordeste (PI A013-2013, A009-2017). CADW also thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil (CNPq- 426334/2018-3, 441760/2020-1, 307125/2020-3) and Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG - APQ-01222-21, APQ-03365-21) for grants and fellowships.

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Appendix

Appendix

Specimens of the Saccharum complex analyzed for the morphometric analysis. The initial number of each sample corresponds to the numbers used in the UPGMA analysis

Saccharum alopecuroides (L.) Nutt. 1. U.S.A., Arkansas, D. Demaree 24038 (MO). 2. U.S.A., Missouri, B. Summers 4733 (MO). 3. U.S.A., Tennessee, T.H. Kearney 939 (MO). S. angustifolium (Nees) Trin. 4. ARGENTINA, Corrientes, M.C. Peichoto 176 (CTES). 5. BRAZIL, Rio Grande do Sul, R. Lerina and G.H. Silveira 94 (CTES). 6. BRAZIL, Rio Grande do Sul, H.M. Longhi-Wagner and C.A.D. Welker 10527 (CTES). 7. PARAGUAY, Presidente Hayes, J.R. Ramírez 275 (BAA). 8. URUGUAY, Durazno, C.A.D. Welker 644 (CTES). 9. URUGUAY, Rocha, C.A.D. Welker 628 (CTES). S. asperum (Nees) Steud. 10. ARGENTINA, Corrientes, S. Tressens et al. 3141 (CTES). 11. ARGENTINA, Misiones, C.A.D. Welker and M.C. Peichoto 583 (CTES). 12. ARGENTINA, Misiones, F.O. Zuloaga et al. 4909 (CTES). 13. BRAZIL, Paraná, E.P. Santos and J. Marzinek 748 (CTES). 14. BRAZIL, Rio Grande do Sul, H.M. Longhi-Wagner and C.A.D. Welker 10673 (CTES). 15. BRAZIL, Rio Grande do Sul, H.M. Longhi- Wagner and C.A.D. Welker 10683 (CTES). 16. BRAZIL, Santa Catarina, A. Krapovickas et al. 23047 (CTES). 17. BRAZIL, Santa Catarina, C.A.D. Welker 366 (CTES). 18. PARAGUAY, Misiones, B. Rosengurtt 5775 (BAA). S. balansae (Hack.) Roberty 19. ARGENTINA, Corrientes, A. Burkart 19731 (CTES). 20. ARGENTINA, Corrientes, A. Schinini and R. Carnevalli 10334 (CTES). 21. ARGENTINA, Misiones, F.O. Zuloaga and O. Morrone 6532 (CTES). 22. PARAGUAY, Amambay, A. Krapovickas et al. 46050 (CTES). 23. PARAGUAY, Caaguazu, B. Balansa 301 (SI). 24. PARAGUAY, Cordillera, E. Hassler 12661 (CTES). 25. PARAGUAY, Misiones, T.M. Pedersen 4408 (CTES). S. baldwinii Spreng. 26. U.S.A., Arkansas, D. Demaree 8401 (MO). 27. U.S.A., Arkansas, J.M. Greenman 4385 (MO). 28. U.S.A., Arkansas, G.W. Letterman 9 (MO). S. brevibarbe (Michx.) Pers. 29. U.S.A., Alabama, H. Eggert s.n. (MO-967128). 30. U.S.A., Arkansas, B. Van Schaack 3625 (CTES). 31. U.S.A., Tennessee, A. Ruth s.n. (MO-775839). 32. U.S.A., Tennessee, A. Ruth s.n. (MO-2967133). S. coarctatum (Fernald) R.D.Webster 33. U.S.A., Alabama, R. Kral 33679 (MO). 34. U.S.A., Alabama, R. Kral 41662 (MO). 35. U.S.A., Georgia, C.L. Quarin 3605 (CTES). S. contortum (Elliott) Nutt 36. U.S.A., Alabama, R. Kral 94643 (MO). 37. U.S.A., Texas, H.E. Moore 1053 (ICN). S. ecklonii (Nees) Steud. 38. SOUTH AFRICA, C.F. Ecklon and C.L.P. Zeyher 621 (MO). 39. SOUTH AFRICA, Kwazulu-Natal, G. Davidse 6923 (MO). 40. SOUTH AFRICA, Kwazulu-Natal, E.A. Kellogg 1144 (SI). S. giganteum (Walter) Pers. 41. U.S.A., Arkansas, P.O. Tinsman 22700 (MO). 42. U.S.A., Louisiana, G.A. Brown 1445 (BAA). 43. U.S.A., Louisiana, L.H. Shinners 22077 (ICN). 44. U.S.A., Delaware, J.R. Churchill 93 (MO). 45. U.S.A., South Carolina, R.K. Godfrey 8192 (MO). 46. U.S.A., Texas, D.J. Stanley and J. Wipff 2105 (MO). S. intermedium Welker & Peichoto 47. ARGENTINA, Corrientes, M.M. Arbo et al. 8052 (CTES). 48. ARGENTINA, Corrientes, C.A.D. Welker and M.C. Peichoto 556 (CTES). 49. ARGENTINA, Misiones, C.A.D. Welker and M.C. Peichoto 584 (CTES). 50. BRAZIL, Rio Grande do Sul, C.A.D. Welker 538 (CTES). 51. BRAZIL, Santa Catarina, C.A.D. Welker 502 (CTES). S. narenga (Nees ex Steud.) Wall. ex Hack. 52. CHINA, Guangdong, L. Deng 10348 (MO). 53. CHINA, Guangdong, Z. Huang 42315 (MO). S. officinarum L. 54. ARGENTINA, Corrientes, M.M. Arbo 352 (CTES). 55. ARGENTINA, Corrientes, R.D. Medina 71 (CTES). 56. ARGENTINA, Corrientes, T.M. Pedersen 12488 (CTES). 57. ARGENTINA, Corrientes, M.C. Peichoto 172 (CTES). 58. ARGENTINA, Corrientes, M.C. Peichoto and E. Lattar 333 (CTES). 59. ARGENTINA, Corrientes, M.C. Peichoto and E. Lattar 334 (CTES). 60. ARGENTINA, Misiones, H. Keller 893 (CTES). 61. ARGENTINA, Misiones, V. Maruñak 122 (CTES). 62. ARGENTINA, Santa Fe, R.D. Medina 70 (CTES). 63. PARAGUAY, Central, C.L. Quarin et al. 1616 (CTES). S. robustum E.W.Brandes & Jeswiet ex Grassl 64. NEW GUINEA, Halalinja, P. Sillitoe and J. Sillitoe 43 (K). 65. NEW GUINEA, Papua, E.W. Brandes 41989 (K). S. spontaneum L. 66. ARGENTINA, Chaco, A.G. Schulz 15388 (CTES). 67. ARGENTINA, Salta, M.C. Peichoto and J. Gonzalez 337 (CTES). S. villosum Steud. 68. ARGENTINA, Corrientes, M.C. Peichoto 183 (CTES). 69. ARGENTINA, Corrientes, C.A.D. Welker and M.C. Peichoto 560 (CTES). 70. ARGENTINA, Misiones, M.C. Peichoto 184 (CTES). 71. ARGENTINA, Misiones, M.C. Peichoto 189 (CTES). 72. ARGENTINA, Misiones, C.A.D. Welker and M.C. Peichoto 575 (CTES). 73. BOLIVIA, Santa Cruz, T. Killeen 1787 (LPB). 74. BOLIVIA, Santa Cruz, T. Killeen 2219 (LPB). 75. BRAZIL, Rio Grande do Sul, C.A.D. Welker 477 (CTES). 76. PARAGUAY, Amambay, G.J. Schwarz 12180 (CTES). Tripidium arundinaceum (Retz.) Welker, Voronts. & E.A.Kellogg 77. CHINA, Hong Kong, H. Shiu-Ying 11199 (K). 78. CHINA, Kiangsi, H.C. Cheo 484 (K). 79. JAPAN, Ryukyu, M. Furuse 1790 (K). T. bengalense (Retz.) H.Scholz 80. INDIA, Uttarakhand, H.H. Haines 3950a (K). 81. TRINIDAD AND TOBAGO, Tobago, W.E. Broadway 4681 (MO). T. kanashiroi (Ohwi) Welker, Voronts. & E.A.Kellogg 82. U.S.A., Florida (original collection from Okinawa, Japan), C. Grassl s.n. (K). T. procerum (Roxb.) Welker, Voronts. & E.A.Kellogg 83. BANGLADESH, Chittagong, J.D. Hooker and T. Thomson 308 (K). 84. CHINA, Hainan, L. Deng 2580 (MO). 85. CHINA, Guangdong, D. Guangoi 1771 (MO). T. ravennae (L.) H.Scholz 86. ARGENTINA, Buenos Aires, Z.E. Rúgolo 2165 (SI). 87. ARGENTINA, La Pampa, Z.E. Rúgolo 2343 (CTES). 88. RUSSIA, Krasnodar, N. Jzevelev 21 (MO). T. strictum (Host) H.Scholz 89. IRAQ, Jebel, H. Field and Y. Lazar 801 (MO). 90. TURKEY (seed from between Istanbul and Corlu, plant grown at College Station), F.W. Gould 206639 (MO).

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Vasquez, S.I., Welker, C.A.D., Solís Neffa, V.G. et al. Morphometric analysis of the Saccharum complex (Poaceae, Andropogoneae). Plant Syst Evol 308, 10 (2022). https://doi.org/10.1007/s00606-021-01801-z

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