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The fern genus Saccoloma (Saccolomataceae) in Brazil, including new species and new combinations

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Abstract:

Saccoloma is a medium-sized genus of ferns with taxonomic uncertainties and pending nomenclatural clarifications. Within the last 150 years, only four names have constantly been used for the Brazilian species. After a bibliographic review and analyses of ca. 270 collections, I present an updated taxonomic revision for those species. Here, nine species are recognized, of which four occur in Amazonia, four in the Atlantic Forest, and one in the Cerrado (= Brazilian savanna). They are: S. chartaceum, S. hieronymi, S. inaequale s.str., and S. membranaceum from Amazonia; S. catharinae, S. distans, S. elegans, and S. nigrescens from the Atlantic Forest; and S. matto-grossense, from the Cerrado. I present a key to species, descriptions, illustrations, maps of distribution, complete type information, nomenclatural clarifications, and full synonymy regarding Brazilian names. The name S. nigrescens (Mett.) A.Rojas, recently applied to plants from Costa Rica to Bolivia, is shown to be based on erroneous basionym and type information; Saccoloma nigrescens(Kunze) A.Rojas is the correct authorship, and it applies to a species endemic to the Atlantic Forest. Saccoloma hieronymi should be used for plants from Costa Rica to Bolivia. In total, 16 spp. are currently recognized for the Neotropics, and 24 worldwide (plus four varieties). I also present a preliminary, maximum likelihood, phylogenetic analysis using three chloroplast genes (atpB, rbcL, and rps4) to test the monophyly of Saccoloma and Orthiopteris. In the phylogeny, the 1-pinnateS. chartaceum (= Saccoloma s.str.) is nested among the 2–4-pinnate species of the genus, to which the name Orthiopteris has sometimes been applied. Orthiopteris could be applied, if desired, solely to the Old World species of the genus, which form a clade that is sister to the New World species. The following names are lectotypified: Davallia distans, D. inaequalis, D. nigrescens, Microlepia nigricans, S. elegans (second step lectotypification), and S. elegans var. costaricense.

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Acknowledgments

I am extremely grateful to Larissa Orlova (LE), who was able to find the type of Davallia distans at LE. I also thank Alexei Oskolski (Univ. Johannesburg) for getting me in touch with Larissa. I thank the curators and staff of the visited herbaria (see Material and Methods), especially Rafaela Forzza (RB), Claudine Mynssen (RB), and Paulo Labiak (UPCB). I thank Michel Boudrie (CAY and P) for sending images of some literature, Reinaldo Pinto for the illustrations (Figs. 2–9), Túlio Pena (VIC) for helping with the composition of Figs. 1 and 10 and field work, Samuli Lehtonen (TUR) and Hanna Tuomisto (TUR) for sending images of specimens, and Jefferson Prado (SP) for discussion of nomenclatural issues of Saccoloma nigrescens. I also thank the editor (Benjamin Torke, NY), the handling editor (Robbin Moran, MO and NY), and the two reviewers (Lana Sylvestre, RBR, and an anonymous reviewer) for improving the quality of the manuscript. The phylogenetic work was made possible by my sabbatical period at the University of Vermont, Burlington, in 2018. For that visit, I thank CNPq (grant no. 204998/2017-4) for financial support, and Michael Sundue (VT) and David Barrington (VT) who served as my hosts. Finally, I thank Raquel Santana for her support in field work and discussions during the writing of the manuscript.

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  1. Laboratory of Systematics and Evolution of Plants, Department of Plant Biology, Federal University of Viçosa, Av. Peter Henry Rolfs s.n., Viçosa, MG, CEP 36570-900, Brazil

    Pedro B. Schwartsburd

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Correspondence to Pedro B. Schwartsburd.

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Appendices

Appendices

A ppendix 1. Index to collectors. Only the senior collector is listed. Types are in bold. Species numbers follow the alphabetic sequence in the text: 1. Saccoloma catharinae, 2. Saccoloma chartaceum, 3. Saccoloma distans, 4. Saccoloma elegans, 5. Saccoloma hieronymi, 6. Saccoloma inaequale, 7. Saccoloma matto-grossense, 8. Saccoloma membranaceum, and 9. Saccoloma nigrescens

Anderson, W. R. 12,270 (6).

Anonymous: ex herb. Mertens (3).

Baber, K.: 42 (4), 248 (4).

Baeta, A.: s.n. (4).

Barros, W. D.: 668 (4).

Beyrich: s.n. (4).

Bigio, N. C.: 798 (6).

Bisby, F.: P18100 (6).

Black, G. A.: 263 (2), 1525 (6), 1736 (2).

Blanchet, B.: 66 (9), 2507 (9), s.n. (9).

Boone, W.: 1127 (4).

Bovini, M. G.: 3401 (6).

Brade, A. C.: 7730 (3), 10,224 (4), 11,328 (9), 14,022 (9), 14,107 (4), 16,458 (4), 18,177 (4), 18,322 (9).

Burchell, W. J.: 2463 (4).

Cadorin, T. J.: 206 (4), 213 (1), 426 (3).

Caglioni, E.: 77 (4).

Camargo, R. F. N.: s.n. (4).

Campos-Porto, P: 2614 (4), s.n. (9).

Canestraro, B. K.: 648 (3).

Cavalcante, P.: 2662a (2).

Christenhusz, M. J. M.: 3022 (Saccoloma dominguense), 4233 (Saccoloma sp.), 4801 (3).

Cid, C. A.: 2574 (2).

Claussen, M.: s.n. (4).

Costa, M. A. S.: 540 (6).

Costa, M. S.: 14 (6), 37 (2).

Croat, T. B.: 85701 (6).

Daly, D. C.: 1851 (2), 1952 (2).

Demuner, V.: 4508 (4).

Duarte, A.: s.n. (4).

Dusén, P. K. H.: 14711 (3).

Edwards, P. J.: 2413 (4).

Egler, W. A.: 46429 (6).

Engelmann, R. A.: 1519 (4).

Engels, M. E.: 2965 (7), 2996 (7), 3179 (7), 4733 (7).

Esteves, G. L.: 1684 (4).

Fernandes, H. Q. B.: 1162 (9).

Figueiredo, J. B.: 25 (3).

Freyreiss, G. W.: s.n. (3).

Fróes, R. L.: 24629 (2), 26,301 (6).

Funez, L. A.: 970 (4).

Gardner, G.: 159 (4).

Gasper, A. L.: 751 (4), 1507 (4), 2152 (1), 2225 (4).

Gaudichaud, C.: 164 (3), s.n. (3).

Glaziou, A. F. M.: 386 (3), 2376 (4).

Goldenberg, R.: 987 (5).

Gomes, M. G.: 19 (3).

Gonçalves, K. S.: 65 (6), 131 (6).

Graçano, D.: 65 (4), 117 (4).

Granville, J. J.: 12333 (6).

Gurgel, E. S.: 83 (3).

Handro, O.: 2209 (4).

Hatschbach, G.: 23304 (3), 49,928 (4), 58,544 (3), 69,118 (4).

Heringer, E. P.: 1281 (4).

Jangoux, J.: 85–100 (2).

Jansen-Jacobs, M. J.: 338 (2).

Jones, M.: 627 (2); 1019 (6).

Kerling, M.: s.n. (1).

Kollmann, L.: 599 (9), 4910 (9), 5801 (4), 6258 (9), 7652 (4), 9078 (4).

Korte, A.: 291 (4), 2435 (4), 4030 (4).

Krause, R. A.: 1 (4), 82 (4).

Krieger, L: 11133 (3), 11,755 (4), 12,313 (6), 17,318 (6).

Labiak, P. H.: 1884 (4), 3594 (3), 3998 (4), 4637 (9), 7162 (6).

Lautert, M.: 486 (7).

Leoni, L. S.: 944 (4).

Lleras, E.: P16887 (5), P17407 (6), P17438 (6).

Lopes, M. S.: 173 (4), 323 (4), 360 (4), 399 (4), 441 (4), 485 (9), 564 (4), 572 (4).

Luetzelburg, P.: 208 (3).

Macedo, T. S.: 153 (4).

Marquete, R.: 1103 (9).

Martius, C. F. P.: Herb. Fl. Bras. 379 (9), s.n. [Ilheús] (9), s.n. [Guyana Brasilianae] (9).

Matos, F. B.: 666 (4), 692 (4), 811 (4), 813 (9), 952 (4), 997 (9), 1070 (9), 1131 (4), 1178 (3), 1289 (3), 1977 (3).

Mazziero, F. F. F.: 811 (4), 926 (4), 1135 (3).

McFarland, K.: 317 (6).

Mello-Barreto, H. L.: 1601 (4).

Mexia, Y.: 4639 (4), 4964 (4), 8429 (2).

Meyer, F. S.: 639 (3).

Milliken, W.: 543 (2), s.n. (2).

Moreira, E. V.: 6 (3).

Motta, J. T.: 4395 (3).

Mynssen, C. M.: 399 (4), 664 (4), 932 (4), 1091 (3), 1215 (9), 1306 (3), 1359 (3).

Müller, F.: s.n. (3).

Müller, I. O.: Rosenst. Filic. Austro-Bras. 54 (3).

Neuwied, M.: s.n. (4).

Nóbrega, G. A.: 326 (3? or 3 × 9?).

Pena, N. T. L.: 1037 (3).

Pereira, O. J.: 1606 (9), 1671 (4).

Pietrobom, M. R.: 4828 (4), 4886 (4), 4995 (4), 7861 (5).

Plowman, T.: 12167 (6).

Poeppig, E. F.: 2113 (6).

Pohl, J. B. E.: 60 [5261] (3).

Poliquesi, C. B.: 728 (3).

Poole, J. M.: 1953 (2).

Prado, J.: 353 (9), 499 (3).

Prance, G. T.: 1784 (2), 9692 (6), 9932 (6), 10,076 (2), 10,821 (2), 12,194 (6), 12,432 (8), 12,574 (6), 13,422 (6), 13,610 (6), 13,611 (2), 14,649 (2), 23,819 (6), 58,897 (2).

Pruski, J.: 3497 (6).

Quesada, F.: 1082 (2).

Quinet, A.: 1392 (6).

Regnell, A. F.: 265 (3).

Reitz, R.: H196 (3), 1797 (4), 3057 (4), 3139 (4), 3168 (3), 3176 (1), 3287 (3), 3454 (4).

Ribeiro, M. S.: 80 (6).

Riedel, L.: 51 (4), s.n. (3), s.n. (4).

Rosa, N. A.: 602 (6).

Salino, A.: 317 (7), 349 (7), 726 (4), 3428 (4), 3441 (4), 6595 (3), 10,329 (3).

Schmitt, J. L.: 401 (1), 1674 (4), 1854 (4).

Schultes, R. E.: 9426 (6).

Schwartsburd, P. B.: 1640 (3), 2266 (3), 2618 (4), 4815 (3), 4816 (3), 4819 (4), 4820 (4), 4824 (3).

Secco, R. S.: 269 (2), 471 (2).

Sellow, F.: s.n. [B. 249. C. 89] (4).

Silva, J. M.: 3602 (3).

Smith, L.: 5683 (4), 6045 (4).

Sota, E. R.: s.n. (3).

Souza, M. G. C.: 215 (6).

Sperling, C. R.: 6380 (2).

Spuce, R.: 2120 (6).

Steward, W. C.: 153 (2).

Stival-Santos, A.: 1108 (4), 2218 (4), 2122 (1).

Stübel, A.: 1129 (5).

Sylvestre, L. S.: 524 (9), 666 (4), 781 (4), 2284 (9).

Taylor, W. C.: 5052 (6).

Thomas, W. W.: 11967 (4).

Tuomisto, H.: 14409 (Saccoloma sp.); 14,948 (2).

Valente, G. E.: 1071 (4).

Vervloet, R. R.: 470 (4), 581 (9), 673 (4), 720 (9), 721 (4), 1191 (9), 1915 (9).

Wacket, M.: 201 (4).

Webb, J. W.: s.n. (3).

Wercklé, C.: s.n. (2).

Wettstein, R.: s.n. (9).

Windisch, P. G.: 6413 (7).

Xavier, S. R. S.: 127 (4).

A ppendix 2. Taxon, voucher information (when available), and GenBank accession numbers of atpB, rbcL, and rps4, respectively. The m-dash means “not available”

Cyathea poeppigii: AF313553, —, —. Cythea poeppigii, COAH 61235: —, KY458194, —. C. poeppigii, Conant 4410: —, —, FN667569. Dennstaedtia punctilobula: U93836, —, —. D. punctilobula, Schwartsburd 4480: —, MT416336, —. D. punctilobula: —, —, AY459159. Histiopteris incisa, Smith s.n.: EF463374, —, —. H. incisa, SG 1331: —MH289644, MH289684. Hiya brooksiae, SG 1730: —, MH289638, MH289710. Hypolepis punctata, FLX 6: —, MG944784, MH289689. Hypolepis tenuifolia, Schuettpelz 286: EF463375, —, —. H. tenuifolia, HN 31: —, MG944786, MH289701. Llavea cordifolia: EF452039, —, —. Llavea cordifolia: —, U27726, —. Llavea cordifolia, RK 1380 273: —, —, DQ914173. Lonchitis hirsuta, MC 3508: KJ716390, —, —. L. hirsuta: —, U05929, —. L. hirsuta: —, —, AY459162. Microlepia hookeriana, WYD 218: —, MH289650, MH289714. Microlepia platyphylla: U93832, —, —. Microlepia platyphylla: —, U18642, —. Microlepia platyphylla: —, —, AY612668. Monachosorum flagellare, ZXL 7419: —, MH289646, MH289702. Paesia radula, SG 1752: —, MH289637, MH289712. Pteridium aquilinum: U93835, —, —. Pteridium aquilinum (subsp. wightianum), SG 1760: —, MG944787, MH289709. Saccoloma chartaceum (as Saccoloma elegans), MJ [Jones] 627: —, KJ628821, —. Saccoloma chartaceum (as Saccoloma elegans), Tuomisto 14,948: HQ157279, HQ157302, GU478639. Saccoloma distans (as Saccoloma brasiliense), CM [Mynssen] 1091: KJ716391, KJ628819, —. Saccoloma dominguense, MC [Christenhusz] 3022: KJ628519, KJ628820, —. Saccoloma ferulaceum, WELT P025294: —, KT983836, —. Saccoloma inaequale, Jones 1019: KM114062, KJ628823, MG561410. Saccoloma cf. inaequale (as Saccoloma inaequale): AY612708, AY612682, AY459165. Saccoloma cf. inaequale (as Saccoloma inaequale): EU352283, EU352310, —. Saccoloma kingii (as Orthiopteris kingii): —, U18644, —. Saccoloma sp. 1 (as Saccoloma inaequale), Tuomisto 14,409: —, —, GU478638. Saccoloma sp. 2 (as Saccoloma inaequale), Christenhusz 4233: EF463520, EF463265, —.

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Schwartsburd, P.B. The fern genus Saccoloma (Saccolomataceae) in Brazil, including new species and new combinations. Brittonia 74, 155–179 (2022). https://doi.org/10.1007/s12228-021-09689-2

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