Panicum (Poaceae) (Flora Neotropica, 124) 3031337670, 9783031337673

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Flora Neotropica

Fernando Zuloaga Sandra Aliscioni

Panicum (Poaceae)

Flora Neotropica Volume 124

Since 1968, these monographic volumes have provided taxonomic treatments of families or other groups of plants or fungi growing in the Americas between the Tropic of Cancer and the Tropic of Capricorn. The monographs are intended to be comprehensive, so most of them also include information on economic botany, conservation, phylogenetic relationships, taxonomic history, ecology, cytology, anatomy, and phytochemistry, among other topics. Each volume is illustrated with line drawings, black and white photographs, and distribution maps. This series is the official publication of the Organization for Flora Neotropica (OFN), an organization founded in 1964 with the mission of producing a published botanical inventory of the American tropics. The current OFN Executive Director is Wm. Wayt Thomas. Flora Neotropica volumes 1–14 (1968–1974) were published by Hafner Publishing Company. Volumes 15–121 (1975  – 2019) were published by The New  York Botanical Garden. All in-print volumes 1—121 are available exclusively through NYBG Shop: https://www.nybgshop.org/nybg-press/books-in-series/flora-­ neotropica. Volumes 122 and going forward are available exclusively from Springer.

Fernando Zuloaga • Sandra Aliscioni

Panicum (Poaceae)

Fernando Zuloaga Labarden 200 Instituto de Botanica Darwinion San Isidro, Buenos Aires, Argentina

Sandra Aliscioni Instituto de Botanica Darwinion San Isidro, Buenos Aires, Argentina

ISSN 0071-5794     ISSN 2330-202X (electronic) Flora Neotropica ISBN 978-3-031-33767-3    ISBN 978-3-031-33768-0 (eBook) https://doi.org/10.1007/978-3-031-33768-0 © The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Paper in this product is recyclable.

List of Taxa

1. Panicum aquaticum Poir. 1a. P. aquaticum var. aquaticum 1b. P. aquaticum var. cartagoense Davidse 2. P. dichotomiflorum Michx. 2a. P. dichotomiflorum subsp. dichotomiflorum 2b. P. dichotomiflorum subsp. bartowense (Scribn. & Merr.) Freckmann & Lelong 3. P. lacustre Hitchc. & Ekman 4. P. sublaeve Swallen 5. P. vaseyanum Scribn. ex Beal 6. P. altum Hitchc. & Chase 7. P. amarum Elliott 7a. P. amarum subsp. amarum 7b. P. amarum subsp amarulum (Hitchc. & Chase) Freckmann & Lelong 8. P. chloroleucum Griseb. 9. P. complanatum Guglieri, Longhi-Wagner & Zuloaga 10. P. deciduum Swallen 11. P. glabripes Döll 12. P. longissimum (Mez) Henrard 13. P. mystasipum Zuloaga & Morrone 14. P. olyroides Kunth 14a. P. olyroides var. olyroides 14b. P. olyroides var. hirsutum Henrard 15. P. racemosum (P. Beauv.) Spreng.

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List of Taxa

16. P. tricholaenoides Steud. 16a. P. tricholaenoides var. tricholaenoides 16b. P. tricholaenoides var. flavomarginatum (Mez) Zuloaga 17. P. urvilleanum Kunth 18. P. virgatum L 19. P. alatum Zuloaga & Morrone 19a. P. alatum var. alatum 19b. P. alatum var. minus (Andersson) Zuloaga & Morrone 20. P. aquarum Zuloaga & Morrone 21. P. aztecanum Zuloaga & Morrone 22. P. bergii Arechav. 22a. P. bergii var. bergii 22b. P. bergii var. pilosissimum Zuloaga 23. P. capillare L. 24. P. capillarioides Vasey 25. P. chaseae Roseng., B.R. Arrill. & Izag. 26. P. decolorans Kunth 27. P. diffusum Sw. 28. P. ephemeroides Zuloaga & Morrone 29. P. exiguum Mez 30. P. furvum Swallen 31. P. ghiesbreghtii E. Fourn. 32. P. hallii Vasey 32a. P. hallii subsp. filipes (Scribn.) Freckmann & Lelong 32b. P. hallii subsp. hallii 33. P. hirsutum Sw. 34. P. hirticaule J. Presl 34a. P. hirticaule var. hirticaule 34b. P. hirticaule var. verrucosum Zuloaga & Morrone 35. P. hispidifolium Swallen 36. P. lepidulum Hitchc. & Chase 37. P. magnispicula Zuloaga, Morrone & Valls 38. P. miliaceum L. 39. P. mucronulatum Swallen 40. P. parcum Hitchc. & Chase 41. P. peladoense Henrard 42. P. quadriglume (Döll) Hithch. 43. P. stramineum Hitchc. & Chase 44. P. tamaulipense F.R. Waller & Morden 45. P. venosum Swallen

List of Taxa

46. P. coloratum L. 47. P. gouinii E. Fourn. 48. P. pedersenii Zuloaga 49. P. repens L. 50. P. campestre Nees ex Trin. 51. P. cayennense Lam. 52. P. cervicatum Chase 53. P. ligulare Nees ex Trin. 54. P. rudgei Roem. & Schult. 55. P. bartlettii Swallen 56. P. brevifolium L. 57. P. haenkeanum J. Presl 58. P. hirtum Lam. 59. P. millegrana Poir. 60. P. sellowii Nees 61. P. trichanthum Nees 62. P. trichoides Sw. 63. P. venezuelae Hack.

vii

List of Exsicatta

Abbott 1027 (27), 16824 (54) Abrams 2164 (17); 12902 (43) Abruzzi 611 (61) Acevedo et al. 10260 (54) Adams & Bhorai 14132 (62) Adams 6 (14b); 7574, 9033 (62) Agniel & Arsène 10269 (26) Agniel 10393 (26) Aguayo 421 (61) Aguilar 378 (55) Aguirre Galviz 877 (54) Ahart 6168 (38) Ahumada & Castellón 4829, 5023 (43) Ahumada 423, 425, 610, 791, 919, 1673, 1790, 1872, 1924 (2a), 2143 (1a), 2352 (61) Alberdi 12269 (38) Alcorn 2766 (55) Aldave & Fernández 5634 (61) Allard 14148 (31); 14803 (62); 15744, 15902 (27); 16060, 16067 (51) Allem 345, 390 (14b), 1194 (11); 1542, 2146, 2185, 2264 (2a); 2145 (43); 2220 (16a); 2233 (29) Allem & Vieira 1001 (51); 1194, 2506 (16a); 1200, 1240, 1281, 1379, 1615, 2497, 2577 (2a); 1479 (43); 1549, 1552 (14b); 1686 (49); 1819, 1844 (47); 1988 (60) Allemao & Cysneiros 1616 (62) Allen 249 (60); 659 (51); 1317 (57) Almeida 355, 1036 (50); 808 (14a) Alonso Paz 3717 (15) Alston 5235, 5298 (31); 5944 (57); 6053(61) Alstroemer s.n. (49) Alva 53-10 (14a) Alvárez 603 (22a)

ix

x

List of Exsicatta

Anderson 897, 899, s.n. (34a); 1156 (60); 1365 (35); 1366 (4); 2096 (23); 6515 (50); 7381 (29) Anderson et al. 35148, 35584 (50); 36654 (52); 37036 (63); 37074 (62) Andersson s.n. (19b) Andrade Lima 71-6363 (29) Angulo 1083 (58) Araujo 151, 186, 527 (25); 216 (11); 289 (22a); 4118 (15) Arbo et al. 785, 981 (2a); 976 (60) Archer 1462 (61); 2360 (62); 2745 (54); 2892 (60); 3400 (26); 3968 (31); 4289 (15); 7587 (51) Arechavaleta 24a (22a); s.n. (16a); (22a) Arenas 668 (61); 1226 (60) Aristeguieta 1617, 2141 (16a); 4048 (54); 3357, 3360 (63); 4301, 4334, 5665, 5755 (35); 5811 (58) Aristeguieta & Foldats 1355 (61) Aristeguieta & Virrueta 7610 (14a) Ariza Espinar 140 (17) Arséne 2417, 2864, 2985 (36) Arzivenco 640 (23); s. n. (60) Asplund 5281, 5575 (62); 5512 (61); 10846, 15816 (60) Atrupo 1723 (32b) Atwood & Seymour 3546 (58) Avendaño et al. 112 (61) Aviña 1248 (34b) Axelrod & Axelrod 2334 (62) Aymard & Delgado 8249, 8361, 8363 (54) Aymard & Ortega 1413, 4930 (62) Aymard et al. 296 (14a); 7078 (62) Azevedo & Filgueiras 950 (41) Bacigalupo & Deginani 104 (16a) Bacigalupo et al. 844, 1041 (11) Badillo 7, 690 (31); 10 (16b); 60 (63); 67 (61); 146 (31); 793 (62); 1410 (54); 3617 (16a); 3696 (14a) Báez s.n. (38) Baker 5 (62) Balansa 10 (16b); 12, 15 (22a); 13, 14, 4357 (41); 17 (48); 18 (16a); 19, 61 (2a); 23 (14b); 24, 24a, 2954, 4358 (14a); 2935 (12); 2941, 2942, 4355 (60); 46, 4356 (61); 47, 48, 48b (60); 51, 52 (60) Baldwin Jr. 14321 (26) Balick 133 (29) Ball 20, 120 (23) Bang 493 (42) Barbour 2145 (62) Barfod et al. 48489 (62) Barkley et al. 701, 7639 (26)

List of Exsicatta

xi

Barlow 12/7A (7a) Barneby 2455 (32b) Baro 513 (7b); 633 (7a) Barreto & Kappel s.n. (25) Barreto s.n. (2a) Barrett s.n. (49) Bartlett 8577 (62); 10549, 10591 (32b); 11452, 12016, 12245 (55); 20313 (61); 20715, 21218 (15); 21345 (22a) Bartlett & Lasser 16990 (2a) Basile 51, 52 (26) Bastos 152 (62) Basurto & Patrón 177 (62) Beck 761 (29); 2654, 3302, 5028, 5050, 5108, 5150, 5321 (16a); 5685 (16a); 8344 (61); 15176 (43) Beck & Haase 9921 (22a); 9976 (51) Beetle 1031 (32a); 1061, 1077 (32b); 1486 (61); 1772 (22a) Beetle & Guzman 5513, 5536 (5) Beetle et al. M-547 (55); M-718 (44); M-917 (31); M-1211 (19a); M-2646, M-2551, M-7030 (34a); M-7324 (36); M-7325 (5) Belanger 389, 438 (62) Belém 1970 (52) Belém & Mendes 77 (14a) Belém & Pinheiro 2121 (54) Belshaw 3338 (62) Berg 208 (17) Bernal & Cárdenas s.n. (32b) Bertero s.n. (27) Bertoni 2710, 5849 (60); 3405, 3551, 4802, 5522 (2a); 4138, 4614 (16a); 5532, 5534 (41) Betancur 1496 (54) Betancur et al. 261 (62) Beyrich s.n. (60) Bianco 4760 (17) Biganzoli 1988 (15) Billiet & Jadin 3281 (61) Black 844b (60); 47-994, 52-15490, 55-18350, 55-18365 (62); 47-1866, 48- 3536, 51-13354, 54-16161 (60); 48-2914, 49-8010, 55-18217 (2a); 51-11072 (52); 51-11086, 51-11686 (42); 51-12507, 51-13176, 51-13323, 52-15586, s.n. (58); 52-15439 (61); 54-16109 (50); 56-18910 (49) Black & Avelino 86C, 88C (29) Black & Cordeiro 52-14754, 52-15178 (57); 52-14836, 52-14880, 52-15315 (16a); 52-15348 (54) Black & Engelhand 50- 9040 (1a); 50-89761/2 (49) Black & Lobato 50-9659 (51) Black & Magalhaes 51-13009 (2a)

xii

List of Exsicatta

Black & Marano 51-11413 (22a) Black & Pires 50-9734 (58) Black et al. 54- 16587 (2a); 54-16594 (43); 57-19425 (60) Blair 731 (29) Blake 7326A (34a) Blanchard s.n. (23) Blanchet 104, 2961 (39) Blanco et al. 512 (19b) Blodgett s.n. (2a) Blum 2076 (57) Blum & Tyson 481 (62) Blumer 1683 (32b) Blydenstein 148 (43); 204 (51); 777, 1524, 1644, 1748 (14a); 969, 1406, 1570, 1658, 1706 (54); 1439 (2a); 1846 (20) Blydenstein & Saravia 1333 (14a) Bobarti s.n. (23) Böcher et al. 689, 776 (17) Boege 848 (62); 3004 (7a) Boelcke et al. 15578, 15797 (17) Boerboom 8769 (60) Bokermann s n. (60) Bolaños s.n. (62) Bommer 54 (51) Bondar 2609 (63) Bono 212 (54); 2681 (62) Boom & Grillo 6106, 6115 (62); 6189 (35) Bordas & Schmeda 4253 (60) Bordo 1 (49) Borjas 94 (62) Borsotti 66 (2a) Bosc s.n. (2a) Boufford 7295 (18) Box 18, 39 (31); 90, 3908 (62); 176, 190, 3703 (33); 3137 (35); 389 (58); 3889, 3937 (57); 3952 Box & Alayón 3761 (63); 3833 (33) Brace 4812 (31); 7099 (2a) Brade 8144 (41); 9163 (60); 12810 (15) Braga 48 (60) Brandegee 4 (40); 8 (19a); 42 (19b) Braun s.n. (23) Breedlove 9183 (51); 10630 (19b); 11920, 52108 (60); 28117 (60); 28837 (33); 36779-A (43); 36852 (4); 36875, 52871 (40); 37573, 37654 (35); 38598 (60); 41540 (34a); 41585, 52930 (62); 56081 (34a)

List of Exsicatta

xiii

Breedlove & Davidse 53971 (5); 54030, 54047, 54080, 54401 (62); 54051 (61); 54125 (60); 54261 (40); 54319 (60); 54452, 54548, 54664 (4); 54908, 55442 (31); 55039 (18); 55245 (33); 55250 (51) Breedlove & Raven 13492, 13833 (40) Breedlove & Thorne 21270 (18) Brescia & Marchesi 4068 (60) Breteler 4039 (60) Brigada de Dunas 577 (7a) Britton 688 (54); 1380, 2356 (27); 2469 (7b) Britton & Brace 493 (7b); 424 (27) Britton & Coker 1754 (58) Britton & Cowell 4064 (27); 4142 (62) Britton & Earle 6566 (2a) Britton & Hazen 1563 (54) Britton & Millspaugh 6102 (2a) Britton & Schaffer 382 (27) Britton & Wilson 314 (27) Britton et al. 649, 6123 (27); 745, 6349 (31); 6520, 7275, 14455 (51); 6859, 13177 (27); 7201, 7284 (60) Brizuela 982 (43) Broadway 348 (54); 501, 5058 (62); 638, 733, 818 (60); 2629, 4586, 4895, 9291 (33); 3068, 5060 (6); 5909 (60); 6450 (58); s.n. (6) Brown & Britton 21 (23); 982 (2a); 1638 (2a) Bruderreck 93 (2a) Brunken & Perino 241 (32b); 327 (60); 360, 368 (40); 434 (26); 444 (36); 448 (5) Brunner 1719 (61); 1780 (29) Buchtien 32, 1159 (54); 53, 7114 (60); 445, 446, 3612 (61); 4174 (42) Bunch et al. 205 (62) Bunting 1681, 5663 (33); 5447, 5568 (43); 9396 (62); 10056 (34a) Bunting et al. 3637 (51); 3957, 6687, 11012 (54); 12029 (62) Burandt Jr. V0120 (44); V0192, V0688 (63); V0238 (33); V0326 (60) Burandt Jr. & Gantaume V0498 (29) Burandt Jr. & Wingfield V0242 (43) Burchell 6769 (42); 6794-2 (41); 6827 (29); 8791 (61); 8813 (20) Burger & Baker 9920 (62) Burger & Burger 7630 (60); 7875 (35) Burkart 997, 1080 (25); 1137, 1210, 1226, 3715, 3793, 7595, 14546, 17289 (2a); 7197, 24109 (38); 9864, 10858, 12239, 12260, 17651, 19203 (17); 12262, 20275 (43); 13325, 24109 (38); 14203, 20311 (61); 14367 (22b); 14455, 17743, 18397, 18643, 19692, 22915 (22a); 15943, 20264 (23); 16389 (62); 16447 (33); 16508, 16870, 17012 (31); 17013 (63); 17190, 17262 (35); 17204, 17206 (34a); 17269 (14a); 18380 (60); 28260 (60); 30232 (11) Burkart & Bacigalupo 21068 (61) Burkart & Gamerro 21593 (2a); 21661 (49) Burkart & Troncoso 26133 (11)

xiv

List of Exsicatta

Burkart et al. 13933 (17); 23252, 25275, 25276, 26815 (61); 23269 (23); 26865 (47) Burman & Filgueiras 416, 421 (60); 432 (14a); 435 (14b) Bush 858, 1156 (32b) Cabanillas et al. 15, 17 (34a) Cabrera 1249 (55); 1511 (54); 1607 (2a); 2006 (54); 2334 (14a); 2484, 7008 (22a); 3220, 5535, 8533 (15); 3418 (6); 3476 (7b); 5677, 6620, 6751, 6919 (17); 7453 (47); 10065 (15); 11739 (31); 28811 (2a) Cabrera & Cabrera 2237, 2461 (33); 2494 (55); 7002, 9877, 12881 (62); 9236 (7b) Cabrera & Cortez 273 (62) Cabrera & Sáenz 29041 (60) Cabrera & Villanueva 16374 (55) Cabrera & Zuloaga 32328 (22a); 32385, 32388 (41); 32416 (22a); 32423 (25) Cabrera et al. 2100 (55); 4447 (55); 21814, 24661, 30778, 31731, 32443, 32465, 32481 (8); 22167 (41); 25988, 28697, 32704 (22a); 26322, 26418, 26537, 27752, 28055, 30379 (61); 26542 (43); 27694 (60); 28691 (22b); 28853 (2a) Cabrera, Corte & Gebhard 86 (61) Callejas 499 (54) Callejas et al. 2197 (61); 4585 (62) Calzada & Arellano 2230 (55) Calzadas 15935 (21) Campelo 2154 (39); 2158, 2177 (49) Canabes 86 (62) Canby s.n. (7a) Cano & Cámara Hernandez 685 (47); 902 (2a) Canoniero 92 (22b) Cantino 636 (17) Cárdenas 213 (34a); 1315 (33); 1647, 3533, 3534 (43); 4503, 4506 (52) Cardona 2224 (54) Cardoso da Silva et al. 282 (16a) Carnevali 2976 (2a); 4095 (11) Carnevali et al. 1518 (54) Carrillo 9 (5) Carter 5016 (34a) Carter & Ferris 4726 (19a) Carvajalino & Díaz 42 (61) Casco 46 (34a) Castellanos 43 (41); 71 (62); 24653 (14a); 25909 (43); s.n. (61) Castillo 583 (43); s.n. (23) Castillón s.n. (8) Castro 69 (34a) Catalán & Terán 92 (40) Cavalcante 828 (14a) Chagas et al. 165 (14a)

List of Exsicatta

xv

Chase 2726, 4986, s.n. (23); 5766 (17); 5900, 7256 (32b); 6308, 6567, 6577 (27); 6547, 6744, 7480 (31); 6778, 7895, 8022, 8036, 8089, 8225, 9756, 10130, 10177, 10770, 10928 (1a); 7089 (36); 7090, 7270 (26); 7492, 7558 (44); 7518, 9471, 12578 (61); 7558 (32a), 7649, 7787, 7835, 11062, 12574 (62); 7675, 7883, 8465, 10745, 12841 (54); 7729, 8943, 9125, 10810, 10916, 10979, 11122, 11398 (60); 7786, 7789, 8066, 8070, 8135 (63); 7907, 11028 (2a); 8105, 8105bis (39); 8317 (60); 8358, 8815, 8856, 8645, 8995, 9354, 10533, 10637, 10790, 11198, 11315 (50); 8407 (7a), 8549, 9138, 10737, 10951, 11167, 11281, 11519, 11713, 11736, 11863, 11959 (52); 8696, 9279, 10534, 10589, 10665, 10697, 10723, 10780, 10786, 10886, 11207, 11280, 11858 (41); 8802 1/2, 8897, 9089, 10450, 10866, 10976, 11049, 11560, 11678, 11924, 11990, 12115 (42); 8804, 9046, 9230, 10625, 10731, 10785, 10890, 11054, 11179, 11324, 11587, 11707, 11756, 11860, 11952 (14a); 9023 (14b); 9982 (15); 10151 (22a); 10238 (9); 9300, 10725, 10846, 11119, 11465, 11482, 11638, 11641, 11791, 12839 (29); 11097 (16a); 11460, 11987 (53); 12518 (51); 12578 (61) Chavelas & Pérez 116 (51) Chavelas & Quero ES-109 (31) Chávez Alfaro 1730 (62) Chiang 191 (31) Chiang et al. 294, 9201 (32b) Christian s.n. (38) Churchill 7 (38) Cialdella et al. 226, 254, 460 (8) Clark 7653 (54) Clayton & Eiten 4675 (60), 4687 (47) Clayton 454 (23); 578 (18); 4204, 4961 (60); 4263 (60); 4366, 4507 (41); 4401 (47); 4526, 4795, 4842, 4875 (50); 4561, 4578 (60); 4598, 4809, 4839 (52); 4832 (2a); 4895 (14b) Clos 245, 1883 (47) Coelho de Moraes 677 (39); 850 (63); 851 (62) Cogollo 622 (62) Coile et al. 3840 (7b) Collins 37 (55); 139 (18); 151 (23) Colonello & Ramia 2808 (62) Colunga & Zizumbo 44 (34b) Comastri Filho 9 (16a) Combs 1220 (2a) Commerson s.n. (15) Conrad 2255 (60); 5973 (18) Conrad & Conrad 2959 (51) Contreras 628 (55); 3079 (6) Conzatti 3603 (36) Cook et al. 78 (27) Cooley 8165 (62) Coradin 80 (60)

xvi

List of Exsicatta

Coradin et al. 1263, 1987 (62); 2659 (58); 3394 (39) Cordini 20 (25) Cornman 638, 712 (57) Correa et al. 4382 (62) Correll 43251, 48142, 51350 (2b) Correll & Correll 50953 (2b); 50957 (7b) Correll & Evans 44024 (7b) Correll & Johnston 17274 (32a) Correll & Popenoe 46671 (2b); 51405 (49) Correll et al. 50462 (2b); 50262 (7b) Cortés 183, 192, 239 (14a) Costa Sacco 21 (15); 269, 361 (2a) Cowan 2590, 3119 (51); 2677, 31486, 38943 (54); 3398 (34a) Cristóbal et al. 1526 (16a) Cristoff 181 (34a) Croat 5254, 17446, 17954, 70846 (61); 21305 (60); 21646, 21701, 24867, 33618, 43887, 47521, 51131, 68576 (62); 23332, 23387 (55); 47699 (34a); 54021 (54) Crockett 7745 (18) Croft 240 (24) Crow & Rivera 7356 (1a) Crutchfield & Johnston 4956 (44); 6068 (32a) Cruz Cisneros 1490 (62) Cuatrecasas 4085, 4207 (58); 4148 (2a); 4216, 25501 (62); 4258 (14a); 7201 (54) Cuatrecasas & García Barriga 1980 (60); 3546 (61); 3729 (16a); 3840 (54) Cuatrecasas & Romero Castañeda 24998 (62) Cuello 418 (54) Cumana 732, 1258, 3790, 4377 (62) Cunquero 166 (17) Curran 250, 261 (33); 277 (54) Curtiss 177 (2a); 267 (51); 384, 494 (27); 598 (61); 714 (62); 3578, 5527 (7b); 6867 (38) Cutler 9084 (42) da Silva 1, 7, 124, 137 (29); 61 (61); 114 (61); 144 (42); 328 (2a) da Silva & Lopes 923 (13) Dalby 44 (22a) Damazio s.n. (50) Daniel 5520 (62) Daniel & Vasquez 64 (54) Daniels s.n. (18) D’Antoni 114, 115 (8) Darrow & Haskell 3419 (32b) Davidse 1854 (18); 2510, 18851 (33); 2806, 3713, 3724 (58); 2841, 3182, 5106, 5325, 5367. 5390 (54); 2925, 2950, 4314 (35); 2972, 3118, 3136, 4213, 18474 (62); 2997 (2a); 3160, 5108, 5132 (14a); 3909, 5416 (16a); 4319 (51); 4467 (60); 4477, 5008, 5031 (57); 4929 (52); 9587 (40); 9657, 9806, 9912 (36); 24106, 24541 (60)

List of Exsicatta

xvii

Davidse & Brant 31985 (57); 32042, 32392 (62); 32401 (33); 32456 (54); 32539, 32704 (55); 32638 (31); 32947 (1a) Davidse & D’Arcy 10132 (57); 10478 (49); 10541 (60) Davidse & Davidse 9396 (18); 9492 (31); 9927 (36); 10057 (32b) Davidse & González 13205, 14778 (2a); 13589, 21562 (62); 13666, 20995 (61); 13989, 19046 (57); 14217, 14667 (54); 14526 (16a); 19047 (58); 19266 (61); 19313 (14a); 21544 (60) Davidse & Huber 14916, 22517, 22530, 23061, 23118, 23125 (54); 15085 (60); 15117 (16a); 15430, 16811 (51); 22508, 22591 (14a) Davidse & Miller 26378 (51); 26918 (58); 27303 (60) Davidse & Pilz 31533 (35); 31651, 31704 (19b); 31665 (34a); 31674, 31697, 31712 (4) Davidse & Pohl 2155a (63) Davidse & Steyermark 18161 (22a) Davidse et al 1129, 4162, 11126, 11173, 12182 (14a); 3930, 20246, 20271, 20528, 20612, 29501, 30063 (33); 4548, 16770 (51); 10544, 10651, 11375 (50); 10590, 11437, 12246 (2a); 10618, 10712, 11145 (41); 11020, 11132, 11261, 11415, 17897 (60); 11221, 12194 (61); 11323, 23765 (60); 11697, 11716 (63); 12034 (1a); 12084, 12192 (52); 12278 (53); 16866, 17482, 17610, 17939, 26581 (54); 18105 (62); 20108, 20156, 20245 (55) Davidson 4824 (61) Davidson & Donahue 8481 (62) Davidson et al. 10692 (51) Davis s.n. (33) Davis & Andre-Lima 61068 (39) Dawson & Pujals 1702 (38) Dawson 424 (22a); 713 (38); 3355 (38); 14354 (52); 14442 (50) de Alba 53-106 (16a) de Bruijn 1473 (54) de Carvalho et al. 3783 (63) de Granville 9138 (38) de Granville et al. 6153 (60); 9385, 9396 (58) de la Cruz 2576, 3634, 4478 (54); 3772 (62); 4045 (60) de la Sagra 8 (61); 10 (33) de Paula 75 (14a); 1766 (60) de Wet & Harlan 4353 (43); 4354, 4732 (19a) Deam 6041 (62); 13899 (18) Deginani & Cialdella 311 (8) Del Puerto 18 (2a); 685 (22a); 1527 (41) Delanova s.n. (14a) Delascio 388, 3457 (62); 11206 (20); 12052 (54) Delascio & Rodríguez 19243 (51) Delascio et al. 8343 (62); 10053, 10511 (58); 10577 (62); 11585 (35) Delgado 133 (62) Deloche 166 (16a)

xviii

List of Exsicatta

Demaree 10788, 13592 (18) Denslow 2267 (62) Deramus 695 (18); 726 (7b) Desvaux s.n. (60) Díaz & Jaramillo 1270 (54) Díaz Luna 2044 (19b); 4438 (5); 4550 (62) Díaz Piedrahita 3292 (60) Díaz Piedrahita et al. 3272, 3292 (60) Dodge et al. 5610 (33) Dodson & Gentry 12506, 12514 (34a) Dodson et al. 7045, 7966 (62) Dombey s.n. (1a) Dombrowski 2368, 2455 (60); 11096 (41) Dorantes 246bis (34a) Dorantes et al. 1510 (19b); 2263 (62) Dore 46-137, 10544, 12887, s.n. (23) Dorsett 189b (52); Drouet 1953 (41) Duarte 1837 (2a); 6352 (50); 7972 (15) Ducke & Kuhlmann s.n. (15) Dudey 775 (62) Dueñas 2 (35) Dugand 2513 (61); 5395, 5877, 5983 (62) Duke & Bristan 303 (33) Duke 4470, 5107, 5181, 5585 (62); 14117 (33) Dunn et al. 21855 (43) Duque-Jaramillo 2024 (2a) Durán & Olmsted 1039 (18) D’Urville s.n. (17) Dusén 16, 3759, 7688, 17606 (60); 388a, 2602 (14b); 5100 (15); 7339, 13333, 15573, 16128 (11); 7709, 16579 (14a); 14574 (47); 16393, 10074 (50); Duss 12, 668, 3184 (31); Duss 536 (27); 674, 1321 (62); 676, 3181 (27); 768 (33); 3178 (1a); 3180 (2a); 3917 (33); s.n. (31) Dutily 30384 (18) Dutra 66 (11); 286 (2a); 290, 632 (11); 588 (49); 632 (16a) Dwyer 2781 (34a); 4292 (61) Dwyer et al. 454 (54) Earle 302 (32b) Ebinger 386 (62) Echeverry 1648 (61); 2304 (54) Eggers 76 (27); 461 (61); 960, 1174, 13006, 14538 (62); 4512 (2a); 13471 (63); 14949, 15419 (31) Eiten 1669 (50); 10154 (51) Eiten & Eiten 2577 (14b); 2676 (60); 3956, 4504 (29); 4474, 10812 (62); 4709 (58); 10579 (43)

List of Exsicatta

xix

Eiten & Mimura 3351 (54) Eiten et al. 1940 (2a); 2116 (41); 2577 (14a); 3157 (60); 5815 (14a) Ekman 502 (1a); 630 (16a); 636 (14a); 648 (60); 702, 12819, 13301, 16801, 17554 (47); 755, 3419, 4392, 5791, 6118, 6183, 8534, 10068, 13275, 16312, 17578 (33); 995, 1200, 1517, 2151, 7483, 7511, 11732, 13390, 16237, H20 (27); 1442, 1987, 2011, 9667, 12036, 12686, 16129 (31); 3060, 7233, 11211, 11212, 13101, 13191, 17878 (2a); 9991 (34a); 10028, 15694 (51); 11271, 17822 (3); 12400, 12602, 13013 (2b); 12608, 13085 (63); 12698, 15500, 17290, 17291 (7b); 14499, 14602, 15661 (63); 16310, 16939 (18), s.n., Amer Grass Herb. 701 (49) Elías 204 (14a); 560 (62) Elias de Paula 3008 (2a) Elliott s.n. (7a) Emygdio 333 (15) Enriquez 347 (33) Equipe IBGE 1467, 2919, 3046, 5946, 6188, 6503, 6937 (50); 1679- A, 2928, 3000, 3049, 4036, 4338 (14a); 3614 (14b); 4509, 5093, 6145 (60) Escobar 162 (62) Eskuche 2538- 5 (41) Esparza s.n. (5) Espejel et al. 331 (7b) Espina et al. 179 (54) Espinal 582 (61) Eugenio 271, 317 (2a); 278 (63) Evans s.n. (34a) Ezcurra s.n. (19b) Faberling & Wibom 326 (61), 2262 (62) Fabris 1080 (23) Falcao et al. 794 (62) Faria 2495 (63) Farias s.n. (2a) Farinas 176 (60) Fariñas 448, 613, 614 (51) Fariñas et al. 572 (54) Fassett 25374, 25534 (62); 25492 (31) Felger 86-339 (19b) Felippone 4904, 4908 (2a); s.n. (23) Fendler 928 (58); 929, 2499 (62); 1643 (61); 2565 (29) Fernald 306 (2a); 519 (23); 13878 (18) Fernald & Long 11255 (7a); 13877 (2a), 17828, 19754 (23) Fernandes 109 (60) Fernández 11, 13229, 13238 (51); 384 (33); 519 (16a); 524 (62); 846 (2a); 884 (11); 889 (14a); 4225 (60); 7697, 21211 (54) Fernández et al. 1485, 4366 (61) Fernández Casas 4366 (61); 6106 (14a) Fernández-Alonso 6844 (62)

xx

List of Exsicatta

Ferreyra 5929 (34a) Ferris 5605 (62) Ferris & Duncan 3175 (33) Ferris & Mexía 5069 (43); 5070 (19b) Feuillet 4442 (38) Fiebrig 178 (4); 436 (14b); 1302 (2a); 561 (60); 4642 (14a); 5095, 5102 (42); 5176 (61); 5373 (60) Figueroa 178 (4) Filgueiras 436, 817, 2509 (14a); 691, 818 (14b); 698 (60); 997 (50); 1060 (41); 1204 (38); 1261 (53); 2282 (29); 2479 (13); Filgueiras & Fonseca 2992 (53) Filgueiras & Rodrigues 1332 (29) Filgueiras & Soejarto 238 (50) Filgueiras & Zuloaga 2002 (13) Fischer 187 (17); 3313 (44) Fisher 35314 (43); s.n. (61) Fishlock 206 (31) Flavio s.n. (2a) Flores 49 (62) Flores & Tenorio 1180 (62); 1197 (40) Flores et al. 400 (62); 1053 (40) Fogg Jr. 7551, 10047 (7a) Foldats 1126 (31); 2220 (62) Folsom 10106 (62) Fonnegra 7937 (62) Fonseca 13 (50) Fontana F94-56 (41) Forero 369 (61); 407 (62) Forero et al. 2366 (61) Forsskal s.n. (46) Fortuna 2 (22a); s.n. (23) Fosberg 27768 (42); 29052 (61) Foster 9772 (62); 9779 (61) Fournier 283 (19b) Fowler 8d (34a) Fredholm 6365 (18) Freire de Carvalho 619 (60) Friedman 280 (11) Friedrichsthal 859 (62) Fries 81 (61); 882 (8) Froes 26084, 28359 (2a); 27045 (61); 28241 (62) Fróes & Black 17322 (51); 24665 (62) Fromm 1377 (2a) Funck 112 (14a) Gail s.n. (23)

List of Exsicatta

xxi

Galeano et al. 904 (54) Gallian & Leake 1067 (61) Gallinal et al. PE-1290, PE-2362 (22a); PE-4473 (25) Gándara 2 (34a) Garaventa 2847 (23) García & Dominguez 1805 (32b) García Barriga 3202, 10613 (61); 4945 (14a); 4984, 4991 (16a); 18723 (54) Gardner 1838 (39); 2457 (58); 3509 (62); 4053 (14a) Garnier 833 (34a) Garófalo 163 (35); 171 (58); 368 (16a); 381 (22a); 480 (52) Garófalo et al. 450 (57); 1230 (52) Garrido 4 (15) Gartner 1600 (60) Garwood et al. 624 (62) Gattinger s.n. (23) Gaudichaud s.n. (1a) Gaumer 895, 2027 (62); 1501 (34a); 2477 (31) Gehrt 31509 (38) Gemtohujnicov s.n. (41) Gemtchujnicov & Janssen 369 (20) Gentle 1601, 6971, 9647 (55); 1635 (1a); 3603 (7b); 3862, 6686, 6893, 6923, 8156, 8510 (54); 3883, 8099 (58); 6351, 9037 (62); 6863 (49); 6873 (18); 7120, 8380 (33); 9339 (60) Gentry 1841 (34a); 6667 (40); 8349 (32b); 8450, 10185 (62); 14353 (19a) Gentry & Berry 14600 (54) Gentry & Díaz 58422 (62) Gentry & Revilla 16127 (62) Gentry et al. 15966, 26929, 27491 (62); 29965 (54) Ghiesbreght s.n. (31) Giardelli 616 (38) Gilbert 530 (18) Gilberto 7 (11) Giler 1756 (62); 1773 (61) Gilly et al. 243 (33) Gilman 37 (34b) Ginzbarg et  al. 375, 2626 (14a); 2513, 2576, 3946 (54); 2603, 3645, 3649 (51); 3484 (2a); 3773 (43) Giussani 370 (17); 678 (15) Glassman 1746 (1a); 2145 (35) Glaziou 1628 (15); 4342, 20123 (2a); 13338 (22a); 17947 (14a); 22423 (16a); 22525 (53); 22931 (14b) Gleason 40 (54); 322 (62) Glocker 208 (39) Godfrey & Fox 51082 (7b)

xxii

List of Exsicatta

Goeldi 51, 235, 248 (1a); 98, 123, 223 (58); 111, 128 (33); 121, 122 (61); 174 (2a); 245 (51); 261 (43); 278 (62); 295 (60) Goes & Dionisio 122 (60); 590, 846 (1a) Goldfarb 618 (1) Gollmers s.n. (63) Gómez 458 (62); 19690 (54) Gomez & Praderio 474 (60) Gómez & Ramia 823, 824 (43) Gómez Kapla 23 (8) Gonçalves s.n. (29) Gonzales 414 (61) González s.n. (36) Goodding 192-45 (43); 4030 (32b) Goodding & Goodding 243 (23); 418 (34a) Goodland 358 (60); 556 (62); 605 (58) Gottsberger 37 (14a); 950-95B (52); 990-11271 (14b) Gouin 4 (47); 6 (33); 19 (61); 21 (31) Gould 1187, 10929, 7579, 7674, 7707, 11075a, 11157, 11282, 11302, 11796 (32b); 4468, 12032 (34a); 7685, 9763, 10292, 10956, 10972, 10973, 11271 (32a); 9558 (19b); 9643, 10796, 12322 (36); 11000, 11456 (24); 11596 (26); 11669, 11676, 12288, 12729, 12757 (40); 12071, 12110 (19a) Gould & Haskel 3192, 4510, 4550 (34a) Gould & Hudson 3748 (34a) Gould & Morrow 7906 (32b) Gould & Robinson 4937 (32b) Gould et al. 2409 (32b) Grayum & Hammel 5641 (61) Greene 258, 13107 (19b) Griffiths & Thornber 28 (34a); 238 (32b) Griffiths 399, 684 (23); 5004, 7808 (32b); 6939 ½, 7194 (34b); 7317 (19a) Grijalva & Aranda 3199 (62) Grinnell & Grinnell 1082a (19b) Grisol 36 (43) Guaglianone et  al. 270, 360, 595, 667 (22a); 335 (61), 359 (2a); 463 (11); 977, 1039 (60) Guánchez 711, 1389 (54); 4156 (16a) Guánchez & Mercado 2813 (52) Guardado s.n. (36) Guglieri 102 (15); 283 (9) Guinand 13 (52) Guízar 1850 (40) Gunther 5 (23) Gutiérrez 36 (62) Gutiérrez & Schultes 731 (54) Guzmán & Castro 1562(62)

List of Exsicatta

xxiii

Haase 525, 836 (2a) Haber & Zuchowski 9656 (62) Haenke s.n. (17); (34a); (57) Hage et al. 1716 (39) Hahn 481, 1047 (62); 1232 (27); 1911 (43) Hahn et al. 4240 (60) Hall 816, 817 (32b) Halley 69 (24) Hammel 4189, 4246 (60); 11794 (61); 11795 (33) Handro 77 (14a) Hans 266, 323 (2a) Hansen 18347 (33) Hansen et al. 7789 (62) Haridasam 091 (13) Harley 17285 (54); 21319 (50); 21497 (63) Harley et al. 15068, 15202, 15376, 15804, 16640 (60); 16203 (63); 16294, 16384 (43); 16356 (62); 16504, 17155, 17297 (1a) Harmon 3351 (6) Harmon & Dwyer 3394 (62) Harmon & Fuentes 2075 (55) Harriman & Jansen 15627 (31) Harrington 3361 (32b) Harris 40, 197 (35); 11170, 12235, 12845 (54); 11299, 11328, 11309 (62); 11691, 11261 (31); 12164 (27); 12226 (51) Harrison 1339 (58) Harrison & Pensant 967 (14a) Hart 1510 (54) Harvey 950, 1186 (32a); 1378, 1600, 1653 (34a); 1516 (32b); 1578 (26); 6610 (62); 6667 (40) Hassler 640, 640b, 8332, 9107, 11444, 11444a, 11462, 11589a, 11644, 12455, 12462 (60); 640a, 12450, 12477, 12579 (61); 1942, 10967, 11682, 12494, 13022 (42); 1957, 8308, 10118, 11988 (14a); 2396, 11443 (16a); 3652, 11451 (48); 8206, 12469, 12537 (2a); 10986 (29); 12099 (41); 12253, 12930 (16b); 9055a, 12462 (60) Hatch 4615 (24); 5060 (18); 5847 (44) Hatschbach 2648, 8444, 35527 (11); 3035, 7939, 14042, 46308 (14a); 4399, 13802, 42679, 46148, 48993 (60); 12994 (14b); 14269, 15839, 15842, 15980 (60); 21499 (61); 23525 (48); 23576, 46144 (42); 26163, 32527 (2a); 27230 (15); 42905 (1a); 43533 (41); 46653, 47654 (47) Hatschbach & Pedersen 35890 (60) Hauff 86 (60) Hayden 8220 (23) Helano 373-33 (14a) Heller 1809 (32a); 6094 (62); 13864 (23) Heringer 16119, 17760, 17558 (60)

xxiv

List of Exsicatta

Heringer et al. 12 (62) Hernández & Mathus 1667 (36); N-1911 (32b) Hernández & Tapia 240 (34b); 382 (5) Hernández 8, 77 (62); 74A (35) Hernández León s.n. (35) Hernández Xolocotzi et al. 2304, 2793 (26), 3379 (33); X-2468 (40); X-2775 (34a); X-2791 (19b) Herrera 504 (61) Herrera Castro 84 (34b) Herrmann 11196 (54) Hers & Méndez 4000 (43) Herter 432 (47); 544 (22a); 1376 (15); s.n. (22a) Hertzog 1445 (14a) Hess 1459 (31) Heyde & Lux 4299 (62) Hicken 22, s.n. (22a) Higgins 6990 (43) Hill 23758 (18) Hilton s.n. (54) Hinton 1062, 6337 (34a); 1424, 1428, 4413 (31); 1996, 6686 (40); 2010 (21) Hinton et al. 6423 (19a) Hitchcock 150 (2a); 220, 5143, 5318, 5385, 5395 (32a); 2798, 5197, 23470 (23); 3494, 3541, 3547, 3573, 3675, 8794, 13333 (19b); 3526 (34b); 3553 (19a); 3604, 3631, 6814, 7089, 7424, 8517 (34a); 3706, 5293, 5537, 5605, 13255, 13411, 13445, 13490, 13517, 13539, 13597, 16219, 5719 1/2 (32b); 5349, 5547 (24); 5712, 5822, 5864, 6057 (26); 5756, 5958, 6063, 7117, 7199, 7238, 7510, 7591, 9014 (36); 6143, 6363, 6391 6418, 7997, 8014, 8065, 8094, 8452, 8601, 8687, 8755, 8993, 16382, 23292, 23397 (31); 6364 (18); 6551 (47), 6782, 6808, 6810, 6811, 7081 (40); 7031, 7957, 7964, 8382, 8397, 8435, 20580, 23396 (33); 7315, 7485, 7491, 7757 (5); 8167 (6); 10053, 10154, 10154bis, 10159, 10233, 16795, 17144, 17268 (6); 8299, 22752 (60); 8372 (51); 9147, 10099, 23290 (1a); 9168, 9206 (57); 9463, 23239 (27); 9645 (2b); 9927 (35); 10083, 10181, 17191 (54); 10177, 10310, 10320, 10363, 21253, 21320 (58); 16598, 17431, 17547, s.n. (62); 22115 (29); s.n. (60) Hitchcock s.n., Amer. Grass Herb. 15 (5); 17 (2b); 29 (26); 32 (24); 33 (32a); 36 (31); 40 (47); 43 (7a); 44 (7b); 70 (62); 204 (2a); 209 (61); 503 (4); 504 (32b) Hno. Ginés 1476 (14a) Hoehne & Gert 17583 (50) Hoehne 9912 (1a); 23272 (14a); s. n. (60, 61) Hoffmannsegg s.n. (54) Hollermayer 229 (15); 1215 (17) Holm & Iltis 286 (34a) Holm-Nielsen 22896 (62) Holm-Nielsen et al. 2630 (62) Holt 561 (62)

List of Exsicatta

xxv

Holway 1571, 1594, 1615, 1717, 1824 ½ (60) Holway et al. 1116 (54) Hoock 119, 129 (60); 133 (61); 134 (62); 302 (58); 1243 (6); s.n. (51) Hopkins et al. 214 (57) Horner et al. 203 (58) Hosseus 43 (22a) Howard 8357 (33); 8590, 8728, 8827, 9076, 9757, 9965 (19b) Howard & Howard 10190 (2b) Huber 702, 1203, 1981, 3524, 4785, 4990, 5004 (54); 862, 961, 1351, 2132, 5268, 10446 (52); 1100 (16a); 3517, 1100 (16a) Huber & Alarcón 7509, 7655, 10470 (14a); 7587 (52); 7807 (52) Huber & Febres 6807 (14a) Huber & Manara 4253 (14a) Huft et al. 2285 (62) Humbert 27815 (54) Humboldt s.n. (61); (62) Humboldt & Bonpland s.n. (14a); (26) Hunt 5999 (60); 6075 (53) Hunt & Ferreira 5739 (52) Hunt & Ferreira Ramos 5606 (53) Hunter & Allen 63 (31); 727 (62) Hunziker 12 (23); 441 (2a); 891 (61); 2039, 9376, 16046 (17) Hunziker et al. 12539 (8) Hurrell et al. 6303 (11) Hutchings & Stachmann s.n. (32b) Hutchison 209 (17) Ibarrola 36, 52, 1717, 1886, 3085 (2a); 173, 1719 (61); 1590, 1870, 2071 (16a); 2889, 3481 (41) Idrobo 6308 (33) Iltis et al. 114 (32b); 946 (60) Infantes 5061 (62) Irwin 14904 (53) Irwin & Soderstrom 5166 (53); 5700 (50); 5711 (52); 6485 (51); 7494 (60) Irwin et al. 964, 10849, 11299, 11871, 13061, 14095, 19413, 23892, 25208, 28045, 28150, 34361 (60); 9213, 11077, 12574, 13268, 14509, 14657, 25582 (52); 9261, 9573, 13875 (2a); 9262, 13006, 21525 (14b); 10354, 11181 (50); 9653, 13699, 13700, 16122, 16386, 21525 (53); 4984, 17157 (62); 15961, 17302 (54); 16060, 17039 (57); 17992 (61); 19000, 23071, 25210, 28730, 28827, 29364, 34386, 34387 (50); 19032, 19740, 23982, 23982 (60); 21569, 25288, 25431, 34363, 34617, 34677 (14a); 24519 (29); 54530 (58) Itaipú Binacional 963 (60) Jack 6295 (33) Jahn 197 (22a) Jameson 560 (61) Janssen & Gemtchujnicov 373 (57)

xxvi

List of Exsicatta

Janzen 12425 (62) Jaramillo et al. 1047 (54) Jelski s.n. (51) Jenman 197, 6676 (60); 2261 (6); Jenman 4403, 7740, 7760 (62) Jiménez 7 (32b); 530, 730, 880 (31); 950 (63); 3620 (27); 4309 (33) Jobert 120 (1a) Joergensen 1583 (14a); 1720 (8); 2430 (16a); 4085 (60); 4785 (14b) Johnson 5 (18); 4080 (17) Johnston 324 (54); 1108 (57); 1614 (60); 4870 (32a); 7170, 8175, 8250, 8250a (32b); 7866 (34a); 7992 (19b); 9104 (36) Johnston & Muller 467, 694, 1017, 1111 (32b) Johnston et al. 10383 (32b); 12292 (34a) Johson 365 (18) Jones 827 (18); 24493 (19a); s.n. (19b, 34a) Jonsson 16a (60); 1124a (11) Jorgenson 12 (2a) Junge 3085 (15); 3104 (17) Jürgens 419 (14a) Kampf & Boldrini 839 (14b); Kappel s.n. (15, 25) Kappel- Froner s.n. (11, 47) Kappler 403 (62); 1495 (51) Kearney & Peebles 12868 (23) Kearney Jr. s. n. (23) Kellerman 7743 (60) Kellog s.n. (38) Kellogg 1217 (18) Kermes 403 (2a); 678 (16a) Kiesling 4259, 10224 (17) Kiesling et al. 5355 (62) Killeen 552, 604, 1892, 1971 (62); 720, 778 (16a); 797, 1486 (51); 823, 2834 (29); 856, 891, 1563, 1770, 1828, 1848, 2397 (42); 868, 1257, 1559, 1769, 1847, 2398 (14a); 966, 1675, 2320 (60); 1053, 1091, 1259, 1556, 2773 (61); 1094 (57); 1702, 1711 (43); 1732 (60) Killip 4100 (31); 4163, 34319 (54); 4183, 4237, 4256, 4282, 4309 (57); 4326 (34a); 12174 (33);35596 (62); Killip & Smith 14108, 14179 (62); 14859 (54); 15238 (14a) Killip & Tamayo 37046 (22a) King 391 (19b); 771 (43); 3791 (31); 4346 (55) King & Soderstrom 4679 (19b); 4831 (40); 4927 (62) Kirkbride Jr. 2224 (60) Kirkbride Jr. & Bristan 1582 (62) Klein & Bresolin 11362 (2a) Klinger 16860 (60) Kneucker 730 (43)

List of Exsicatta

xxvii

Koch 4899 (62); 7461 (47) Kornelius s.n. (15) Korstanje 3 (8) Krapovickas 1251, 2906, 6681 (22a); 2480 (61); 3693 (17); 23280 (14a) Krapovickas & Cristóbal 11895, 30922 (61); 21720 (49) Krapovickas & Schinini 30990 (62); 31194 (61); 35068 (54) Krapovickas et  al. 14068 (14b); 15705, 27527 (2a); 17861 (41); 19192, 19389, 24552, 24955 (61); 22405 (17); 24632, 24669 (48); 25132 (60); 25201 (22b) Kuhlmann 72 (61); 1195, 1754 (62); 1667, 1755, 2689, 2690 (60); 1701 (51); 1743 (14a); 1745 (53); 1755, 1947, 1949 (60); 1786 (57); 2414 (2a); 2530 (29); 4110 (54); s.n. (15); s.n. (42) Kuntze s.n. (16a); (17); (61) Labat 1186 (26) Laegaard 52215 (62) Laferriere 2725 (34a) Lanfranchi 186 (23); 453, 481 (38); 1818 (15) Langenheim 3180 (61) Langlassé 263 (43) Langlois 49 (47) Langsdorff s.n. (2a), (14a), (60), (60) Lanjouw 42 (61) Lanjouw & Lindeman 1955 (60) Larez 605 (35) Laser & Vareschi 4260 (62) Lasseigne 2812 (38); 2844, 6260 (18) Lasser & Vareschi 4260 (58) Lathrop 5558 (62) Laughlin 1113 (19b) Lawesson 3143 (19b) Le Doux et al. 106, 2125 (51) Ledru s.n. (1a) Le Sueur 032 (19b); 173 (40) Leal & Da Silva 6 (62) Leavenworth 199 (33); 442a (43); 481 (45); 966 (34a) Leavenworth & Hoogstraal 1371 (19a); 1335 (34a); 1453 (43) Leavenworth & Laevenworth 742 (32a); 1335, 1875 (19b); 1839 (26) Leblond s.n. (58); (60) Ledru s.n. (31); (62) Legaspi s.n. (23) Legrand 980 (2a); 1455 (22a) Lehmann 5268, 5269 (51); 8745 (31) Leino 76105 (33) Leitao Filho et al. 10 (60) Lemes 4125 (52) Lenaz s.n. (29)

xxviii

List of Exsicatta

Lenz Cesar 185, 385, 449, 611 (13) León 190, 923, 2532, 2559, 3449, 9651 (27); 296, 563 (47); 751, 805, 2775, 9507 (2a); 1184 (54); 2243 (61); 2674, 3747, 6429, 8834, 9138, 9314, 15339 (31); 3913, 14181 (33); 9470 (1a); 10190 (63); 12607 (60); 16027, 20614 (7b) León & Cazañas 5755 (18) León & Garteiz 5681 (27) León & Loustalot 9507 1/2 (2a); 9521, 9575 (18) Leonard 3493, 4233 (2a); 4901, 7745, 8344, 9417, 9607 (31); 7449 (27); 9207 (62) Leonard & Leonard 11576, 11875, 11900, 12237, 12590, 13609 13828, 13147, 15090 (27); 11720, 12556 (62); 13744, 14338, 15754 (31) Leonard & Radford 2170 (7a) Leopold 109 (33) Leprieur 41, 42 (2a); s.n. (51); s.n. (58); s.n. (62) Levy 37, 1018 (33), 1166 (35) Lewis 526 (61); 629 (22a); 963 (16a); 35344 (42) Lewis & Pire 565 (22a) Lewis et al. 2601, 3416 (62) L’Herminier s.n. (33) Liesner 3703 (54); 17268 (58) Liesner & González 5479 (60); 5482 (14a); 13298 (54) Liewere 137 (18) Lima s.n. (14a) Lindheimer 733 (18); 1266 (32a) Lindig 102 (31); 1032 (62); 1057 (58); 1067 (54); 1074 (61) Lindman A.705 (15); A1209 (47); 1185, A-1187 (41); 1477 (60) Liogier 17408 (51); 20856 (62); 21371 (38) Lix 738 (23) Lofgren 5 (15); 212 (50); 230 (14a); 295 (60); 492 (62) Longhi Wagner et al. 9646 (9) López 2338 (54) Lopez Figueiras & Acuña 341 (31) López-Forment 1417 (62) Lorentz 445 (8); s.n. (16a) Lot 1227 (1a) Lot et al. 1948 (31) Loustalot 9507 ½ (2b) Lozano et al. 5870 (62) Luces 17 (31); 64, 96 (51); 152 (63); 188 (33) Luederwaldt s.n. (2a) Luetzelburg 26703A (60); 6002 (56) Lundel 6557, 6734 (60); 6903 (6) Lundell & Lundell 7427, 7791 (31); 7622 (33); 7657 (55); 8719, 14683 (24); 10657 (32a) Lundell 989 (62); 1181 (55); 1355, 15918 (33); 3868, 3876, 15668, 17770, 17920 (55); 6585 (51); 6938, 18662 (18); 17857 (61)

List of Exsicatta

xxix

Lurvey 313 (2a) Lyonnet 2639 (40) Lyons Jr. 59, 83 (19b) Maas et al. 7199 (58) Macedo 2102 (14a); 2169 (62); 2192, 2194 (29); 4240 (60); 4482, 4876 (50); 4527 (2a); 4561 (16a) Machado de Campos 209 (41) Macías 6 (62) Mackenzie 4452 (7a) Macnab 184 (27) Macuszek 271 (23) Magalhaes 3 (60) Magana & Zamudio 746 (54) Magaña 427 (33) Maguire 23912, 29424, 29060, 34667 (54); 30517 (10) Maguire et al. 31601 (6); 46249, 53979, 54208 (58); 54053 (2a); 54237 (60) Mahecha 4122 (62) Maia 18 (50) Mally s.n. (24) Malme 1460 (1a); 3044 (29) Manara s.n. (60) Mann 60 (32b) Marchesi 4383 (60); s.n. (11) Marie-Victorin 8152 (18) Marsh 1278 (32b) Mársico 1217 (62) Martin s.n. (54); (58); (60) Martínez & Luyando F-1709, F-2275 (32b); F-2161 (44) Martínez 1313 (19a); 2265 (62); 6077 (22a); 11359 (55); 14651 (61) Martínez Calderón 221, 1565, 1786 (61); 568 (33); 1676 (51); 1798 (33) Martínez Crovetto 1994 (2a); 2532 (47); 3519, 3524 (60); 8178 (14a); 8978 (41); s.n. (47) Martínez et al. 2413 (62) Martínez Martínez & Borja Luyando F-2303 (32b); F-2333, F-2364 (44) Martius 3800 (53); s.n. (1a); (2a); (16a); (41) Martorell & Medina 429 (62) Marulanda 1257 (14a) Matthei 275, 302 (17) Mattos 3587, 14535 (60); 10000 (60); 12255 (52) Matuda 749 (18); 1799 (51); 3087 (54); 3577 (33); 3722 (18); 19747 (5); s.n. (31) McBride 332 (23); 3526 (34a) McCarthy s.n. (23) McDaniel 6807 (51); 11772 (2a); 12933 (62) McDaniel & Rimachi 20250 (61); 23032 (62) McDaniel & Tyson 14702, 14709 (57)

xxx

List of Exsicatta

McGregor et al. 840 (19a); 870 (44) McKenzie 183 (7b) McVaugh 16326, 19117, 19333 (34a); 16819, 17705, 19651, 19578 (36): 19291 (19a); 20345 (21); 21107 (40) Mearns 738 (19b); 2294 (34a) Medina 763, 953, 1246 (33); 847 (62); 1243 (61) Medrano et al. F-1495 (34a) Méndez 6220 (33) Mendonça et al. 4507 (11) Mendoza et al. 2720 (62) Mengies 8023 (38) Mercado B-499 (2a) Mereles 715 (43) Mertens s.n. (54) Metcalfe 1138 (23); 1442 (34a); 5511 (38) Mexía 747 (62); 2740 (36); 5543a (42); 5567 (50); 5622 (29); 5921, 5975 (54); 6190 (61) Meyer 80, 380, 2554 (2a); 110 (18); 321 (61); 6329 (41); 11017 (16a) Michaux s.n. (2a) Mick & Roe 44 (32b) Miers 3429 (22a) Milano 5100 (47) Mileski 106 (14a); 221 (16a); 353 (2a) Miller 12001 (7a) Millspaugh 126 (2a) Mimura 96 (14b) Miranda 128 (53); 130 (58); 2496 (2a); 3238 (47) Mizoguchi 572 (60) Mohr s.n. (22a); (49) Molas & Vera 1382 (61) Moldenke 6377, 1372a (23) Molina & Barkley 53 (58) Molina 242, 3905 (31); 2219, 15281, 15426 (33); 2591 (60); 3591 (62); 3831 (61); 15533 (51); 27395 (54) Molina & Molina 25789 (60) Mondolfi s.n. (16a) Montaldo & Ramia 3304 (35) Monteiro s.n. (1a) Montero 1953 (17); 2102 (15) Montes 186, 1933, 11057, 15390, 14889, 15319 (60); 329 (2a); 434, 752, 1788 (60); 1885, 10938, 27649 (14a); 2018 (35), 2580 (58); 2584 (62); 11238 (41); 14642 (22a); 15373, 15374, 15381 (61) Monticelli 168 (38) Montilla 26 (33) Moore Jr. 2885 (55); 3924 (32b); 4218 (36)

List of Exsicatta

xxxi

Moraes 590 (51) Moran 13664, 18878 ½ (19b); 16483, 29255 (23) Morel 764, 3502, 5005 (61) Morello 841, 1234 (8) Moreno & Lopez 10 (62) Moreno 830, 3861, 10741, 11157, 18081, 18117, 18380 (62); 9838 (19b); 21790, 21833 (34a) Moreno Casasola et al. 175 (47) Mori & Anderson 243 (35) Mori et al. 9674 (54) Morillo 2533, 7789, 10563 (62) Morillo & Manara 1427, 1432 (60); 1729 (14a) Morley 696 (35) Morong 317, 1571 (61); 538 (16a); 543, 1002 (2a) Morrone 5918, 5919, 5921, 5926 (15); 5930 (17) Morrone & Pensiero 529 (52) Morrone et al. 4698 (35); 4688 (51); 4801 (62) Morton & Acuña 3507 (62) Mosén 4469 (42); 4571 (29) Muelhenbach 667 (38) Mueller 404 (43) Mulford 1078 (34a) Muller 2177 (47) Muller 804, 987, s.n. (35); 814 (14a); 815 (62); 2177 (47); s.n. (51) Müller 2002 (18) Muñoz & Rollins 1006 (23) Murguia 85 (62) Mutis 2151 (33); 5339, 5378, 5498, 6110 (51) Myers 450 (7b); 4189 (33) Nascimento et al. 194 (54) Nash & Taylor 1297 (31) Navarro s.n. (38) Nealley 634 (24) Nee 183, 17751, 6783 (33); 8485 (57); 23649, 34549 (62); 22491 (60); 23829, 39458 (61); 34647 (16a); 37618 (22a); 43735 (18) Nee & Mori 3783 (54) Nee & Taylor 29458 (60) Nee & Téllez 28161 (62) Neill 1115 (35); 8550 (18) Nelson 995 (18); 4819 (6); 5467 (62); 6355 (34b); s.n. (33) Nelson & Cruz 9452 (61) Nelson & Romero 4470 (62) Nelson & Vargas 4987, 5021 (54); 5077, 5110 (57) Nelson et al. 3186 (62); 5621 (35) Nicora 2120, 6512 (2a); 2724 (22a); 6094 (16a); 6511, 6806 (23); s.n. (15); s.n. (43)

xxxii

List of Exsicatta

Nicora & Cámara Hernández 219 (2a) Nicora et al. 8361 (38); 9166 (8); 9737 (43) Nienstedt 207 (16a) Nino 99 (14a) Noblick 3728 (43) Noblick & Lemos 3955 (39) Novaes 1269 (50) Novara 8436 (38); 9744 (8) Novelo & Ramos 3823 (5) Nozawa et al. 190 (52); 280 (54) Nuñez & Munos 5298 (2a) Nuñez & Ortíz 12809 (61) Núñez & Silva 3196 (62) Occhioni s.n. (50) O’Donell 2629 (43) Ojasti 67, 83 (2a) Oliva 49 (18) Oliva & Ramírez 882 (62) Oliveira2929 (2a) Opler 1578, 4515 (62); 4197 (34a); 4687 (40) Orihuela 92, s.n. (25) Ornelas et al 1167 (34a) Oropeza s.n. (33) Orozco et al. 767 (54) Ortega 5760 (2a) Orth 3, 850 (16a); 2717 (60); s.n. (47) Ortiz & Ramia 2127, 2135, 2767, 3559, 4212, 4219, 3083 (20); 3532 (58) Ortíz 188, 228, 1093 (62); 900 (7b); 1049 (33); 1693 (55) Osten 4512 (47) Otavio s.n. (60) Otero 48 (43) Pabón et al. 224 (54) Paget 152 (23) Palacios 1856 (43) Palacios & Bravo 642 (8) Palacios & Cuezzo 4368 (22a) Palmer 1c, 143, 947 (34a); 168a, 206, 270, 1538 (43); 208, 249, 251, 346 (19a); 510 (18); 525, 533 (36); 554, 1388, 13277, 14147 (32b); 750, 1554 (19b); 1657 (40); 14214 (32a) Palmer & Riley 802 (27) Parish & Parish 887 (17) Parodi 61, 3939 (11); 590, 1750 (23); 760, 5736, 5808, 8269, 8324, 8340, 9347, 10593, 11096 (2a); 4490 (22b); 4512, 4513, 6935 (41); 4847 (17); 5150, 7059, 8307 (61); 8218 (23); 9505 (15); 12209 (1a); 12363 (22a); 12890 (16a) Parodi & Horowits 44 (2a)

List of Exsicatta

xxxiii

Pase 988, 1048 (32b); 1084 (34a) Pastore 849 (38) Patoni s.n. (18) Pavetti & Rojas 9714 (60) Paz Zambrano & Ramos Marchena 303 (32b) Peck 123 (6) Pedersen 1030, 4313, 5456, s.n. (14a); 1418 (47); 1430, 1710 (61); 1882, 6112, 10286 (11); 2591, 4818, 11679 (60); 4792, 4805, 11693 (2a); 3046 (16a); 5428, 5457, 5482, 5812, 11109 (41); 5868, 13473 (48); 5904, 13675 (60); 15316 (8) Pedra do Cavalo 475 (63) Peebles10366 (34b) Peebles et al. 7417 (19b) Penn s.n. (32b) Pennell 1445, 1832 (14a); 1598 (60); 3358, 3702 (62); 3509 (54); 3849, 3866 (33); 3918 (61) 20033 (21; 20221 (43) Pennell & Killip 7253 (60) Pereira 157 (60); 5201 (14a) Pereira & Mendonca 401 (14b) Pérez 8 (22a) Peréz Arbelaéz 598 (61) Peterson & Annable 6453, 6600, 6885 (62); 6729 (33); 6943 (61) Pfister 60 (23) Philcox et al. 3631 (51) Philipson et al. 1329 (54) Piccinini 1983 (17) Pickel 1393 (39); 1403 (22a); 1663, 2469, 3848, s.n. (63); 2275, 2280, 4566, 4640 (60); 3137 (54); 3799 (29); 5154, 5795 (41); 5887 (52); s.n. (60) Piergentili 3578 (17) Pineda 16 (62) Pinheiro 1855, 7855 (63) Pinheiro & Carvalho 36, 55, 74, 182, 218, 265, 273, 297, 336, 362, 382, 385, 483, 530, 561, 512, 581, 587, 596, 603, 634 (29) Pinto 121, 306 (39); 132/80 (60); 307 (63); 345, 933 (62); 1021 (51); 1548 (57) Pinto & Bischler 345 (62) Pinto & Sastre 933 (14a); 1063 (14a) Piper 5260, 52656 (57); s.n. (24); s.n. (32a) Pire 1410 (16a) Pires 818 (61); 1828 (2a); 1936 (57); s.n. (60) Pires & Black 355, 409 (60); 850, 1204 (2a); 1698a, 1710a (57); 1951, 1991 (58) Pires & Santos 16272 (2a); 16681 (16a) Pires et al. 6302, 9176 (52); 50973 (54) Pittier 2169, 2636, 3636, 3640, 3661, 4626, 5247, 12011 (57); 2633 (31); 3441, 3634, 6332, 6943, 10534, 10620 (33); 4526 (4); 7224, 8004, 12789, 14433 (14a); 8214 (62); 9054, 13734 (60); 10588, 10594, 12064 (54); 10830, 12511 (35); 12231 (58); 12581 (63); 12617, 12763, 13069 (61); 14479 (51)

xxxiv

List of Exsicatta

Plank 38 (23); 57 (32b) Plowman et al. 6550, 9658 (62); 8226 (42); 9269 (29); 9697 (60); 12574 (2a) Pohl 2435, 3885 (29); 12651 (1a); 12674 (34a); 12707 (40); 12774 (4); 12914 (61); 12924 (62); 14182 (4); 15441 (60) Pohl & Calderón 10049 (60); 10216 (34a) Pohl & Davidse 10556, 12210 (31); 10653, 11872, 12177, 12228, 12369 (34a); 10761, 10979 (51); 10831, 11073, 12257, 12460 (60); 10941 (4); 10944, 11291, 14172 (35); 11013, 11305 (62); 11330, 11537 (57); 11437 (40); 11727, 12078 (61); 12386, 11547 (1a); 11700 (1b); 12458 (63) Pohl & Gabel 13322, 13382 (62); 13521 (60); 13609, 13665 (34a); 13689, 13728 (57); 13703 (35); 13871 (51) Pohl & Lucas 13042 (62) Pohl et al. 13116 (54) Ponce & Rodríguez 16 (35) Popenoe 34 (33) Porta 103 (2a) Porter s.n. (23) Pott et al. 444, 3862, 3915 (29) Prado 260 (61) Prance 3386, 13791, 4035, 4274, 9899 (54); 3517, 8177, 8600, 8831 (51); 24993 (52) Pringle 197, 464, 13940 (19b); 376 (32b), 497 (36); 1415 (5); 1688 (5); 3828 (62); 5569 (47); 5573 (33); s.n. (43) Proctor 2571 (43); 26301 (54) Pruski 1560 (54) Purpus 6212 (34a); 9199 (18) Pursell et al. 8096 (62) Quarín 110, 1639 (22a); 1158 (60); 1186 (61); 1321 (38); 1657 (49); 2866, 2909 (41); 3188 (11); 3324 (47) Quarín & Schinini 1098 (2a); 1215 (61) Quarín & Tressens 1337 (61) Quarín et al. 1766 (48); 1862 (22a); 1869, 2156 (61); 1969 (41); 3373 (60); 3877 (11) Quero et al. 2855 (7b) Questel 1439 (31) Quevedo 93 (62) Quezada & López 11 (15) Raddi s.n. (54) Ragonese 48 (15); 2751 (11); s.n. (23) Ragonese & Covas s.n. (62) Ramalho 37 (62) Rambo 1937 (38); 28796, 40524, 42068, 45722 (60); 9845 (22b); 9875 (25); 40614, 40742, 41130, 41798 (2a); 41286, 44477 (14a); 43391, 43692 (41) Ramia 1121, 1184, 1423, 1359, 1789, 3622, 3596, 4006, 5654, 5580, 5581 (16a); 1209, 1724, 2442, 3604, 3620, 3864, 3874, 4770, 7195 (2a); 1260, 1967, 2559, 3637 (4); 1757, 1807, 2167 (51); 3330 (33); 3884 (14a); 4444, 4452 (43); 6481, 7201 (35); 7195 (1a); 8778 (58)

List of Exsicatta

xxxv

Ramia & Doreste 599 (33) Ramia & Marchena 5345, 8491 (43) Ramia & Montes 4730, 4830 (51); 4826 (2a); 4831 (35); 5345 (43) Ramírez 51 (43); 211 (14a); 2868, 2869, 3341, 3344 (35) Ramírez & Cárdenas López 1733 (62) Ramírez et al. 418 (51) Ramírez Reyes 1399 (60); 1715, 1773 (35); 2402 (58); 2048, 2428, 2752 (31); 2051, 2054 (33); 2066, 2460 (62); 2184, 2485, 2488 (34a); 2405 (51) Ramírez Reyes & López 4210 (63) Ramos 190 (62) Ramos & Paz 914 (61) Rangel 2071 (16a) Ratter 2069 (54) Ratter & Fonseca 3295 (62) Ratter & Ramos 240 (61) Ratter et al. 2090 (53) Redfield 10941 (18) Redmond 1002 (7a) Reed 517 (31) Reeder & Reeder 609, 2082, 2430 (19b); 1439, 2975, 7042 (26); 1936 (31); 1962 (33); 1976 (62); 1979 (60); 1996 (18); 2152, 2164, 2288 (40); 2300, 3814 (36); 2579, 3526, 4849 (5); 2967, 3928 (32b); 3517 (34a); 6111 (51); 6718, 7045 (19a) Regnell III-1359 (60); III 1366 (29); III 1369 (52); III 1406 (42) Reis 32 (60) Reitz & Klein 287 (1a); 8492 (1a); 17008 (38) Reitz 1307, 1987 (15); 2015 (47); 5927 (60) Renvoize 3640 (22a); 4687 (62) Renvoize & Cope 3897 (60); 3898, 3890 (61); 3912 (43); 3932, 4002 (62); 4053 (14a); 4253 (60) Renvoize et  al. 3290 (2a); 3583 (22a); 3637, 3933, 4044 (41); 3639 (14a); 3663 (16a); 3954 (51) Reverchon 2230 (32a); 3527 (32b) Revilla 450 (2a); 1234 (62) Reyes 17 (55); 34 (33) Richard s.n. (2a), (27) Ricketson 677 (38) Ridoult 1919 (34a) Riedel 2155 (41); s.n. (1a), (14a); s.n. (16a); s.n. (2a); s.n. (42); (60) Rimachi 4657 (2a); 6572 (62) Robbin 860 (23) Robleto 980, 1269 (34a) Rodrigues & Coêlho 4749 (58) Rodriguez 99 (22a); 433, 839 (15); 541, 1234, 1981 (35); 694, 3056, 3220, 27191 (31); 853 (60); 878 (63); 1011, 1341, 3443, 3492 (4); 1392 (60); 3126 (33); 3512 (38)

xxxvi

List of Exsicatta

Rodway s.n. (33) Rogers 4491, 4756, 6119 (32b); 6609 (24) Rohweder 1327 (62); 1497 (34a); 1648 (19b); 1996 (60); 2486 (4); 2507 (35) Roig s.n. (19b) Rojas 90 (61); 105, 2358, 2762 (22a); 237, 237b, 237c, 13191 (2a); 535 (16b); 2745, 6654, 12741, 12819 (42); 2784 (41); 5476a, 11027, 11047 (43); 6434, 6750b (14b); 10491 (16a) Romanczuk et al. 464 (2a); 507 (60) Romero 190 (33) Romero Castañeda 8948 (33) Rondón 2566 (52); 2585 (42) Rosa 3697 (49) Rosa Mato 1913 (2a) Rosales & Dezzeo 867 (52) Rosales & Valles 143 (51) Rose 1878, 1883, 1889 (43); 18052 (32a) Rose et al. 4410 (27); 9555 (36) Rosengurtt B-242, B-7128 (25); B-480, B-681B, B-2228, 5411, B-5623 (22a); B-942, B-3313, 11327 (16a); B-3802, B-5242, B-7373 (2a); B- 5772, B-5794, B-5897 (60); B-5607, B-5783 (14a); B-6281 (15); 10804 (25) Rosengurtt & Del Puerto 8893 (25) Rotman et al. 100 (22a) Roybal 867 (32b) Rudd 293 (14a) Rúgolo 675 (23); 721 (17); 2057 (15) Ruiz 6 (23); 2543 (63); 3081 (61) Ruíz Leal 1551 (31); 5576 (38) Rusby 233 (60) Rusell 186 (50) Russi 57 (62) Rzedowski 565, 796, 3775, 32145, 49933 (36); 2047, 17367, 17423, 17449 (21); 4618 (24); 4931, 6547, 6869, 5174, 50000 (32b); 38973, 39002, 49884 (26) Rzedowski & McVaugh 836 (5) Sagástegui 10912 (34a) Sagot 1338 (58) Salinas 6163 (62) Salzmann s.n. (1a); (54); (56) Sampaio 2971 (22a); 5333 (2a); 5570 (14a); 5995 (60); 6249 (50) Samuels 233 (54); 455 (62) Sánchez et al. 34 (54)6 Sánchez Vega 2338, 2363 (34a) Sánchez-Ken & Nieto M. 430 (19b) Sánchez-Ken et al. 328 (32b) Sanders et al. 9297 (34a); 9687 (61) Sandino 3038 (34a); 3831 (62)

List of Exsicatta

xxxvii

Santana & Cervantes 752 (62) Santana Michel et al. 1782 (5); 1850 (18) Santos Rubio s.n. (49) Sanzin 900 (17) Saravia 468, 2360 (62); 2664 (14a); 2685 (54) Saravia Toledo 1243, 1301, 1305 (43) Saravia Toledo & Nelson Joaquin 10412 (43) Sastre 830 (16a); 1639 (58); 2132 (54) Sauer & Gade 3264 (7b) Saver et al. 3350 (7b) Schaffer 2735, 2780 (7b) Schaffner 301 (2a) Scharwz 2285 (60) Schellenger 3 (19a) Schiede 895 (47); s.n. (60) Schikendantz 107 (8) Schimpff 532 (61) Schinini 2474, 4285, 4393, 6195, 8445 (60); 6580 (2a); 8598, 8694, 8715, 22718, 24028 (61); 9356, 23973 (60); 13782, 16095 (16a); 16081 (14a); 23924 (48) Schinini & Bordas 15188 (61); 16368 (43); 25204 (2a) Schinini & Cristóbal 9775 (14a); 9776 (14b) Schinini & González 9293 (22a) Schinini & Martinez Crovetto 12911 (61) Schinini & Miranda 9578 (2a) Schinini & Quarín 8524 (2a) Schinini & Vanni 25990, 26047 (60) Schinini et al. 8190 (41); 11044, 16978, 17098 (60); 11090, 16857 (16a) Schlegel 2394 (15) Schnell 11424 (2a) Schnetter 193 (62) Schomburgk 259, 456, 653, 656,701 (22a) Schott 16 (19a) Schreiter 4941, 10252 (8) Schroeder s.n. (16a) Schultes 8528 (62) Schultes & Cabrera 17192 (54) Schultes & Reko 565 (55) Schultes et al. 17936 (54) Schulz 1799, 3424 (16a); 3261, 10903, 15899, 17777 (2a); 12138, 12349 (33) Schunke Vigo 6991 (62); 7096 (61) Schwacke 62 (50); 507 (60) Schwarz 1376, 6685, 8995 (14a); 1979 (60); 6205, 6840, 9452, 10204 (2a); 6484, 6499, 9492 (41); 9144, 9192 (48); 9595 (25) Schwindt 1154 (42) Scoffield 60 (17)

xxxviii

List of Exsicatta

Scolnik et al. 19jAn327 (62) Segadas Vianna 60 (15) Seidel & Beck 355 (61) Seidel & Vargas 2695 (62) Seijo 2661 (15); Seiler 3825 (38) Sellow 130 (11); 960 (16a); s.n. (15); (41); (50); (60) Sendulsky 11 (14a); 18, 37, 148, 169 (52); 126, 208, 311, 637, 716, 725 (50); 154, 298, 575, 727, 1101 (60); 162 (14b); 236 (2a); 565, 661, 668, 1477, 1869 (60); 1078, 1299 (49); 1434A (39) Senn 4315, 4374 (17) Sessé s.n. (61) Seymour 2829, 6275 (34a); 2895 (1a); 4665 (57); 4977 (54); 17670, 22235, 22920 (18) Shafer 385, 1437, 1512 (27); 561 (60); 1361 (31); 10510 (51); 10927 (18); s.n. (18) Shafer & León 13682A (60) Shear 436 (23 Sherff s.n. (34a) Shreve 9821 (32b) Shreve & Tinkham 9897 (32b) Sieber 28 (62) Silva 173 (62); 287 (14a); 365 (61); 608 (60); 1239 (2a); 2782 (54); 4876 (16a); s.n. (25) Silva & Pinheiro 4101 (52) Silva et al. 223 (60); 371 (14a); 373 (14a) Silverstone 6512 (61) Silveus 625 (33); 3461 (43); 7293, 7302 (31) Simpson 70 (58) Simpson & Schunke V. 829 (43) sin colector 39 (14b); 229 (62); sin colector s.n. (17), (20), (31), (62) Sintenis 160 (62); 4983 (31) Skorepa 116 (19a) Skutch 3065 (54); 3977 (62); 4053 (57) Skvortzov 283 (60); 304 (60) Smart 112 (33) Smith 164 (33); 165 (31); 166 (6); 167 (62); 585, 632 (55); 913 (47); 2146, 168 (60); 2151 (61); 2152 (35); 2261, 2398 (58); 8042, 9169 (63); s.n. (32b) Smith V5562, V5563 (58); V6410 (14a) Smith & Klein 10774, 11467, 11832, 14951, 16093 (60); 11453, 11986 (14a); 15616 (14b) Smith & Reitz 6038 (60); 9661 (60) Smith et al. 14724 (14a); 14726 (41) Snethlage 150 (62) Snodgrass & Heller 454, 492, 548, 565, 612, 648, 722 (19b)

List of Exsicatta

xxxix

Soderstrom 108, 1125, 1405 (60); 160 (34b); 445 (26); 531, 819 (32b); 647, 896 (19b); 812 (36); 1010 (54); s.n. (50) Soderstrom et al. 1045 (33) Sohns 210 (26); 253, 285, 288, 504, 1066, 1114 (36); 399 (5); 791, 797 (31); 834, 836, 905, 922, 945, 960 (40); 1094, 1215, 1248, 1344 (32b); 1430 (24); 1431 (44); 1448 (62); 1566 (18); 1602, 1651, 1714 (55) Solomon 2741 (32b); 3305, 9881A (62); 7919, 8930, 9881, 14188 (61); 14724 (16a); 14771 (2a) Solomon & Urcullo 14188 (61) Sonia de Campos 3 (14a); 191 (14b) Soreng & Soreng 7006 (17) Soto 53 (60) Sousa 11207 (1a) ; 11219 (7b) Spegazzini s.n. (23); s.n. (38); s.n. (62) Spruce 11, 12, 677 (60); 13 (60); 466 (62); 788 (2a); 1431 P. 29 (2a); 2687 (58) St. Hilaire 2235 (15) Standley 3983, 6441 (32b); 5289 (18); 12634, 22020, 22483, 23356 (60); 12793, 14492, 24654, 29252 (35); 14567, 20874, 21272, 21640, 27498, 29263, 29327, 59580, 80411, 80463, 80630 (63); 19527, 23187, 24175, 24577 (62); 23298, 26154 (61); 25277, 27407, 27960, 74383, s.n. (34a); 28598 (54); 53428, 53692, 55711 (33); 54701 (51); 72464 (60); 74761, 75102a, 75925 (40); 76308 (1a) Stanford et al. 2338 (32b) Starry 102 (62) Steere 1632 (31) Stehle s.n. (62) Steibel 2451 (22a); 3473 (23); 4971 (2a) Stein et al. 1466 (62) Steinbach 242, 1213 (61); 784 (62); 1906, 3180 (2a); 2204, 5299, 5639, 6979, 7076bis (42); 3220, 6900, 6901 (16a); 5378, 6978 (14a); 6935 (51); 7086bis (29); 9095 (60) Steller & Jacobs 335 (23) Stergios 6935 (22a) Stergios & Aymard 7342 (54) Stergios et al. 9809, 19391 (54) Stern et al. 497, 1894 (61) Stevens 2672 (34a); 4346, 11138, 19948 (62); 7982 (33); 8561, 12849 (57); 10006, 20198, 21647 (31); 23096 (40) Stevens & Grijalva 15518, 16351 (62) Stevens & Martínez 25218 (60); 25813 (33); 25887 (61) Stewart 1211, 2395 (32b); 1293 (19b) Steyermark 29053, 29310, 30064 (40); 29326, 47766, 47859 (34a); 33708, 37138, 51834, 86911 (60); 39673, 59429 (54); 39802, 45998, 49430 (33); 44244, 57679, 61825, 90562, 97479, 99139 (60); 45889 (55); 51627 (30); 59173 (52); 59240, 88378 (14a); 72470 (23); 80666 (18); 90508, 90559 (58) Steyermark & Braun 94643 (62)

xl

List of Exsicatta

Steyermark & Bunting 103118 (10) Steyermark & Carreño 106943, 112723 (62) Steyermark & Davidse 116427 (62) Steyermark & Fernández 99665 (62) Steyermark & Liesner 120970 (62) Steyermark & Nilsson 810 (62) Steyermark & Wurdack 22 (54) Steyermark et  al. 100143, 127074 (62); 107780 (14a); 114619A (2a); 114879, 114898 (6); 117119 (51); 119536 (60); 131327 (35); 131495 (16a) Stork & Horton 11361 (34a) Stoupy s.n. (51) Strang 1348 (2a) Strudwick et al. 4389 (58) Stuckert 14434, 14798, 14799 (22a); 15213, 15336, 15337 (17); 15511 (22a) Suárez 4 (31); 24 (62) Suazer 1361 (31) Sucre 142 (62); 2192, 8578 (60); 2196, 2518 (60); 7852 (15); 9236 (58) Sucre et al. 8854 (1a) Suksdorf 2330 (34a) Svenson 236 (34a); 8780 (18) Swallen 1079, 1506, 1716, 1742, 1743, 10242, 10290 (32a); 1555, 1642, 1693, 1780, 10039, 10102, 10197, 10254 (24); 1618 (44); 1717, 2381, 2404, 2621, 11147, 11150, 11263 (31); 3149 (2a); 3221, 3255, 3461, 3462, 3486, 3520, 3622 ½ , 4416, 4891, 5037, 5077, 6914 (60); 3559, 3895, 4162, 4709, 4955 (62); 3569, 4266, 4400, 4414, 4830 (2a); 3608, 4313, 10783 (60); 3648 (52); 3674 (51); 3721, 4974 (54); 3822, 3998, 4106, 4859 (29); 3844, 9390 (42); 3889 , 6940(57); 4094 (53); 4130, 8312, 8415, 8773, 8884, 8989 (14a); 4192, 4396 (43); 4813, 11329, 11333, 11377 (63); 4915, 10803 (1a); 5393, 10300 (22a); 7467 (25); 8203, 8311, 8682, 8771, 9097, 9156, 9308, 9427 (41); 8678, 9598 (50); 9574 (16a); 10745, 10839, 10982, 11145 (4); 10765, 10909, 11250, 11254, 11371 (40), 10838, 11264, 11344 (35) Swartz s.n. (27); (33); (62) Systma & D’Arcy 3150 (57) Takaki s.n. (18) Tamayo 110, 1449, 3743 (63); 732 (62); 1000 (43); 1379 (14a); 1469 (29); 1638, 2919, 3949 (60); 1768, 3824, 4159 (33); 1862 (61); 1866, 2932 (14a); 2134, 2151 (57); 2674, 2806A, 2808, 2873 (54); 2907, 2964 (52) Tamberlik s.n. (52) Taylor 34 (51) Tellez & Cabrera 2630 (33) Tellez & Rico 3414 (33) Téllez 1413, 1671, 1678, 1982 (55); 3911, 9906 (62) Téllez et al. 3811, 5214 (62) Tenorio & Flores 16264 (40) Tenorio et al. 3378 (62)

List of Exsicatta

xli

Tharp 43066, 49042, 49235 (24); 43120 (32b) Thieme 532 (6) Thomas s.n. (18) Thomas et al. 1579, 5567 (14a) Thomber 176 (19b) Thompson 338 (18) Thornber 208 (34a); s.n (32b) Tirel 156 (41) Tollersey 2626 (22a) Tonduz 3679, 4875 (54); 3685 (51); 8600 (61); 14586 (33) Torres 2774, 2912 (60); 2864 (62) Torres & Cabrera 6322 (62) Torres & Hernández 3097 (31) Torres C. et al. 7860 (34a) Torres et al. 750 (60) Torrey s.n. (7a) Tovar 4585 (42) Tracy 6508 (7b); 8200 (32b); 8908 (32a); 9018, 9116 (31); 9073, 9093 (51); 9111 (27); 9338 (32b); 9342 (2a) Tressens et al. 414 (11) Triana 40 (14a); 351 (16a); 2179 (58); s.n. (33); s.n. (58); s.n. (61) Trinta 1163 (47) Troiani & Steibel 2827 (22a); 3902 (23); 4090 (2a); 4094 (23) Troncoso et al. 1650 (16a); 2638 (22a); 2721 (11) Trujillo & Fernández 472 (62); 691 (35) Trujillo 3816, 587213904 (62); 5936 (51); 9022 (31); 11172, 11399, 11492 (33); 11761 (2a); 12906 (58); 14194 (54) Trujillo et al. 17095 (58) Tür 1205 (2a) Türpe 3009 (38); 3064 (8) Tyson 1396 (34a); 2681 (57); 5494, (2a); 6018 (62); 6791 (33) Tyson et al. 3030 (4); 3034 (2a) Ugent & Flores 1893, 5801 (36) Ule 6857, 8014 (22a); 8027 (58) Uribe 3967 (54) Usteri 9820 (50); s.n. (60) Valdés Reyna 1280, 1548 (18) Valdés Reyna & Riskind 1230 (23) Valenzuela & de la Sagra 6 (60) Valerio 601, 605, 1062 (51) Valeur 7 (51) Vali & Pott 150 (61) Vallejos 259 (2a) Valls 261, 2562, 11678 (60); 649 (52); 1188 (61); 1645, 1929 (41); 1646, 2112 (25); 2309 (15);

xlii

List of Exsicatta

Valls & Arzivenco 1352 (14a); 2026 (25); 2033 (41) Valls & Mundstock 860 (25) Valls & Werneck 9867 (28) Valls et al. 2443 (14a); 2583 (60); 2769 (25); 4689 (2a); 7664 (42); 8379 (29); 8433 (6); 11521 (37); 11951 (13) van de Venne 38 (2a) van der Werff 1373 (2a) van der Werff & Wingfield 3268 (63) Van Hermann 355, 2444 (27) Vanni et al. 2047 (61) Vareschi 3745 (19b); 6569, 8158 (54) Vargas 17225 (42) Vargas et al. 401 (62) Vasey s.n. (7a), (7b) Vasquez & Jaramillo 9869 (2a) Vázquez 3592 (36) Vásquez et al. 446 (62); 7025 (33) Vázquez Avila 195 (60) Vazquez Yañes 635 (47) Velasquez 830 (2a) Velez 2734 (16a) Ventura & López 503 (62) Venturi 644, 2539, 7530 (22a); 6755 (8); 8255 (61) Vera Caletti et al. 97 (33) Vera Carnielo 6762 (60) Vera Santos 1832, 2256 (31); 1836, 1842, 1850 (34a); 2275, 3372 (51); 2826 (55) Vernier 1505 (40) Vervoorst et al. 4524 (61); s.n. (47) Viana 514 (9) Vianna 1468 (60) Vibrans 2453 (5) Vidal II-210 (15) Viegas s.n. (1a); (14a) Viereck 857 (24) Villamil 1994 (22a); 2740, 2747 (17); 2932 (62); 3554, 5369 (15); 5889 (23) Villamil & Nicora 2154 (17) Villamizar-Jaramillo s.n. (38) Villarreal 6661 (34b) Villeda 30 (62) Virlet 1300 (47); 1305, 1371 (32b); 1306 (31) von Sneidern 162 (62); 529 (60) von Wedel 1449, 1998 (61) Wagner & Solomon 4341 (36) Walker 1414 (55) Wallace et al. 292 (34a)

List of Exsicatta

xliii

Wallen 109 (7a) Waller & McAden 2643 (22a) Waller 1872, 1983, 1994, 2163, 2425, 2434 (32b); 2008, 2017, 2193, 2413, 2419, 2987a (32a); 2410, 2620 (22a) Wallies 13294 (32b) Walter 161 (2a) Walther 63 (19b) Ward s.n. (23) Warming s.n. (52) Warnock s.n. (32b) Wasum et al. 5471 (60) Waterfall 5340 (32b) Webster 3246 (44); 3291 (18) Weddell 2531 (42); 3081 (52) Weigelt 936 (1a) Wendt et al. 9788 (34a) Werner 4 (8) Weston et al. 3309 (62) Wheeler et al. 122 (19b) Widgren s.n. (50) Williams 3090 (7b) White 42 (18); 1209 (16a); 3542 (36) Whitefoord 1699, 2111 (55); 2090 (33) Whiting 608 (32b) Widgren 908 (50); 1213 (60) Wiggins 6023 (34b); 15097, 15135, 15160 (19a); 15179, 15510 (19b); 15359 (62); 18766 (2a) Wiggins & Demaree 4793, 4833 (23) Wiggins & Rollins 137 (34a) Wilbur & Teeri 12997 (62) Wilcox 303 (19b) Williams 1010, 5800 (42); 1020, 13879 (54); 1198 (18); 12152 (31); 13085 (51); 13221 (52) Williams & Molina 16751 (34a) Wilson 1405, 9443 (27) Wilson-Browne WB-61 (58) Wingate G-12 (18) Wingfield 5548, 5651 (33); 5581, 5599, 6840, 7361 (43); 5913, 13206 (31); 7288, 7333 (62); 9987, 15428 (63) Wood 3555, 3673, 4947 (60); 4091 (34a); 4192, 4752 (54); 4241 (62); 4992 (2a) Woodard s.n. (7b) Woodson & Schery 865 (62) Woodson, Jr. 1402 (51) Woodson Jr. et al. 1621 (62) Woolston G.29, G.62 (60); G.36, 258 (61); G-110 (2a)

xliv

List of Exsicatta

Wooton s.n. (19b); s.n. (24); s.n. (32a); s.n. (32b) Woronow et al. 4691 (33); s.n. (58) Wright 183 (18); 753 (61); 758 (31); 1538 (62); 1865 (2a); 3456 (1a); 3455, 3855 (60); 3877, 3852, 3860, 1540 (27) Wullschaegel 621, 622 (31); s.n. (60) Wunderlin et al. 8318 (2a) Wurdack & Monachino 41399 (16a) Wynd & Muller 174 (32b) Yepes-Agredo 1192 (62) Youngstrom 6 (18) Yuncker et al. 8777 (62) Zambrano 293 (62) Zambrano & Gutiérrez 1693 (54) Zambrano et al. 2091 (54) Zamora et al. 1184 (54) Zardini & Aquino 31413, 31692 (61) Zardini & Cuevas 5328 (61) Zardini & Soria 3894 (60) Zardini & Tilleria 29780, 32673 (61) Zardini & Velásquez 10516, 11843a, 11852, 13193, 13449, 26697 (60); 12203, 12704, 12943, 13546, 13554, 14578, 19873, 22272 (61) Zardini et al. 23582, 24200, 27533 (22a); 27353, 27536, 27610 (2a) Zelaya 229 (62); 2307 (31) Zoellner 409 (17) Zuloaga 1889 (22a); 1890 (38); 2994 (47); 3360, 7227 (15); 3554 (8); 3867 (2a); 3940 (60); 3981 (16a); 4020, 4244 (60) Zuloaga & Deginani 137, 412, 3702 (43); 457, 477, 514, 535 (22a); 547 (22b); 3447 (41); 3626 (23); Zuloaga & Londoño 4104 (14a); 4110 (61) Zuloaga & Morrone 3039 (62); 3060 (23); 3851 (16a) Zuloaga & Ortíz 4300, 4301 (31); 4524 (62) Zuloaga & Sáenz 573 (22a); 574 (47) Zuloaga et al. 661, 1880, 1899, 2219, 2267, 4506, 12232, 14624, 14974, 14978, 15260, 16052 (17); 4493 (60); 879 (47); 924, 2323, 3072, 3151, 3273 (22a); 1412, 1952, 2074, 2217, 3291 (61); 1976, 1984, 2119 (60); 2262, 3277, 3293, 4349 (2a); 2304, 4324 (16a); 2464, 3977, 4434 (54); 2473 (56); 2479 (1a); 3271, 3478 (22b); 3978, 4093, 4337, 4348 (51); 2515, 4340, 4350, 4506 (43); 4354 (4); 4363, 4504 (35); 4368 (57); 4380, 4481, 4502 (14a); 3837, 4444 (52); 4507 (33); 4508, 4509 (34a); 5906, 8450, 9201, 11875, 12109, 13401, 13581, 13705, 13706, 13713, 13777, 16090, 16116, 16401 (8); 9031 (53); 9060, 9064 (63); 9687 (24)

Acknowledgments

We thank the Agencia Nacional de Promoción Científica y Técnica (ANPCyT) for the support of the project 2418-16, the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) for the support through project PID-785, the National Geographic Society, # 3964-88, 4594-91, 5534-94, 5765-96, 6698-00, 7425-03, 8365-07, 8662-2010-9398-2013, and Vladimiro Dudás, Marcelo Moreno, and Francisco Rojas for preparation of most of the illustrations. To the curators of the herbaria consulted and to the anonymous reviewers of this contribution for their valuable suggestions and corrections.

xlv

Contents

Introduction������������������������������������������������������������������������������������������������������    1 Materials and Methods������������������������������������������������������������������������������������    3 History of the Genus����������������������������������������������������������������������������������������    5 Morphology������������������������������������������������������������������������������������������������������    9 Habit, Sheaths, Ligule, and Blades��������������������������������������������������������������������    9 Inflorescences����������������������������������������������������������������������������������������������������   10 Spikelets������������������������������������������������������������������������������������������������������������   11 Texture and Ornamentation of the Upper Anthecium����������������������������������������   12 Leaf Anatomy ��������������������������������������������������������������������������������������������������   17 Chromosome Numbers������������������������������������������������������������������������������������   23 Phylogeny and Evolution��������������������������������������������������������������������������������   29 Geographic Distribution and Habitat������������������������������������������������������������   33 Taxonomic Treatment��������������������������������������������������������������������������������������   37 Key to Ungrouped Panicum Sections and Species��������������������������������������������   38 Ungrouped Species��������������������������������������������������������������������������������������������  224 Species Excluded����������������������������������������������������������������������������������������������  275 Literature Cited�����������������������������������������������������������������������������������������������  281 Scientific Index ������������������������������������������������������������������������������������������������  289 Subject Index����������������������������������������������������������������������������������������������������  293

xlvii

Introduction

The genus Panicum L. comprises approximately 165 species of tropical and subtropical distribution, with few species reaching temperate regions of the world. It includes annual or perennial herbs belonging, within the Poaceae, to subfamily Panicoideae A.  Braun, characterized by having biflowered spikelets, with two glumes shorter or equal than the anthecia, membranous, the lower anthecium is staminate or neutral and the upper anthecium is perfect, hardened and with the lemma embracing the upper palea. Since the establishment of the genus by Linnaeus, it included a large number of species, many of which have been transferred, mainly in the twentieth and twenty-­ first centuries, to other genera of the subfamily. The present revision includes, for the Flora Neotropica, a total of 54 species within the genus and 9 “incertae sedis” species, which still remain to be classified within the Paniceae or Paspaleae. A total of 35 species were studied in the field; this revision includes 78 illustrations and distribution maps of all taxa.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_1

1

Materials and Methods

Morphological studies were based on herbarium specimens from the following institutions: ANSM,  B, BA, BAA, BAB, BAF, BLA, BR, C, CEN, CEPEC, CHAPA, COL, CORD, CTES, F, G, GH, IAN, IBGE, ICN, ISC, K, LA, LE, LIL, LP, LPB, M, MA, MEXU, MO, MY, NY, P, PORT, R, RB, S, SI, SP, UB, US, VEN, and WIS (Thiers 2021), complemented by the study of species in the field. For the histofoliar study, transverse sections were obtained at the height of the middle third of the penultimate blade of a fertile innovation. Material from herbarium specimens, previously treated with Contrad 70 (Schmid and Turner 1977) for 24 to 48 h at 20 °C, or fresh material fixed in FAA, was used. Transverse sections were made freehand, after treatment with 5% HF for 24 h. Sections were stained with 1% methylene blue and 1% safranin in 80° alcohol or with safranin-alcian blue and mounted in gelatin-glycerin. For histofoliar descriptions, the terminology proposed by Ellis (1976, 1979) was adopted. Anatomical observations were made with a Wild M20 microscope with a drawing camera. Dissections were studied with a Wild M5 stereo microscope with a drawing camera. The photomicrographs were taken with a Nikon FXA automatic equipment, with a DX-DB2 35 mm camera, and Kodak T-MAX 100 ASA film was used. A Jeol JSM-25 SII scanning electron microscope, belonging to the Faculty of Odontology (National University of Buenos Aires, Argentina), was used to obtain photomicrographs of abaxial epidermis of the lemma and palea of the upper anthecium. In this contribution we used a diagnostic criterion (Davis and Nixon 1992) for species circumscription, which has also been formulated as a phylogenetic species concept (Cracraft 1983; Nixon and Wheeler 1990; Davis and Nixon 1992; Davis 1997, 1999; Snow 1997). One or more fixed characters, or character state, were accordingly used for specific delimitation.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_2

3

History of the Genus

The name Panicum, used by ancient Latin authors, referred to what is now called Setaria italica (L.) P. Beauv., and the concept of Panicum of medieval botanists also corresponds to this species, commonly cultivated as a cereal. Tournefort (1700) gave the genus a more precise range and described 15 species, one of which he illustrated as a form of Setaria italica, which according to the author’s concept would be the type of the genus. The old name Milium corresponds to Panicum miliaceum L. Tournefort followed his predecessors and included within Milium the millets, P. miliaceum, and the sorghums, Holcus sorghum L., illustrating the former. Linnaeus (1737), in the Genera Plantarum, ed. 1, recognizes the genera Panicum and Milium, basing the former on “Panicea Scheuch. 2: 2” and the latter on “Tournef. 298.” The illustration in Scheuchzer’s work, cited by Linnaeus, corresponds to the spikelets of Setaria viridis (L.) P. Beauv. and Echinochloa crus-gallii (L.) P. Beauv. According to his description, Linnaeus seems to consider S. viridis as the type species of the genus Panicum, indicating that in some species the glume ends in an awn, referring to E. crus-gallii. He uses the name Milium in the same sense as Tournefourt. In 1753, Linnaeus, in Sp. Pl., ed. 1, describes 20 species and 5 forms under Panicum, divided into two groups, Spicata and Paniculata, including in the latter P. miliaceum. Later, Linnaeus (1762) describes 28 species of Panicum, including 18 of those mentioned above. Nees von Esenbeck (1829) divides the genus into nine sections, including species currently considered within the genera Digitaria Haller, Mesosetum Steud., Isachne R. Br., Paspalum L., Echinolaena Desv., Setaria P. Beauv., and Echinochloa P. Beauv. among others. Steudel (1853-1855) includes in the genus 17 sections, with species currently treated, for example, in Axonopus P.  Beauv., Alloteropsis J.  Presl, Ichnanthus P. Beauv., Oplismenus P. Beauv., and Tricholaena Schrad.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_3

5

6

History of the Genus

Bentham and Hooker (1883) divide the genus into 11 sections and Hackel (1887) into 9; both treatments consider species currently classified under different genera of the tribes Paniceae and Paspaleae. With the study of the Panicum species of North America (Hitchcock and Chase 1910, 1915), the current concept of the genus was initiated, which was divided into three subgenera: Eupanicum, Paurochaetium (= Setaria), and Dichanthelium, the latter also a genus segregated by Gould (1974) and Gould and Clark (1978). They treat within the subgenus Eupanicum 15 groups. They use for the delimitation of the groups characters such as shape and size of inflorescences, size of spikelets, pilosity, length of the lower glume, presence of lower palea and lower flower, ornamentation of the upper anthecium, etc. Stapf (1920), in his study of the African species of the genus, divides it into 14 sections, also using as differential characters the morphology of inflorescences and spikelets. In a similar way, Pilger (1931, 1940) divides the genus into 8 subgenera and the subgenus Eupanicum into 11 sections. However, analysis of a larger number of species in the genus, and others in the Paniceae, shows that the morphological characters of such a diverse genus are found in other genera of the tribe. Hsu (1965) studies the genus worldwide and divides it into five subgenera, using exomorphological characters of ligule, blade, leaf epidermal pattern, nervation and consistency of lodicules, and epidermal pattern of the fertile lemma. More recently, cytological, anatomical, and physiological characters were found to be diverse in Panicum s.l. Thus, two basic chromosome numbers were cited for the genus, including species with x = 9 and others with x = 10. From the anatomical point of view, numerous species were classified as non-Kranz and others as Kranz, including within the latter all the anatomical types found in Poaceae, correlated with the photosynthetic patterns Kranz-NAD-me, Kranz-NADP-me, and Kranz-PEPck; in addition, there are species of the genus with intermediate Kranz and non-Kranz anatomy (Brown 1977; Ellis 1988; Acosta et al. 2014). Recent phylogenetic studies in Panicum, based on morphological and molecular characters (Zuloaga et al. 2000; Gómez-Martínez and Culham 2000; Giussani et al. 2001; Duvall et al. 2001, 2003; Aliscioni et al. 2003), have proven that the genus is polyphyletic. Consequently, several subgenera were raised, or considered again at the generic level, such as Dichanthelium (Hitchc. & Chase) Gould, Phanopyrum Raf., Steinchisma Raf., Megathyrsus (Pilg.) B.K.  Simon & S.W.L.  Jacobs, and Louisiella C.E.  Hubb. & J.  Léonard (Zuloaga et  al. 1998, 2003; Aliscioni et  al. 2003; Freckmann and Lelong 2003a; Simon and Jacobs 2003; Barkworth 2006; Scataglini et al. 2014). At the same time, Panicum was restricted to the type subgenus (Aliscioni et al. 2003; Zuloaga et al. 2018) and the rest of the species classified, at the time, as “incertae sedis” within the Paniceae. Of these “incertae sedis” species, several have been transferred to existing genera, or new genera such as Aakia J.R.  Grande, Adenochloa Zuloaga, Apochloa Zuloaga & Morrone, Batochloa Salariato & Zuloaga, Canastra Morrone, Zuloaga, Davidse & Filg, Cnidochloa Zuloaga, Coleataenia Griseb. (= Sorengia Zuloaga & Morrone), Cyphonanthus Zuloaga & Morrone, Hopia Zuloaga & Morrone, Janochloa Zuloaga & Delfini, Morronea Zuloaga & Scataglini, Ocellochloa Zuloaga & Morrone, Osvaldoa

History of the Genus

7

J.R.  Grande, Parodiophyllochloa Zuloaga & Morrone, Renvoizea Zuloaga & Morrone, Stephostachys Zuloaga & Morrone, and Zuloagaea Bess, Doust, Davidse & Kellogg (Bess et  al. 2006; Grande Allende 2014; Delfini et  al. 2023a, 2023b; Morrone et al. 2007, 2008; Zuloaga et al. 2006, 2007, 2010, 2015, 2020; Sede et al. 2008, 2009; Soreng 2010; Scataglini and Zuloaga 2013). These contributions have undoubtedly established the polyphyly of Panicum, highlighting differential exomorphological, anatomical, and physiological characters previously considered within the genus. Among these we can stress differences in the Kranz syndrome, correlations of this character with anatomical structures of the leaves, such as the distance between vascular bundles, presence of chloroplasts in the parenchymatous or mestomatic sheaths, also types of inflorescences, nervation of bracts, length of the lower glume, presence of lower palea and lower flower, and texture and ornamentation of the upper anthecium, including pilosity and presence or absence of simple or compound papillae. As a conclusion, Panicum should be restricted to its type subgenus, i.e., Panicum s.s., which is in turn the type genus of the tribe Paniceae and the subtribe Panicinae (Morrone et al. 2012; Zuloaga et al. 2018; see phylogeny of the genus); thus, the genus strictly speaking includes only C4 species. On the other hand, the “incertae sedis” species still need to be classified in other subtribes of the Panicoideae.

Morphology

Habit, Sheaths, Ligule, and Blades The genus includes annual or perennial species, when perennials are short to long rhizomatous, with strong rhizomes in species of section Hiantes Stapf, such as P. glabripes Döll and P. tricholaenoides Steud. (Figs. 12 and 19 in chapter “Taxonomic Treatment”). The size of the plants is variable, ranging from species about 15 cm tall, such as P. furvum Swallen, to others reaching approximately 2–3 m tall, such as P. hirsutum Sw. (Fig. 37 in chapter “Taxonomic Treatment”). Species are cespitose, usually with hairy cataphylls, branching at the base of the plant (species of section Panicum), or they have erected to decumbent and rooting culms, densely branching or not at the lower nodes, as in species from humid environments of section Dichotomiflora (Hitchc.) Hitchc. & Chase ex Honda, such as P. dichotomiflorum Michx. (Fig. 2 in chapter “Taxonomic Treatment”). Culms are either unbranched or branching at the basal and middle nodes, exceptionally branching in the upper portion of culms, as in species of section Rudgeana (Hitchc.) Zuloaga (P. cayennense Lam. and P. rudgei Roem. & Schult., Figs. 54 and 58 in chapter “Taxonomic Treatment”); culms are many noded, with very short internodes toward the base of the plant and elongate in the floriferous branches, or few noded. The internodes are cylindrical, hollow, exceptionally solid, and hairy to glabrous. Subwoody, conspicuously branched culms are observed in P. deciduum Swallen, P. glabripes Döll, and P. tricholaenoides Steud. (Figs. 11 and 19 in chapter “Taxonomic Treatment”). The nodes are usually compressed, brown to dark brown with variable pilosity. The sheaths are open and striate and vary from glabrous to densely pilose with variable pilosity, with or without tuberculate hairs, urticating hairs are observed in P. hirsutum (Fig. 37 in chapter “Taxonomic Treatment”), and sheaths are glaucous in P. hallii subsp. hallii Vasey, P. hallii subsp. filipes (Scribn.) Freckmann & Lelong, and P. tamaulipense F.R. Waller & Morden. Moreover, less frequently sheaths are spongy, with aerenchyma, in P. aquarum Zuloaga & Morrone © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_4

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Morphology

(Fig. 24 in chapter “Taxonomic Treatment”). Ligules are membranous in the basal portion and short to broadly ciliate toward the distal portion. Leaf blades are flat, or with involute margins, and vary in shape from oblong-lanceolate, lanceolate, linear-­ lanceolate, and linear to filiform; pilosity on the leaf blade is variable, from hispid or hirsute, with tuberculate hairs, to glabrous, the margins being scabrous, the basal ones with or without deciduous tuberculate hairs. In P. hallii subsp. hallii, P. hallii subsp. filipes, and P. tamaulipense, the basal leaves, at senescence, were persistently enrolled at the base of the plant. The persistence of enrolled basal leaves was previously reported in Panicum mystasipum Zuloaga & Morrone (Zuloaga and Morrone 1992); these authors suggest that this character is possibly related to the protection of buds from external environmental factors. Among the “incertae sedis” species discussed here, the C3 species grow in the interior or edges of forests, are gracile plants, with decumbent, geniculate culms, rooting at the lower nodes and leaning on the vegetation, for example, P. millegrana Poir., P. sellowii Nees, and P. trichanthum Nees. In these species ligules are membranous and blades usually ovate-lanceolate, i.e., P. bartlettii Swallen, P. brevifolium L., P. haenkeanum J.  Presl, P. hirtum Lam., P. millegrana, P. trichanthum, P. trichoides Sw., and P. venezuelae Hack.; in addition, blades are subcordate to cordate and amplexicaul leaf blade, often pseudopetiolate at base, as in P. bartlettii, P. trichanthum, and P. trichoides.

Inflorescences The species of the genus usually have terminal panicles, and in sections Panicum (P. aquarum, Fig. 24 in chapter “Taxonomic Treatment”) and Rudgeana (as in P. cayennense and P. rudgei, Figs. 54 and 58 in chapter “Taxonomic Treatment”), there are conspicuous terminal and axillary inflorescences. The inflorescences are lax and diffuse, with spikelets solitary or in pairs, in most species, to contracted, with the branches appressed to the main axis, as in P. decolorans Kunth, P. chaseae Roseng., B.R.  Arrill. & Izag., P. longisssimum (Mez) Henrard, P. magnispicula Zuloaga, Morrone & Valls (Fig. 42 in chapter “Taxonomic Treatment”), and P. racemosum (P. Beauv.) Spreng (Fig. 17 in chapter “Taxonomic Treatment”). The lower branches of the inflorescence are verticillate in P. bergii Arechav. (Fig. 28 in chapter “Taxonomic Treatment”) and occasionally verticillate in specimens of P. capillare L. and alternate to subopposite in the rest of the taxa. Furthermore, in P. bergii and P. capillare, the inflorescences disarticulate as a whole at maturity and are dispersed by the wind, the panicle acting as a dispersal unit (Zuloaga and Morrone 1996). Vegetti and Pensiero (1993) study the typology of several species of section Panicum, concluding that the inflorescence in this group is polytelic, with a developed, non-truncate main inflorescence. The inflorescences of the “incertae sedis” species are lax and open, with the exception of P. venezuelae, which has spikelets arranged in unilateral racemes. It

Spikelets

11

should be noted that conspicuous glands are present on the inflorescence axes of P. hirtum (Fig. 62 in chapter “Taxonomic Treatment”).

Spikelets The spikelets are biflowered, with a sterile or staminate lower flower and a perfect upper one, exceptionally triflowered, with two sterile lower anthecia and a perfect upper one in P. quadriglume (Döll) Hitchc. (Fig. 47 in chapter “Taxonomic Treatment”). Species have ovoid to long ellipsoid spikelets, closed to open at maturity, as in species of section Hiantes, glabrous to hairy, for example, in P. chloroleucum Griseb. (Fig. 8 in chapter “Taxonomic Treatment”), P. racemosum, and P. urvilleanum Kunth. (Figs. 17 and 20 in chapter “Taxonomic Treatment”) or with tuberculate-based hairs (P. deciduum, Fig. 11 in chapter “Taxonomic Treatment”); spikelets are pale or tinged with purple. The lower glume varies in size from 1/4 to 3/4(−4/5) the length of the upper glume and lower lemma; it is truncate (P. repens L., Fig. 49l in chapter “Taxonomic Treatment”), acute to acuminate, nerveless to 3–5 nerved, occasionally 7–9 nerved, with the midrib often scabrous. The upper glume and lower lemma are usually subequal and (5-)7–9(−13–15) nerved. In all C3 species, treated here as “incertae sedis,” the upper glume and lower lemma are 3–5 nerved. The lower and upper glumes are separated by a distinct internode in species such as P. aquarum (Fig. 24 in chapter “Taxonomic Treatment”), P. aztecanum Zuloaga & Morrone (Fig. 26 in chapter “Taxonomic Treatment”), and P. parcum Hitchc. & Chase. The upper glume, but not the lower glume, is early deciduous in a group of species that includes P. exiguum Mez (Fig. 35 in chapter “Taxonomic Treatment”), P. furvum, P. peladoense Henrard (Fig. 46 in chapter “Taxonomic Treatment”), and P. quadriglume. This character is correlated with the upper anthecium being blackish at maturity, which is exposed by the fall of the upper glume; at the same time, these characters are correlated with the upper anthecium being early deciduous. A similar correlation to the American species previously described is found to be P. atrosanguineum A. Rich., a species inhabiting Africa and NW India (Clayton 1989). Van der Pijl (1982) indicates that the presence of this striking color, and stiffness at maturity of the upper anthecium, would function as a mechanism of attraction for granivorous birds that would act as dispersal agents. The lower palea varies from species in which it is null or not very evident to species in which it reaches the same length as the lower lemma. The lower flower is absent or staminate. The upper anthecium encloses a perfect flower; the lemma and palea are hardened and pale to brownish in most species, or less frequently blackish, as mentioned above for P. exiguum, P. peladoense, and related species. The presence of an indurated lemma and palea, enclosing the caryopsis at maturity, was correlated to dispersal by granivorous or herbivorous animals, which ingest the diaspores accidentally or intentionally (Ridley 1930; van der Pijl 1982). Thomasson (1985) suggests that the hardened bracts protect the caryopsis from insect attack or during passage through the digestive tract of animals. The lemma encloses the palea and is

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Morphology

5–7 nerved; the palea is 2 nerved. Inside the upper anthecium are two truncate, cuneate lodicules, which embrace the lower edges of the upper palea, three stamens and an ovary with two free styles from the base and feathery stigmas. Finally, the caryopsis varies from ovoid to ellipsoid or fusiform and has a punctiform to oblong hilum, and the embryo reaches about 1/3 to 1/2(−3/4) the length of the caryopsis. The spikelet species of the genus have the structural plan of the Paniceae, in which the spikelet falls as a whole at maturity. As indicated above, in some species the upper glume is early deciduous, with a primary articulation zone at the base of the upper anthecium, the latter falling naked at maturity, subsequently disarticulating the remaining bracts at the base of the lower glume together with the pedicel. The presence of primary disarticulation at the base of the upper anthecium has been previously noted in the tribes Paniceae and Paspaleae for Axonopus, Brachiaria (Trin.) Griseb., Echinolaena Desv., Ichnanthus, Melinis P.  Beauv, Panicum, Paspalum L., Pennisetum Rich., Sacciolepis Nash, and Tricholaena (Lazarides and Webster 1984; Silberbauer-Gottsberger 1984; Davidse 1987; Webster 1987; Zuloaga 1987a; Morrone and Zuloaga 1992).

Texture and Ornamentation of the Upper Anthecium The upper anthecium is ovoid to broadly ellipsoid (obovoid in P. alatum Zuloaga & Morrone), dorsiventrally compressed, crustaceous, with the edges of the lemma rolled up and covering 2/3 of the surface of the palea. The abaxial epidermis of the lemma and palea has long rectangular cells distributed in longitudinal rows, more than three times longer than wide, with wavy longitudinal anticlinal walls. The abaxial epidermis can present, particularly on the palea, different ornamentations, including simple papillae, verrucose papillae, compound papillae (Fig.  1g, h), microhairs, and prickle hairs. The epidermis of the palea commonly lacks ornamentation, with simple papillae regularly distributed over the entire surface in Panicum alatum var. alatum (Fig. 1f), P. hirticaule var. verrucosum Zuloaga & Morrone (Fig. 1e), and P. magnispicula, or only along the distal portion of the palea in P. quadriglume (Fig. 1d). Irregularly distributed simple papillae, at the apex of the palea, were observed on P. alatum var. minus Zuloaga & Morrone, P. aztecanum, P. bergii, P. capillare, P. chaseae, P. diffusum Sw. (Fig.  1a), P. exiguum, P. furvum (Fig.  1b), P. ghiesbreghtii E.  Fourn., P. hallii subsp. hallii (Fig.  1c), P. hirsutum Sw., P. hirticaule J.  Presl var. hirticaule, P. hispidifolium Swallen, P. lepidulum Hitchc. & Chase, P. mucronulatum Mez, P. peladoense, P. quadriglume (Fig. 1d), and P. stramineum Hitchc. & Chase. Verrucose papillae, irregularly grouped at the apex of the palea, were found in P. aquarum, P. capillarioides Vasey, P. decolorans, P. ephemeroides Zuloaga & Morrone, P. hispidifolium (Fig. 2a), P. miliaceum, and P. parcum, as well as in species of sect. Rudgeana, in P. cervicatum (Fig. 2b) and P. ligulare.

Fig. 1  Scanning electron microscope (SEM) photomicrographs of upper anthecia of Panicum species. (a) P. diffusum (From Eggers 76, MO), simple papillae toward the apex of the palea. (b) P. furvum (From Steyermark 51627, US), simple papillae toward the apex of the palea. (c) P. hallii subsp. hallii (From Hall 816, US), simple papillae toward the apex of the palea. (d) P. quadriglume (From Chase 8897, MO), simple papillae toward the apex of the palea. (e) P. hirticaule var. ­verrucosum (From Colunga & Zizumbo 44, MEXU), detail of apex of upper anthecium with papillae regularly distributed over the entire surface. (f) P. alatum var. alatum (From Wiggins 15160, MO), detail of apex of upper anthecium with papillae regularly distributed over the entire surface. (g) P. cervicatum (From Chase 10737, US), apex of upper anthecium with compound papillae at apex of palea. (h) P. ligulare (From Irwin 14904, US), apex of upper anthecium with compound papillae at apex of palea. Scales: (a, e–h) 50 μm; (b–d) 20 μm

Fig. 2  Scanning electron microscope (SEM) photomicrographs of upper antheciums of Panicum species. (a) P. hispidifolium (From Rodríguez 1981, US), detail of verrucose papillae at the apex of the palea. (b) P. cervicatum (From Chase 10737, US), detail of compound papillae at apex of palea. (c) P. urvilleanum (From Zuloaga et al. 14624, SI), base of upper anthecium with long unicellular macrohairs. (d) P. campestre (From Chase 8645, US), base of upper anthecium with stipe with membranous portion reduced towards base of palea. (e) P. cervicatum (From Steyermark 59173, US), base of upper anthecium with membranous appendages. (f) P. alatum var. alatum (From Wiggins 15160, MO), base of upper anthecium with two fleshy expansions on lemma. (g) P. hispidifolium (Rodriguez 1981, US), base of upper anthecium with a basal ring. (h) P. hirticaule var. verrucosum (From Colunga & Zizumbo 44, MEXU), base of upper anthecium with two scars on lemma. Scales: (a) 10 μm; (b) 20 μm; (c–h) 50 μm

Texture and Ornamentation of the Upper Anthecium

15

Prickle hairs are present at the apex of the lemma and next to the palea in Panicum alatum var. minus, P. bergii, P. ghiesbreghtii, P. lepidulum, P. miliaceum, and P. mucronulatum. Panicum complanatum, P. mystasipum, and P. olyroides present flattened unicellular macrohairs at the base of the upper lemma; moreover, linear macrohairs are conspicuous at the base of the upper lemma in P. chloroleucum, P. racemosum, and P. urvilleanum (Fig. 2c). The base of the upper anthecium differs in the species studied; a conspicuous stipe, at the base of the upper anthecium, is observed in species of sect. Rudgeana. This stipe consists of two well-defined segments: (a) a membranous portion toward the base of the upper palea (reduced in P. cayennense Lam. and P. campestre Nees ex Trin, Fig. 2d) and (b) a hardened, smooth, and shiny tissue toward the lemma of the upper anthecium; in P. ligulare Nees ex Trin. and P. cervicatum Chase, this conspicuous stipe may extend in two wings toward the base of the upper lemma (Fig. 2e). Several Australian species with a similar and conspicuous stipe were considered within the genus Yakirra Lazarides & R.D. Webster (Lazarides and Webster 1984) and are currently treated within the genus Panicum in sect. Yakirra (Lazarides & R.D. Webster) Zuloaga (Zuloaga et al. 2018, see phylogeny of the genus). Davidse (1987) mentioned that the stipe present in species of Rudgeana is an elaiosome associated with myrmecochory; therefore, the upper anthecium functions as a dispersal unit by ants; similarly, Berg (1985) describes this dispersal mechanism for P. australiense Domin, a species of sect. Yakirra. In addition, two winged and fleshy expansions on the basal margins of the lemma, olivaceous at maturity, were observed in Panicum alatum var. alatum (Fig. 2f); this character is also present in species of the genera Ichnanthus and Echinolaena. Histochemical analysis with Sudan IV (Jensen 1962) determined the presence of oils in the fleshy appendages of P. alatum. Other species of sect. Panicum, such as P. aquarum, P. aztecanum, P. decolorans, P. ephemeroides, P. hispidifolium (Fig.  2g), P. lepidulum, P. magnispicula, and P. parcum, present a disk composed of cells less thickened than the rest of the lemma surface, noticeably brown at maturity. In Panicum bergii, P. capillarioides, P. diffusum, P. exiguum, P. furvum, P. ghiesbreghtii, P. hallii, P. hirsutum, P. hirticaule var. verrucosum (Fig. 2h), P. miliaceum, P. mucronulatum, P. peladoense, P. quadriglume, and P. stramineum, two basal scars are found at both margins of the lemma, composed of thin-walled cells, less thickened than the rest of the lemma surface, and markedly brown at maturity. The species of Panicum “incertae sedis” treated here have a hardened, pale to whitish, obtuse to apiculate, smooth or transversely rugose upper anthecium, with bicellular microhairs on its entire surface in P. millegrana, P. sellowii, P. bartlettii, P. hirtum, P. trichanthum, and P. trichoides, together with simple papillae, or stomata, prickle hairs, and bicellular microhairs toward the apex. Panicum venezuelae is distinguished by its upper anthecium shortly stipitate.

Leaf Anatomy

The photosynthetic patterns C3 and C4 are a fundamental characteristic of variation within plants, with importance both at the level of biomes and genomes; knowing the distribution of these patterns, and their taxonomic implication, among wild species is a priority, to understand the patterns and processes of photosynthetic evolution and their role in large-scale ecological processes (Osborne et al. 2014). In the Poaceae, more than 50% of the taxa are Kranz or C4 (Sage et al. 1999); the species currently considered in the genus Panicum (Zuloaga et al. 2018) are characterized by including Kranz or C4 species, of the anatomical subtype PS (XyMS PLUS_SPI) (Hattersley and Watson 1976; Brown 1977), with two sheaths around the vascular bundles, the internal mestomatic and the external Kranz, with specialized chloroplasts. The chloroplasts centripetally positioned (group A) are characteristic of species of the anatomical NAD and physiological NAD-me subtype (Gutiérrez et al. 1974). The centrifugally positioned chloroplasts (group B) correspond to the anatomical subtype PCK, which has traditionally been associated with the physiological type PEP-ck (Gutiérrez et  al. 1976). However, this structural and functional correlation should be taken with caution, given the exceptions found so far within Panicum. Thus, several authors have cited Panicum species with PCK anatomy and NAD-me photosynthetic subtype (Ohsugi and Murata 1980; Ohsugi et  al. 1982; Oguro et al. 1985; Prendergast et al. 1987). Within section Panicum, for example, P. bergii and P. capillare, species included in group B and corresponding to the anatomical subtype PCK have been physiologically described as NAD-me type (Gutiérrez et  al. 1974; Prendergast et  al. 1987). Likewise, within section Dichotomiflora, species were described as physiologically NAD-me, but the chloroplasts are centrifugally located within the Kranz sheath, i.e., P. dichotomiflorum (Ohsugi and Murata 1985, 1986) and P. aquaticum (Fig. 1f) (Guglieri et al. 2008). The presence of centrifugal chloroplasts in the Kranz parenchyma sheath was also mentioned for species of section Repentia such as P. gouinii, P. pedersenii, and P. repens and of section Hiantes such as P. altum, P. glabripes, and P. tricholaenoides (Guglieri et al. 2008). © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_5

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Leaf Anatomy

Fig. 1  Cross section of the leaf blade of Panicum species. (a, b) P. stramineum (From Zuloaga et al. 2515, SI). (c, d) P. racemosum (From Zuloaga 3360, SI). (e) P. peladoense (From Zuloaga & Deginani 3447, SI). (f) P. aquaticum var. aquaticum (From Davidse & Brant 32947, MO). (g) P. cayennense (From Zuloaga 3978, SI). (h) P. cervicatum (From Zuloaga 3837, SI). (i, j) P. tricholaenoides (i, From Zuloaga s.n., SI; j From Zuloaga et al. 2304, MO). Scales: (a) 200 μm; (b, d–j) 50 μm; (c) 100 μm

Leaf Anatomy

19

Neotropical species of Panicum s. str., physiologically C4 and anatomically Kranz, of the anatomical subtype PS, present a variable leaf anatomy. The leaf blade transverse section can be flat, expanded as in P. aquaticum, P. altum, P. campestre, P. exiguum, and P. stramineum (Fig. 1a, b) or involute in “U” or “V” shaped as in P. ephemeroides, P. gouinii, and P. racemosum (Fig. 1c, d), symmetric on both sides of the midrib. Some species, such as P. hirsutum, P. miliaceum, P. mucronulatum, and P. stramineum (Fig. 1a, b), may have a central keel with adaxial colorless parenchyma, with different degrees of development; in other cases, as in P. bergii (Fig.  2c), P. cayennense, P. peladoense, and P. racemosum (Fig.  1c, d), the

Fig. 2  Cross section of the leaf blade of Panicum species. (a) P. hispidifolium (From Davidse & Pilz 31533, MO). (b) P. coloratum (From Burandt Jr. V0120, MO). (c, d) P. bergii (From Zuloaga et al. 2323, SI). (e) P. brevifolium (From Zuloaga et al. 2473, SI). (f) P. millegrana (From Zuloaga 2219, SI). (g) P. bartlettii (From Davidse & Brant 32539, SI). (h) P. sellowii (From Zuloaga et al. 2119, SI). Scales: (a, b, d–h) 50 μm; (c) 200 μm

20

Leaf Anatomy

parenchymatous central keel is absent. The adaxial and abaxial surfaces may show ribs and furrows with different degrees of development; the abaxial surface lacks furrows or these are very little evident; on the contrary, the adaxial surface varies from almost smooth as in P. exiguum, P. miliaceum, and P. peladoense (Fig. 1e) to poorly developed furrows present, 1/4–1/3 of the thickness of the transverse section in P. aquaticum (Fig. 1f), P. cayennense (Fig. 1g), P. cervicatum (Fig. 1h), P. dichotomiflorum, P. hirsutum, P. stramineum (Fig. 1a, b), and P. tricholaenoides (Fig. 1i, j), to deep adaxial furrows, 1/2 or more the thickness of the transverse section, e.g., in P. aztecanum, P. ephemeroides, P. mucronulatum, and P. racemosum (Fig. 1c, d). Vascular bundles are distinguished into first-order bundles (e.g., with up to 30 first-­ order vascular bundles in P. hirsutum), subcircular to elliptical in outline, with angularly contoured metaxylem vessels; first-order vascular bundles alternate with (2–)3–7 second-order vascular bundles, also angular in outline, with distinguishable xylem and phloem; in some cases, third-order bundles are differentiated, the latter distributed below the bulliform cells as in P. chloroleucum, P. racemosum (Fig. 1c, d), and P. urvilleanum. The vascular bundles are surrounded by two sheaths, the inner mestomatic sheath composed of small cells with uniformly thickened walls and the outer parenchymatous sheath containing specialized chloroplasts. The sclerenchyma develops in discontinuous subepidermal clusters forming girders, usually abaxially interrupting the parenchymatous sheath in first-order bundles as in P. racemosum (Fig. 1c, d), P. bergii (Fig. 2d), or more frequently in adaxial and abaxial strands, for example, in P. cervicatum (Fig. 1h) and P. tricholaenoides (Fig. 1i, j). In some species, an extension of the parenchyma sheath toward both epidermis is distinguishable, as in P. racemosum (Fig. 1c, d). The mesophyll is conspicuously radiate, compact, continuous, or interrupted by sclerenchyma strands or girders, with tabular chlorenchyma cells; 2–3(−4) chlorenchyma cells are present between contiguous vascular bundles and in some cases interrupted by 1–3-seriate columns of colorless parenchyma cells in contact with both epidermes, as in P. alatum, P. aquarum, P. capillare, P. decolorans, P. hirsutum, P. hirticaule, P. hispidifolium (Fig. 2a), P. lepidulum, P. magnispicula, P. parcum, and P. tricholaenoides (Fig. 1i, j). The epidermis has bulliform cells located in the adaxial furrows and between the vascular bundles, in small groups and fan-shaped, composed of 2–7 cells, the central one usually larger than the lateral ones, occupying up to 1/3 of the width of the transverse section. In those species where third-order bundles are differentiated, bulliform cells may be located above them; the remaining adaxial and abaxial epidermal cells are regular in shape; in some cases, for example, P. cervicatum (Fig. 1h), prickle hairs and hooks may be present in both epidermes. In some species, as in P. aquarum, P. coloratum (Fig. 2b), P. dichotomiflorum, P. gouinii, P. pedersenni, P. repens, and P. stramineum (Fig. 1a, b), adaxial papillae are observed; less frequent is the presence of papillae on both epidermes, for example, in P. aquaticum (Fig. 1f). Macrohairs are present or absent, when present with bulbous base, associated with raised epidermal cells or at the same level as the rest of the surface, as in P. capillare and P. quadriglume. Regarding the species considered here as “incertae sedis” of the genus, P. bartlettii (Fig.  2g), P. brevifolium (Fig.  2e), P. haenkeanum, P. hirtum, P. millegrana

Leaf Anatomy

21

(Fig. 2f), P. sellowii (Fig. 2h), P. trichanthum, P. trichoides, and P. venezuelae are C3 species, of non-Kranz anatomy, with two sheaths around the vascular bundles, the outer parenchymatous and the inner mestomatic, both without specialized chloroplasts and with more than five chlorenchymatous cells between contiguous vascular bundles (Brown 1977; Osborne et al. 2014).

Chromosome Numbers

The basic chromosome number in Panicum is x = 9, being uniform for the genus (Dubcovsky and Zuloaga 1991; Aliscioni et  al. 2003). However, different non-­ multiple numbers of 9 were also reported, for example, in P. gouinii 2n = 20, 40 (Núñez 1952), P. repens 2n = 40 (Krishnaswamy 1941), and 2n = 45 (Tateoka 1955, 1956), P. virgatum 2n = 25, 30 (Brown 1948). Some of these numbers could have evolved as cytological aberrations (Aliscioni et al. 2003). As for the species not yet grouped into sections, most of them also report a basic chromosome number x = 9, with the exception of P. haenkeanum which is 2n = 20 (Pohl and Davidse 1971). The presence of polyploid series is also common in Panicum, for example, it was reported for P. virgatum associating this character with its distribution area (Brunken and Estes 1975). Likewise, P. repens has been reported as diploid (Ashan et  al. 1994) and also forming a polyploid complex including tetraploids, hexaploids, and aneuploids (Freckmann and Lelong 2003b) (Fig. 1 and Table 1).

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24

Chromosome Numbers

Fig. 1  Photograph of species in the field. (a, b) P. chloroleucum. (c) P. racemosum. (d) P. glabripes. (e) P. olyroides var. olyroides. (f) P. urvilleanum. (g) P. trichoides

Chromosome Numbers

25

Table 1  Chromosome numbers in Panicum Section Species Dichotomiflora P. aquaticum

P. dichotomiflorum

Number n = 36 2n = 36 2n = 72 n = 18 2n = 36 2n = 54

Hiantes

P. sublaeve P. vaseyanum P. amarum subsp. amarum P. glabripes

n = 18 2n = 36 2n = 36

Reference Davidse and Pohl (1972a) Davidse and Pohl (1974), Mehra and Chaudhary (1974) Pohl and Davidse (1971) Davidse and Pohl (1974) (under P. chloroticum); Dubcovsky and Zuloaga (1991) Brown (1948) Church (1929), Gould (1958, 1968), Probatova and Sokolovskaya (1983), Hamoud et al. (1994) Dubcovsky and Zuloaga (1991) Reeder (1971) Brown (1948)

2n = 36

Parodi (1946), Núñez (1952) (under P. subjunceum) P. olyroides var. n = 18 Dubcovsky and Zuloaga (1991) olyroides 2n = 36 Norrmann et al. (1994) P. racemosum 2n = 36 Parodi (1946), Núñez (1952) P. tricholaenoides 2n = 36 Parodi (1946), Núñez (1952) P. urvilleanum 2n = 36 Parodi (1946), Núñez (1952) P. virgatum 2n = 21, 25, Brown (1948) 30, 32 2n = 36 Love and Love (1981), Hamoud et al. (1994) 2n = 36, 54 Ridley and Vogel (1982) (continued)

Chromosome Numbers

26 Table 1 (continued) Section Panicum

Species P. alatum var. minus P. bergii var. bergii

Number 2n = 18

Reference Gould (1965) (under Panicum hirticaule)

n = 18 2n = 36

P. capillare

n = 9

Dubcovsky and Zuloaga (1991) Parodi (1946) (under P. pilcomayense); Núñez (1952) (under P. pilcomayense). Spellenberg (1967) (under P. capillare var. occidentale Rydb. ex Fernald); Vahidy et al. (1987) Avdulov (1931), Gould (1968, 1975), Spellenberg (1970), Fairbrothers (1953), Tzvelev (1976) (under P. capillare subsp. barbipulvinatum (Nash) Tzvelev); Vahidy et al. (1987), Hamoud et al. (1994), Murín et al. (1999) Gould (1960) Gould (1968) Gould (1966) Gould (1960) Brown (1951), Gould (1958, 1968), Waller (1976) Gould (1958) Gould (1958, 1960) Gould (1958, 1965, 1966, 1968),Reeder (1971), Waller (1976), Gould (1958, 1965, 1966, 1968), Reeder (1971); Waller (1976) Waller (1976) Gould (1975), Waller (1975, 1976) Parfitt (1981), Ahsan et al. (1994) Avdulov (1931), Krishnaswamy (1951), Frey et al. (1981), Probatova and Sokolovskaya (1983), Sinha et al. (1990), Hamoud et al. (1994), Haroun (1995), Warwick et al. (1997) Krishnaswamy (1951) Arenkova (1940), Krishnaswamy (1951) Davidse and Pohl (1972a), Morrone et al. (2006) Fairbrothers (1954), Gould (1975) Dubcovsky and Zuloaga (1991) Gould (1966), Gould and Soderstrom (1970) Hunziker et al. (1998) Fairbrothers (1954)

2n = 18

P. capillarioides P. decolorans P. hallii subsp. filipes

n = 18 2n = 36 2n = 36 n = 9 2n = 18

P. hallii subsp. hallii

2n = 36 n = 9 2n = 18

P. hirsutum P. miliaceum

P. hirticaule

P. hispidifolium P. parcum P. stramineum

2n = 36 2n = 36 n = 18 2n = 36

2n = 54 2n = 72 n = 9 2n = 18 n = 27 2n = 36 n = 18II 2n = 36

(continued)

Chromosome Numbers

27

Table 1 (continued) Section Repentia

Species P. coloratum

Number 2n = 36

P. gouinii

2n = 36 2n = 20 2n = 40 n = 9 n = 27

P. repens

n = 18, 30 2n = 36

Rudgeana

Incertae sedis

P. cervicatum P. rudgei

2n = 40 2n = 45 2n = 54 2n = 18 n = 9 n = 27 n = 9 n = 9

P. brevifolium

2n = 18 n = 18

P. campestre P. cayennense

P. haenkeanum P. sellowii

P. trichanthum

P. trichoides

2n = 20 n = 27 2n = 36 2n = 54 n = 18 n = 18II 2n = 18 2n = 36 2n = 18

Reference Núñez (1952), Hamoud et al. (1994), Haroun (1995), Haroun (1995) Parodi (1946) Núñez (1952) Núñez (1952) Ahsan et al. (1994) Gould (1960), Bir and Sahni (1986), Devesa et al. (1991) Dujardin (1979), Mehra and Chaudhary (1981), Mehra (1982) de Wet and Anderson (1956), Jauhar and Joshi (1966), Haroun (1995) Krishnaswamy (1941) Tateoka (1955, 1956) Gould (1960), Fernandes and Queiros (1969) Dubcovsky and Zuloaga (1991) Dubcovsky and Zuloaga (1991) Davidse and Pohl (1974) Dubcovsky and Zuloaga (1991) Davidse and Pohl (1974), Dubcovsky and Zuloaga (1991) Pohl and Davidse (1971) Dujardin (1979), Mehra and Chaudhary (1981) Pohl and Davidse (1971) Davidse and Pohl (1972a, 1974) Gould and Soderstrom (1970) Pohl and Davidse (1971) Davidse and Pohl (1972a, b) Morrone et al. (2006) Davidse and Pohl (1972a, b) Pohl and Davidse (1971), Honfi et al. (1990), Norrmann et al. (1994) Pohl and Davidse (1971), Reeder (1984)

Phylogeny and Evolution

As previously mentioned (see history of the genus), numerous contributions in the last 20 years have corroborated the polyphyly of Panicum, highlighting exomorphological, anatomical, and physiological characters, as well as basic chromosome number data of the species. These characters also include differences in the Kranz syndrome, species C3 and C4, or intermediate C3–C4 previously grouped into subgenera, sections, or species groups of Panicum (Zuloaga 1987a), correlation of physiological characters with blade structures, such as distance between vascular bundles, presence of specialized chloroplasts in the mestomatic or parenchymatous sheath, type of inflorescence, bract nervation, length of the lower glume, presence and/or absence of lower palea and lower flower, ornamentation and texture of the upper anthecium including pilosity, and presence or absence of simple or compound papillae and their distribution on the lemma and upper palea. As a consequence, Zuloaga et al. (2018) conclude that Panicum should be restricted to the type subgenus (Figs. 1 and 2), its type species being P. miliaceum L. The genus is in turn the type of the subtribe Panicinae (Soreng et al. 2017), which it integrates together with the genus Louisiella C.E. Hubb. & J. Léonard, the latter with two species, L. elephantipes (Nees ex Trin.) Zuloaga, from Tropical America, and L. fluitans C.E.  Hubb. & J.  Léonard from Africa. In its current circumscription, Panicum includes ca. 165 pantropical and temperate species, grouped into seven sections: Arthragrostis (Lazarides) Zuloaga, Yakirra, Dichotomiflora, Hiantes, Panicum, Repentia and Rudgeana, of which the last five occur in America and the Neotropical region. A total of 159 species have been excluded from the genus (see list of excluded species) and included in three subtribes of the Paspaleae, Arthropogoninae, Paspalinae, and Otachyriinae, and in the five subtribes of the Paniceae: Boivinellinae, Cenchrinae, Dichanthelliinae, Melinidinae, and Panicinae. There are still 74 “incertae sedis” species to be classified, 9 of which grow in America and are treated here with this criterion.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_7

29

30

Phylogeny and Evolution

100 98 1

1

Andropogoneae o

86 96 1

1

99 1

76

1

100 1 100 1

Panicoideae e

98

99

1

99 1

Paspaleae e

1

91

99 1

0 0,99

72

88

97 1

85 1

1

76

100 1

100

100 1

1

99 1

82 1

92 1

87

73

0,99

93 1 100 1

83 1 98 1 72

96 1

79

0,99

9 0,96

100

91 0,99

85 1

1

98 1

Paniceae P

100

93 1

100 1

1

97 1 100 1 100 1

92 1

0,98

84 1

82 1

0,93

100 1

0,94

0 0,94

82 1

99 1

100 1

99 1

90

100 1

1

99 1 97 1

85 1

96 1

84 1

86

T hysanolaena maxima Zeugites pittieri Zea mays Cymbopogon flexuosus Phacelurus digitatus Streptostachys asperifolia Renvoizea saccharoidea Renvoizea trini Echinolaena inflexa Ocellochloa chapadense Anthaenantia lanata Hopia obtusa Paspalum conjugatum Rugoloa hyaleica Hymenache pernambucensis Hymenachne donacifolia Plagiantha tenella Otachyrium versicolor Steinchisma laxa Steinchisma hians Oplismenopsis najada Phanopyrum gymnocarpon Coleataenia prionitis Coleataenia tenera Cyphonanthus discrepans Homolepis glutinosa Mesosetum chaseae Altoparadidium chapadense Arthropogon villosus Dichanthelium acuminatum Adenochloa adenophora Adenochloa claytonii Echinochloa colona Parodiophyllochloa cordovensis Panicum andringitrense Acroceras zizanioides Panicum pygmaeum Panicum robynsii Oplismenus hirtellus Pseudechinolaena polystachya Lasiacis sorghoidea Panicum comorense1 Panicum comorense2 Panicum gardnerii Panicum trichanthum1 Panicum trichanthum2 Panicum trichanthum3 Panicum notatum Panicum monticola Panicum bartlettii Panicum millegrana Panicum sellowii Panicum trichoides Panicum laticomum Panicum pleianthum Panicum hirtum Panicum brevifolium Panicum glandulopaniculatum Panicum issongense Panicum capuronii Panicum subhystrix Panicum heterostachyum Panicum pusillum Kellochloa brachyantha Kellochloa verrucosa Sacciolepis angustissima Sacciolepis africana Panicum bisulcatum1 Panicum bisulcatum2 Sacciolepis myuros Sacciolepis indica Sacciolepis vilvoides Sacciolepis chevalieri Trichanthecium natalense Trichanthecium gracilicaule Trichanthecium margaritiferum Panicum aequinerve Panicum perrieri Panicum inaequilatum Panicum eickii Panicum chionachne Panicum calvum Panicum acrotrichum Trichanthecium dinklagei Trichanthecium brazzavillense Trichanthecium lindleyanum Trichanthecium nervosum Trichanthecium polycomun Trichanthecium cyanescens Trichanthecium wettsteinii Trichanthecium subulatum Trichanthecium parvifolium Trichanthecium schwackeanum Trichanthecium caaguazuense Trichanthecium auricomun Trichanthecium machrisianum Trichanthecium pseudisachne Trichanthecium rivale Trichanthecium granuliferum Trichanthecium micranthum Trichanthecium arctum Trichanthecium pyrularium Melinis repens Moorochloa eruciformis Urochloa plantaginea Eriochloa punctata Panicum trichocladum Megathyrsus maximus Zuloagaea bulbosa Setaria lachnea Setaria viridis Stenotaphrum secundatum Paspalidium geminatum Panicum antidotale Cenchrus ciliaris Cenchrus setaceus Whiteochloa cymbiformis Whiteochloa capillipes1 Whiteochloa capillipes2 Whiteochloa semitonsa Whiteochloa airoides Whiteochloa biciliata Whiteochloa multiciliata

1

EOCENE

40

OLIGOCENE

35

30

E. MIOCENE

25

20

M. MIOCENE

15

L. MIOCENE

10

PLIO

5

Fig. 1  Phylogeny of Panicum (reproduced from Zuloaga et al. 2018)

PLEI

0 MYA

Paspalinae

Otachyriinae

Arthropogoninae

Dichantheliinae

Boivinellinae

Incertae sedis genera

Melinidinae

Cenchrinae C

Panicinae P

31

Phylogeny and Evolution Thysanolaena_maxima Zeugites pittieri

A

Andropogoneae + Paspaleae Paniceae 1 Paniceae 2 Cenchrinae + Melinidinae

1

99 1

0,99

100 1

B

94 1

Louisiella el

Panicinae

90 1 78 1

87 1

85 1

90 9 0,99

Louisiella

0,95

0,97

86 1

74 1

76 1

Panicinae n

70 1 0,9

96 1

Panicum

90 1

Sect. Rudgeana

0,96 0,98

96 1

0,95

Panicum Sect. Hians 0,92 1

96 1

Sect. Panicum

99 1

1

Sect. Dichotomiflora

89 1 0.96

Sect. Repentia 94 1 70 1

Sect. Yakirra

74 1

Sect. Arthragrostis 71 0.99

25

20

10

15

5

EOCENE 40

35

83 1

0 MYA

OLIGOCENE 30

E. MIOCENE 25

20

Fig. 2  Phylogeny of Panicum (continued)

M. MIOCENE 15

PLIO

L. MIOCENE 10

5

Louisiella fluitans 1 Louisiella fluitans 2 Louisiella elephantipes 2 Louisiella elephantipes1 Panicum voeltzkowii 1 Panicum pinifolium Panicum cinctum Panicum voeltzkowii 2 Panicum queenslandicum Panicum rudgei Panicum cervicatum 2 Panicum cervicatum 1 Panicum ligulare Panicum urvilleanum Panicum curviflorum Panicum mystasypum Panicum olyroides Panicum paucinode Panicum fluviicola 1 Panicum afzelii Panicum grifonii Panicum pilgeri Panicum deciduum Panicum pansum 1 Panicum pansum 2 Panicum fluviicola 2 Panicum callosum Panicum nigerense Panicum phragmitoides 1 Panicum turgidum 2 Panicum turgidum 1 Panicum amarum 1 Panicum ruspolii Panicum virgatum Panicum anabaptistum Panicum phragmitoides 2 Panicum tricholaenoides Panicum chloroleucum Panicum racemosum 2 Panicum amarum 2 Panicum racemosum 1 Panicum nephelophilum Panicum fauriei Panicum carteri Panicum bergii Panicum stramineum Panicum alatum Panicum venosum Panicum miliaceum 2 Panicum miliaceum 1 Panicum parcum 2 Panicum parcum 1 Panicum capillaroides Panicum pampinosum Panicum hirticaule Panicum ghiesbregtii Panicum aquaticum 1 Panicum perangustatum Panicum schinzii Panicum impeditum Panicum psilopodium Panicum sumatrense Panicum subalbidum 1 Panicum subalbidum 2 Panicum aquaticum 2 Panicum paludosum Panicum dichotomiflorum Panicum laetum 3 Panicum laetum 2 Panicum laetum 1 Panicum buncei Panicum decompositum Panicum coloratum 1 Panicum latzii Panicum repens Panicum laevinode Panicum larcomianum Panicum coloratum 2 Yakirra australiensis 2 Yakirra australiensis 1 Yakirra muelleri Yakirra majuscula Panicum mitchelii Panicum pilgerianum Panicum seminudum Panicum chillagoanum Arthragrostis deschampsioides Arthragrostis aristipicula Arthragrostis brassiana Arthragrostis clarcksoniana Panicum robustum

PLEI 0 MYA

Rudgeana

Hiantes

Panicum

Dichotomiflora

Repentia

Yakirra

Arthragrostis

32

Phylogeny and Evolution

Regarding the species here placed as “incertae sedis,” seven of them, P. bartlettii, P. brevifolium, P. hirtum, P. millegrana, P. sellowii, P. trichanthum, and P. trichoides, are placed in the “grex” Sacciolepis (Zuloaga et al. 2018; Soreng et al. 2022), related in this clade to the genera Kellochloa Lizarazu, Nicola & Scataglini, Sacciolepis, and Trichanthecium Zuloaga & Morrone. Panicum haenkeanum has not yet been classified within the subtribes C4 of the Panicoideae, and finally, P. venezuelae is related to the genus Oedochloa C. Silva & R. P. P. Oliveira within subtribe Paspalinae Griseb.

Geographic Distribution and Habitat

Panicum species are found approximately from 50° north latitude to 40° south latitude, from southeastern Canada and the USA to central Chile and Argentina; they grow from sea level to, exceptionally, 3500 m asl (P. chloroleucum Griseb.). The species of section Dichotomiflora, mostly annuals (with the exception of P. aquaticum Poir.), are typical of open and humid environments, being common on the margins of lakes, swamps, or rivers, including aquatic or marsh plants that reach up to 2700 m asl. Within this section, P. dichotomiflorum is the species with the widest distribution (Fig. 3 in chapter “Taxonomic Treatment”), being found from southern USA, Mexico, Mesoamerica, the Caribbean, and being frequent throughout South America, from Colombia and Venezuela to central Argentina. Likewise, P. aquaticum also has a wide distribution (Fig. 1 in chapter “Taxonomic Treatment”), from Mesoamerica and the Caribbean, extending from Venezuela and the Guianas to Brazil and northeastern Argentina, where it is sporadic. In contrast, P. lacustre is endemic to Cuba (Fig. 5 in chapter “Taxonomic Treatment”), P. vaseyanum is restricted to Mexico (Fig. 5 in chapter “Taxonomic Treatment”), and P. sublaeve to Mesoamerica, having also been reported from Venezuela (Fig. 5 in chapter “Taxonomic Treatment”). Regarding species of section Hiantes, annuals or perennials, these are found in dry and open environments, forming populations in coastal dunes or are found as grasslands in humid places next to watercourses. A group of three species is distributed mainly from the USA through Mexico, Mesoamerica, and the Caribbean: P. virgatum L. in grasslands and forest margins from Canada to the Caribbean and Nicaragua (Fig. 21 in chapter “Taxonomic Treatment”), P. amarum Elliott along beaches and coastal dunes from the USA to Belize and the Caribbean (Fig. 7 in chapter “Taxonomic Treatment”), and P. altum Hitchc. & Chase in coastal dunes from Mesoamerica and the Caribbean to northern South America. Panicum chloroleucum Griseb., a species endemic to northwestern Argentina (Fig. 7 in chapter “Taxonomic Treatment”); P. racemosum (P. Beauv.) Spreng, a coastal dune species present on the

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Geographic Distribution and Habitat

Atlantic coast from northern Brazil to Argentina and disjunct on the coasts of Chile (Fig. 18 in chapter “Taxonomic Treatment”); and P. urvilleanum Kunth, a species with a disjunct distribution in the southern USA and in Chile and Argentina (Fig. 21 in chapter “Taxonomic Treatment”), are three species with unicellular, filiform macrohairs at the base of the upper lemma, useful as dune fixers. Then, three species with bambusoid canes, typical of sandy soils and humid places, are P. deciduum Swallen, endemic to Venezuela (Fig. 10 in chapter “Taxonomic Treatment”); P. tricholaenoides Steud., distributed from Colombia and Venezuela to southern Brazil, Paraguay, and Argentina (Fig. 18 in chapter “Taxonomic Treatment”); and P. glabripes Döll, a species restricted to southern Brazil, Uruguay, and Argentina (Fig. 10 in chapter “Taxonomic Treatment”). P. longissimum (Mez) Henrard is endemic to Paraguay (Fig. 10 in chapter “Taxonomic Treatment”). Finally, there is a group of three species, all of which have in common flattened macrohairs at the base of the upper anthecium and inhabit savannas or dry fields of South America; of these, P. complanatum Guglieri, Longhi-Wagner & Zuloaga and P. mystasipum Zuloaga & Morrone are exclusive to “cerrados” and “campos rupestres” of central Brazil (Figs. 10 and 15 in chapter “Taxonomic Treatment”), while P. olyroides Kunth grows from Venezuela to Argentina (Fig. 15 in chapter “Taxonomic Treatment”). All species of section Panicum usually inhabit fields or savannas, on clayey, sandy to rocky soils, on roadsides, and also on mountain slopes; some species in Central America are found in deciduous forests. An exception is P. aquarum, a species of low-flooded fields in northern South America. Species of this section grow from sea level to about 1300 m asl, with P. lepidulum up to 2150 m asl in Mexico and P. hirticaule up to 2500 m asl in Peru. Among the more widely distributed taxa of section Panicum are Panicum stramineum, from southwestern USA and the Caribbean to Argentina (Fig. 32 in chapter “Taxonomic Treatment”); P. alatum var. minus from the southwestern USA to Venezuela and Ecuador, in the Galapagos Islands (Fig. 29 in chapter “Taxonomic Treatment”); P. hirticaule var. hirticaule (Fig. 39 in chapter “Taxonomic Treatment”) from southwestern USA to Venezuela, Ecuador, and Peru; and P. ghiesbreghtii from southwestern USA, Mexico, Caribbean Islands to Venezuela, Colombia, and Ecuador (Fig. 25 in chapter “Taxonomic Treatment”). Panicum hirsutum ranges from the southwestern USA, Mexico, and the Caribbean to Ecuador, Colombia, Venezuela, and northern Brazil, having been occasionally collected in northeastern Argentina (Fig. 38 in chapter “Taxonomic Treatment”). Finally, P. hispidifolium grows from southern Mexico to Colombia and Venezuela (Fig. 34 in chapter “Taxonomic Treatment”). Panicum capillare has been introduced in South America, in Brazil, Uruguay, Argentina, and Chile (Fig. 29 in chapter “Taxonomic Treatment”). Panicum alatum var. alatum, P. capillarioides, P. hallii subsp. hallii, P. hallii subsp. filipes, and P. hirticaule var. verrucosum grow from southern and southwestern USA to northeastern and central Mexico. Finally, P. aztecanum, P. decolorans, and P. tamaulipense are endemic to Mexico (Figs. 25, 27, and 32 in chapter “Taxonomic Treatment”). Panicum lepidulum and P. parcum are exclusive to Mexico and Mesoamerica (Figs. 34 and 36 in chapter “Taxonomic Treatment”), and P. furvum and P. diffusum are endemic to Guatemala and the Caribbean Islands, respectively (Fig. 32 in chapter “Taxonomic Treatment”).

Geographic Distribution and Habitat

35

Nine species are exclusive to South America. Among these, the most widely distributed are P. exiguum and P. bergii var. bergii (Figs. 25 and 32 in chapter “Taxonomic Treatment”), the latter species having been introduced in the USA; P. aquarum is found in Venezuela, Colombia, and Brazil (Fig. 25 in chapter “Taxonomic Treatment”); P. peladoense in southern Brazil, Bolivia, Paraguay, and Argentina (Fig. 38 in chapter “Taxonomic Treatment”); P. quadriglume in Peru, Bolivia, Paraguay, Brazil, and Argentina (Fig. 34 in chapter “Taxonomic Treatment”), P. bergii var. pilosissimum in northeastern Argentina and southern Brazil (Fig. 34 in chapter “Taxonomic Treatment”); P. chaseae in eastern Argentina, southern Brazil, and Uruguay (Fig. 32 in chapter “Taxonomic Treatment”); and P. mucronulatum, P. ephemeroides, and P. magnispicula are endemic to Brazil (Figs. 34 and 38 in chapter “Taxonomic Treatment”). Species of section Repentia are usually found on sandy soils (P. repens L. has been introduced as a dune-fixing species), dry or in coastal dunes, or low fields along river and stream banks. Of the two native species in the section, P. gouinii E. Fourn. has a disjunct distribution, being found in the USA and Mexico and in South America in southern Brazil, Paraguay, Uruguay to Argentina (Fig. 50 in chapter “Taxonomic Treatment”), while P. pedersenii Zuloaga grows in southern Brazil, Paraguay, and northeastern Argentina (Fig. 50 in chapter “Taxonomic Treatment”). The species of section Rudgeana, exclusively American, inhabit open and sunny environments, on sandy or clayey soils, in savannas, fields or enclosed areas, from sea level to 1500 m asl. Panicum cayennense Lam., an annual species, and P. rudgei Roem. & Schult. are the most widely distributed, occurring in Mesoamerica, the Caribbean, and South America from Colombia and Venezuela to Bolivia and Brazil (Figs. 53 and 56 in chapter “Taxonomic Treatment”); P. campestre Nees ex Trin. and P. ligulare Nees ex Trin. are restricted to cerrados of Brazil (Figs. 53 and 56 in chapter “Taxonomic Treatment”), while P. cervicatum Chase has a disjunct distribution, growing in savannas of Venezuela and cerrados of Bolivia and Brazil (Fig. 56 in chapter “Taxonomic Treatment”). The C3 species of Panicum commonly grow in the interior or margins of tropical and subtropical forests and jungles and have in general a wide distribution, ranging from Mexico and Mesoamerica, the Caribbean Islands and South America, from Colombia and Venezuela to Brazil or northern Argentina. This group includes P. bartlettii (Fig. 59 in chapter “Taxonomic Treatment”), P. hirtum (Fig. 61 in chapter “Taxonomic Treatment”), P. millegrana (Fig. 64 in chapter “Taxonomic Treatment”), P. sellowii (Fig. 66 in chapter “Taxonomic Treatment”), P. trichanthum (Fig. 66 in chapter “Taxonomic Treatment”), and P. trichoides (Fig. 69 in chapter “Taxonomic Treatment”). On the other hand, Panicum haenkeanum and P. venezuelae are distributed from Mesoamerica and the Caribbean to northern South America, P. venezuelae in Venezuela and northern Brazil (Fig. 71 in chapter “Taxonomic Treatment”), while P haenkeanum reaches Bolivia and Brazil (Fig. 61 in chapter “Taxonomic Treatment”).

Taxonomic Treatment

Panicum L., Sp. Pl. 1: 55. 1753. Type: Panicum miliaceum L. Eatonia Raf., J. Phys. Chim. Hist. Nat. Arts 89: 104. 1819. Type: Eatonia purpurascens Raf. Monachne P. Beauv., Ess. Agrostogr. 49. 1812. Type: Monachne racemosa P. Beauv. Thalasium Spreng, Syst. Veg. 4 (2): 22, 30 (“Thalasium”). 1827. Type: Thalasium montevidense Spreng. Eriolytrum Kunth, Revis. Gramin. 1: 219. 1829, nom. inval. as a synonym of Panicum reptans Kunth. Panicum subgen. Eupanicum Pilg., Oesterr. Bot. Z. 51: 369. 1901, nom. inval. Chasea Nieuwl., Amer. Midl. Naturalist 2: 64. 1911, nom. nud. Plants annual or perennial, usually cespitose, short to long rhizomatous, with erect to, less frequently, decumbent, rooting, branching culms at lower, middle, and upper nodes; internodes hollow, occasionally solid, with or without aerenchyma, hairy to glabrous; nodes pale to brownish, hairy to glabrous. Sheaths open, striate, pilose to glabrous, with or without aerenchyma. Ligule membranous-ciliate to ciliate. Blades lanceolate to linear-lanceolate, flat to involucrate, base rounded to cordate, apex acute to acuminate, hairy to glabrous. Inflorescence an open to contracted terminal panicle, exerted or partially included in the upper leaves; axillary inflorescences occasionally present. Spikelets on second- or third-order branches, solitary or in pairs, biflowered, exceptionally triflowered (P. quadriglume), hairy to glabrous, with conspicuous rachilla or not between bracts; lower glume ¼ to 4/5 as long as spikelet, (0–)3–9 nerved, membranous, hairy to glabrous, truncate or with the apex acute; upper glume and lower lemma usually subequal, upper glume (5–)7–11(–13) nerved, membranous, lower lemma (5–)7–11 nerved, membranous; lower palea similar in length to lower lemma or reduced, 2 nerved, hyaline; lower flower staminate or absent; upper anthecium stipitate or not, hardened, smooth, pale to blackish, shiny, with simple, verrucose or compound papillae over the entire

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_9

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Taxonomic Treatment

surface or, more frequently, toward the apex of the lemma and palea. Caryopsis with punctiform to oblong hilum, embryo 1/3 to ½ the length of the caryopsis. Pantropical genus with approximately 165 species grouped in seven sections, one of them endemic to Australia, another one present in Australia and Southeast Asia, one restricted to the Neotropics, and the remaining four pantropical.

Key to Ungrouped Panicum Sections and Species 1. Spikelets with upper glume and lower lemma 7–9(–15) nerved, upper anthecium smooth, shiny, with simple or compound papillae toward apex of palea; plants cespitose, with or without elongate rhizomes, exceptionally with decumbent culms; blades linear-lanceolate to lanceolate, flat or with margins involuting. Plants usually of open, dry places 2. Plants with a heterogeneous stipe at the base of the upper anthecium............................................................................. Sect. Rudgeana 2. Plants without a stipe at the base of the upper anthecium, if present homogeneous 3. Annual or perennial plants, shortly rhizomatous when perennials, without elongated rhizomes 4. Lower glume 1/3 or less the spikelet length, with a truncate, obtuse, or subacute apex. Aquatic or marshy plants, usually occurring in open, moist environments.................................. Sect. Dichotomiflora 4. Lower glume ½ or more the length of the spikelet, the apex acute. Plants of open, dry places, growing in fields, savannas or enclosed ................. ..................................................................... Sect. Panicum 3. Perennial plants, with manifest, robust, short to elongated rhizomes 5. Lower glume ¼ to 1/3, exceptionally ½ of the length of the spikelet; spikelets not open at maturity........................................Sect. Repentia 5. Lower glu ½ to 4/5 of the length of the spikelet; spikelets open at maturity……...................................................................Sect. Hiantes 1. Spikelets with the upper glume and lower lemma 3–5 nerved; upper anthecium transversely rugose to smooth, with simple papillae all over; blades ovate-lanceolate to lanceolate, flat; plants decumbent, rooting and branching at the lower nodes, of moist, shady places, frequent in forest margins and jungles 6. Upper anthecium transversely rugose 7. Plants annual, stoloniferous; spikelets 0.9–1.4 mm long, not open at maturity................................................................................62. P. trichoides 7. Plants perennial, non-stoloniferous; spikelets 1.6–2.5  mm long, open at maturity

Key to Ungrouped Panicum Sections and Species

39

8. Panicle contracted, few-flowered, up up to 11 cm wide, branchess naked toward the base, spikelets congested toward the apex; blades lanceolate, 6–13 × 0.7–1.4 cm .......................................60. P. sellowii 8. Panicle lax, multiflowered, up to 30 cm wide; spikelets scattered along the branches, not congested; blades ovate-lanceolate, 6–14 × 1–3.8 cm. ......................................................................59. P. millegrana 6. Upper anthecium smooth, not transversely rugose. 9. Axillary inflorescences present, spikelets arranged unilaterally on the branches; panicles with cleistogamous and chasmogamous spikelets, usually with glands present on the lower lemma............................................... ...........................63. P. venezuelae 9. Axillary inflorescences absent, spikelets not unilaterally arranged; cleistogamous spikelets absent, without glands on the lower lemma 10. Lower glume ¼ to ½ the length of the spikelet; plants perennial 11. Spikelets 2.3–2.6 mm long; upper glume and lower lemma without vesicles on their surface................57. P. haenkeanum 11. Spikelets 1–1.9 mm long; upper glume and lower lemma with minute vesicles on their surface 12. Lower glume ½ or slightly less than spikelet length, 1–3 nerved, acute; lower palea absent…......... 55. P. bartlettii 12. Lower glume ¼ or less than spikelet length, nerveless, obtuse to truncate; lower palea present............................. ......................... ....................................61. P. trichanthum 10. Lower glume 4/5 or the equal length of the spikelet; plants annual 13. Inflorescences with conspicuous glands on primary and secondary axes; spikelets obliquely arranged on the pedicels; upper anthecium stipitate.....................................58. P. hirtum 13. Inflorescences without glands on the primary and secondary axes; spikelets not oblique; upper anthecium not stipitate.........................................................56. P. brevifolium Panicum Sect. Dichotomiflora (Hitchc. & Chase) Hitchc. & Chase ex Honda, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 9: 244, 246. 1930. Panicum (unranked) Dichotomiflora Hitchc., N.  Amer. Fl. 3(2): 200, 202. 1915. Panicum group Dichotomiflora Hitchc. & Chase, Contr. U.S. Natl. Herb. 15: 28, 47. 1910, nom. inval. Type: Panicum dichotomiflorum Michx. Annual plants, less frequently shortly rhizomatous perennials, with erect to decumbent culms, branched and rooting at the lower nodes; internodes hollow, with or without aerenchyma. Ligule membranous-ciliate. Blades lanceolate to linearlanceolate or linear, flat, hairy to glabrous. Inflorescence a terminal panicle, open, diffuse to contracted, exserted to shortly exserted; axillary inflorescences present or absent. Spikelets narrow to long-ellipsoid or long-ovoid, glabrous, pale to greenish to purplish tinged, not open at maturity, solitary or in pairs; lower glume 1/5 to 1/3,

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Taxonomic Treatment

or slightly more the length of the spikelet, 1–3(–7) nerved, the apex ovate to obtuse or truncate; upper glume and lower lemma subequal, upper glume 7–9(–13) nerved, membranous, lower lemma 7–9(–13) nerved; lower palea present or absent; lower flower staminate or absent; upper anthecium ellipsoid to broadly ovoid, glabrous, not stipitate, smooth, shiny, hardened, whitish to pale and dark brown at maturity, early drooping (P. vaseyanum). Caryopsis ellipsoid; hilum ovate to oblong, embryo 1/3 to ½ or slightly more than the length of the caryopsis. Section with approximately 17 species worldwide (Zuloaga et  al. 2018), of which 5 grow in the Neotropics, distributed from the southern USA, Mexico, Mesoamerica, and the Caribbean to Uruguay and Argentina. It includes aquatic or marsh plants, usually present in open and humid environments. Key to the Species 1. Perennial plants, with succulent culms, forming reservoirs in watercourses 2. Spikelets 2 mm long; upper glume and lower lemma 7 nerved; lower palea and lower floret absent..............................................................3. P. lacustre 2. Spikelets 2.7–3.5 mm long; upper glume and lower lemma, 9 nerved; palea and lower flower present.......................................................1. P. aquaticum 1. Annual plants, with non-succulent culms, growing in moist soils, not in watercourses 3. Spikelets 3.5–4 mm long; lower glume 5–7 nerved, 1/3 or more the length of the spikelet; upper glume and lower lemma 11–13 ner ved......................................................................................4. P. sublaeve 3. Spikelets 2–3.2 mm long; lower glume 1–3 nerved, ¼ to 1/3 of spikelet length; upper glume and lower lemma 7–9 nerved 4. Panicles contracted, up to 1.5  cm wide; upper anthecium early caducous..........................................................................5. P. vaseyanum 4. Panicles open, lax, up to 20  cm wide; upper anthecium not early caducous.... ............................................................2. P. dichotomiflorum 1. Panicum aquaticum Poir. Plants perennial, rhizomatous, culms decumbent and rooting at lower nodes, when in water with elongated internodes with numerous adventitious roots, then erect, 80–200 cm tall; internodes cylindrical, glabrous, hollow; nodes brown, glabrous. Sheaths striate, 8–15  cm long, usually larger than the internodes, densely papillose-pilose over the entire surface or densely hispid toward the upper portion, or glabrous, the margins membranous, upper ones with long whitish hairs or without. Ligule membranous-ciliate, 1–4 mm long, membranaceous portion 0.4–0.8 mm long, collar pale to brownish, pilose or glabrous. Blades linear, 9–25 × 0.3–0.6 cm, flat, ascending, rounded at the base, the apex acuminate, the adaxial surface papillose-pilose, more densely pilose toward the base, abaxial surface sparsely pilose, glabrous toward the distal portion, or glabrous on both faces, the margins glabrous, the basal ones ciliate to glabrous, midrib manifest. Inflorescence terminal exerted,

Key to Ungrouped Panicum Sections and Species

41

peduncle cylindrical, glabrous; panicles lax, 15–26(–30)  ×  8–15  cm; main axis cylindrical, glabrous, pulvini brownish, glabrous, first-order branching alternate, the upper ones verticillate or alternate, divergent, branching axes and pedicels triquetrous, scabrous, spikelets in pairs and congested on second-order branches, pedicels 2–4 mm long. Spikelets long ovoid, 2.7–3.5 × 1 mm, greenish or tinged with purple, glabrous, upper glume and lower lemma subequal, acute, exceeding the upper anthecium in length; lower glume acute to obtuse, 0.8–1.4 mm long, ca. 1/3 of spikelet length, 1–3-nerved, the nerves anastomosed; upper glume and lower lemma 9 nerved; lower palea elliptic, 2.4 × 0.8 mm, glabrous, hyaline; lower flower staminate, stamens 3, anthers 1.2–1.4  mm long; upper anthecium long ovoid, 1.9–2.2 × 0.8 mm, whitish, smooth, shiny, glabrous, hardened. Caryopsis ellipsoid, 1.3 × 0.6 mm, whitish; hilum ovate, embryo 1/3 as long as the caryopsis This species is morphologically related to P. dichotomiflorum, the latter being an annual plant, with spikelets 2–3 mm long and lower flower absent. Key to the Varieties 1. Sheaths densely hispid; panicles less than 25 cm long........................................... .....1a. P. aquaticum var. aquaticum 1. Sheaths densely papillose, panicles larger, ca. 30 cm long..................................... ...........1b. P. aquaticum var. cartagoense 1a. Panicum aquaticum Poir. var. aquaticum, Encycl., Suppl. 4: 281. 1816. Type: Puerto Rico, without locality, 1797, M. Ledru s.n. (lectotype, P 00371641, designated as “type” by Hitchcock & Chase, 1910: 48; isolectotypes, FI n.v., US 00148147) Panicum chloroticum var. sylvestre Nees ex Trin. Gram. Pan.: 236. 1826. Type: Brazil. Bahia. Almada, L.  Riedel s.n. (holotype, LE-TRIN-0633.06; isotype, US-974404) Panicum chloroticum Nees ex Trin. var. sylvestre Nees, Fl. Bras. Enum. Pl. 2(1): 164. 1829, nom. illeg. hom., non Trin. 1826. Type: Brazil. “Habitat in sylvis ad Almada, Ferradas et in via Felisbertia districtus Insulanorum prov. Bahiensis; (Martius et Maximil. Princ. Neovid.)”, C. F. P. von Martius s.n. (lectotype M, designated, as “type,” by Hitchcock & Chase, 1910: 49; isolectotype, US 00148356) Panicum hygrophilum Salzm. ex Steud., Syn. Pl. Glumac. 1: 71. 1855]1853]. Type: Brazil. Bahia: without locality, 1827, P. Salzmann s.n. (lectotype designated by Zuloaga, Darwiniana n.s. 10(1): 330. 2022, P 00740831; isolectotypes, G 00099659, G 00099660, GH, HAL 0133162, K 000003808, K 000003809, MO-105084, MPU 024441, MPU 024442, MPU 024443, P 00740830, P 00740831, P 00740832, W 18890239825, W 19160024808) Panicum proliferum var. strictum Griseb., Cat. Pl. Cub.: 232. 1866. Type: Cuba. Without locality, 1860-1864, C.  Wright 3456 (holotype unknown; isotypes, G 00099640, G 00099641, GH 00135362, K 000309158, K 000309159, P 00740938, MO-321061, NY 01767173, S04-321, US 00139898)

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Taxonomic Treatment

Distribution and ecology (Fig. 1). Variety distributed from Mexico and the Caribbean to Argentina; grows in margins of lakes and rivers, from sea level to 1300 m above sea level (asl); it flowers and fruits between August and April Additional material examined. Mexico. Quintana Roo: en brecha a Vallarta, a 5 kms al oeste de Puerto Morelos, 31 August 1980, Sousa 11207 (MEXU). Veracruz: a 7  km de Minatitlán rumbo a Coatzacoalcos, 21 January 1971, Lot 1227 (MEXU) Belize. Gracie Rock, Sibun River, 21 April 1935, Gentle 1635 (MO, USA); 9 mi west of Belize on the Western highway to Belmopan, 50 m, 21 March 1987, Davidse & Brant 32947 (MO, NY) Guatemala. Puerto Barrios, Hitchcock 9147 (SI, USA). Jutiapa: vicinity of Jutiapa, 850 m, 24 October 1940, Standley 76308 (MO) Honduras. El Paraiso, 32 km west of Danli along highway 4, 700 m, 21 July 1970, Pohl & Davidse 12386 (F, MO). Francisco Morazán: región of Las Mesas, in water, edge of stream, 16 October 1951, Swallen 10803 (MO, USA); drainage of the Río Yeguare, 24 June 1948, Glassman 1746 (MO) Nicaragua. Zelaya: Puerto Isabel, scrubby growth near seabeach, 3 January 1970, Seymour 2895 (GH, MO)

Fig. 1  Distribution of Panicum aquaticum

Key to Ungrouped Panicum Sections and Species

43

Costa Rica. Guanacaste: road to Hacienda Inocentes, 12 km from CIA, 330 m, 27 July 1971, Pohl 12651 (F, MO); 2 km E of Carretera Interamericana, on road to Las Animas, 200 m, Pohl et Davidse 11547 (F, US); Laguna on road to Cuajiniquil, ca. 200 m from intersection with Interamerican Highway, Ruta 1, 300 m, 14 October 1989, Crow & Rivera 7356 (F, MO) Cuba. Habana: Puentes Grandes, Ekman 502 (G). Pinar del Río: 13 km south of Pinar del Río, 25 November 1926, Hitchcock 23290 (US). Santa Clara: Sabana de Motembo, León 9470 (NY) Puerto Rico. Floresta del rio Piedras, vicinity of San Juan, northeast of Río Piedras, Chase 6778 (RB, US) Guadalupe. Environs de la Basse Terre, Duss 3178 (NY) Venezuela. Apure: without locality, Ramia 7195 (VEN) Trinidad and Tobago. Pitch Lake, 7 December 1912, Hitchcock 10099 (G, MO, NY, US) Suriname. Near Dohsen savanna, Krukoff 130 (NY), without locality, Weigelt 936 (G, MO) Peru. Without locality, Dombey s.n. (P), Gaudichaud s.n. (P) Brazil. Bahia: 10 km east of Seabra, along Highway BR-242, 770 m, 6 April 1976, Davidse et al. 12034 (MO); near Salvador, low moist ground near tidal marsh, 23 December 1924, Chase 8022 (F, MO, NY, US); Salvador, Chase 7895 (F, NY, US), 8036 (F, USA); estrada Olivenca-Maruim, entre los kms 7-10, restinga arbórea, 50 m, 19 May 1985, Zuloaga et al. 2479 (MO); Belmonte, road to Itapebi, Harley et  al. 17297 (CEPEC, MO); 5  km  S of Sta. Cruz Cabralia, Harley et  al. 17155 (CEPEC, MO, NY, RB); Cachoeira, 10 km north of Cachoeira, 29 December 1924, Chase 8089 (F); Serra de Jacobina, 8  km  N of Senhor do Bonfim, 600  m, 27 February 1974, Harley et  al. 16504 (MO, RB). Mato Grosso do Sul: between Campo Grande and Dourados, 400–450 m, 14 February 1930, Chase 10928 (US); Tres Lagoas, 300 m, 4 February 1930, Chase 10770 (RB, US). Minas Gerais: Bem Querer, 900  m, 23 March 1980, Hatschbach 42905 (MO). Pará: Fazenda Camburupy, near Soure, Ilha do Marajó, Swallen 4915 (RB); Ilha do Marajó, Fazenda Menino Deus, Black & Engelhand 50- 9040 (IAN, US); suburbs of the city of Belem, June 1914, Goeldi 51 (MO); Marajó Island, Estate Gavinho, Goeldi 235 (NY, RB, US), 248 (US); Marajó, Jobert 120 (P). Rio Grande do Sul: Porto Alegre, Canoas, Malme 1460 (US). Rio de Janeiro: Ararauma, 10 December 1929, Chase 10177 (US); Petrópolis, Caetitu, Goes et Dionisio 846 (RB); Leblon, near Lago de Freitas, 13 January 1925, Chase 8225 (MO), 9 May 1925, Chase 9756 (F, MO); Carangola, Goes et Dionisio 590 (RB); San Pedro, northwest of Cabo Frío, 9 December 1929, Chase 10130 (US); a margem da estrada de Plazolandia, Monteiro s.n. (RB -68470, US); Serra do Medanha, Sucre et  al. 8854 (RB, SI). Santa Catarina: Rio Oratório, Lauro Muller, Reitz & Klein 8492 (US); Rio Tavares, Ilha de Santa Catarina, Reitz & Klein 287 (G, NY). São Paulo: Alto da Serra, Hoehne 9912 (US); Campinas, Fazenda Campo Grande, Viegas s.n. (US); Santos, Parodi 12209 (USA). Rio Tavares, Reitz & Klein 287 (NY); Serta Ozinho, Viegas s.n. (IAN) Argentina. Corrientes: 3 km E de Ituzaingó, ruta 12, 7 April 1978, Ahumada 2143 (MO)

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Taxonomic Treatment

1b. Panicum aquaticum Poir. var. cartagoense Davidse, Novon 2: 103. 1992. Type: Costa Rica. Cartago: 0.5 km E of Planta Radiográfica along railroad, 2 km W of Paraíso, 1300 m, 6 February 1969, plants 1–2 meters tall, rooted in a dense tangle of herbage, R. W. Pohl & G. Davidse 11700 (holotype, ISC-v-0000567; isotypes, F, MO-183078) Distribution and ecology (Fig. 1). Variety restricted to Costa Rican rainforests It can be distinguished from the type variety by the characteristics indicated in the key. 2. Panicum dichotomiflorum Michx. Plants annual, 0.10–1.40 m tall, cespitose, herbaceous; culms simple to branched, decumbent, rooting and densely branching or not at lower nodes to erect, not succulent; internodes cylindrical, glabrous, hollow, 3–16  cm long; nodes glabrous, brownish, compressed, 0.5–3 mm. Sheaths open, striate, 4–15 cm long, shiny, pale or tinged with purple, glabrous to densely hirsute, the margins membranous. Ligule 2 mm long, membranous-ciliate; collar brown, glabrous. Blades oblong-lanceolate to lanceolate, 8–45 × 0.3–1.4 cm, flat, subcordate at the base, the apex attenuate, herbaceous, glabrous, pale green, the margins scaberulous. Inflorescence a lax, diffuse panicle, 4–30 × 2–20 cm, with opposite or alternate, less frequently verticillate, divergent, and distant second-order branches, spikelets solitary or paired on short third-order branches, those adpressed to second-order branches; main axis scaberulous, pulvini glabrous; pedicels triquetrous, scabrous, 2–7 mm long; axillary panicles usually present, similar to the terminal one. Spikelets long ovoid, (2–)2.4–3  ×  0.9–1.1  mm, glabrous, acuminate, pale or tinged with purple, upper glume and lower lemma subequal and exceeding the upper anthecium in length; lower glume ovate, 0.7–1.2 mm long, 1/4 to 1/3 as long as spikelet, glabrous, 1–3 nerved, the apex truncate to subacute; upper glume 2.3–3.3  mm long, glabrous, acuminate, 9 nerved; lower lemma 2.2–3.3 mm long, glumiform, 7–9 nerved; lower palea elliptic, 2.1–2.5 × 0.5–0.7 mm, hyaline, or absent, lower flower absent; upper anthecium ellipsoid, 2–2.5 × 0.7–1.1 mm, pale, dark brown at maturity, glabrous; upper lemma 7 nerved. Caryopsis ellipsoid, 1.5–1.6  ×  0.9–1  mm, brown; hilum oblong, embryo half as long as the caryopsis Key of the Subspecies 1. Sheaths glabrous; spikelets 2.4–3 mm long............... .........................................2 a. P. dichotomiflorum subsp. dichotomiflorum 1. Sheaths densely hirsute; spikelets 2.2–2.4 mm long..................... 2b. P. dichotomiflorum subsp. bartowense 2a. Panicum dichotomiflorum Michx. subsp. dichotomiflorum, Fl. Bor.-Amer. 1: 48. 1803. Type: USA. “Hab. in occidentalibus montium Alleghanis,” A. Michaux s.n. (holotype, P-MICHX; isotype, P 00740942). Fig. 2 Panicum miliaceum Walter, Fl. Carol.: 72. 1788, nom. illeg. hom., non L., 1753. Type: USA. South Carolina (holotype unknown)

Key to Ungrouped Panicum Sections and Species

45

Fig. 2  Panicum dichotomiflorum. (a) Habit. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Lower palea. (e) Upper anthecium, dorsal view. (f) Upper anthecium, ventral view. (From Ahumada 610, CTES)

46

 

Taxonomic Treatment

Panicum geniculatum Muhl, Cat. Pl. Amer. Sept.: 9. 1813. Type: USA, Muhlenberg Hb. Fol. 181 (holotype, PH 00047147; isotype, US 00148518, probable isotype, BM 001042409) Panicum multiflorum Poir., in Lamarck, Encycl. Suppl. 4: 282. 1816. Type: USA: “cette plante croit a la Caroline; elle m’a ete ete communiquee par M. Bosc.,” L. A. G. Bosc s.n. (lectotype, P 00740941, designated as “type,” by Hitchcock & Chase, 1910: 48; isolectotypes, US 00139776, US 00139777) Panicum brachiatum Bosc ex Spreng. Syst. Veg. [Sprengel] 1: 321. 1825, nom. illeg. hom., non Poiret, 1816. Type: Bermuda. “Ins. Bermud” (holotype unknown; isotype, US 00148189, fragment ex Willd. herb.) Panicum chloroticum Nees ex Trin, Gram. Panic.: 236. 1826. P. chloroticum var. agreste Nees ex Trin., Gram. Panic.: 236. 1826. Panicum proliferum Lam. var. chloroticum (Nees ex Trin.) Hack., Repert Spec. Nov. Regni Veg. 6(21-26): 343. 1909. P. aquaticum var. chloroticum (Nees) R.C. Foster, Rhodora 68: 320. 1966. Type: Brazil. “V. utriusque spp. Brazil (N. ab Esenb. Langsdorff)”, G. H. von Langsdorff s.n. (holotype, LE-633.1) Panicum chloroticum Nees ex Trin. var. agreste Nees, Fl. Bras. Enum. Pl. 2(1): 164. 1829. Type: Brazil: “Habitat in graminosis cultis prope Soteropolin et Oeiras provinciae Bahiensis et Piauhiensis”, C.  F. P. von Martius s.n. (lectotype, M, designated as “type”, by Hitchcock & Chase, 1910: 49; isolectotype, US 00148354) Panicum chloroticum Nees ex Trin. var. pingue Nees, Fl. Bras. Enum. Pl. 2(1): 164. 1829. Type: Brazil. “Habitat in cultis ad Soteropolin. Provinciae Bahiensis,” C. F. P. von Martius s.n. (lectotype, M, designated as “type,” by Hitchcock & Chase, 1910: 49; isolectotype, US 00148355) Panicum retrofractum Delile ex Desv., Mém. Soc. Agric. Angers 1: 200. 1831. Type: USA, “America borealis.”, collector unknown s.n. (lectotype, P 00661391, designated as “type,” by Hitchcock & Chase, 1910: 49; isolectotypes, LINN-HS 118-84, US 00139933) Panicum chloroticum Nees ex Trin. var. luxurians Döll, in C. Martius, Fl. Bras. 2 (2): 198. 1877. Type: holotype unknown Panicum proliferum Lam. var. richardii Döll, in C.  Martius, Fl. Bras. 2(2): 200. 1877. Type: Brazil. Pará: “prope urbem Para, in littore arenoso (L. C. C. Richard), L. C. Richard s.n. (holotype unknown; isotype, W 19040013871) Panicum amplectens Chapm., Bot. Gaz. 3(3): 20. 1878. Type: USA. “South Florida,” J. L. Blodgett s.n. (holotype unknown) Panicum francavillanum E.  Fourn., Mexic. Pl. 2: 25. 1886. P. francavillanum E. Fourn. ex Hemsl., Biol. Cent.-Amer. Bot. 3: 489. 1885. nom. nud. Type: Mexico. Tabucaya, J. W. Schaffner 301 (lectotype, P 00740939, designated as “type,” by Hitchcock & Chase, 1910: 50; isolectotype, W 0021637) Panicum proliferum Lam. var. geniculatum Alph. Wood, Amer. Bot. Fl. 2: 392. 1871. Panicum dichotomiflorum Michx. var. geniculatum (Alph. Wood) Fernald, Rhodora 38(455): 387, pl. 441, Fig. 2. 1936. Type: USA (holotype unknown) Panicum dichotomiflorum Michx. var. imperiorum Fernald, Rhodora 44(526): 380. 1942. Type: USA. Virginia: Greensville County, Fontaine Creek, southwest of

Key to Ungrouped Panicum Sections and Species

47

Haley’s Bridge, 14 October 1941, Fernald & Long 13877 (holotype, GH 00023012; isotypes, GH 00023013, NY 00413977, PH 00018649) Panicum dichotomiflorum Michx. var. purinatorum Svenson, Rhodora 22(261): 154, Figs. 1–5. 1920. P. dichotomiflorum Michx. subsp. purinatorum (Svenson) Freckmann & Lelong, Sida 20(1): 171. 2002. Type: USA. Maine: Barnstable, 13 September 1919, M. L. Fernald 306 (holotype, GH 0002404098; isotypes, BR 0000008408792, NEBC 00028645, NY 00413974, P 00740940, PH 00018647, PH 00018648, US 00147806) Distribution and ecology (Fig. 3). Subspecies widely distributed in temperatewarm America, from the USA to Argentina; common in open and humid places, being found on riverbanks and streams and in low places; also found as a weed in cornfields and other crops; flowers and fruits throughout the year. Local names. “Fall panicum,” “smooth witchgrass” (Hafliger and Scholz 1980), “blue grass” (Smith et al. 1982), “kapi’í setaria” (“kapi’í = straw”) (Zuloaga 1994), “gramalote de agua” (Colombia, Zuloaga and Giraldo Cañas 2013) Panicum chloroticum was described by Trinius in 1826 on the basis of a collection of Langsdorff in Brazil. In 1829, Nees again described the species with three

Fig. 3  Distribution of Panicum dichotomiflorum

48

 

Taxonomic Treatment

varieties, citing different specimens collected by Martius. Hitchcock and Chase (1910) indicate that they did not find, in the Trinius herbarium in LE, any specimens of this species and consider that the specimens in the Munich herbarium could constitute the types of the described varieties. Several authors have considered P. chloroticum as a good species, taking into account the absence of the lower palea in this species and the presence, in contrast, of the same in P. dichotomiflorum. The analysis of the type material and abundant material of the species has allowed us to verify that there is a great variation in the presence or absence of lower palea, reason for which P. chloroticum has been included in the synonymy of P. dichotomiflorum. This is an extremely polymorphic species, ranging from very small plants, approximately 10  cm high, very branched, with numerous axillary panicles and spikelets of variable size. On the contrary, in places with higher humidity and different types of soil, more erect and vigorous specimens are observed, reaching up to about 1.40 m high, or larger, plants that are also branched and with abundant axillary panicles toward the upper portion. There are also adventitious specimens in corn and alfalfa crops that have somewhat larger growth habit and spikelets, reaching more than 1 m in height. Panicum aquaticum is a species related to P. dichotomiflorum, synonymized by Renvoize in 1984, differing from the latter by being a perennial and rhizomatous plant. There are numerous specimens lacking the basal part, so this character is often difficult to determine exactly. Histofoliar studies conducted on species of the section Dichotomiflora have shown that in P. dichotomiflorum, there are papillae on both epidermis, while they are absent in the abaxial epidermis of the specimens examined of P. aquaticum. Additional material examined. Mexico. Puebla: Laguna Epatlán, 11 December 1942, Miranda 2496 (MEXU). Sinaloa: Mazatlán, Ortega 5760 (US) Panama. Zona del Canal: Gatún, Hitchcock s.n. (Amer. Gr. Hb. 204, US). Los Santos: 5 km SW of Chitre, 22 January 1966, Tyson et al. 3034 (MO). Panama: vicinity of La Jagua, 7 September 1940, Bartlett & Lasser 16990 (MO); Macapale Island in Madden Lake, 8 June 1969, Tyson 5494 (MO) Bermuda. Hamilton, Millspaugh 126 (USA); roadside, Hungry Bay, Brown & Britton 1638 (NY); roadside, west end of Causeway, Brown & Britton 982 (NY) Bahamas. Near Nassau, 25 April 1903, Curtiss 177 (MO, US); Hog Island, Eggers 4512 (US); Andros, Mastic Point and vicinity, Brace 7099 (NY); Watling’s Island, Cockburn town and vicinity, Britton & Millspaugh 6102 (NY); Great Abaco Island, ca. 5 mi. north of Dundas Town along main road, 14 December 1979, Wunderlin et al. 8318 (MO) Cuba. Batabano, 15–20 March 1906, Hitchcock 150 (MO, US). Habana: Habana, near Ciénaga, Ekman 13191 (NY, US); Laguna del Ariguanabo, León 2775 (US); Laguna de Ariguanabo, near Cayo La Rosa, 15 August 1921, Ekman 13101 (NY, US). Pinar del Río: Remates, Ekman 11211 (NY, US); Remates, edge of small laguna, Ekman 11212 (USA); vicinity of Herradura, 28 August 1910, Britton & Earle 6566 (NY, US). Santa Clara: Ciénaga de Zapata, Cayo de los Pavos, 14 August 1920, León 9507 (NY, US); Herradura, 15 August 1907, Tracy 9342 (MO, US)

Key to Ungrouped Panicum Sections and Species

49

Haiti. Vicinity of Etang, Etang Saumatre, 5 April 1920, Leonard 3493 (NY), 4233 (US); Plaine Cul-de-Sac, Croix des Bouquets, Manneville, 20 January 1925, Ekman 3060 (US); Massif da la Hatte, Miragoane, 15 November 1926, Ekman 7233 (US) Guadalupe. Without locality, 1892, Duss 3180 (MO) Colombia. Amazonas: trapecio amazónico, entre los ríos Loretoyacu y Hamacayacu, orilla del Loretoyacu, November 1945, Duque-Jaramillo 2024 (COL). Antioquia: orillas del río Negro, sitio El Tablazo, sin fecha, Mercado B-499 (HUA). Arauca: 13  km al sur de Arauca, Jorgenson 12 (COL). Cesar: Playines de Chiriguaná, 50 m, 16 July 1985, Wood 4992 (COL). Meta: de CuMarchal a San Nicolás, 10 km del desvío a San Nicolás, bosque secundario junto a arroyo y sabana, 480 m, 11 June 1989, Zuloaga 3867 (COL, SI) Vichada: Rio Meta, Macoya Guafa, Cuatrecasas 4148 (F) Venezuela. Anzoátegui: Pariaguan, Laguna de los Patos, 17 July 1946, Burkart 17289 (SI, VEN). Apure: Sabanas del Alto Apure, February 1980, Ramia 7195 (VEN); cercanías de Mantecal, 7 August 1970, Ramia 3604 (VEN); El Trompillo, frente a La Urbana, Trujillo 11761 (MY); Hato La Trinidad de Arauca, Dtto. Muñoz, 5 September 1966, Ojasti 83 (VEN); Hato El Frío, October 1969, Velasquez 830 (VEN); sabanas del Hato El Frío, entre El Saman y Mantecal, Ramia 3620, 3864, 3874, 4770 (VEN); Bajo Apure, 22 August 1972, Ramia & Montes 4826 (VEN); Hato San Juan de Rio Claro, Borsotti 66 (US, VEN); sabanas entre Achaguas y San Fernando, cerca de Bethel, 6 August 1989, Zuloaga et  al. 4349 (SI, VEN); San Fernando de Apure, 11 November 1971, Davidse 2997 (MO, VEN); Dtto. Munoz, Caño Caicara, 11 airline west of Mantecal, 3 March 1978, Davidse & González 14778 (MO, VEN); Dtto. San Fernando, mouth of the Rio Arauca at its intersection with the Rio Orinoco, 14 May 1977, Davidse & González 13205 (MO). Barinas: Los Mangos, costa del Río Caparo, Ramia 1724 (VEN); Hato Corozal, orilla sur del Río La Yegua, 6 August 1966, Ojasti 67 (VEN), carretera Calabozo-Camaguan, Hato Flores Moradas, 6 August 1957, Ramia 1209 (VEN). Delta Amacuro: a lo largo del Caño Araguato, entre Los Rastrojos, El Mosquero y Yaguaraco, 14 October 1977, Steyermark et al. 114619A (MO). Monagas: Llanos, Ramia 2442 (VEN) Suriname. 2 km above confluence with Lucie River, 3 July 1963, Maguire et al. 54053 (F, MO, NY) French Guiana. Pres de Loca-Loca, sur le Maroni, Schnell 11424 (P); without locality, Leprieur s.n. (P), 41, 42 (P) Ecuador. Archipielago de Colón: Isla Santa Cruz, 3 August 1974, van der Werff 1373 (MO), Wiggins 18766 (LIL) Peru. Loreto: Dtto. Mazán, slight above Varadero de Mazán, 11 October 1979, Rimachi 4657 (MO); Ramón Castilla, Pevas, Río Ampiyacu, 15 October 1987, Vasquez & Jaramillo 9869 (MO); Maynas, Iquitos, 22 June 1969, Mc Daniel 11772 (MO); Rio Nanay, frente a Santa Clara, 30 March 1976, Revilla 450 (F, MO). Puno: between Rio Azata-Colorado, 26 June 1986, Nuñez & Muñoz 5298 (MO) Brazil. Amapá: Braco do Rio Macacoari-Macapa, Farias s.n. (MO); Rio Oiapoque, Froes 26084 (IAN). Amazonas: Lago Tefe, mouth of Rio Bavaria, Plowmann et al. 12574 (MO); cercanias de Manaos, Spruce 1431 Panicum 29 (G,

50

 

Taxonomic Treatment

GH, K); Manaos, Ponta Negra, Fromm 1377 (MO, SI); Paraná do Ramos, Pires & Black 1204 (IAN); Barulos, Rio Negro, Froes 28359 (IAN); km 21 da rodovia Manaus-Itacoatiara, Oliveira 2929 (IAN); Benjamin Constant, Pires & Black 850 (IAN). Bahia: Joazeiro, near Rio San Francisco, 13 December 1924, Chase 7907 (F, MO, NY); Ceará: Serra de Baturité, sitio São Inácio de Azevedo, Eugenio 271, 317 (RB); Iguatu, Swallen 4400 (K), 4414 (IAN, K, SP); Picos, Piaui to Campo Salles, Swallen 4266 (RB, SP); Quixada, beira do acude do Cedro, Black 55-18217 (IAN). Distrito Federal: west margin of Lago Paranoa, Brasília, 11 March 1966, Irwin et al. 13875 (F, MO, NY, UB); Universidade de Brasília, 26 October 1965, Irwin et  al. 9573 (F, MO, UB), Clayton 4832 (K); 5  km northeast of Planaltina, Irwin et al. 9261 (UB). Goiás: 31 km N of intersection of highway BR-153 and 60-54, 10 April 1976, Davidse et  al. 12246 (MO, SP); entre o Rio Javaes e Rio Formoso, 28 July 1978, Pires & Santos 16272 (MO); Mun. de Araguatins, 5°21′S, 48°45′W, 17 November 1983, Mileski 353 (RB). Maranhao: Caxias to Barra do Corda, Swallen 3569 (RB); Perizes, Black et al. 54- 16587 (IAN). Mato Grosso: Fazenda Descalvados, Mun. Caceres, Allem 1542 (CEN); Mun. Barao de Melgaco, Fazenda Santa Lucia, Allem & Vieira 2497 (CEN); Mun. Poconé, Porto Cercado, 8 February 1978, Allem & Vieira 1615 (CEN, MO); Mun. Poconé, Fazenda Santa Isabel, Allem & Vieira 2577 (CEN); Fazenda São Bento, 17 November 1977, Allem & Vieira 1281 (MO); margem do Guaporé, Kuhlmann 2414 (SI); south of Rio SuiaMissu Ferry, Harley 11218 (IAN, RB, UB); Porto XV, Hatschbach 32527 (K); Rio Brilhante, Hatschbach 26163 (K). Mato Grosso do Sul: Fazenda Rio Negro, 29 October 1978, Allem et  al. 2264 (CEN, MO); Mun. Corumbá, Fazenda Santo Estevao, 14 November 1977, Allem & Vieira 1200 (CEN, MO); Mun. Corumbá, Fazenda Nhumirim, 20 November 1977, Allem & Vieira 1379 (CEN, MO); Mun. Corumbá, Fazenda São Bento, 16 November 1977, Allem & Vieira 1240 (CEN, MO); Aquidauana, Chase 11028 (RB); Fazenda Bodoquena, 25 October 1978, Allem et  al. 2146 (CEN), 2185 (CEN, MO). Minas Gerais: Juiz de Fora, 20 February 1925, Chase 8549 (F, GH, MO, NY); 12 km southwest of São Gonçalo de Sapucay, 25 February 1976, Davidse et al. 10590 (MO); 19 km south of intersection of highways BR-116 and BR-262, N of village of Sao Joao do Manhuacu, 27 March 1976, Davidse et al. 11437 (MO); Fazenda Santa Terezinha, Capinopolis, Macedo 4527 (UB). Pará: Bõa Vista, Rio Tapajos, Swallen 3149 (RB, SP); Marajó Island, Estate Gavinho, Goeldi 174 (F, MO); Rio Cumina, Sampaio 5333 (IAN, R); Rio Trombetas, Silva 1239 (SP); Belem, Black 48-2914 (IAN); Jacundazinho, Black 49-8010 (IAN); Fazenda Cacaval Grande, proximo a Santarem, Pires 1828 (IAN); Santarem., Spruce 788 (K). Paraná: Parque. Nac. de Iguazu, Porto Franco, Duarte 1837 (RB). Pernambuco: Rio São Francisco, Elias de Paula 3008 (UB). Rio Grande do Norte: Nova Cruz to Montanhas, Swallen 4830 (SP). Rio Grande do Sul: Mun. Itaqui, BR-472, margem Direita do Rio Ibicui, Valls et al. 4689 (CEN); Cristal, Rambo 40742 (IAN, LIL); km 69, BR-158, Livramento-Rosario, Esteio, Rambo 40614, 41798 (LIL); Gral. Vargas, 28 April 1956, Barreto s.n. (SI); São Leopoldo, Dutra 286 (SI); Pelotas, Costa Sacco 269 (F), 361 (BAA, F, IAN, NY, RB); Porto Alegre, Ilha da Pintada, Rambo 41130 (LIL, MO); Uruguaiana, Fazenda Paraiso, 12 May 1964, Flavio s.n. (SI). Rio de Janeiro: Copacabana, Glaziou 4342

Key to Ungrouped Panicum Sections and Species

51

(P); Impetiba, Glaziou 20123 (P); Macae, Strang 1348 (SI). Roraima: beira do Rio Branco, capoeira perto de Bõa Vista, Black & Magalhaes 51-13009 (CEN, IAN, US). Santa Catarina: Sitio Elias Baskerotto, São Ludgero, Klein & Bresolin 11362 (R); Joinville, Hans 266 (RB, SI), 323 (SI). São Paulo: Alto da Serra, November 1910, Luederwaldt s.n. (SP); Mogi-Guacu, Eiten et al. 1940 (SP); S. Bernardo, da Silva 328 (SP); São Paulo, Sendulsky 236 (SP). Without locality, Riedel s.n. (GH, NY, P, W) Bolivia. Beni: Estancia El Porvenir, 50 km east of the Rio Mariqui on the road to Trinidad, 17 November 1985, Solomon 14771 (MO); Prov. Ballivian, Espiritu, en la zona de influencia del Río Yacuma, 200 m, 13 October 1980, Beck 5028, 5108 (LPB, SI), 5050, 5321 (LPB), 5150 (K). La Paz: Prov. Iturralde, Luisita, Haase 525 (SI), 836 (K). Santa Cruz: Buena Vista, 25 November 1916, Steinbach 3180 (GH, LIL); Dolores, Steinbach 1906 (LIL); Prov. Velasco, 13 km de San Ignacio, aprox. 400 m, 15 December 1987, Bruderreck 93 (SI) Paraguay. Alto Paraguay: Chaco, Fiebrig 1302 (G, P). Alto Paraná: Puerto Bertoni, Bertoni 3405, 3551 (W) Amambay: Pedro Juan Caballero, 1888-1890, Morong 543 (MO, NY). Central: Bords du Rio Paraguay a L’Assomption, Balansa 61 (G, K, P); bords du Rio Paraguay, pres de L’Assomption, Balansa 19 (G, P); cerca del lago Ipacaray, Hassler 12469 (F), 12537 (BAF, G, GH, K, LIL, MO, NY). Concepción: curso superior del rio Apa, Hassler 8206 (G, GH, K, LIL, MO, NY). Cordillera: In regione lacus Ipacaray, S.  Bernardino, Hassler 12469 (F), 12537 (BAF, G, GH, K, LIL, MO, NY). Guairá: Colonia Independencia, Schinini & Bordas 25204 (G). Itapúa: Carmen del Paraná, April 1981, Lurvey 313 (MO) Neembucú: Alonso #1, Walter 161 (K). Paraguarí: Río Paraguarí, Balansa 19 (P); Caapucú, Rosengurtt B-5242 (P), Rojas 13191 (BAA). Presidente Hayes: in regiónis cursus inferioris fluminis Pilcomayo, Rojas 237 (K), 237a, 237b, 237c (G). San Pedro: San Pedro, Alto Paraguay, Woolston G-110 (K, NY, SI, SP) Argentina. Buenos Aires: Pdo. Avellaneda, Isla Maciel, Parodi 760 (BAA). Pdo. La Plata, La Plata, Parodi 10593 (BAA), Pdo. Capitan Sarmiento: Capitán Sarmiento, León 751 (BAA). Pdo. Coronel Vidal: Coronel Vidal, Parodi 9347 (BAA). Pdo. San Isidro: San Isidro, Parodi 5736, 5808 (BAA). Pdo. Tigre: Tigre, Parodi 11096 (BAA), Hunziker 441 (LP). Delta del Paraná, Burkart 1210, 1226, 3715, 3793 (SI); Delta del Paraná, La Serna y Paraná de las Palmas, 4 March 1936, Burkart 7595 (SI); Paraná Miní, Cabrera 1607 (F, LP, NY); Isla Santa Catalina, 29 January 1939, Nicora 2120 (SI). Chaco: Dpto. 1° de Mayo: Colonia Benitez, 1 December 1959, Schulz 10903 (SI), 15899 (BAB). Dpto. San Fernando: Resistencia, Meyer 380 (LIL, SI), Parodi 8269 (BAA, CTES); Fontana, Meyer 80 (LIL); Dpto Libertad, Gral. Donovan, 1 March 1942, Schulz 3261 (MO). Corrientes: Dpto. Mburucuyá, Estancia Santa Teresa, 5 January 1958, Pedersen 4792 (MO), 4805 (P); Dpto. Bella Vista: Est. Exp. INTA, Schinini & Miranda 9578 (CTES, LIL, SI, US). Dpto. Berón de Astrada: Valencia, 46  km oeste de Itá Ibaté, 14 January 1977, Ahumada 423 (CTES, MO, SI), 425 (CTES, F); Arroyo Santa Isabel, ruta 12, Schinini & Quarín 8524 (CTES, SI). Dpto. Capital: ruta 12, 25 km este de Corrientes, Ahumada 1673 (CTES, F); Riachuelo, Arrocera Argentina, Ahumada 610 (CTES, F, SI). Dpto. Concepcion: Tabay, Arbo et al. 981 (CTES); 6 km este de Santa Rosa, Estancia Millán, Arbo et al. 785 (CTES, SI). Dpto. Empedrado: Campo Cooperadora

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Taxonomic Treatment

INTA, Fernández 846 (SI); Laguna El Tabaco, Carnevali 2976 (CTES); El Sombrerito, Nicora & Cámara Hernández 219 (BAA), Vallejos 259 (BAA); Manuel Derqui, Schwarz 10204 (CTES, LIL); Estancia La Yela, 16 March 1977, Pedersen 11693 (MO, SI). Dpto. Gral. Alvear: Torrent, Ibarrola 1717 (LIL, NY). Dpto. Gral. Paz: 6 km sudoeste de Caa Catí, ruta 13, Ahumada 1924 (CTES). Dpto. Itati: Ruta Nac. 12, 47 km E de Itatí, Ahumada et al. 791 (CTES, F, SI), 60 km este de Itatí, Ahumada et al. 919 (CTES, F, SI, US). Dpto. Goya: ruta 12, 40 km sur de Goya, Krapovickas et al. 27527 (CTES). Dpto. Ituzaingó: 35 km del limite con Misiones, Krapovickas et al. 15705 (CTES). Dpto. Lavalle: Colonia Echeverria, 1 km norte de Santa Lucía, 25 May 1973, Schinini 6580 (CTES, MO). Dpto. Mercedes: Colonia Carlos Pellegrini, Quarín & Schinini 1098 (CTES, LIL). Dpto. Paso de los Libres: Estacion Yapeyú, Ibarrola 52 (LP). Dpto. Saladas: Saladas, Ibarrola 3085 (LIL, NY); San Lorenzo, Schwarz 9452 (LIL). Dpto. San Marchtin: Yapeyú, Ibarrola 1886 (LIL, NY, SI). Dpto. San Miguel: 12  km norte de San Miguel, ruta 17, 15 March 1978, Ahumada 1872 (MO); 19 km sur de Caa-Catí, ruta, 5, Ahumada 1790 (SI). Dpto. San Roque: Arrocera Passetto, Cano & Cámara Hernández 902 (BAA, CTES). Dpto. Santo Tomé: Cuay Grande al sur, Ibarrola 1590 (LIL, NY, US); boca de ríos Cuay Grande y Uruguay, Ibarrola 36 (LP). Entre Ríos: Dpto. Federación: Salto Grande, Burkart & Gamerro 21593 (SI). Dpto. Gualeguaychú: Holt, Martínez Crovetto 1994 (BAB). Dpto. Uruguay: Isla Cambacuá, December 1957, Nicora 6512 (BAA, SI). Formosa: Dpto. Formosa: Formosa, Parodi 8324, 8340 (BAA), Kermes 403 (BAA, BAB); ruta 11 vieja, al norte de Arroyo Francesa Cué y sur de Estancia La Emilia, 6 Ene 1980, Guaglianone et al. 359 (SI, US). Dpto. Patiño: Las Lomitas, Parodi & Horowits 44 (BAA); Bartolomé de las Casas, Schulz 17777 (LIL). La Pampa: Dpto. Conhelo: Estancia Castex, Troiani & Steibel 4090 (SI). Dpto. Utracán: Ruta 35, Steibel 4971 (SI). Misiones: Puerto España, Schwarz 6205 (NY); Isla del Toro, Río Paraná, Montes 329 (NY); Dpto. Candelaria: Ruta 12, 15 km noreste de Candelaria, Romanczuk et al. 464 (SI); Bonpland, van de Venne 38 (SI). Dpto. Capital: Posadas, Bertoni 4802 (LIL). Dpto. Eldorado: Puerto Eldorado, 7 December 1943, Burkart 14546 (SI), 17 October 1977, Cabrera et al. 28853 (SI), Zuloaga et al. 3293 (SI); Dpto. Libertador Gral. San Martín: Puerto Rico, Cabrera 28811 (SI), Zuloaga et al. 3277 (SI). Dpto. Montecarlo: Puerto Piray, Schwarz 6840 (LIL, MO), 27 December 1978, Renvoize et al. 3290 (K, MO, NY, SI). Dpto. San Ignacio: camino al Peñón del Teyu Cuaré, 11 April 1985, Zuloaga et al. 2262 (MO, SI). Santa Fe: Dpto. Constitución: Juncal, León 805 (BAA). Dpto. Gral. Obligado: Estero El Chajá, Meyer 2554 (LIL). Dpto. La Capital: Arroyo Leyes, Tur 1205 (LP, SI). Dpto. Vera: Caraguatay, Porta 103 (CTES) Uruguay. Artigas: Bella Unión, Rosengurtt B-7373 (US). Montevideo: Carrasco, Legrand 980 (F); Parque Centenario, Rosa Mato 1913 (LIL) Paysandú: Queguay, Felippone 4904, 4908 (SI); Paysandú, Rosengurtt B-3802 (BAA). Salto: Salto, February 1927, Burkart 1137 (SI); Espinillar, del Puerto 18 (F) 2b. Panicum dichotomiflorum Michx. subsp. bartowense (Scribn. & Merr.) Freckmann & Lelong, Sida 20(1): 171. 2002. P. bartowense Scribn. & Merr., Circ. Div. Agrostol. U.S.D.A. 35: 3. 1901. P. dichotomiflorum Michx. var. bartowense (Scribn. & Merr.) Fernald, Rhodora 38(455): 387. 1936. Type: USA.  Florida: Polk County, Bartow, 29 September 1898, R.  Combs 1220 (lectotype, US

Key to Ungrouped Panicum Sections and Species

53

00133099, designated as “type” by Hitchcock & Chase, 1910: 52; isolectotype, ISC-v-0000569) Panicum proliferum Lam. var. pilosum Griseb., Cat. Pl. Cub.: 232. 1866. Type. Cuba, Ad lagunas, C.  Wright 1865 (lectotype, GOET 006760, designated by L. Catasús Guerra, 2004: 85; isolectotypes, GOET 006761, GOET 006762, US 00139897) Distribution and ecology (Fig. 3). Subspecies restricted to the southern USA and the Caribbean; found in marsh edges and flooded sites; flowers and fruits from July to December It is distinguished from the type subspecies by its densely hirsute sheaths and smaller spikelets, usually 2–2.2 mm long. Additional material examined. USA.  Florida: Miami, Hitchcock s.n., Amer. Gr. Hb. 17 (US, W) Bahamas. Grand Bahama: about Freeport dump north of Airport, 25 January 1976, Correll & Popenoe 46671 (MO); in rocky open pinelands along Midshipman Road, Freeport, Correll & Correll 50953 (NY); Bimini Island, North Bimini, Howard & Howard 10190 (NY); North Andros: in depression on edge of Casuarina grove at Forfar Station, Blanket Sound, Correll 51350 (NY); South Bimini: in disturbed moist soil about airport, Correll et al. 50462 (NY); Great Exuma, in open area on south end of road in eastern part of island, Correll & Correll 48142 (NY) Cuba. Havana: Barabanó, in coastal swamps, 10 December 1920, Ekman 12602 (G, US). Isla de Pinos: Isla de Pinos, Ekman 12400 (US). Pinar del Río: Guanajay, Laguna de Brito, in grassy, sometimes flooded places, 9 July 1921, Ekman 13013 (G, US). Santa Clara: Ciénaga de Zapata, north of Bahía de Cochinos, León & Loustalot 9507 1/2 (NY) Jamaica. Near the town of Black River, in a swamp along the Black River, 22 October 1912, Hitchcock 9645 (MO, NY, US) Caicos Island. Pine Cay, in clumps in drying open pond of coppice on northeast side, Correll 43251 (NY) 3. Panicum lacustre Hitchc. & Ekman, Man. Grasses W. Ind.: 253, Fig. 205. 1936. Type: Cuba. Pinar del Río: Laguna de la Máquina, 31 October 1923, E. L. Ekman 17878 (lectotype designated by Zuloaga, 2002: 337, US 00132924; isolectotypes, B_10_0248983, BAA 00000387, BR 0000008408877, DAO 000465583, G 00099619, GH 00135365, ISC-v-0000574, LD 1010829, LIL 000085, MICH 1127266, MO 128363, MVFA 0000436, NY 00071083, NY 00803749, P 00740958, R 000076339, S13-12944, SI 002808, US 00132923, W 1959000033). Fig. 4 Perennial plants, shortly rhizomatous, with erect culms 1–1.5 m tall, succulent, with adventitious roots at the lower nodes; basal internodes short, hollow, cylindrical, glabrous; nodes brown, glabrous. Sheaths striate, with anastomosed veins, glabrous. Basal leaf blades reduced and reflexed, drooping, basal sheaths shorter than internodes, the upper ones longer, pale to purplish, with membranous margins, shortly hairy or glabrous. Ligule membranous-ciliate, ca. 2  mm long, the membraous portion reduced, less than 0.5  mm long; collar brown, glabrous. Blades

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Taxonomic Treatment

Fig. 4  Panicum lacustre. (a) Habit. (b) Ligular region. (c) Portion of the inflorescence. (d) Spikelet, lateral view. (e) Spikelet, ventral view. (f) Spikelet, dorsal view. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Upper lemma, lodicules and gynoecium. (j) Upper lemma, lodicules and androecium. (From Ekman 17878, USA)

Key to Ungrouped Panicum Sections and Species

55

linear-lanceolate, the upper ones ca. 20  ×  0.2  cm, margins involute toward basal portion, then flattened, base narrow and apex long subulate, the adaxial surface densely hispid at the base, sparsely hispid toward the apical portion, abaxial surface scabrous, the margins scabrous. Inflorescence terminal, exerted, peduncle cylindrical, scabrous; panicles lax, multiflowered, 20–30  cm long; main axis wavy, scabrous, pulvini brown, glabrous, first-order branches alternate or opposite, divergent, branch axes and pedicels triquetrous, scabrous, spikelets congested on short pedicels toward the upper portion of branches, pedicels in pairs. Spikelets narrowly ellipsoid, 2 × 0.8 mm, glabrous, greenish or tinged with purple, upper glume and lower lemma subequal or the upper glume smaller and in this case obtuse, acute; lower glume 0.5 mm long, 1/4 the length of the spikelet, truncate to ovate, 3 nerved; upper glume 7 nerved; lower lemma 7 nerved; lower palea and lower flower absent; upper anthecium ellipsoid, whitish, 1.8  ×  0.7  mm, smooth, glabrous, shiny. Caryopsis not seen. Distribution and ecology (Fig. 5). Species endemic to the west of the island of Cuba, where it is found on the edges of lagoons or swamps; it is rare; flowers and fruits between June and October

Fig. 5  Distribution of Panicum lacustre, P. sublaeve, and P. vaseyanum

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Taxonomic Treatment

This species is characterized by being a perennial plant, with reflexed and drooping basal blades, a lax and exserted terminal panicle and spikelets with the upper glume and lower lemma 7 nerved, the lower palea and flower absent. Additional material examined. Cuba. Pinar del Río: Pinar del Río, in a small Laguna at km 12 of the carretera to La Colona, 28 October 1923, Ekman 17822 (G); La Grifa, at a Laguna north of town, 14 June 1920, Ekman 11271 (G) 4. Panicum sublaeve Swallen, Contr. US Natl. Herb. 29(9): 424. 1950. Panicum rigidum Swallen, Ann. Missouri Bot. Gard. 30(2): 215. 1943, nom. illeg. hom., non Balfour, 1884. Type: Panama. Sabana de Juan Corso, near Chepo, 1 October 1911, H. Pittier 4526 (holotype, US 00147999). Fig. 6 Annual plants, cespitose, with geniculate to erect culms, 15–65 cm tall, branched; internodes cylindrical, hollow, glabrous, nodes brown, glabrous. Sheaths larger than internodes, striate, glabrous. Ligule membranous-ciliate, ca. 1.6 mm long, the membranous portion 0.3  mm long; collar brown, glabrous. Blades linear-lanceolate, (3–)13–24 × (0.2–)0.5–1.2 cm, base rounded to subcordate and apex attenuate, flat, the adaxial surface sparsely hairy, the abaxial one glabrous, midrib manifest. Inflorescence terminal exserted, peduncle cylindrical, glabrous, 5–20 cm long; panicles lax, diffuse, 15  ×  10  cm; main axis angulate, scabrous to glabrous, pulvini glabrous, branches alternate, branch axes and pedicels triquetrous, scabrous, spikelets solitary and scattered on the branches; axillary panicles present. Spikelets plano-convex, long-ovoid, 3.5–4  ×  0.7–1.2  mm, glabrous, greenish, upper glume and lower lemma subequal, exceeding the upper anthecium in length; lower glume ovate, 1.2–1.5 mm long, subacute, 5–7 nerved; upper glume acute, 11–13 nerved; lower lemma acute, 11–13 nerved; lower palea elliptic, 2.7–3.3 × 1 mm, hyaline, glabrous; lower flower staminate, stamens 3; upper anthecium long ovoid, 2.5–2.8 × 0.9–1.2 mm, whitish, smooth, hardened, shiny, lemma 7 nerved, apiculate and with a slightly incurved apex, with conspicuous prickle hairs toward the apex, and papillae over its entire surface; anthers orange, 1.4–2 mm long, stigmas purplish. Caryopsis long ellipsoid, 1.9 × 0.8 mm, whitish; hilum oblong, embryo more than half the length of the caryopsis. Distribution and ecology (Fig. 5). Species distributed from Mexico and Mesoamerica to Venezuela; it grows in open and humid environments, from sea level to 1100 m asl; flowers and fruits between August and January. This species is distinguished by being an annual plant, with a lax and terminal panicle, spikelets with the upper glume and lower lemma 11–13 nerved, exceeding in length the upper anthecium, the latter apiculate and with conspicuous prickle hairs at the apex, the flower with orange anthers. Additional material examined. Mexico. Chiapas: flats with savanna adjacent to Río San Pedro, 86 km south of Mexican Highway 190 on road to Nueva Concordia near El Brillante, 3 November 1981, Breedlove & Davidse 54548 (MO); small forested stream and adjacent savanna 2  km southeast of Arriaga, Mun. Arriaga, 27 August 1974, Breedlove 36852 (MEXU, MO); Mun. Angel Albino Corzo, near Finca Mérida along road from Vicente Guerrero to Finca Tuxtepec, 4 November

Key to Ungrouped Panicum Sections and Species

57

Fig. 6  Panicum sublaeve. (a) Plant. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Lower palea. (e) Upper anthecium, dorsal view. (f) Upper anthecium, ventral view. (g) Apex of upper anthecium. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (From Zuloaga 4354, SI)

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Taxonomic Treatment

1981, Breedlove & Davidse 54664 (MO); Mun. Villa Corzo, 58 km south of Mexican Highway 190 on road to Nueva Concordia, 2 November 1981, Breedlove & Davidse 54452 (MO). Colima: Alzada, 21 September 1910, Hitchcock s.n. AGH 503 (F, MO, US) Honduras. Francisco Morazán: camino San Antonio, 21 October 1943, Rodriguez 1341 (F); Zamorano, 24 September 1943, Rodriguez 1011 (F, GH), 16 October 1943, 1228 (F); vicinity of El Zamorano, 17 October 1951, Swallen 10839 (MO, US); vicinity of El Zamorano, in pasture of Jaragua, 29 October 1951, Swallen 11145 (MO, US); region of Las Mesas, 14 October 1951, Swallen 10745 (US); Río del Gallo, near El Jicarito, about 2 km from El Zamorano, 900–1000 m, 22 October 1951, Swallen 10982 (US); alrededores de Colonia Kennedy, 1100 m, 30 January 1985, Figueroa 178 (NY). Valle: San Lorenzo, 13 September 1945, Rodriguez 3443, 3492 (F, US); just south of Jícaro Galán along the Interamerican Highway, 60 m, roadside borrow-pit marsh dominated by Nymphaea in the center, water ca. 40 cm deep, edge of marsh, 5 October 1986, Davidse & Pilz 31697 (MO); 3 km east of San Lorenzo along the road to the new sea harbor, 2 m, open marsh along road next to Conocarpus mangroves, edge of marsh, 5 October 1986, Davidse & Pilz 31674 (MEXU, MO); 3 airline km west of San Lorenzo along the road to Amapala, 30  m, Crescentia savanna on heavy black clay; heavily grazed and disturbed, 5 October 1986, Davidse & Pilz 31712 (MEXU, MO) El Salvador. La Libertad: Along CIA, km 38, ca. 10 km west of La Libertad, open roadside near Pacific Ocean, 10 m, 17 August 1971, Pohl 12774 (F, MO). San Miguel: San Miguel-La Unión, km 151, 12 September 1951, Rohweder 2486 (MO) Costa Rica. Guanacaste: Nicoya Peninsula, 5  km south/southwest of Veinte Siete, 17 August 1968, Pohl & Davidse 10941 (F); Finca América, Anderson 1366 (US); Hacienda Murciélago, roadside in pasture, 20 October 1982, Pohl 14182 (MO) Panama. Zona del Canal: Monagre Beach 5 mi SE of Chitre, 22 January 1966, Tyson et al. 3030 (MO) Venezuela. Apure: Sabanas del Hato El Frío, entre El Saman y Mantecal, 9 August 1970, Ramia 3637 (VEN). Guárico: Región Corozopando-Camaguan, en sitio cubierto por agua, San Fernando de Apure, 27 August 1959, Ramia 1967 (VEN); carretera entre Calabozo y Camaguan, Hato El Trompillo, en bajo, común, Ramia 1260 (VEN); en sabana aguachinosa entre Corozopando y Camaguan, 27 August 1961, Ramia 2559 (MY, VEN); Distrito Miranda, 10 km al N de Camaguan, 6 August 1989, Zuloaga et al. 4354 (SI, VEN) 5. Panicum vaseyanum Scribn. ex Beal, Grass. N. Amer. 2: 140. 1896. Type: Mexico. Chihuahua. Base of Sierra Madre, 30 September 1887, C. G. Pringle 1415 (lectotype, US 00148082, designated as “type,” by Hitchcock & Chase, 1910: 47; isolectotypes, COLO 00392639, COLO 00392621, E 00373819, F 0046899F, GH 00024127, K 000309146, MEXU 00003724, MICH 1108751, MIN 1001936, MO-128362, MSC 00922516, NDG 06888, NY 00381781, NY 00381782, PH 00018737, RSA 0000465, US 00148081, US 00037314, UVMVT 027972, W 19160022600)

Key to Ungrouped Panicum Sections and Species

59

Annual plants, cespitose, with geniculate culms, decumbent and rooting at the lower nodes, then erect, 15–70 cm tall, branching toward the upper nodes; internodes compressed, with aerenchyma, pale to purplish, glabrous, hollow, nodes brownish, glabrous. Sheaths striate with anastomosed veins, pale to purple, glabrous, the margins membranaceous. Ligule membranous-ciliate, the membranous portion 0.1–0.6 mm long, ciliate portion 0.4–1.5 mm long; collar brown to purple, glabrous. Blades lanceolate, 3–25 × 0.2–0.6 cm, flat or with involute margins toward the base, base rounded, apex attenuate, greenish to violaceous, glabrous, the margins scabrous. Terminal and axillary inflorescences present, similar, shortly exserted; panicles contracted, 2–9 × 1–1.5 cm; main axis wavy, scabrid to glabrous, pulvini glabrous, first-order branches alternate to opposite, ascending and adpressed, spikelets in pairs or solitary on short second-order branches, adpressed on first-order branches; pedicels short, claviform, scabrous. Spikelets long ellipsoid, 2.2–3.2 × 1.1–1.4 mm, acute, greenish, glabrous; lower glume 1/4 to 1/3 as long as the spikelet, 1 nerved, ovate, hyaline, amplexicaul, the ape obtuse to truncate; upper glume 7 nerved, acute; lower lemma glumiform, 7 nerved; lower palea and lower flower absent; upper anthecium long ovoid, 2–2.7 × 1–1.2 mm, smooth, shiny, glabrous, pale, brown at maturity, with verrucose papillae toward the apex of the palea, easily drooping at maturity before the bracts. Caryopsis ellipsoid; hilum oblong, embryo 1/2 the length of the caryopsis. Distribution and ecology (Fig. 5). Species endemic to Mexico, where it grows in humid environments between 1700 and 2700 m asl; flowers and fruits between October and November Panicum vaseyanum is defined as an annual plant, with contracted panicles and conspicuous axillary inflorescences, spikelets with the upper glume and lower lemma 7 nerved, lower palea and lower flower absent and upper anthecium brown at maturity, and early drooping before the bracts. Additional material examined. Mexico. Aguas Calientes: Aguas Calientes 2 October 1910, Hitchcock 7491, 7485 (US); near Aguas Calientes, 2 October 1910, Hitchcock s.n., AGH 15 (F, MEXU, MO, NY, P); 12 km al sudoeste de La Congoja, Mun. de San José de Gracia, 2700 m, Rzedowski & McVaugh 836 (US); Puente Las Arboleas, Mun. de Aguascalientes, November 1978, Esparza s.n. (MEXU-272166); Chapingo, 15  km de V.  Guerrero Santo Tomas, Hernández & Tapia 382 (US). Chiapas: Mun. San Cristóbal de las Casas, west edge of San Cristóbal de las Casas, plant in marsh, 2195  m, 29 October 1981, Breedlove & Davidse 53971 (MO). Chihuahua: 11 miles west of Cuauhtémoc, in wet depression, 7000 ft, Reeder & Reeder 4849 (US); shallow water, base of Sierra Madre, Pringle 1688 (NY, P); Miñaca, 13 October 1910, Hitchcock 7757 (USA); Pine-Oak region, Sierra Madre Occidental, west of Casas Grandes, five miles south of Hernandez, 18 September 1960, Reeder et al. 3526 (MEXU, US). Durango: Reserva La Michila, 80 km al sudeste de Durango, 2500 m, 6 October 1980, Carrillo 9 (MEXU); ten miles east of el Salto, 2530 m, 2 October 1953, Reeder &. Reeder 2579 (MEXU). Guanajuato: about 6  km west of San Felipe, on dry slopes approaching Cerro del Fraile, 24 October 1952, Sohns 399 (US); Santa Cruz de Juventino Rosas, junto a Guanajuato, 2 September 1981, Beetle M-7325 (MO); cerca de Coroneo, presa y zona inundada,

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Taxonomic Treatment

2640  m, 28 September 1999, Novemberelo & Ramos 3823 (MEXU). Jalisco: Guadalajara, road to Barranca Oblato, 6100 ft, 27 September 1910, Hitchcock 7315 (US); Pegueros, between Tepatitlán and Valle de Guadalupe, 21 September 1980, Beetle & Guzman 5513 (MO); Villa Obregón, suelos calcáreos, 1780  m, 21 September 1980, Beetle & Guzmán 5536 (MO); 25 km al noroeste de Bolaños por la brecha a Los Asmoles, bosque de pino y encino, 2470 m, 9 August 1986, Santana Michel et al. 1782 (MEXU); 20 km al S de Matanzas, barranca del Río San Juan, Ojuelos, 20 October 1973, 1700 m, Díaz Luna 4438 (MEXU). México: ciudad de Progreso, 2700 m, 30 September 1950, 2700 m, Matuda 19747 (MEXU, MO, US). Michoacán: 25  km east of Morelia, on highway 15, 2000  m, 20 October 1976, Brunken & Perino 448 (MO). Puebla: autopista de Puebla a Orizaba, al noreste de Puebla, 1.4 km al este de la Puente Amalucan, 2260 m, Vibrans 2453 (MEXU) Panicum Sect. Hiantes Stapf, Fl. Trop. Afr. 9: 640, 644. 1920. Type: Panicum phragmitoides Stapf. Panicum sect. Durae Stapf, Fl. Trop. Afr. 9: 640, 648. 1920. Type: Panicum turgidum Forssk., lectotype designated by F.  O. Zuloaga et  al., PLOS One 13(2):29. 2018. Panicum sect. Urvilleana (Hitchc.) Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 11(104): 244. 1931. Panicum group Urvilleana Hitchc. & Chase, Contr. US Natl. Herb. 15: 28, 132. 1910, nom. inval. Panicum [no rank] Urvilleana Hitchc., N. Amer. Fl. 3(2): 200, 205. 1915. Type: Panicum urvilleanum Kunth Panicum sect. Virgata Hitchc. & Chase ex Pilg., Nat. Pflanzenfam. (ed. 2), 14e: 22. 1940. Panicum group Virgata Hitchc. & Chase, Contr. U.S. Natl. Herb. 15: 29, 84. 1910, nom. inval. Panicum [no rank] Virgata Hitchc., N. Amer. Fl. 3(2): 200, 203. 1915. Type: Panicum virgatum L. Perennial, cespitose, rhizomatous plants with short to elongated rhizomes, hairy to glabrous cataphylls, hollow internodes, occasionally solid in the basal portion. Sheaths membranaceous to rigid. Ligule membranous-ciliate. Blades lanceolate to linear-lanceolate or linear, flat to involute, hairy to glabrous. Inflorescence terminal, an exserted panicle, open to contracted or spiciform, oblong to ovate or elliptic. Spikelets solitary or in pairs, ellipsoid to ovoid, long-ovoid to long-ellipsoid or lanceolate, open at maturity, villous to hairy or glabrous, pale or tinged with purplish, rachilla manifest or not between bracts; lower glume (1/2–) ¾ to 4/5 as long as the spikelet, ovate, (3–)5–7(–11) nerved; upper glume equal in length to the spikelet, 7–9(–15) nerved, membranous, acuminate; lower lemma slightly shorter than the upper glume (5–)7–9(–11) nerved, acuminate; lower palea present, hairy to glabrous; lower flower staminate or absent; upper anthecium shorter than the upper glume and lower lemma, whitish to pale or olivaceous at maturity, pilose to glabrous, ovoid to broadly ovoid or broadly ellipsoid, smooth, shiny, hardened; stipe present or absent. Caryopsis ellipsoid to broadly ellipsoid; hilum punctiform, elliptic, oblong or obovate, embryo slightly less than or more than half the length of the caryopsis.

Key to Ungrouped Panicum Sections and Species

61

This section includes 13 species in the Americas and about 24 in the Old World, with a total of 5 annual species. They usually grow in open and dry places or in mesophytic environments. Key to the Species 1. Upper anthecium hairy on the lower edges of the upper lemma 2. Upper anthecium with flattened macrohairs at the base of the upper lemma 3. Spikelet with conspicuous internodes, upper anthecium stipitate.............. ...........................................................13. P. mystasipum 3. Spikelet without conspicuous internodes, upper anthecium not stipitate 4. Upper anthecium with a continuous row of macrohairs at the base of the upper lemma; spikelets 4.7–5.3 mm long; panicles 12–20 cm long, few flowered, non-deciduous at maturity.................................................................. 9. P. complanatum 4. Upper anthecium with macrohairs only at the lower margins of the upper lemma; spikelets 5–8.5 mm long; panicles 20–45 cm long, multiflowered, drooping as a whole at maturity.......................................................................14. P. olyroides 2. Upper anthecium with filiform, not acuminate, macrohairs at the base 5. Spikelets sparsely hairy to glabrous, 3.5–4.9 mm long; lower glume 5–7 nerved, upper glume 7–9 nerved.................................8. P. chloroleucum 5. Spikelets densely hairy, 5–7.5  mm long; lower glume (7–)9–11 nerved, upper glume (9–)11–13(–15) nerved 6. Panicle lax, diffuse, with divergent branches; spikelets broadly ovoid, 1.8–2.6 mm wide........................................... 17. P. urvilleanum 6. Panicle contracted, with branches appressed to the main axis; spikelets broadly ovoid, 2.5–3.4(–4) mm wide....................................................... .....................15. P. racemosum 1. Upper anthecium glabrous 7. Glumes and lower lemma densely pilose, covered with tuberculate-based hairs; lower palea densely pilose; upper anthecium shortly stipitate......................................................................... 10. P. deciduum 7. Glumes, lower lemma and lower palea glabrous; upper anthecium not stipitate 8. Panicle spiciform, 1 cm wide. Endemic to Paraguay.................................. ................................12. P. longissimum 8. Panicle lax, if contracted not spiciform, more than 1 cm wide 9. Culms hard, bambusoid, with sheaths conspicuous, pale and aphyllous toward the basal portion of culms; pedicels with long persistent whitish hairs..................................... 16. P. tricholaenoides 9. Culms herbaceous, not bambusoid, without pale and aphyllous sheaths toward the basal portion of culms; pedicels without long whitish hairs, if present deciduous

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Taxonomic Treatment

10. Spikelets (4.4–)5.2–6 mm long……………............. 7. P. amarum 10. Spikelets 2.4–4 mm long 11. Spikelets with pedicels with long deciduous hairs, exceptionally glabrous; upper anthecium with a dark disk at the base and two black spots along the upper palea. Brazil, Uruguay, and Argentina................................ 11. P. glabripes 11. Spikelets with glabrous pedicels; upper anthecium without a dark disk at the base and upper palea without black spots. Canada and the USA, Caribbean, Mesoamerica, and northern South America 12. Culms geniculate, decumbent, branching and rooting at the lower nodes; culms 2–3 m high................... P. altum 12. Culms simple, erect, 1–2 m tall………. 18. P. virgatum 6. Panicum altum Hitchc. & Chase, Contr. US Natl. Herb. 17(6): 488, Fig. 57. 1915. Type: Panama. Panama: Pt. Chame, 18 September 1911, A. S. Hitchcock 8167 (holotype, US 00133066; isotypes, BM 000938684, BR 0000006883669, BR 0000006883997, F 0046883F, G 00099661, K 000309148, K 000307788, MO-105078, NY 00381755, P 00371643, PH 00018617, RM 0001849, SI 002792, S-R-3936, W 19220009762). Panicum lundellii Swallen, J.  Wash. Acad. Sci. 28(1): 9. 1938. Type: Belize. El Cayo: along Mahogany Creek, Mountain Pine Ridge, 7 August 1936, C. L. Lundell 6903 (holotype, US 00074269, US 00074270, holotype labeled as I and II; isotypes, K 000309148, MICH 1108741, SI 002809, US-1722486). Panicum vigoratum Swallen, Mem. New York Bot. Gard. 9(3): 257. 1957. Type: Venezuela. Amazonas: abundant and dominant around savanna margins, Sabana de Santo, Río Guaviarito, Cerro Guanay, 23 January 1951, B.  Maguire, K. D. Phelps, C. B. Hitchcock & G. Budowski 31601 (holotype, NY 000414075) Plants perennial, cespitose, shortly rhizomatous; culms 2–3 m tall × 3–7 mm in diameter, herbaceous, non-bambusiform, geniculate, decumbent and rooting at lower nodes; internodes reddish, 10–20  cm long, geniculate, hollow, cylindrical, glabrous; nodes dark to glaucous, glabrous. Sheaths shorter than internodes, glabrous or with a hairy margin toward the top, somewhat inflated in the basal portion. Ligule membranous-ciliate, 1.5–3 mm long; collar brown, glabrous. Blades linear, 25–45 × 0.8–2 cm, flat, rigid, rounded at base, the apex subulate, glabrous, the margins scabrous and with hairs toward the base or without hairs, the adaxial surface glaucous. Peduncle up to 8 cm long. Inflorescence an elliptic, lax, open, terminal panicle, 20–45  cm long; main axis scabrous, pulvini shortly hairy, first-order branches opposite or alternate, ascending and diverging, branch axes triquetrous, pedicels glabrous, spikelets congested toward the apex of branches. Spikelets ovoid, 2.8–4 × 1–1.2 mm, open at maturity, glabrous, pale or tinged with purple, rachilla conspicuous between glumes and lower lemma; lower glume 2–3 mm long, 3/4 as

Key to Ungrouped Panicum Sections and Species

63

long as the spikelet, ovate, (3–)5–7 nerved, acuminate to attenuate, glabrous; upper glume as long as the spikelet, ovate, 7–9 nerved, midrib scabrous, otherwise glabrous, attenuate to slightly recurved at apex; lower lemma ovate, 5–7 nerved, glabrous; lower palea elliptic, 2.4–2.6  ×  0.9  mm; lower flower staminate; upper anthecium ovoid, 2–3 × 0.9–1 mm, indurate pale, glabrous, smooth and shiny, without a dark disk at the base and palea without black spots. Caryopsis not seen. Distribution and ecology (Fig. 7). This species is distributed from Mesoamerica and the Caribbean to Venezuela, Colombia, and northern Brazil; it grows on coastal dunes, marsh edges and river beaches, flowers and fruits between August and March. Additional material examined. Mexico. Quintana Roo: Vigía Grande, coastal dune, 26 August 1982, Cabrera 3418 (MEXU, MO) Guatemala. Guatemala city: 1 km south of Amatitlán, 26 July 1970, Harmon 3351 (MO). Petén: Dolores, in pineland, about 3  km northwest of village, 18 October 1961, Contreras 3079 (MEXU) Belize. Swamp near Manatee Lagoon, Peck 123 (GH) Honduras. San Pedro Sula, near Santa Anna, Thieme 532 (USA). Gracias a Dios: Puerto Lempira, west shore of Caratasca Lagoon, 5 September 1978, Nelson 4819 (MO) Colombia. Magdalena: Santa Marta, 1898–1901, Smith 166 (K, NY, P) Venezuela. Delta Amacuro: Dpto. Pedernales, sandy beach bordering open sandy mud flats and wet depressions, mouth of Caño Guiniquina, between Punta Araguabisi and Punta Baja, at Barra Guiniquina, 18 October 1977, Steyermark et al. 114879 (F, MO, NY, VEN); Pedernales, boundary with Dpto. Tucupita, 50 m, mouth of Caño Guiniquina, between Punta Araguabisi and Punta Baja, 18 October 1977, Steyermark et al. 114898 (MO, VEN) Trinidad and Tobago. Icacos, Cedros, along seashore, 12 December 1912, Hitchcock 10154 (NY, US), 10154bis (US); sea beach, 3  miles south of Cedros, Hitchcock 10159 (US); Port of Spain, sandy sea beach, 10 December 1912, Hitchcock 10053 (NY, US); Dabadie, Broadway s.n. (NY, P, US); Mount St. Benedict, 2 November 1923, Broadway 5060 (G); Pigeon Point, 11 October 1909, Broadway 3068 (US); along sea beach, Plymouth, 18 December 1912, Hitchcock 10233 (US); without locality, Broadway s.n. (NY, US) Guyana. Itooribisci Lake, August 1886, Jenman 2261 (US); Parika, 19 November 1919, Hitchcock 16795 (US); Penal Settlement, 3 December 1919, Hitchcock 17144 (US); Bartica, 16 December 1919, Hitchcock 17268 (US); Essequibo, Mission St. Edward, 29 August 1956, Hoock 1243 (US) Brazil. Maranhao: Loreto, 20 March 1985, Valls et al. 8433 (CEN, SI) 7. Panicum amarum Elliott Plants perennial, strongly rhizomatous; culms cespitose, herbaceous, simple, or sometimes branching at the upper nodes, (30–)100–200(–300) cm tall; basal internodes short, glaucous, cylindrical, glabrous, hollow; nodes thickened, brown to pale, glabrous. Sheaths striate, pale, smaller or larger than internodes, overlapping, glabrous or shortly hairy toward upper margins. Ligule membranous-ciliate,

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Fig. 7  Distribution of Panicum altum, P. amarum subsp. amarum, P. amarum subsp. amarulum and P. chloroleucum

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2–3 mm long, the membranous portion much reduced, 0.3–0.5 mm long, with long hairs behind at base of lamina, collar brownish, glabrous. Blades lanceolate to linear-lanceolate, 10–30(–40) × 0.4–1 cm, ascending to divergent, rigid, rounded at the base, the apex long subulate, margins involute, smooth, glabrous, glaucous. Inflorescence terminal, shortly exserted to partially included in the upper leaves, peduncle cylindrical, glabrous; panicle contracted, oblong, few to multiflowered, 30–40  ×  ca. 3  cm wide; main axis wavy, glabrous, pulvini glabrous, first-order branches alternate, adpressed, branch axes and pedicels triquetrous, scaberulous, spikelets congested on short third-order branches. Spikelets lanceolate to longovoid, solitary or in pairs (4.4–)5.2–6  ×  1–2  mm, glabrous, pale or tinged with purple; lower glume long-ovate, (3–)4.4–5 mm long, 2/3 to 4/5 as long as spikelet, ovate, acuminate, clasping, (3–)5–7-nerved, with anastomosing nerves; upper glume as long as the spikelet, 7–9 nerved, acuminate; lower lemma glumiform, 5–7-nerved, acuminate; lower palea long-elliptic, (3.6–)4 × 1–1.5 mm, hyaline, glabrous; lower flower staminate, stamens 3; upper anthecium ellipsoid to ovoid, (2.8–) 3.1–3.5 × 1–1.5 mm, shorter than the upper glume and lower lemma, smooth, glabrous, shiny, indurate, whitish, deciduous at maturity, lemma 5 nerved. Caryopsis not seen. Key to the Subspecies 1. Spikelets 5.2–6 mm long; panicles few flowered, with deflexed branches............. .....................................................7a. P. amarum subsp. amarum 1. Spikelets 4.4–5.1 mm long; panicles multiflowered, with ascending branches...... .........................................................7b. P. amarum subsp. amarulum 7a. Panicum amarum subsp. amarum Sketch, Bot. S.  Carolina 1(2): 121. 1816. Chasea amara (Elliott) Nieuwl., Amer. Midl. Naturalist 2: 64. 1911. Type: USA. South Carolina. “Grows among the sandhills on the seashore, S. Elliott s.n. (lectotype, PH 00018618, designated by P.  Palmer, Brittonia 27: 148. 1975; isolectotypes, K 000674438, PH 000377574, US 00148131) Panicum amarum Elliott var. minor Vasey & Scribn., Dept. Agric. Bot. Div. Bull. 8: 38. 1889. Panicum amaroides Scribn. & Merr., Circ. Div. Agrostol. U.S.D.A. 29: 5-7, Fig. 1. 1901. Type: USA. Virginia: Fort Monroe, 1879, G. Vasey s.n. (lectotype, US 00133067, designated as “type,” by Hitchcock & Chase, 1910: 95; isolectotypes, MO-992999, MO-993000, NY 00381578, US 01117845  W 19160024694, WISv0254154WIS) Distribution and ecology (Fig. 7). It is a frequent subspecies on beaches and coastal dunes, growing from the southern USA to Belize and the Caribbean; flowers and fruits all year round. Additional material examined. USA.  North Carolina: Seacoast east of Wilmington, 28 August 1905, Hitchcock s.n., Amer. Gr. Hb. 43 (MO, P). South Carolina: Charleston Co, Dunes of Sullivan’s Island, 18 October 1968, Leonard & Radford 2170 (MO). Connecticut: seacoast of Conneticut, Torrey s.n. (P). Delaware: Wilmington, October 1864, Canby s.n. (MO). Maryland: Worcester

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Co, Pocomoke, Redmond 1002 (P); Chesapeake Bay, Scientists Cliff, Calvert Co, 1 September 1946, Chase 8407 (MO). New Jersey: Cape May, 18 September 1909, Mackenzie 4452 (MO); Salem Co, sandy beach along Delaware River, near Kellys Point, S of Pennsville, 3 October 1934, Fogg Jr. 7551 (MO). New York: Long Island, Wading River, 11 September 1879, Miller 12001 (MO). Texas: Matagorda Beach, at end of Farm Road 2031, 6.6 miles south of Matagorda, 5 November 1972, Wallen 109 (MO). Virginia: Northampton Co, sandy beach, along Chesapeake Bay, 13 October 1935, Fogg Jr. 10047 (MO); Charles City Co, sandy beach of James River, southeast of Tettington, 19 September 1939, Fernald & Long 11255 (MO) Mexico. Campeche: km 11 de Isla Aguada, 1 April 1982, Brigada de dunas 577 (MEXU). Tabasco: Chiltepec, January 1964, Barlow 12/7A (MEXU). Tamaulipas: Playa Bagdad, 16  km north of Playa Lauro Villar, coastal dune vegetation, 13 February 1985, Baro 633 (MO). Veracruz: Tecolutla, December 1973, Boege 3004 (MEXU) 7b. Panicum amarum subsp. amarulum (Hitchc. & Chase) Freckmann & Lelong, Sida 20(1): 171. 2002. Panicum amarulum Hitchc. & Chase, Contr. US Natl. Herb. 15: 96, Fig. 87. 1910. Panicum amarum var. amarulum (Hitchc. & Chase) P. Palmer, Brittonia 27(2): 149. 1975. Type: USA. Virginia: Virginia Beach, 24 September 1900, T.  A. Williams 3090 (holotype, US 00133068; isotypes, B_10_0366174, ILL 00009931) Distribution and ecology (Fig. 7). It grows in coastal dunes from the USA to the Caribbean, Mexico, and Belize. Additional material examined. USA.  Alabama: Dauphin Island, Cadillac Ave., near Quivira St., 26 August 1965, Deramus 726 (MO). North Carolina: sandy shore of Currituck Sound at Point Harbor, 5 October 1950, Godfrey & Fox 51082 (MO). Florida: Santa Rosa, 29 August 1899, Tracy 6508 (MO); Broward Co., F. Lauderdale-corner Sunset Blvd. & A-la Dr., sandy soil, 14 August 1985, Mckenzie 183 (MO); shores of the ocean and Indian river, Curtiss 3578 (MO); beach ridges near Lake Worth Inlet, 2 September 1895, Curtiss 5527 (MO). Georgia: St. Catherines Island, north end of island, growing on first dunes, 18 November 1983, Coile et  al. 3840 (MO). Oklahoma: Stillwater, 19 July 1897, Myers 450 (MO). Texas: Cameron County, Padre Island, Andy Bowie County Park, 29 October 1970, Woodard s.n. (MO). Virginia: Near Norfolk, 5 August 1890, Vasey s.n. (MO); Lynnhaven, in low sand dunes, seacoast, 2 September 1905, Hitchcock s.n., Amer. Gr. Hb. 44 (MO) Mexico. Campeche: 40  km west of Isla Aguada, Saver et  al. 3350 (USA). Quintana Roo: Isla Mujeres, near north end of island, 17 July 1962, Sauer & Gade 3264 (US); camino rumbo a Punta Molas, 28 August 1985, Ortíz 900 (MO); Puerto Morelos, duna costera, 7 September 1982, Cabrera 3476 (MEXU, MO); a 5 km al sur de la zona urbana de Isla de Mujeres, a 2 km al norte del Parque Natural El Garrafón, 18 September 1985, Cabrera & Cabrera 9236 (MO); 2 km al sur de las ruinas de Tulum, vegetación de dunas costeras, 16 September 1979, Quero et al. 2855 (MO); Punta Nizuc, a 11  km al Este del aeropuerto de Cancún, 31 August 1980, Sousa 11219 (MEXU). Tamaulipas: El Mezquital, 61  km al este de la

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67

carretera a Matamoros, suelo arcilloso-arenoso, 3 October 1984, Baro 513 (MEXU, MO). Yucatán: 3 km al W de Sisal, hacia Celestun por el camino costero antiguo, 12 November 1981, Espejel et al. 331 (MEXU) Belize. Toledo District: Monkey River, near river in broken ridge, 25 August 1941, Gentle 3603 (MO, US) Bahamas. New Providence, Southwest Bay, Britton & Brace 493 (NY) Grand Bahama: in disturbed soil behind sea wall at west end, Correll & Correll 50957 (NY). North Andros: on beach sands along shore road between Nicholls Town and Morgans Bluff, Correll & Evans 44024 (NY). South Andros: along sandy beach area beneath coconut trees just north of Congo Beach Hotel, 23 September 1978, Correll et al. 50262 (NY, US) Cuba. Habana: Río Mayabeque, 6 March 1942, León 20614 (US) Camaguey: Cayo Paredón Grande, Schaffer 2735 (NY, US); eastern part of Cayo Cruz, Schaffer 2780 (NY); Cayo Tabinal, on the open seashore east of Faro Maternillos, 13 October 1922, Ekman 15500 (US). Oriente: Bahía de Manabí, in sand Playa de Muertos, 21 December 1932, León 16027 (US). Pinar del Río: Bahía Honda, sandy beach at the mouth of the bay, 10 June 1929, Ekman 17290, 17291 (US); Bahía Honda, on the sandy seacoast east of Punta Gobernadora, 28 December 1920, Ekman 12698 (G, US) Jamaica. Saint Ann’s Bay and vicinity, coastal sands, Britton 2469 (NY) 8. Panicum chloroleucum Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 262. 1874. Panicum urvilleanum Kunth var. chloroleucum (Griseb.) Kuntze, Revis. Gen. Pl. 3(3): 364. 1898. Type: Argentina. Catamarca: Laguna Blanca, 1872, P. G. Lorentz 445 (lectotype, GOET 006791, here designated; isolectotypes, BAA 00002357, CORD 00004658, CORD 00004659, G 00099649, GOET 006792, US 00133141, US 00148352, W 19160024809). Fig. 8 Plants perennial, cespitose, with elongate rhizomes, cataphylls hairy; culms erect, 0.40–0.70(–1) m tall, internodes cylindrical, 3.4–10 long × 0.2–0.3 cm diameter, hairy, with antrorse hairs, to glabrous, nodes brown, hairy. Sheaths 5–12 cm long, larger than internodes, membranous, pilose to glabrous. Ligule membranousciliate, 1.5–2.5 mm long, collar brown, pilose. Blades lanceolate, 15–30 × 0.1–0.4 cm, densely pilose on both sides, with appressed whitish hairs, the margins involute, scabrous. Inflorescence terminal, a lax, exserted panicle, open, diffuse, 10–33 × 4–14 cm; main axis triquetrous, densely to sparsely hairy, slightly wavy at top, pulvini densely hairy, first-order branches divergent, alternate, rarely opposite, pedicels claviform, densely hairy, 1–6  mm long. Spikelets long ovoid, 3.5–4.9 × 1.2–1.7 mm, sparsely pilose to glabrous, pale or tinged with purple, open at maturity; lower glume ovate, 2.5–4 mm long, 3/4 of spikelet length, pilose toward the apex, 5–7 nerved; upper glume 3.4–4 mm long, pilose toward the apex and over midrib, 7–9 nerved; lower lemma 3.6–4 mm long, glumiform, pilose at the apex and in lower margins, 5–7 nerved; lower palea elliptic, 3–3.8 × 1.1–1.3 mm, membranous, sparsely pilose at the lower margins, apex acuminate; lower flower staminate; upper anthecium broadly ellipsoid, 2.5–3 × 1.2–1.6 mm, smooth, whitish, indurate, with filiform macrohairs on the lower margins of the upper lemma, lemma 7 nerved; stipe absent. Caryopsis ellipsoid, 2.2–2.3 × 1.3–1.5 mm; hilum punctiform, embryo more than 1/2 the length of the caryopsis.

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Fig. 8  Panicum chloroleucum. (a) Plant. (b) Spikelet, lateral view. (c) Lower lemma and dorsum of upper anthecium. (d) Palea and lower flower. (e) Upper anthecium, ventral view. (f) Upper palea and flower. (g) Lodicules. (From Cabrera et al. 32481, SI.)

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Distribution and ecology (Fig. 7). This species is endemic to northwestern Argentina, growing in the provinces of Catamarca, Jujuy, La Rioja, Salta, and Tucumán. It inhabits open and dry places, in sandy and brackish soils, between 1900 and 3500 m asl; flowers and fruits from January to June. Local name: “Jaboncillo” Species previously placed in section Dura Stapf (Zuloaga 1979), related to P. racemosum and P. urvilleanum, from which it differs by having sparsely hairy to glabrous spikelets; additionally, P. racemosum differs by its contracted panicles and larger spikelets, 5.8–7.5  mm long, while P. urvilleanum has spikelets 5–7.3  mm long, with the upper glume and lower lemma 9–15 nerved. Additional material examined. Argentina. Catamarca: Barranca Larga, Schreiter 10252 (GH); Dpto. Belén, Laguna Blanca, 24 February 1981, Cabrera et  al. 32481 (SI, US), 1 April 1982, Gómez Kapla 23 (SI); Nacimientos de San Antonio, Cabrera et al. 32443 (SI, US), November 1998, Korstanje 3 (SI); al pie de la Cuesta de Randolfo, 23 February 1981, Cabrera et al. 32465 (SI). Campo del Arenal, Cabrera et  al. 21814 (LP, MO), 24661 (LP), Joergensen 1720 (MO, SI, US); 21 January 1976, Palacios & Bravo 642 (SI), Morello 881 (LIL); Werner 4 (LP); D’Antoni 114, 115 (LP); Schikendantz 107 (CORD); Ruta Nacional 43, de Villavil a Nacimientos, 8 March 2018, Zuloaga et al. 16116 (SI). Dpto. Santa María, Valle de Santa María, Venturi 6755 (US); al sur de Santa María, 22 January 2012, Zuloaga et al. 13713 (SI); de Ruta Nacional 40 a Fiambalasto, 26 February 2010, Zuloaga et  al. 11875 (SI) Dpto. Tinogasta: between Carrizal and Andaluca, 23 March 1989, Pedersen 15316 (SI), Ruta Nacional 60, de Aimogasta a Tinogasta, 7 March 2018, Zuloaga et  al. 16090 (SI). Jujuy: Dpto. Humahuaca: Cuesta Azul Pampa, 30 March 1995, Deginani & Cialdella 311 (SI); Ruta Nacional 9, Esquinas Blancas, 11 January 2012, Zuloaga et al. 13401 (SI), cerca de Uquía, 12 February 2007, Zuloaga et  al. 9201 (SI); Ruta Nacional 9, 1  km pasando San Roque, 15 February 1997, Zuloaga et al. 5906 (SI). Dpto. Tumbaya: Tres Morros al límite con Salta, 15 February 1980, Cabrera et  al. 31731 (SI, US); Fries 882 (BAA); Ruta Nacional 40 y entrada a El Moreno, 17 January 2012, Zuloaga et al. 13581 (SI); de Tres Morros a El Moreno, 17 January 1988, Zuloaga 3554 (SI). La Rioja: without locality, Morello 841 (LIL). Dpto. Coronel Felipe Varela: Ruta Provincial 76, de Pagancillo a entrada al Parque Nacional Talampaya, 25 January 2012, Zuloaga et al. 13777 (SI), 21 February 2012, Zuloaga et al. 12109 (SI). Salta: Dpto. Cachi: Ruta Provincial 42, Parque Nacional Los Cardones, camino hacia Seclantás, 14 February 2002, Cialdella et al. 254 (SI); Ruta Provincial 42, de recta de Tin-Tin a Seclantás, 21 January 2012, Zuloaga et  al. 13705 (SI); Ruta Nacional 40, Angastaco, 21 January 2012, Zuloaga et al. 13706 (SI). Dpto. Cafayate: Ruta Nacional 68 al norte de Cafayate, 18 March 2018, Zuloaga et  al. 16401 (SI). Dpto. La Poma: Ruta Nacional 40, 57 km de San Antonio de los Cobres, 3 km antes del límite con Jujuy, 19 February 2002, Cialdella et al. 460 (SI); Ruta Nacional 40, entre Pueblo Viejo y Rodeo, 25 February 1987, Nicora et al. 9166 (SI). Dpto. San Carlos; 9 km al S de Angastaco, 15 February 1979, Türpe 3064 (US); entre Angastaco y San Carlos, Paso de las Flechas, 28 March 1979, Cabrera et al. 30778 (SI, US); Ruta Nacional 40, de Cafayate a Cachi, Sierra de Quilmes, 12 February 2002, Cialdella et al. 226

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(SI); 4 km al sur de Angastaco, 19 March 1993, J. H. Hunziker et al. 12539 (SI); Amblayo, río y lomadas al este del pueblo, 1 April 1990, Novara 9744 (MCNS, SI); Ruta Nacional 40 hacia Cachi, al sur de Angastaco, 2 March 2005, Zuloaga et al. 8450 (SI). Tucumán: Dpto. Tafí del Valle: entre Amaicha y Fuerte Quemado, Castillón s.n. (US); Tiopunco, 7 February 1927, Schreiter 4941 (US); Yasamayo, Morello 1234 (SI) 9. Panicum complanatum Guglieri, Longhi-Wagner & Zuloaga, Syst. Bot. 31(3): 506, Figs. 1–4. 2006. Type: Brazil. Minas Gerais: Brumadinho, Parque Estadual do Rola Moça, 6 February 2004, A. Guglieri, H. M. Longhi-Wagner, P. L. Vianna & E. M. Ferreira 283 (holotype, ICN 00029374; isotype, SI 001174). Fig. 9 Plants perennial, cespitose, shortly rhizomatous, (35–)40–60 cm tall; culms simple, few noded; internodes cylindrical, hollow, hairy, with adpressed whitish hairs; nodes hairy. Sheaths longer than the internodes, pale, hirsute, hairs 2–3 mm long, pale. Ligule membranous-ciliate, 0.2–1  mm long, with long hairs beneath at the base of the blade; collar pilose. Blades linear to linear-lanceolate, 14–24 × 0.2–0.4 cm, mostly basal, hirsute, flat to involute, base attenuate, apex subulate. Inflorescence terminal, few flowered, not deciduous at maturity; panicle lax, open, elliptic, 12–20  ×  5–12  cm; peduncles long exserted, ca. 25  cm long, cylindrical, hirsute; main axis hirsute, scaberulous, triquetrous, first-order branches 4-5, alternate, hirsute, divergent from the main axis, pulvini pilose; pedicels solitary, claviform, 3–8  mm long, pilose, more densely so toward the apex. Spikelets ellipsoid, 4.7–5.3 × 1–1.5 mm, solitary or in pairs, glabrous, open at maturity; lower glume 3.5  mm long, 2/3 to 3/5 the length of the spikelet, acute, 5–7 nerved, glabrous; upper glume, subulate, 9 nerved, glabrous, scabrous at apex; lower lemma glumiform, acuminate, 9 nerved; lower palea ovate-lanceolate, (1.3–)2.3–2.5 × 1.2–1.6 mm, glabrous, hyaline; lower flower absent; upper anthecium long ovoid, 3–3.1 × 1–1.2 mm, smooth, hardened, with a continuous row of flattened macrohairs at the base of the upper lemma, not stipitate. Caryopsis ellipsoid, 2 × 1 mm; hilum elliptic, embryo ca. ½ the length of the caryopsis. Distribution and ecology (Fig. 10). Species endemic to Brazil, where it grows in rock fields and savannas in the state of Minas Gerais, at elevations between 900 and 1400 m asl; flowers and fruits between December and March. Panicum complanatum is related to P. olyroides, the latter differing in having flattened macrohairs only on the margins of the upper lemma, panicles deciduous as a whole at maturity and spikelets larger 5–8.5 mm long. Additional material examined. BRAZIL. Minas Gerais: Brumadinho, Retiro das Pedras, Serra da Calçada 5 February 2002, Viana 514 (BHCB); Parque Estadual de Rola Moç, 16 March 2005, Longhi Wagner et al. 9646 (ICN, SI): Ponte Nova, Estaçao Ferroviaria Hargreaves, 21 December 1929, Chase 10238 (MO; US) 10. Panicum deciduum Swallen, Mem. New York Bot. Gard. 9(3): 257. 1957. Type: Venezuela. Amazonas: northwest slopes of Cerro Yapacana, Río Orinoco, 8001000 m, 1 January 1951, B. Maguire, R. S. Cowan & J. J. Wurdack 30517 (holotype, US 00147801; isotype, NY 00381734). Fig. 11

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Fig. 9  Panicum complanatum. (a) Habit. (b) Spikelet, lateral view. (c) Spikelet, dorsal view. (d) Lemma and lower palea, ventral view. (e) Upper anthecium, dorsal view. (f) Upper anthecium, ventral view. (g) Caryopsis, hilum view. (h) Caryopsis, embryo view. (From Guglieri et  al. 283, ICN)

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Fig. 10  Distribution of Panicum complanatum, P. deciduum, P. glabripes and P. longissimum

Plants perennial, shortly rhizomatous, cataphylls glabrous; culms erect, bambusoid, up to 4 m tall, ascending, densely branched at base, with a bud about 3 mm above each node, covered by an ovate, hairy bract; internodes 2–16 cm long, cylindrical, solid in the basal portion, then hollow, glabrous, lustrous, dark green except near the nodal region where they are pale; nodes thickened, forming a membranous disk, pilose toward the basal portion, with whitish hairs, glabrous toward the apex. Sheaths ca. 5 cm long, smaller than internodes, deciduous, pale, glabrous, the lower ones pilose. Ligule membranous-ciliate, reduced, ca. 0.7  mm long; collar pilose, with short, dense hairs. Blades lanceolate, 8 × 0.6 cm, flat, acuminate, subcordate, glabrous, the margins scabriform, blades of the floriferous culms small. Inflorescence terminal, exserted, peduncle cylindrical, glabrous; panicle lax, elliptic, 12–15 × ca. 7 cm; main axis wavy, scabrous to glabrous, branches divergent from the rachis, axis of the branches triquetrous, scabrous, spikelets congested in pairs on scabrous, short pedicels. Spikelets lanceolate, 2.9–3 × 1 mm, pale, densely pilose, glumes and lower lemma acute, rachilla conspicuous; lower glume about equal in length to the upper glume and lower lemma, ovate-lanceolate, 3 nerved, midrib scabrous, glabrous on the inner face and with long hairs toward the apex on the external face;

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Fig. 11  Panicum deciduum. (a) Habit. (b) Ligular region. (c) Portion of inflorescence. (d) Spikelet, lateral view. (e) Lower glume. (f) Upper glume. (g) Spikelet dorsal view without the lower glume, exposing the upper anthecium. (h) Lower lemma and upper anthecium, lateral view. (i) Upper anthecium, ventral view. (j) Upper anthecium, dorsal view. (k) Upper lemma, lodicules, gynoecium, and stamen. (From Maguire et al. 30517, US)

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Taxonomic Treatment

upper glume equal in length to spikelet, 5 nerved, midrib scabrous, with thick tuberculate-based hairs; lower lemma glumiform, 5 nerved, midrib scabrous, with thick tuberculate-based hairs; lower palea lanceolate, hyaline, 2.6–2.8  ×  0.5–0.6  mm, densely pilose, lower flower absent; upper anthecium ellipsoid, 1.5–1.6 × 0.5 mm, shortly stipitate, glabrous, smooth, shiny, about 1/2 the length of the spikelet. Caryopsis not seen. Distribution and ecology (Fig. 10). Species endemic to the Amazonas department, Venezuela, where it grows on Cerro Yapacana, between 800 and 1200 m asl; flowers between January and May Species is known from few collections from the Amazonas department in Venezuela. It is characterized by its bambusoid culms up to 4  m tall, with solid internodes in the basal portion and densely hairy spikelets, covered with thick tuberculate hairs, with the lower glume reaching approximately the same length as the spikelet, the glumes and lower lemma with a manifest rachilla, the upper glume and lower lemma 5 nerved and the upper anthecium stipitate. Additional material examined. Venezuela. Amazonas: Cerro Yapacana, 5 May 1970, Steyermark & Bunting 103118 (MO, P, VEN) 11. Panicum glabripes Döll, Fl. Bras. 2 (2): 216. 1877. Type: Uruguay. Montevideo, F.  Sellow 130 (holotype, B_10_0158003; isotypes, BAA 00000904, US 00148529f, possible isotype, K 000309192). Fig. 12 Panicum subjunceum Ekman, Ark. Bot. 10(17): 16, Table 5, Fig. 2, Table 6, Fig. 8. 1911. Panicum glabripes Döll var. subjunceum (Ekman) Roseng., B.R. Arrill. & Izag., Gram. Urug.: 335, Fig. 142. 1970. Type: Brazil. Paraná: Serrinha, in campo subhumido, 9 December 1908, P. Dusén 7339 (lectotype, S-R-4014, here designated; isolectotypes, B_10_0366243, BAA 00000929, S-05-9526, S-05-9527, US 00140033, US 00148040, US 00148041, W 19160039078) Plants perennial, cespitose, rhizomatous, with elongated rhizomes, cataphylls hairy; culms erect, 0.8–1.60 m tall, herbaceous, non-bambusoid, simple, internodes rigid, cylindrical, hollow, pale, shortly pilose, 4–32 cm long; nodes glabrous, dark. Sheaths not pale, larger than internodes, 8–23  cm long, the basal ones 9–41  cm long, not rigid and achillate, upper margins ciliate, glabrous on the rest of the surface, not keeled, membranous. Ligule membranous-ciliate, 0.8–3 mm long; collar glabrous, with purplish tints. Blades lanceolate, (6–)16–45 × 0.2–0.4(–1) cm, rigid, keeled, narrowed toward the base, apex subulate, the margins involute and scabrous, adaxial surface densely pilose, with caducous hairs, abaxial surface glabrous. Inflorescence terminal, exserted, panicle lax, pyramidal to contracted, multiflowered, 15–33 × 3–12 cm, not deciduous at maturity; main axis glabrous to scabrous, cylindrical to wavy, scabriform, pulvini glabrous, first-order branches alternate or opposite, branch axes scabrous; pedicels 0.8–2.5 mm long, with one or two long, deciduous hairs, to glabrous. Spikelets ovoid, 2.4–3.3 × 0.7–1.2 mm, solitary, glabrous, open at maturity, pale and tinged with purple; lower glume ovate, 1.4–2.4 mm long, (1/2–) 3/4 the length of the spikelet, (3–)5–7 nerved, separated from the upper glume by a short internode; upper glume ovate, equal in length to the spikelet, 7–9

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Fig. 12  Panicum glabripes. (a) Plant. (b) Detail of ligular region. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Spikelet, lateral view. (f, g) Lower palea. (h) Upper anthecium, dorsal view. (i) Upper anthecium, ventral view. (j) Upper palea with lodiculae. (k) Caryopsis, embryo view. (l) Caryopsis, hilum view. (From Bacigalupo et al. 1041, SI)

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Taxonomic Treatment

nerved, acuminate; lower lemma glumiform, 7–9 nerved, acuminate; lower palea elliptic, 1.8–2.5 × 0.7–1 mm, membranous, glabrous; lower flower staminate; upper anthecium long-ovate, 1.7–2.1 × 0.7–1(−1.2) mm,, glabrous, smooth, shiny, indurate, pale, with a dark disk next to its insertion and two dark spots along the palea; stipe absent; lemma 7 nerved. Caryopsis broadly ellipsoid, 1.5  ×  1  mm, brown; hilum oblong, embryo more than 1/2 the length of the caryopsis. Distribution and ecology (Fig. 10). Species restricted to southern Brazil, Uruguay, and Argentina. It is found forming grasslands in humid places next to watercourses. It flowers from October to January. When describing P. subjunceum, Ekman cites Dusén 7339 as the type collection, without indicating the herbarium where it was deposited. In the herbarium S, the base of this author’s work, we located three specimens of this collection, of which the specimen S-R-4014 was selected as lectotype, for being a complete specimen, coinciding with the protologue and for possessing the original label, in Ekman’s handwriting, indicating “Panicum subjunceum Ekman Orig. spec.!” Panicum glabripes is morphologically related to P. tricholaenoides Steud, the latter species being distinguished by its densely branched, subwoody, bambusoid culms, with aphyllous sheaths at the base of the culms or with reduced blades and spikelets on pedicels with long, persistent, whitish hairs. Additional material examined. Brazil. Mato Grosso do Sul: Corumbá, 14 November 1977, Allem et al. 1194 (CEN). Paraná: Pinhais, Jonsson 1124a (F, G, NY, SI); Serra do Mar, Ypiranga, 8 November 1911, Dusén 13333 (G, MO, NY, SI); Serrinha, Dusén 15573 (F, G, MO, SI); Banhado, Dusén 16128 (G); São José dos Pinhais, 25 December 1951, Hatschbach 2648 (SI), 8444 (MBM); Rio Taquarí, Hatschbach 35527 (MBM). Rio Grande do Sul: Pelotas, 12 December 1962, Kappel & Froner s.n. (RSPF-686); Pelotas, Escola Agrotecnica, Gilberto 7 (F, MO), Araujo 216 (SI); São Leopoldo, Dutra 66 (IAN, R), 290 (R), 632 (ICN). Tocantins: Ilha do Bananal, Parque Nacional do Araguaia, 27 March 1999, Mendonça et al. 4507 (SP) Argentina. Buenos Aires: Archipiélago Solís, Isla Sur, 16 December 2006, Hurrell et  al. 6303 (CTES). Corrientes: Dpto. Concepción: Tatacuá, Pedersen 10286 (NY, P); Colonia Tatacuá, banquina alta Ruta 117, 20 October 1979, Carnevali 4095 (CTES); 7 km E de Saladas, 18 November 1977, Quarín et al. 3877 (MO). Dpto. Lavalle: Ruta 22, Estancia Sr. Sandiano, 26 October 1983, Goldfarb 618 (CTES). Dpto. Mercedes: Estancia “Ita Caabo,” Pedersen 6112 (P); 75  km norte de Mercedes, Laguna Trin, Estancia Culantrillar, 12 October 1975, Quarín 3188 (CTES, G, LP). Dpto. San Roque: cruce de Ruta 9 de Julio, 18 October 1982, Fernández 884 (CTES, MO, SI); Ituzaingó, Ruta Nacional 12, 5  km de Arroyo Itaembé, 24 October 1974, Tressens et  al. 414 (CTES, SI). Entre Ríos: Dpto. Colón: Colón, Parque Nacional El Palmar, Arroyo El Palmar, 3 December 1973, Burkart 30232 (F, SI, US); Dpto. Concordia: cerca de Salto Grande, Arroyo Gualeguaycito, Parodi 61 (BAA, BAF), 3939 (BAA); Pedermar, Friedman 280 (SI); Concordia, Ayuí, 14 November 1979, Troncoso et al. 2721 (SI, US); Colonia Ayuí, 29 November 1988, Bacigalupo et  al. 1041 (SI); Concordia, vías ferreas, 11

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December 1969, Burkart & Troncoso 26133 (SI); Concordia, Calabacilla, desvío a Nueva Escocia, 24 November 1988, Bacigalupo et al. 844 (SI). Dpto. Federación: Prefectura Naval Santa Ana, 12 December 1986, Guaglianone et al. 463 (SI). Santa Fe. Dpto. San Javier: entre Alejandra y San Javier, 26 November 1937, Ragonese 2751 (BAA) Uruguay. Río Negro: Tres Bocas, Estancia El Rosario, 4 November 1994, Marchesi s.n. (MVFA) 12. Panicum longissimum (Mez) Henrard, Meded. Rijks-Herb. 54: 380. 1927. Sacciolepis longissima Mez, Repert. Spec. Nov. Regni Veg. 15: 122. 1918. Type: Paraguay. Paraguarí: Yaguarón, dans les prairies, 1879, B. Balansa 2935 (lectotype, B_10_0841262, here designated; isolectotypes, BAA 00001056, BAA 00003963, BM 000524507, C 100007855, G 00030926, G 00030297, G 00030298, G 00009730, K, MVFA 0000604, P 00740905, P 00740902, P 00740903, P 00740904, US 00141334, US 00141333). Fig. 13 Plants perennial, strongly rhizomatous, with elongated rhizomes, culms erect 1.60–1.80 m tall, with hairy cataphylls toward the edges, culms rigid, simple in the basal portion, branching at the upper nodes; internodes cylindrical, hollow, glabrous; nodes glabrous. Sheaths striate, rigid, keeled, basal sheaths larger than the internodes, upper sheaths shorter, glabrous, the margins shortly ciliate. Ligule membranous-ciliate, 2 mm long, the membranous portion 0.5 mm, shortly hairy on the abaxial side, ciliate portion 1.5  mm long, collar brown, glabrous. Blades linearlanceolate, 65–80 × 0.6–0.9 cm, reduced at the upper nodes, membranous, the margins involute, markedly scabrous, base rounded and apex long subulate, adaxial surface densely hispid, the abaxial surface glabrous. Inflorescence terminal, long exserted, peduncle cylindrical, glabrous, 40–50  cm long; panicle spiciform, 10–20 × 1–3 cm, main axis wavy, scaberulous, pulvini shortly hairy to glabrous, brownish, branches alternate, the lower ones divergent from the rachis, upper ones adpressed, or the entire panicle contracted with adpressed branches; spikelets congested on short pedicels on third-order branches, branches and pedicels scabrous, pedicels with or without long whitish hairs, solitary or in pairs. Spikelets narrowly ellipsoid, open at maturity, 3 × 1.2 mm, glabrous, greenish or tinged with purple; lower glume 1.5  mm long, 1/2 the length of the spikelet, ovate, acute, 5 nerved, clasping, midrib scabrous toward the apex; upper glume and lower lemma subequal, or the upper glume sometimes somewhat smaller; upper glume ovate, acute, 7–9 nerved, midrib scabrous the toward apex; lower lemma 7–9 nerved, strongly nerved; lower palea elliptic, hyaline, 2.5 × 1 mm, glabrous; staminate flower present; upper anthecium ovoid, 2.4 × 1 mm, glabrous, smooth, shiny, pale, not stipitate, drooping at maturity before the glumes. Caryopsis not seen. Distribution and ecology (Fig. 10). Species endemic to eastern Paraguay, where it grows in humid fields Local names: “Kapi’í takuarí” (kapi’í = straw, takuarí = tacuarilla), “takuari ñu” (ñu = field) (Zuloaga 1994)

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Fig. 13  Panicum longissimum. (a) Inflorescence and leaf. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Spikelet, lateral view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (From Balansa 2935, G)

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Species is only known from the type collection. Originally described in the genus Sacciolepis, this rare species is characterized as a robust plant, with spiciform inflorescences bearing glabrous spikelets, with the lower glume half the length of the spikelet and lower palea present, with a staminate lower flower. 13. Panicum mystasipum Zuloaga & Morrone, Iheringia, Bot. 42: 14, Figs.  1–4, 9–14. 1992[1993]. Type: Brazil. Distrito Federal: rodovia BR-020, campo limpo entre Sobradinho e o Centro de Pesquisas Agropecuarias do Cerrado (CPAC/ Embrapa), 28 November 1988, J. F. M. Valls, Faraco & F. O. Zuloaga 11951 (holotype, CEN; isotype, SI 002811). Fig. 14 Plants perennial, cespitose, shortly rhizomatous, cataphylls glabrous; culms erect, 0.45–1 m tall, few noded, short, internodes cylindrical, finely striate, hollow, glabrous; nodes glabrous. Sheaths longer than the internodes, 6–14 cm long, hirsute toward the distal portion, glabrous elsewhere, margins membranaceous, ciliate toward the apex. Ligule membranous-ciliate, 0.8–1.2 mm long, with long whitish hairs behind at the base of the blade; collar pilose, with long whitish hairs. Blade linear-lanceolate, 20–45 × 0.3–0.5 cm, rigid, pungent, flat, the basal ones senescent and in rolled, the adaxial surface sparsely pilose, the abaxial one scaberulous, base narrowed and apex long attenuate, more or less pungent, the margins involute, ciliate toward the basal portion, otherwise scaberulous. Inflorescence terminal, exserted, peduncle glabrous, up to 70  cm long, wavy, glabrous to sparsely hairy; panicle lax, open, ovate, 9–20 × 1–6 cm, few flowered with 10–20 spikelets, not deciduous overall at maturity; main axis triquetrous, glabrous to sparsely hispid, wavy, scaberulous on the margins; pulvini glabrous to hairy, with whitish hairs; first-order branches alternate, appressed or slightly divergent from the main axis, the lower ones up to 12 cm long; pedicels solitary, wavy, scabrous, appressed to the branches. Spikelets long ellipsoid, 7.2–9.2  ×  2–2.4  mm, open at maturity, glabrous, attenuate toward the base, pale, tinged with purple toward the apex of bracts; rachilla internodes conspicuous between glumes and upper anthecium, hairy; upper glume and lower lemma subequal, longer than the upper anthecium, with manifest nerves; lower glume ovate, 4.8–5.8 mm long, ½–3/4 the length of the spikelet, acute, 7–9 nerved, midrib scabrous toward the upper portion; upper glume as long as the spikelet, abruptly acuminate, 9–11 nerved, midrib scaberulous toward the apex; lower lemma glumiform, 7–9 nerved; lower palea lanceolate, 4.8–5.2 × 1 mm, glabrous, hyaline, with ciliate margins; lower flower absent; upper anthecium ellipsoid, 4.8  ×  1.6  mm, hardened, pale and tinged with black, smooth, stipitate, stipe ca. 0.5 mm long; upper lemma 7 nerved, with long flattened macrohairs toward the basal margins; upper palea 2 nerved. Caryopsis ellipsoid, 3.5 × 1.5 mm, pale; hilum punctiform, embryo ½ or less the length of the caryopsis. Distribution and ecology (Fig. 15). Species endemic to Brazil, where it grows in the Federal District and Goiás, in cerrado subject to periodic fires; it flowers and fruits between November and May. Panicum mystasipum is a species related to P. olyroides Kunth, from which it is separated by having smaller, few flowered panicles, 9–20  ×  1–4  cm, with 10–20 spikelets arranged on the first-order branches, the plants being usually smaller and with senescent leaves rolled at the base; in addition, P. olyroides is distinguished by

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Taxonomic Treatment

Fig. 14  Panicum mystasipum. (a) Plant. (b) Spikelet, lateral view. (c) Upper anthecium, ventral view. (d) Upper anthecium, lateral view. (From Valls et al. 11951, SI)

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Fig. 15  Distribution of Panicum mystasipum and P. olyroides

its drooping inflorescences at maturity, its spikelets lacking rachillae with conspicuous internodes between the bracts and upper anthecium not stipitate. Panicum mystasipum is a cespitose plant, with very short basal internodes, and sheaths persisting at the base of the plant. Blades in this species are rigid and pungent, and when aged they persist enrolled at the base of the plant. The persistence of enrolled leaves at the base of the plants is probably related to the protection of the buds from adverse environmental factors. Panicum mystasipum grows in open fields subject to periodic fires. The cerrado has, according to Eiten (1978), an annual rainfall of 750–2000 mm, with a marked dry season (between May and September) with 10–30 mm or no rainfall in some years. Panicum olyroides shares the same habitat as P. mystasipum, with both species growing frequently in open and closed fields and savannas, in sandy soils, with a predominance of grasses and low xeromorphic trees. Additional material examined. Brazil. Distrito Federal: Fazenda Agua Limpa, cerrado, sensu stricto, April 1983, Haridasam 091 (UB); Fazenda Agua Limpa, 6 February 1980, Lenz Cesar 185 (IBGE), 6 March 1980, Lenz Cesar 385 (IBGE), 15 April 1980, Lenz Cesar 449 (IBGE), 22 May 1980, Lenz Cesar 611 (IBGE, UB); Fazenda Agua Limpa, UnB. ca 1 Km do cato sul da cerca da Reserva

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Taxonomic Treatment

Ecológica do IBGE, 8 January 1990, da Silva & Lopes 923 (IBGE); rodovia BR-020, campo limpo entre Sobradinho e o CPAC, Filgueiras & Zuloaga 2002 (IBGE, SI). Goiás: Parque Nacional das Emas, Mineiros, campo limpo, 22 May 1993, Filgueiras 2479 (SI) 14. Panicum olyroides Kunth Plants perennial, cespitose, shortly rhizomatous, with simple culms 0.40–1 m tall; internodes cylindrical, pale, glabrous or sparsely hairy below the nodes; nodes glabrous, purplish. Sheaths commonly larger than the internodes, densely papillose-pilose toward the upper region to sparsely pilose, with few hairs scattered on their surface, or glabrous. Ligule membranous-ciliate 0.7–2.5 mm long; collar usually pilose. Blades linear-lanceolate, (10–)18–40 × 0.7–1 cm, flat or with slightly involute margins, membranous, glabrous or with the abaxial surface pilose to densely pilose, the margins smooth toward the upper portion and ciliate toward the base. Inflorescence terminal, lax, pyramidal, multiflowered panicle, 20–45 × 7–23 cm, deciduous at maturity; main axis scabrous to ciliate or densely pilose, pulvini pilose to glabrous, lower branches whorled to opposite, the upper alternate, axes of the branches and pedicels pilose to scabrous, spikelets oblique on pedicels. Spikelets long ovoid, 5–8.5  ×  1.7–2.2  mm, solitary and in pairs, open at maturity, glabrous, greenish to brownish, glumes and lower lemma subequal, larger than the upper anthecium, with manifest nerves; lower glume 2.5–5.5 mm long, (1/2–)3/4 the length of the spikelet, ovate, acute, 5–7 nerved; upper glume as long as the spikelet, 9–11 nerved, acuminate to attenuate at the apex, midrib scabrous at the apex; lower lemma 9–11 nerved, acuminate to attenuate; lower palea elliptic, 1.4–4 × 0.7–1 mm, hyaline, glabrous or with the margins shortly ciliate; lower flower absent; upper anthecium ovoid, 2.5–4.3 × 1.5–2 mm, indurate, smooth and shiny, pale, not stipitate, with flattened macrohairs toward the basal margins of the upper lemma. Caryopsis ovoid, 1.8–3 × 1.6–1.9 mm; hilum punctiform, embryo more than half the length of the caryopsis. Local names: “Paja quebradiza,” “trigo sabanero” This species is characterized by the presence of flattened macrohairs on the lower margins of the upper lemma. In addition, the panicle falls as a whole at maturity and is thus dispersed by the wind, a character also present in Panicum bergii Arechav. Key to Varieties 1. Sheaths, blades, and panicles glabrous or with few hairs scattered on their surfac e......................................................................14a. P. olyroides var. olyroides 1. Sheaths, blades, and panicles densely pilose.............14b. P. olyroides var. hirsutum 14a. Panicum olyroides Kunth var. olyroides, Nov. Gen. Sp. (quarto ed.) 1: 102. 1815[1816]. Panicum olyroides var. fimbriatum Döll, in C.  Martius, Fl. Bras. 2(2): 230. 1877, nom. illeg. superfl. Type: Venezuela. “Panicum olyroides H.B.K. in locis aridis Nova-Andalusia in demisis Montis Impossible,” Sep, F. W. H. H. A. von Humboldt & A. J. A. Bonpland s.n. (lectotype, P 00371639, designated as “holotype,” by E.  Judziewicz, 1990: 407; isolectotypes, B_10_0158001, BAA 00000914, US 00139814). Fig. 16

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Fig. 16  Panicum olyroides var. olyroides. (a) Plant base and inflorescence. (b) Spikelet, lateral view. (c) Spikelet, dorsal view. (d) Lower palea. (e) Upper anthecium, ventral view. (f) Caryopsis, embryo view. (g) Caryopsis, hilum view. (From Krapovickas 23280, CTES)

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Taxonomic Treatment

Panicum proboscideum Trin, Gram. Panic.: 184. 1826. Type: Brazil: “in campis siccis graminosis,” G.  H. von Langsdorff s.n. (L.  Riedel s.n.) (holotype, LETRIN-0902.01; isotypes, P 00740768, P 00740767, US 00147979, US-00177979; probable isotype, L. Riedel 2150, K 000309190) Panicum funckianum Steud, Syn. Pl. Glumac. 1: 77. 1855[1853]. Type: Venezuela. Guanaguana, Caracas, 1843, N. Funck 112 (lectotype, P 00740769, here designated; isolectotypes, G 00099720, G 00099721, P 00740770, P 00740771, US 00148511, US 00148512, W 18890112358, W 19160024382) Distribution and ecology (Fig. 15). It usually inhabits savannas or fields, in open and dry places. It is found from Venezuela, Guyana, and Colombia to Bolivia, Paraguay, southern Brazil, and Argentina, from sea level to 1600 m asl. It flowers throughout the year; it is favored by cattles. Local names. Paja peluda, jiwuasinaman, tosé (Guahiba language), yakani yĩñagopeha (Sáliva language) (Zuloaga and Giraldo Cañas 2013) There are three collections of Funck 112 at P, of which P 00740769 is designated as lectotype of P. funckianum; it agrees with the protologue and is a full specimen. Additional material examined. Colombia. Arauca: 55 km SE of Arauca, 19 August 1963, Blydenstein 1524 (US). Casanare: margen derecho del río Casanare, caserío indígena Tsamani, 31 July 1977, Silva 287 (COL); Morichito, 10 July 1977, Niño 99 (COL); Aguas Claras, 9 July 1963, Saravia 2664 (COL); Llanos Orientales, al Sur de El Yopal, 12 July 1963, Blydenstein & Saravia 1333 (COL, MO) Cundinamarca: Monte Redondo, south of Quetame, Pennell 1832 (NY). Guaviare: municipio San José del Guaviare, 290  m, November 1995, Giraldo-Cañas 2626 (COAH, HUA), s.d., Marulanda 1257 (HUA). Meta: 2 km pasando Villa Nueva, 600 m, campo natural arenoso, 19 June 1989, Zuloaga & Londoño 4104 (COL, MO, SI); Cumaral y Restrepo, Garcia Barriga 4945 (COL); alrededores del Hato San Jorge, km 67 entre El Fundo y La Cuerera, Pinto & Sastre 1063 (COL); Reserva Nacional de La Macarena, Savanna at El Refugio, right bank of rio Guayabero, Thomas et al. 1579 (COL); Llanos Orientales, Mesa de San Juan de Arama, 17 April 1963, Blydenstein 777 (COL); 15 km este de San Martín, al sur de Caño Camoa, Blydenstein 1644 (COL); 5  km al sudoeste de Puerto López, Blydenstein 1748 (COL); Los Llanos, Matabubosa, 29 October 1938, Cuatrecasas 4258 (COL, US); 43  km NEE of Puerto López, along road to Puerto Gaitán, 24 December 1973, Davidse 5108 (COL, MO); Villavicencio, Pennell 1445 (NY, US). Santander: Mesa de los Santos, Killip & Smith 15238 (US). Vichada: ca. 8  km E of Las Gaviotas, along the unimproved dirt road to Santa Rita, 25 December 1973, Davidse 5132 (COL, MO, NY); cerca del campamento Las Gaviotas, 22 June 1972, Cabrera 2334 (COL). Llano de San Martín, Triana 40 (COL) Venezuela. Anzoátegui: carretera El Tigre a Soledad, Muller 814 (VEN); Distr. Bolívar, just S of El Zamuro, Fila El Purgatorio, 9 airline km northeast of Bergantín, 1100 m, November 1981, Davidse & González 19313 (MO, NY, VEN); sabanas al este de Santamé, Pittier 14433 (VEN); Las Piedritas, 16 July 1946, Burkart 17269 (VEN); near Santamé, Adams 6 (US); 12  km al oeste de El Tigre, carretera a Pariaguán, 250 m, 22 August 1989, Zuloaga et al. 4502 (MO, SI, VEN). Aragua:

Key to Ungrouped Panicum Sections and Species

85

Guacamaya, Ramia 3884 (VEN); Parque Nacional Henry Pittier, Rancho Grande, alrededor del km 12 carretera Maracay-Ocumare, 31 August 1979, Huber & Manara 4253 (MO, VEN) Barinas: 16 km southwest of the Mérida intersection just outside of Barinas, 250 m, hilly savanna with a few stunted Curatella trees, 18 November 1971, Davidse 3160 (MO). Bolívar: Sur de Ciudad Bolívar, Alva 53-10 (US); Gran Sabana, South of Mount Roraima, Steyermark 59240 (F, US); 5  km del Hato de Nuria, al este de Miamo, Steyermark 88378 (NY, US, VEN); Sabanas del Hato Divina Pastora, Tamayo 2932 (US, VEN); Puerto Ordaz, Aymard et al. 296 (PORT); Distr. Roscío, sabanas secundarias al sur del caserío Chirimata, 22 km al este/noreste de San Ignacio de Yuruaní, Huber & Alarcón 7655 (VEN); Distr. Roscío, Valle medio del Río Aponguao, approx. 8–10 km al noroeste del Caserío Chinikén, 950 m, Huber & Alarcón 10470 (NY); Distr. Roscío, approx. 3  km al noroeste de San Ignacio de Yuruaní, 900 m, June 1983, Huber & Alarcón 7509 (MO, NY); Distr. Roscío, sabanas en los alrededores de San Ignacio de Yuruaní, 920  m, October 1982, Huber & Febres 6807 (MO, NY); entre Guasipati y El Cintillo, 200 m, 16 August 1989, Zuloaga et al. 4481 (MO, SI, VEN); San Pedro de las Dos Bocas, west bank of river, forest along river, adjacent savanna and slopes, 200  m, July 1978, Liesner & González 5482 (MO, NY, VEN); Distr. Piar, trail between Karamata and Uruyan, 480  m, 23 November 1982, Davidse & Huber 22508 (MO, VEN); Distr. Piar, Río Ocoine, Pie de la Roca, southeastern base of Auyan-tepui, 500 m, 24 November 1982, Davidse & Huber 22591 (MO). Carabobo: Chirgua valley, Rudd 293 (VEN); Hacienda de Cura, cerca de San Joaquín, 15 August 1918, Pittier 8004 (US, VEN). Distrito Federal: en sabanas de la falda del Ávila, Tamayo 1379 (F, VEN); Caracas, Elías 204 (F); cerca de Caracas, 24 June 1917, Pittier 7224 (US). Guárico: 10 km NWN of Altagracia de Orituco along hwy. to Caucagua, 440 m, 18 November 1973, Davidse 4162 (MO). Lara: Terepaima camino sur, Barquisimeto, Smith V6410 (VEN). Mérida: cerca de Tovar, 31 January 1928, Pittier 12789 (US). Miranda: Silla de Caracas, bajada desde La Loma hasta Quda.Quintero, 1200–1300  m, Morillo & Manara 1729 (VEN). Monagas: Sabana de Jusepín, Aristeguieta & Virrueta 7610 (VEN); Hato de los hermanos Palacios, 8 km al S de Maturín, Badillo 3696 (VEN); Jusepín, 150 m, August 1989, Zuloaga et al. 4380 (MO, SI, VEN). Sucre: Distr. Sucre, sabana pedregosa, La Lomita entre La Sabana y Zurita, 500–600 m, Steyermark et al. 107780 (NY, VEN). Trujillo: cercanías de La Cristalina, 1600–1900  m, Tamayo 1866 (VEN). Zulia: Perijá, 30 December 1950, Hno. Ginés 1476 (USA) Guyana. Guaranama Ranch, 4 June 1958, Harrison & Pensant 967 (US). Brazil. Distrito Federal: Acampamento do D.V.O., Belem & Mendes 77 (CEPEC); Brasília, Bacia do rio Sao Bartolomeu, Equipe IBGE 2928, 3000, 3049, 4036 (IBGE); Campo no lago Norte, Filgueiras 817 (IBGE); Planaltina, CPAC, Chapada entre Sobradinho e CPAC, Almeida 808 (IBGE); Reserva Ecologica do IBGE, Equipe IBGE 4338 (IBGE); Reserva, area de coleta n. 1, entre o Corrego Escondido e o Cristo Redentor, Silva et al. 371 (IBGE); DF 3 rodovia, 6 November 1976, Allem 390 (MO); Ponte Alta-Gama, November 1976, Allem 345 (MO); without locality, Riedel s.n. (P). Goiás: 18 km SW of Abadiânia along highway BR-060, 9 April 1976, Davidse et al. 12182 (MO, SP); Serra do Facão, 35 km northeast of

86

 

Taxonomic Treatment

Catalão, 24 January 1970, Irwin et al. 25288 (F, NY, US); ca. 58 km northeast of Catalão, 26 January 1970, Irwin et  al. 25431 (F, MO, NY, US); Corumbá, Sa. Pireneus, Burman & Filgueiras 432 (IBGE); Edeia, Fazenda Paraiso, Local Da. Linha, Helano 373-33 (IBGE); Goiandira, 26 March 1930, Chase 11587 (US); 12  km  S of Guará, 20 March 1968, Irwin et  al. 21569 (F, MO, RB, USA); Niquelândia, Irwin et  al. 34617 (SP); gravelly hillside and sandy cerrado, ca. 15 km S of Niquelândia, 21 January 1972, 21 January 1972, Irwin et al. 34677 (F, MO, NY, US); 17 km west of Rio Verde, Chase 11707 (USA); Santo Antonio do Descoberto, Equipe IBGE 1679-A (IBGE); Planaltina, São Gabriel, margem da Rodovia São Gabriel, Filgueiras 436 (IBGE); São João D’Alianca, Fazenda Corrente, Chagas et al. 165 (IBGE); Serra dos Pirineus, 20 km east of Pirenópolis, 1000 m, 16 January 1972, Irwin et al. 34363 (MO, US); between Viannapolis and Ponta Funda, Chase 11324 (US); between Yatai and Araguaia river, Chase 11756 (US); without locality, Gardner 4053 (US); Parque Nacional das Emas, Mineiros, campo limpo, 22 May 1993, Filgueiras 2509 (IBGE, SI). Maranhão: Carolina to San Antonio do Balsas, Swallen 4130 (R, RB, US). Mato Grosso: Aquidauana, Chase 11054 (US); Chapada dos Guimaraes, Mileski 106 (RB); entre Diamantina y Cabeceira do Lobo, Kuhlmann 1743 (RB); Santa Rita do Araguaia, Chase 11860 (US); northwest of São Lourenço, Chase 11952 (US); Tres Lagoas, Chase 10731 (US). Mato Grosso do Sul: Campo Grande, Chase 10785 (US); between Campo Grande and Dourados, Chase 10890 (US); Antonio João, arredores, March 1985, Hatschbach 49308 (MO). Minas Gerais: Chapeu do Sol, Serra do Cipó, Chase 9230 (US); Lagoa Santa, 42 km north of Belo Horizonte, Chase 9046 (US); Lavras, 10 March 1925, Chase 8804 (F, MO, NY, USA), Matosinhos, Handro 77 (SP); Paraopeba, de Paula 75 (SP); Pilhoes, 3 February 1950, Macedo 2102 (MO, USA); Poços de Caldas, Chase 10625 (US); Serra do Sendeiro, Glaziou 17947 (P); between Sucupira and Omega, S of Uberlandia, Chase 11179 (US). Pará: Cuminá, Campos Gerais, Sampaio 5570 (R); Rio Paru de Oeste, Cavalcante 828 (US). Paraná: Campo Morao, Swallen 8989 (US); 43  km west of Guarapuava, along highway BR-277 to Foz do Iguacú, 14 March 1976, Davidse et  al. 11298 (MO, SP); Guarapuava, Swallen 8884 (US); by Jaguariava boundary, 17 January 1975, Smith et  al. 14724 (F, MO); Ponta Grossa, Vila Velha, Pereira 5201 (RB, US); Ponta Grossa, Hoehne 23272, Swallen 8312, 8773 (US); 10  km este de Vila Velha, Krapovickas et  al. 23280 (CTES); Barra do Perdizes, Hatschbach 7939 (MBM, US); Rio Passa Dois, Hatschbach 14042 (MBM, US); Tamandua, Dusén 7709 (US); Tibagí, Fazenda Monte Alegre, Harmonia, Hatschbach 3035 (MBM, SI); Jaguariaiva, in campo, 740 m, Dusén 16579 (US). Rio Grande do Sul: Furma do Tigre, Rambo 44477 (US); Morro Santa Teresa, cerca de Porto Alegre, Rambo 41286 (US); 11  km southeast of Sarandi along Highway BR-386 to Carazinho, 620 m, March 1976, Davidse et al. 11173 (MO, SP); Soledad, Jürgens 419 (US); Vacaría, Valls & Arzivenco 1352 (US). Santa Catarina: entre Abelardo Luz e Clevelandia, Castellanos 24653 (R); N of Abelardo Luz, Smith & Klein 11453 (US); 20 km south of Campos Novos, Smith & Klein 11986 (R, US); 10 km south of Lages along Highway BR-116 to Porto Alegre, 930  m, March 1976, Davidse et al. 11126 (MO); Vacaría, BR-116, no km 205,6, 15 November 1972, Valls et al.

Key to Ungrouped Panicum Sections and Species

87

2443 (CEN, SI). São Paulo: Botucatú, Delanova s.n. (SP-123458); 18 km north of Botucatú, along the S. Manuel-Piracicaba highway, Gottsberger 37 (SP); Campinas, Viegas s.n. (US 1761518); Ibaté, 290  km a lo largo de ruta Washington Luiz, Sendulsky 11 (SP, US); Itapetininga, Sonia de Campos 38 (SP, US), Lima s.n. (RB 60685); São Jose dos Campos, Lofgren 230 (RB); Moji-Guaçu, Eiten et al. 2577 (SP); Pirassununga, Rachid 15 (SP); Mun. Itapetininga, along highway from Sorocaba to Itapetininga, at ca. km 157, 4 February 1965, Eiten et al. 5815 (MO) Bolivia. Santa Cruz: entre los ríos Piraí y Cuchi, Hertzog 1445 (US); Buena Vista, Steinbach 5378 (NY), 11 March 1925, 6978 (MO); Florida, Samaipata, 1600  m, on rocky slopes, 23 March 1981, Renvoize & Cope 4053 (MO), Prov. Velasco, Serranía de Huanchaca, Parque Nacional Kempff Mercado, 800  m, 3 December 1987, Thomas et al. 5567 (MO); Ñuflo de Chavez, Concepción, laguna west of town, 25 March 1986, Killeen 1847 (MO); Ñuflo de Chavez, Estancia Las Madres on road to Monte Verde, February 1986, Killeen 1769 (MO); Warnes, Pampa Viru Viru, October 1985, Killeen 1257 (MO); Andrez Ibañez, 1 km east of Int. airport Viru Viru, 16 km N of Santa Cruz, 400 m, 1 January 1986, Killeen 1559 (MO); Estancia El Recreo, 2  km north of Concepción, 480  m, 21 March 1987, Killeen 2398 (MO, SI), Ñuflo de Chavez, 8 km S of Concepción on road to Lomerio, 480 m, February 1985, Killeen 868 (MO) Paraguay. Alto Paraná: Ñacunday, Montes 10938 (US). Amambay: cerca del Parque Nacional Cerro Corá, Cerro Tuyá, 300  m, 9 February 1982, Fernández Casas 6106 (MO, NY); in regione cursus superioris fluminis Apa, Hassler 8308 (MO, NY, RB, US). Caaguazú: Caaguazú, Balansa 2954 (BAF, P). Canindeyú: Sierra de Amambay, Hassler 10118 (US). Concepción: Villa Sana, Fiebrig 4642 (G) Guairá: pres de Villa Rica, Balansa 24 (P). Misiones: Estancia La Soledad, November 1956, Pedersen 4313 (US); San Ignacio, Ramírez 211 (US), 16 February 1950, Rosengurtt B-5783 (MO, US). Paraguarí: Barrerito, Rossengurt B-5607 (US); Cerro Peró, cerca de Paraguarí, Balansa 4358 (G, P) Argentina. Corrientes: Bella Vista, ruta 27, 10 km S de Bella Vista, Toropí, 13 October 1974, Schinini & Cristóbal 9775 (SI); Ensayo Prod. Nat. Capital, Fernández 889 (SI); Empedrado, Estancia Las Tres Marias, Pedersen s.n. (SI); Empedrado, Río Paraná, near Arroyo Sombrero, 27 November 1978, Renvoize et al. 3639 (K, MO, SI, US); Empedrado, Estación Experimental INTA, El Sombrerito, Schinini 16081 (CTES, SI); Mburucuyá, Estancia Santa Teresa, 24 February 1951, Pedersen 1030 (MO, US); Santo Tomé, Estancia Garruchos, Pedersen 5456 (US); Pago de los Deseos, Schwarz 8995 (US). Dpto. Ituzaingó, Ruta 12, 22  km este del acceso a Ituzaingó, lado N de ruta, 21 July 1987, Ginzbarg et  al. 375 (MO). Misiones: Apóstoles, Martínez Crovetto 8178 (SI); Dpto. Candelaria, Bonpland, Joergensen 1583 (US); Loreto, Ekman 636 (US). Dpto. Cainguás, Mineral, 31 March 1958, Montes 27649 (F, MO, US). Dpto. San Ignacio, Jardín América, Schwarz 6685 (US); San Ignacio, Montes 1885 (F, US); Puerto Cazador, Schwarz 1376 (US) 14b. Panicum olyroides Kunth var. hirsutum Henrard, Meded. Rijks-Herb. 40: 52. 1921. Type. Paraguay. Caaguazú: campos situes au NE de Caaguazú, 9 September 1874, B.  Balansa 23 (holotype, L 1300329; isotypes, G 00030347, G 00009727, P 00740766, US 00139815)

88

 

Taxonomic Treatment

Distribution and ecology (Fig. 15). This variety grows in Brazil, Paraguay, and Argentina, in environments similar to those of the type variety, from sea level to 1300 m asl; flowers and fruits throughout the year. This variety is characterized by densely hairy sheaths, blades, and panicles, with a sericeous pilosity, which may or may not reach the pedicels; the ligule is ciliate. Additional material examined. Brazil. Distrito Federal: Brasilia, São Bartolomeu, Equipe IBGE 3614 (IBGE); 25 km SW of Brasilia, 950 m, 19 February 1966, Irwin et al. 13006 (NY, US); adjacencias da Cachoeira do rio Pipiripau, Pereira & Mendonca 401 (IBGE); Campo, no Lago Norte, Filgueiras 818 (IBGE); on road to São Gabriel de Goiás, 1200 m, ca. 5 km N of Planaltina, 16 October 1965, Irwin et al. 9262 (MO, NY, US); Reserva Ecologica do IBGE, area de coleta n. l, entre o Corrego Escondido e o Cristo Redentor, Silva et al. 373 (IBGE); Brasília, University site, 2 March 1965, Clayton 4895 (K, NY, US); Brazlândia, January 1978, Allem & Vieira 1549, 1552 (MO). Goiás: Corumbá de Goiás, Serra dos Pirineus, 10 km NW Cocalzinho, 1150–1250 m, 7 June 1979, Burman & Filgueiras 435 (IBGE, MO); Guara, Irwin et al. 21525 (SP); Lagoa Formosa, Glaziou 22931 (P); Santo Antonio do Descoberto, Filgueiras 691 (IBGE). Mato Grosso: Mun. Ponta Porá, 22 km de Ponta Porá, camino a Bela Vista, 19 February 1968, Krapovickas et al. 14068 (CTES, US). Mato Grosso do Sul: Fazenda Modelo de Campo Grande, s. col. n. 39 (US 1385906). Minas Gerais: Lagoa Santa, 42 km N of Belho Horizonte, 900 m, Chase 9023 (US). Paraná: Mun. Campo Mourao, Campo Mourao, 14 October 1965, Hatschbach 12994 (F, US); Jaguariaiva, in campo cerrado, 720  m, 19 May 191, Dusén 388a (MO); Ponta Grossa, 11 December 1903, Dusén 2602 (MO, R), Estaçao Experimental, Swallen 8415 (US). Rio Grande do Sul: Tupanciretá, Kampf & Boldrini 839 (SI). Santa Catarina: Mun. Abelardo Luz, 6 km N of Abelardo Luz, 3 December 1971, Smith & Klein 15616 (R, US). São Paulo: Mun. Moji-Guaçu, Fazenda Campininha, just north of Rio Moji-Guaçu, 6 km NNWW of Padua Sales, April 1961, Eiten & Eiten 2577 (MO, NY, US); Fazenda Sta. Luzia do Campo Largo, SSE da cidade de Itapetininga, Sonia de Campos 191 (NY, SP, US); Mun. Porto Ferreira, from Porto Ferreyra along road to São Carlos, Sendulsky 162 (SP, US); Mun. São José dos Campos, São Jose dos Campos, 14 November 1961, Mimura 96 (NY, SP, US); Mun. Botucatu, 18  km north of Botucatu, along the S.  ManuelPiracicaba highway, 11 November 1971, Gottsberger 990-11271 (MO, US) Paraguay. Amambay: Cabecera Estrella, Sierra de Amambay, October 1933, Rojas 6434 (BAA); Cerro Corá, Sierra de Amambay, March 1934, Rojas 6750b (BAA). Cordillera: Cordillera de Altos, Fiebrig 436 (G, P). Estancia Primavera, Joergensen 4785 (US) Argentina. Corrientes: Bella Vista, ruta 27, 10 km S de Bella Vista, Toropí, 13 October 1974, Schinini & Cristóbal 9776 (SI) 15. Panicum racemosum (P.  Beauv.) Spreng., Syst. Veg. [Sprengel] 1: 313. 1825[1824]. Monachne racemosa P.  Beauv., Ess. Agrost. 1: 168, Table  10, Fig. 10. 1812. Panicum racemosum (P. Beauv.) Raspail, Ann. Sci. Nat. (Paris) 5: 299. 1825, comb. superfl. Type: Uruguay: “De Monte Video, Monachne, Hb. Palisot de Beauvois.”, Montevideo, 1 November 1767, P. Commerson s.n. (lectotype, P 003781644, here designated; isolectotypes; P 018666649, US 00140544). Fig. 17

Key to Ungrouped Panicum Sections and Species

89

Fig. 17  Panicum racemosum. (a) Plant. (b) Spikelet, ventral view. (c) Upper anthecium, ventral view. (From Cabrera 10065, LP)

90

 

Taxonomic Treatment

Saccharum reptans Lam., Tabl. Encycl. 1: 155. 1791. Panicum reptans (Lam.) Kunth, Révis. Gramin. 2: 219, Table 21. 1830, nom. illeg. hom., non Linnaeus, 1759. Eriolytrum junceum Desv. ex Kunth, Révis. Gramin. 1: 35. 1830, nom. inval. Type: Uruguay. Montevideo: Montevideo, P. Commerson s.n. (lectotype, B_10_0367288, here designated; isolectotypes, BAA 00002403, MPU 024563) Thalasium montevidense Spreng, Syst. Veg. [Sprengel] 4. Curt. Post.: 30. 1827. Type: Uruguay. Montevideo: Montevideo, F.  Sellow s.n. (holotype unknown; possible type, K) Perennial, rhizomatous plants, with elongated rhizomes, culms erect, rooting at the lower nodes, 0.40–1 m tall, simple, internodes 2–18 cm long, cylindrical, glabrous, nodes purplish, hairy. Sheaths 10–36  cm long, larger than the internodes, pilose, with adpressed whitish hairs or glabrous, pale. Ligule 2–3.5 mm long, shortly membranous at the base then long ciliate, collar brown or purplish, hairy to glabrous. Blades lanceolate, 24–70 × 0.2–0.5 cm, glabrous, narrowed at the base, apex enrolled, the margins smooth and involute. Inflorescences terminal, exserted; panicles contracted, multiflowered, 13–45 × 3–10 cm; main axis glabrous, smooth, pulvini glabrous, first-order branches alternate, rarely subopposite, adpressed to main axis; pedicels smooth, 1–6 mm long, glabrous or with spreading hairs, solitary or in pairs. Spikelets broadly ovoid, 5.8–7.5 × 2.5–3.4(−4) mm, open at maturity, densely pilose, pale, upper glume and lower lemma subequal; lower glume ovate, 4–6.5 mm long, 3/4 of spikelet length, pilose, more so toward the margins, 7–9(−11) nerved; upper glume ovate, 5.2–7.3 mm long, densely pilose on the outer face and shortly pilose at the apex of the inner surface, 9–13(−15) nerved; lower lemma glumiform, 5.4–6.6 mm long; lower palea elliptic, 5–6.5 × 1.7–2.3 mm, membranous, sparsely pilose on both surfaces; lower flower staminate; upper anthecium long ovoid, 4.5–5.8 × 1.9–2.5 mm, smooth, shiny, hardened, pale, upper lemma 7 nerved, with hairy margins toward the base, macrohairs filiform, stipe absent; anthers 2–3.3 mm long. Caryopsis ovoid, 3.3 × 2 mm wide; hilum oblong, embryo less than 1/2 the length of the caryopsis. Distribution and ecology (Fig. 18). Species restricted to coastal, fluvial, or maritime dunes in Brazil, from the state of Ceará, Uruguay, and Argentina and also present, with a disjunct distribution, in Chile, in coastal dunes; it is found in fluvial or maritime dunes and flowers from October to January. Local names: “Ajo macho,” “tupe” It is a characteristic South American species and excellent fixer of coastal dunes, present on the Atlantic Ocean from northern Brazil to Argentina, in the province of Buenos Aires, with a disjunct distribution in Chile on the Pacific coast. Two types of specimens of the species were examined in the herbarium P, of which specimen P 003781644 is selected as lectotype because it coincides with the protologue and is a well-preserved specimen. When describing Saccharum reptans, Lamarck cited “Monte-video, Comers.” The specimen B_10_0367288 agrees with the protologue and is here designated as lectotype of the species.

Key to Ungrouped Panicum Sections and Species

91

Fig. 18  Distribution of Panicum racemosum and P. tricholaenoides

Panicum racemosum differs from P. urvilleanum in having contracted panicles, with spikelets 5.8–7.5 mm long (vs. panicles lax and spikelets 5–7.3 mm in the latter species). Additional material examined. Brazil. Ceará: Dunas de Fortaleza, Lofgren 5 (R). Rio de Janeiro: Restinga do Grumari, December 1971, Sucre 7852 (SI); Copacabana, Glaziou 1628 (K, P); Leblon, Ducke & Kuhlmann s.n. (RB 16321, SI), Brade 12810 (IAN, RB); Ipanema, Dusén 5100 (G), Chase 9982 (IAN, NY, SP); Recreio dos Bandeirantes, Emygdio 333 (R), Ilha Grande, Praia do Sul, Araujo 4118 (K); restinga de Jacarepaguá, 3 March 1963, Duarte 7972 (RB); Recreio dos Bandeirantes, restinga, 30 March 1946, Segadas Vianna 60 (R). Rio Grande do Sul: Torres, Hatschbach 27230 (K, NY); Imbé, Santa Teresina, Guglieri 102 (ICN); estrada Passinhos a Osorio, margem da Lagoa dos Barros, Valls 2309 (CEN, CTES); Pelotas, Costa Sacco 211 (NY, RB); São Pedro do Rio Grande, Lindman A.705 (P); Jucao, Archer 4289 (SP); Praia do Arroio Teixeira, Kornelius s.n. (SI ex BLA 9091); Tramandai, Kappel s.n. (SI ex BLA 12023); Baia de Tramandai, Vidal II-210 (SI). Santa Catarina: Sombrío, Reitz 1987 (R), 1307 (IAN, NY)

92

 

Taxonomic Treatment

Chile. Bío Bío: Caleta Lenga, 29 October 2008, Morrone 5919 (SI), 5926 (SI). Cautín: Puerto Saavedra, December 1919, Hollermayer 229 (CONC); Puerto Saavedra, Moncul, December 1934, Montero 2102 (CONC); Talcahuano, November 1994, Quezada & López 11 (CONC); Coronel, Puchoco, January 1958, Junge 3085 (CONC); Coronel, November 1973, Garrido 4 (CONC); Laraquete, February 1959, Schlegel 2394 (CONC); de Coronel a Lota, Playa Grande, 12 January 2001, Zuloaga 7227 (SI); Ruta 160, entre Lota y Colcura, October 2008, Morrone 5918, 5921 (SI) Argentina. Buenos Aires: Pdo. La Plata, Isla Martín García, arenal central, 19 March 1996, Rúgolo 2057 (SI); Pdo. Coronel Dorrego, Oriente, balneario Marisol, 4 November 2008, Biganzoli 1988 (SI); Pdo. General Alvarado, Miramar, 10 November 1939, Cabrera 5535 (NY, SI); dunas de Miramar, 7 December 1971, Lanfranchi 1818 (SI); Pdo. General Lavalle, La Margarita, 1 December 1944, Cabrera 8533 (SI); Pdo. de la Costa, San Bernardo dunas sobre el mar, 8 November 1987, Zuloaga 3360 (SI); Pdo. Lobería, Quequén, Rodriguez 839 (NY); Pdo. Necochea, Las Grutas, Villamil 3554 (NY); Pdo. General Pueyrredón, Punta Mogotes, Rodriguez 433 (NY); Pdo. Monte Hermoso, límite con la Reserva Natural de Pehuen-Co, 20 December 2017, Giussani 678 (SI); Pdo. Necochea, Necochea, Nicora s.n. (SI); Pdo. Tres Arroyos, Claromecó, 14 November 1987, Villamil 5369 (BBB, SI); Pdo. Pinamar, Pinamar, January 1946, Cabrera 10065 (LP). Entre Ríos: Gualeguaychú, Médanos, Parodi 9505 (BAA, SI); Islas del Ibicuy, Paranacito, December 1949, Ragonese 48 (SI) Uruguay. Colonia: near cost, Riachuelo to Artilleros, Bartlett 20715 (NY); Río San Juan, dunas, Cabrera 3220 (NY); Arroyo de Pintos, Artilleros, near Puerto Platero, Bartlett 21218 (F, GH). Montevideo: Montevideo, Herter 1376 (RB), Kuhlmann s.n. (RB). Rocha: Cerro Verde, Alonso Paz 3717 (MVFQ). San José: Barra del Rio Santa Lucía, Rosengurtt B-6281 (P); Balneario Kiyú, 2 December 2001, Seijo 2661 (CTES, SI). Without departament, Banda oriental del Uruguay, St. Hilaire 2235 (P) 16. Panicum tricholaenoides Steud. Perennial, rhizomatous plants, with elongated rhizomes, covered with short hairy cataphylls at the edges; culms erect, 1.5–2  m tall, in fascicles of numerous congested, bambusoid, subwoody, densely branched culms; internodes 10–30 cm long, cylindrical, hollow, pale, glabrous; nodes glabrous, dark. Sheaths 7–14  cm long, smaller than the internodes, aphyllous at base of culms or with extremely reduced blades, sparsely papillose-pilose to glabrous, persistent when dry, pale. Ligule membranous-ciliate, 0.5–1.5 mm long; collar glabrous, brown to purplish. Blades lanceolate, 12.5–40 × 0.2–0.5 cm, apex subulate, base rounded, flat or with margins partially involute, scabrous, ciliate towards the basal portion, adaxial surface densely pilose toward the base, abaxial surface sparsely pilose to glabrous. Inflorescence a lax, open, multiflowered terminal panicle, 20–40  ×  5–20  cm, erect or drooping; main axis cylindrical toward the base, then wavy, scabrous, pulvini pilose, brownish, lower branches subverticillate, upper branches alternate or opposite; pedicels 0.5–3.4 mm long, smooth, solitary or in pairs, with long persistent whitish hairs. Spikelets long ovoid, 2.2–2.9(−4) × 0.8–1 mm, open at maturity, glabrous, pale or tinged with purple; lower glume ovate-lanceolate, 1.5–2.9 mm long, ¾ to 4/5 the

Key to Ungrouped Panicum Sections and Species

93

length of the spikelet, glabrous, apex acuminate to acute, 5–7-nerved, the midrib scabrous toward the apex; upper glume ovate-lanceolate, acuminate to acute, 7–9-nerved, glabrous; lower lemma glumiform, acuminate, 7–9-nerved, glabrous; lower palea elliptic, 1.6–2.5 × 0.8–1 mm, membranous, glabrous; lower flower staminate; upper anthecium narrowly ovoid, 1.6–2.5  ×  0.7–0.9  mm, glabrous, pale, with blackish spots at maturity, shiny smooth, obtuse, not stipitate; lemma 7 nerved, with a dark disk at its base. Caryopsis ellipsoid, 1.3–1.9 × 0.8–1 mm, brown; hilum obovate, embryo more than half the length of the caryopsis. Key to the Varieties 1. Spikelets 2.2–2.9 mm long 16a. P. tricholaenoides var. tricholaenoides 1. Spikelets 3.2–4 mm long 16b. P. tricholaenoides var. flavomarginatum 16a. Panicum tricholaenoides Steud. var. tricholaenoides, Syn. Pl. Glumac. 1: 68. 1855[1853]. Type: Uruguay. Montevideo: Montevideo, Deloche 166 (holotype, W). Fig. 19 Panicum junceum Nees, Fl. Bras. Enum. Pl. 2(1): 159. 1829, nom. illeg. hom., non Trin., 1826. Panicum junceum Nees var. subnutans Döll, in C. Martius, Fl. Bras. 2 (2): 216. 1877. Type: Uruguay. Montevideo: Montevideo, F. Sellow 960 (holotype, B_10_0248986; isotypes, BAA 00000909, 00000910, fragments ex B; LE-768.01) Panicum junceum Nees var. strictius Döll, in C. Martius, Fl. Bras. 2 (2): 216. 1877. Type: Brazil. “Habitat in campis editis ventosis Parannam in confinibus P. Goyaz et prov. Min. Ger. Martius Fl. Bras.”, C F. P. von Martius s. n. (lectotype, M, here designated; isolectotype, US 0148596) Panicum bambusoides Speg. ex Arechav., Annals. Mus. Nac. Montevideo 1: 128, Figs. 9–10. 1894, nom. illeg. hom, non Desv. ex Ham., 1825. Type: Uruguay. “Orillas del Río Negro, near Mercedes,” February 1892, J.  Arechavaleta s.n. (lectotype, W 19160024645, here designated; isolectotypes, W 19160024642, W 19160024643, W 19160024644) Panicum sempervirens Kuntze, Revis. Gen. Pl. 3(3): 364. 1898. Type: Bolivia. OstVelasco, 200 m, July 1892, C. E. O. O. Kuntze s.n. (lectotype, NY 00414049, NY 00414050, part 1 and 2, here designated; isolectotype, W 19160022692) Panicum pilgeri Herter, Herb. Corn. Osten Com. 1: 2. 1925, nom. illeg. hom., non Mez, 1904. Type: Uruguay. “Orillas del Río Negro near Mercedes, Uruguay” (holotype not known) Distribution and ecology (Fig. 18). This variety inhabits from Colombia and Venezuela to Bolivia, southern Brazil, Paraguay, Argentina, and Uruguay. It is usually found in sandy soils, next to watercourses or lagoons, leans on shrubs and forms dense clumps, and flowers from December to April or June. Local names: “Tacuarita”; “takuari ñu” (takuari  =  tacuarilla, ñu  =  field) (Zuloaga 1994)

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Fig. 19  Panicum tricholaenoides var. tricholaenoides. (a) Habit. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (From Rojas 10491, SI)

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When describing P. junceum, Nees cites two syntypes: Uruguay, Montevideo, F. Sellow s.n. (B_10_0248986, BAA 00000909, 00000910, fragments), and Brazil. “Habitat in campis editis ventosis Parannam in confinibus P. Goyaz et prov. Min. Ger. Martius Fl. Bras.”, C F.  P. P. von Martius s. n. (syntype, M; isosyntype, US-80713, fragment). Of these, the Berlin specimen of Sellow s.n., B_10_0248986 corresponds to the holotype. It should be noted that this specimen was also the one used by Döll to describe P. junceum var. subnutans, together with Deloche 166, the type material of P. tricholaenoides. On the other hand, Döll based P. junceum var. strictius on a specimen collected by Martius in Brazil. Additional material examined. Colombia. Casanare: Caño Cancia, 23  km sudoeste de Hato Yarumito, 36  km sur de Mani, 23 February 1971, Sastre 830 (COL). Meta: 2  km east of Puerto Gaitán in the flood plain of Rio Manacacias, 150 m, 31 December 1973, Davidse 5416 (COL, MO, NY); Los Llanos, Río Meta, María, 130  m, 17 October 1938, Cuatrecasas & García Barriga 3729 (COL, F, USA); Llano de San Martín, Triana 351 (P, USA); Puerto Gaitán, Río Manacacías, a orillas del río, 250 m, 14 June 1989, Zuloaga 3981 (COL, MO, SI). Sabanas de Barranca de Upía, García Barriga 4991 (COL); Sabana del Vergel, 31 January 1937, García Barriga 4984 (COL) Venezuela. Amazonas: Dpto. Atures, raudales de Atures, 10 km south of Puerto Ayacucho, downstream along rio Orinoco, 75  m, 6 September 1985, Steyermark et al. 131495 (MO, NY, SI, VEN); Dpto. Atures, 150 m, San Juan de Manapiare, sabanas temporalmente inundadas en la margen izquierda del Río Manapiare, 13 October 1977, Huber 1100 (MO, NY, VEN); Dpto. Atures, sabana inundable en el valle del cano Guanay, a la altura del cerro El Santo, 28 February 1979, Huber 3516 (MO), 3517 (VEN). Apure: Distr. Pedro Camejo, rocks by raudal de Marimara, 27 airline km NE of Puerto Páez, 40 m, 25 February 1978, Davidse & González 14526 (MO, VEN); región Mantecal-Elorza, Ramia 1359 (VEN); Capanaparo, Hato La Coromoto, Mondolfi s.n. (VEN 126125); sabanas de Hato El Frío, entre El Saman y Mantecal, Ramia 4006 (VEN); Palmarito, Ramia 1789 (VEN); Puerto Páez, Velez 2734 (US); between río Orinoco and Piedra La Villa, Wurdack & Monachino 41399 (F, NY, VEN); sur de San Fernando de Apure, Capanaparo river, de Alba 53-106 (US); 42 km northeast of Mantecal along hwy. to San Fernando de Apure, 80 m, 9 November 1973, Davidse 3909 (MO, PORT). Barinas: sabanas de los alrededores del Río Paguey, Abr 1954, Aristeguieta 2141 (MO). Brazil. Goiás: proximo a Ilha do Bananal, Mileski 221 (RB); Ilha do Bananal, Rio Javaés, cerrado sujeto a inundaçao periódica, 20 August 1978, Silva 4876 (MO, NY); Mun. Santa Izabel, Ilha do Bananal, Parque Nacional do Araguaia, Cardoso da Silva et  al. 282 (IBGE, SP); Pantanal Seco, a esquerda do Rio Formoso, 24 August 1978, Pires & Santos 16681 (MO); without locality, Glaziou 22423 (US). Mato Grosso: Puerto Esperanza, sobre el río Paraguay, Chase 11097 (F, RB, US); Ponta Porá, Puerto Esperanza, Swallen 9574 (US); Mun. Barão de Melgaço, Fazenda Santa Lucia, Allem & Vieira 2506 (CEN). Mato Grosso do Sul: Mun. Corumbá, Fazenda Bodoquena, Allem et al. 2220 (CEN); Mun. Corumbá, Fazenda Santana, regiao do Paiguas, Comastri Filho 9 (CEN); Campo Grande, Nienstedt 207 (RB, US); Mun. Corumbá, Fazenda Santo Estevão, 14 November 1977, Allem & Vieira 1194 (MO). Rio Grande do Sul: Porto Alegre, Orth 3 (US); São Leopoldo,

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Quinta São Manoel, Dutra 632 (IAN, R); São Leopoldo, 13 November 1934, Orth 850 (MO). Rondonia: Rio Guaporé, Fazenda Santa Rosa, Black & Cordeiro 52-14880 (IAN); Rio Guaporé, Santo Antonio de Manoelito, Black & Cordeiro 5215315 (IAN); Rio Guaporé, Fazenda Santa Rosa, Black & Cordeiro 52-14836 (US). Without state and locality, Riedel s.n. (G) Bolivia. Pampas near Lake Rogagua, Hacienda Rosario, White 1209 (NY). Beni: Prov. Itenez, S side of Río Guaporé, ca. 15 km east/southeast of Costa Marques, 150 m, 31 March 1987, Nee 34647 (MO); Prov. Ballivián, Espíritu, Beck 2654 (LPB), 3302 (K, LPB), 5685 (K); Prov. Ballivián, Estancia El Porvenir, 50 km E of the Río Maniqui on the road to Trinidad, vicinity of Laguna Normandía, 250  m, 14 November 1985, Solomon 14724 (K, MO). Santa Cruz: Buena Vista, 22 January 1925, Steinbach 6900 (K, MO, US), 3220 (G, K), 6901 (G); Velasco, 10 km east of Comunidad Santa Rosa de la Roca, 400 m, 22 January 1985, Killeen 778 (MO, SI); Ñuflo de Chavez, 10 km southeast of Concepción, Estancia Salta, 490 m, 7 January 1985, Killeen 720 (F, MO, SI) Paraguay. Alto Paraná: Puerto Bertoni, January 1907, Bertoni 4138, 4614 (W). Amambay: Sierra de Amambay, 1912/1913, Hassler 11443 (BAF, G, K, MO, NY, P, US). Central: Central Paraguay, Morong 538 (MO). Presidente Hayes: Prairies du Gran Chaco, en face de l’Assomption, March 1874, Balansa 18 (G, P, US), Gran Chaco, ad ripam occidentale fluminis Paraguay, Hassler 2396 (G, K, NY). San Pedro: Colonia Nueva Germania, orillas Río Aguaray-Guazú, Rojas 10491 (SI, US) Argentina. Chaco: Dpto. 1° de Mayo, Antequera, Schulz 3424 (CEN); Colonia Benitez, Schulz 1799 (BAB); Las Palmas, Joergensen 2430 (MO, US). Corrientes: Empedrado, Estancia Las Tres Marías, Pedersen 3046 (G, P), 28 November 1978, Schinini 16095 (F, MO, SI); Río Uruguay, 29 January 1952, Nicora 6094 (MO). Dpto. San Roque, Chavarría, 6 November 1978, Fernández 519 (MO); Paraná River, Estancia Las Tres Marías, 28 November 1978, Renvoize et al. 3663 (MO, SI). Isla Apipé, Puerto Vizcaino, 18 November 1977, Schinini 13782 (CTES); Dpto. Ituzaingó, desembocadura del Arroyo Garapé en el río Paraná, 45  km al este de Ituzaingó, 24 April 1975, Schinini et al. 11090 (SI); Puesto de Prefectura, 42 km al este de Ituzaingo, 12 April 1985, Zuloaga et al. 2304 (MO, SI); Dpto. San Cosme, Paso de la Patria, costa del Río Parana, 29 May 1977, Cristóbal et al. 1526 (CTES, F, SI); San Martin, 6 km sudoeste de La Cruz, sobre rio Uruguay, 8 February 1979, Schinini et al. 16857 (SI); Yapeyú, Parodi 12890 (US), Ibarrola 1870 (NY); Paso de los Libres, al sur de la ciudad, Ibarrola 2071 (NY). Entre Rios: Dpto. La Paz, Paso Yunque, Río Guayquiraró, 14 February 1985, Bacigalupo & Deginani 104 (MO); Concordia, Parque Mitre, 18 March 1996, Zuloaga & Morrone 3851 (MO); Federación, Santa Ana, Troncoso et  al. 1650 (SI); without locality, 17 February 1876, Lorentz s.n. (BAF); San Carlos, 12 December 1946, Meyer 11017 (MO, US). Formosa: Puerto Bermejo, Kermes 678 (US). Misiones: Candelaria, Loreto, Río Yabebiry, Ekman 630 (CORD, US). Santa Fe: Obligado, campo Isla Tacuarendi, Lewis 963 (SI), 10 November 1983, Pire 1410 (SI) Uruguay. Artigas: orilla del Río Uruguay, Las Cachuelas, al norte de Belén, Rosengurtt 11327 (NY). Durazno: Río Negro, San Gregorio del Polanco, Schroeder s.n. (ex herb. Corn. Osten 18459, US-1297228). Paysandú: Río Uruguay, Isla

Key to Ungrouped Panicum Sections and Species

97

Queguay, Rosengurtt B-3313 (US). Salto: Río Uruguay y Arroyo San Antonio Grande, Rosengurtt B-942 (US); without locality, Arechavaleta s.n. (US 974616) 16b. Panicum tricholaenoides var. flavomarginatum (Mez) Zuloaga, Candollea 46(2): 540. 1991. Panicum flavomarginatum Mez, Bot. Jahrb. Syst. 56, Beibl. 125: 2. 1921. Type: Paraguay. Caaguazú: Caaguazú, dans les forets, 19 November 1874, B.  Balansa 10 (lectotype, P 007400869, here designated; isolectotypes, BAA, BAF 00000257, BR 0000006883478, G 00009719, G 00030573, GOET 006770, LD 1226359, P 00740868, S05-9131, US 00148497) Distribution and ecology (Fig. 18). The variety is restricted to the departments of Caaguazú Central and Paraguarí in Paraguay, where it grows on the vegetation in sandy soils; it flowers from August to November. It can be distinguished from the type variety by the characteristics indicated in the key. Additional material examined. Paraguay. Central: cerca del Lago Ipacaraí, August 1913, Hassler 12253 (BAF, G, K, MO, NY, US) Paraguarí: cerca de Sapucay, October 1913, Hassler 12930 (BAF, G, K, MO, U). Presidente Hayes: in regione cursus inferiores fluminis Pilcomayo, Aug 1906, Rojas 535 (G) 17. Panicum urvilleanum Kunth, Révis. Gramin. 2: 403, Table 115. 1831. Monachne urvilleana (Kunth) Herter, Revista Sudamer. Bot. 9: 118. 1953. Type: Chile. Concepción: Concepción, 1815, J.  S. C.  C. D.  D’Urville s.n.(lectotype, B_10_0366225, designated as “type” by Hitchcock & Chase, 1910: 132; isolectotypes, BAA 000024201, P 00128866, P 00128867). Fig. 20 Panicum megastachyum J. Presl, Reliq. Haenk. 1(4-5): 305. 1830, nom. illeg. hom., non P. megastachyum Nees ex Trin., 1826. Panicum preslii Kunth, Enum. Pl. 1: 121. 1833. Type. [Chile]: “Hab. in montanis Peruvia huanocensibus,” T. Haenke s.n. (holotype PR, n.v.; isotype, US 00148288) Panicum patagonicum Hieron., Bol. Acad. Ci. Córdoba 3: 377. 1881. Type: Argentina. “En los médanos de la boca del Río Negro y de la costa del Océano Atlántico, 10 October 1874, C.  A. Berg 208 (lectotype, CORD 00001745, here designated; isolectotypes, BAA 00002380, LP 001606) Panicum urvilleanum f. subpilosum Kuntze, Revis. Gen. Pl. 3(3): 364. 1898. Type: Argentina, Pampas, January 1892, C. E. O. Kuntze s.n. (lectotype, NY 00414071, here designated) Panicum urvilleanum var. longiglume Scribn., Bull. Div. Agrostol., U.S.D.A. 17(ed. 2): 49. 1901. Type: USA.  California: San Jacinto, June 1882, S.  B. Parish & W. F. Parish 887 (lectotype, US 00148077, here designated; isolectotype, US 00782151) Perennial, rhizomatous plants, with elongated, robust rhizomes, culms erect, 40–70 cm tall, simple, internodes cylindrical, rigid, glabrous, hollow toward the upper portion; nodes pilose, covered with whitish antrorse hairs. Sheaths 8–20  cm long, longer than the internodes, pale, densely to sparsely hairy, with retrorse hairs. Ligule shortly membranous in the lower portion, then long ciliate, 1–3 mm long; collar hairy or glabrous. Blades lanceolate, 24–50  ×  0.2–0.5  cm, narrowed and with involute

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Fig. 20  Panicum urvilleanum. (a) Plant. (b) Spikelet, ventral view. (c) Upper anthecium, ventral view. (From Cabrera 6919, LP)

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margins towards the base, apex attenuate, rigid, densely villous on both faces, with antrorsal hairs on the adaxial surface and retrorsal ones on the abaxial surface, or glabrous. Inflorescence terminal, exerted, multiflowered, pyramidal; peduncle 15–30 cm long; panicles lax, open, 20–35 × 5–20 cm; main axis smooth, glabrous or with spreading or densely hairs, pulvini hairy, first-order branches alternate or opposite, divergent, branch axes and pedicels densely to sparsely hairy, pedicels 2.5–10 mm long, solitary or in pairs. Spikelets long ovoid, 5–7.3 × 1.8–2.6 mm, densely pilose, straw-like or reddish tinged, open at maturity; lower glume ovate-lanceolate, 3.5–6.3 mm long, acuminate, ¾ to 4/5 the length of the spikelet, 7–9(–11) nerved, pilose, more so toward the base and margins; upper glume ovate-lanceolate, 11–13(–15) nerved, as long as the spikelet, densely pilose, acuminate; lower lemma glumiform, 9–11(–13) nerved, densely pilose, acuminate; lower palea elliptic, 4.1–5.6  ×  1.4–2  mm, membranous, sparsely pilose; lower flower staminate; upper anthecium long ovoid, 3.5–5.3 × 1.4–2.2 mm, pale, smooth, shiny, indurate, not stipitate; lemma 7 nerved and with lower margins with long filiform macropels. Caryopsis not seen. Distribution and ecology (Fig. 21). A species of disjunct distribution, occurring in the southern USA and Mexico and in Argentina and Chile. It inhabits dunes and dry, sandy soils. It reaches up to about 2000 m asl. It is a very useful species as a dune consolidator, thanks to its powerful rhizomes and the ability to branch at the lower nodes, when the plant is partially covered by sand. Additional material examined. USA. California: Barstow, Chase 5766 (US); Hesperia, Abrams 2164 (US) Mexico. Sonora: on tall dunes at proximal end of sandspit at El Socorro, 23 March 1979, Johnson 4080 (MEXU) Chile. Bío Bío: Mulchen, December 1963, Zoellner 409 (CONC); road between Río Claro and Yumbel, November 1963, Matthei 275 (CONC); NW of Cabrero, near Río Itata, Soreng & Soreng 7006 (CONC, US); Ñuble, 5  km E of Quillon, 150 m, Hutchison 209 (F); El Roble bridge, December 1967, López & Marquez 81 (CONC); San Vicente; November 1963, Matthei 302 (CONC); Cerro La Pólvora, Laguna Galindo, March 1958, Junge 3104 (CONC); Ñuble, General Cruz, campos del Itata, November 1934, Montero 1953 (CONC); Arenales de Itata, February 1905, no collector s.n. (MO-1211903). Cautín: Puerto Saavedra, February 1942, Hollermayer 1215 (CONC) Argentina. Buenos Aires: Bahía Blanca, Punta Alta, 5 November 1940, Cabrera 6620 (LP, SI); Coronel Dorrego, Monte Hermoso, 8 November 1940, Cabrera 6751 (LP, SI); General Villegas, 22 December 1939, Cabrera 5677 (LP, SI); Pellegrini, 28 November 1940, Cabrera 6919 (LP, SI); Trenque Lauquen, Piergentili 3578 (SI); Villarino, Balneario Chapalcó, 20 November 1981, Villamil & Nicora 2154 (SI); Pedro Luro, Marahué, Scoffield 60 (BBB). Catamarca: 21 km S of Andalgalá, 9 February 1973, Cantino 636 (SI). Chubut: Península de Valdes, Punta Pardelas, 1 December 1982, Villamil 2740 (BBB, SI); Puerto Pirámides, 4 December 1982, Villamil 2747 (MO, SI). Córdoba: Río Cuarto, 20 December 1905, Stuckert 15336 (CORD), 15337 (MO), 9 December 1905, Stuckert 15213 (MO); between Del

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Fig. 21  Distribution of Panicum urvilleanum and P. virgatum

Campillo and Huinca Renancó, 14 November 1951, A. T. Hunziker 9376 (CORD, MO); General Roca, Villa Valeria, Bianco 4760 (RIOC). La Pampa: La Pastoril, on ditch ruta 143, 6 December 1964, Cunquero 166 (SI); Utracán, Estancia Chuni Malal, potrero Zubeldía, on ruta 148, 20 February 1971, Rúgolo 721 (SI); Utracán, General Acha, 3 November 1953, Burkart 19203 (SI); Maracó, General Pico, 10 November 1939, Burkart 9864 (SI); Utracán, Ruta Nacional 35, before Padre Buodo, 14 November 2013, Zuloaga et al. 14624 (SI); Utracán, from Parque Luro toward Ataliva Roca, Ruta Nacional 35, 22 November 2017, Zuloaga et al. 16052 (SI). La Rioja: Vinchina, 23 February 1941, Burkart 12239 (MO, SI), 12260 (SI); edge of the Bermejo or Jaguel River, J. Hunziker 2039 (SI). Mendoza: Malargüe, Sierra del Nevado, Boelcke et al. 15797 (SI); between Malargüe and Fortín Malargüe, Burkart et al. 13933 (SI); San Rafael, Sierra del Nevado, Puesto Los Zainos, Boelcke et  al. 15578 (SI); El Sosneado, 18 November 1948, Senn 4374 (MO); Colonia Alvear, 25 December 1922, Parodi 4847 (BAA, MO); Río Diamante, riverbed near San Rafael, 600 m, 30 October 1955, Böcher et al. 689 (MO); Atuel valley near El Sosneado at irrigation canal, 1600 m, 3 November 1975, Böcher et al. 776 (MO); Ruta Nacional 188, km 712, road to General Alvear, 24 November 2008, Morrone 5930 (SI); Las Heras, San Ignacio, Sanzin 900 (MOY); Malargüe, Fortunoso, 6

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December 2014, Zuloaga et al. 15260 (SI). Neuquén: 60 km north of Zapala, 16 November 1948, Senn 4315 (MO); road from Zapala to Las Lajas, Arroyo Covunco, Krapovickas 3693 (BAB); Ruta Nacional 22, east of Zapala, 27 November 2014, Zuloaga et al. 14978 (SI). Rio Negro: vicinity of General Roca, Fischer 187 (F); J.J. Gomez, banks of the Río Negro, 3 November 1972, Krapovickas et al. 22405 (CTES, MO); médanos de El Pescadero, mouth of the Río Negro, 3 February 2009, Giussani 370 (SI). Salta: Cafayate, 22 October 1948, Burkart 17651 (SI). San Juan: 25 de Mayo, Ruta Nacional 20 de Villa Borgias al Refugio, 22 August 2014, Zuloaga et  al. 14974 (SI); Caucete, E of Caucete, dunes, Kiesling 10224 (SI); Chimbas, Villa Paula A. de Sarmiento, Finca Ariza, Ariza Espinar 140 (CORD); Iglesia, Rodeo, Kiesling 4259 (SI); Valle Fértil, upper part of Salina de Mascasín, Piccinini 1983 (BAB). San Luis: General Pedernera, Nueva Escocia, 12 November 1940, Burkart 10858 (SI); General Pedernera, Esperanza, heading to La Portada, 24 November 1962, A. T. Hunziker 16046 (CORD); Gobernador Dupuy, Ruta Nacional 188, de Unión a La Maroma, 19 November 2010, Zuloaga et al. 12232 (SI) 18. Panicum virgatum L., Sp. Pl. 1: 59. 1753. Chasea virgata (L.) Nieuwl., Amer. Midl. Naturalist 2: 64. 1911. Milium virgatum (L.) Lunell, Amer. Midl. Naturalist 4: 212. 1915. Type: USA. Virginia. “Habitat in Virginia,” J. Clayton 578 (lectotype, LINN-80.61, designated by A. S. Hitchcock, Contr. U.S. Natl. Herb. 12: 118. 1908; isolectotypes, BM 001042408, US, fragment ex BM). Panicum virgatum L. var. cubense Griseb., Cat. Pl. Cub.: 233. 1866. Panicum virgatum subsp. cubense (Griseb.) Borhidi, Bot. Közlem. 58(3): 175. 1971. Type: Cuba. Havana, low savannas, 19 May 1865, C.  Wright 183 (lectotype, GOET 006795, designated as “type” by Hitchcock & Chase, 1910: 92) Panicum ichnanthoides E. Fourn., Mexic. Pl. 2: 30. 1886. Panicum ichnanthoides E.  Fourn. ex Hemsl., Biol. Centr.-Amer., Bot. 3: 490, 1885, nom. nud. Type: Mexico. Veracruz: Orizaba, J.  K. Müller 2002 (lectotype, LE; designated as “type” by Hitchcock & Chase, 1910: 88; isolectotypes, NY 00381764, USA 00074268) Panicum buchingeri E. Fourn., Mexic. Pl. 2: 30. 1886. Panicum buchingeri E. Fourn. ex Hemsl., Biol. Cent.-Amer., Bot. 3: 486. 1885. Type: Mexico. Veracruz: Orizaba, Thomas s.n. (holotype not known) Perennial, rhizomatous plants, with elongated rhizomes, culms erect, many noded simple, non-bambusoid culms 1–2 m tall, with short internodes and fascicled leaves at the base, cylindrical; internodes, 17–30 cm long, glaucous, smooth, glabrous, the lower ones solid, hollow toward the upper portion; nodes brownish, glabrous. Sheaths shorter than internodes on the floriferous culms, striate, rigid, with purplish tints, glabrous to hairy toward the upper portion. Ligule membranous-ciliate, 2–4 mm long, hairy beneath at the base of the blade; collar brown, glabrous. Blades lanceolate, 10–60 × 0.2–1.3 cm, ascending, narrowed toward the base, apex attenuate, flat, glabrous or hairy toward the base on the adaxial surface margins scabrous. Inflorescence terminal, long exserted, pyramidal; peduncle smooth, glabrous, cylindrical; panicles lax, diffuse, open to contracted, not spiciform, multiflowered 20–45 × 5–25 cm; main axis cylindrical, glabrous, pulvini hairy, first-order

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Taxonomic Treatment

branches divergent from the main axis, alternate or opposite, branch axes wavy, scabrous, spikelets scattered on the branches on scabrous pedicels, solitary. Spikelets 3.6–4 mm long, lanceolate, pale or tinged with purple, glabrous, open at maturity, acuminate; lower glume 2.1–3 mm long, 2/3 to 3/4 the length of the spikelet, ovate, acute, 5 nerved, midrib scaberulous toward the apex; upper glume as long as the spikelet, ovate, attenuate, 7–9 nerved; lower lemma glumiform, 7–9 nerved, ovate; lower palea long-elliptic, hyaline, glabrous; lower flower staminate; upper antecium long-ovoid, 2.6–2.8 mm long, shorter than the upper glume and lower lemma, smooth, shiny, indurate, glabrous, pale to olivaceous at maturity, with a dark disk and two black spots along the palea, not stipitate, early deciduous; lemma 7 nerved. Caryopsis oblong in outline, olivaceous; hilum oblong, embryo little more than 1/2 the length of the caryopsis. Distribution and ecology (Fig. 21). This species grows in humid open environments, is frequent in grasslands, and also on the margins of Pinus and Quercus forests; it is distributed from Canada and the USA, where it is very abundant, to the Caribbean, Mexico, and Mesoamerica, being present in Belize and Nicaragua and flowers and fruits between July and October. Local name: “Switchgrass” Additional material examined. Canada. Ontario, Dutily 30384 (MO); Quebec, Marie-Victorin 8152 (MO) USA.  Alabama: Mobile, Deramus 695 (MO). Arkansas: Ashley, Demaree 13592 (MO). Colorado: Denver, Jones 827 (MO). Connecticut: Hartford, Seymour 22920 (MO). Delaware: New Castle, Kellogg 1217 (MO). Florida: Hillsborough, Fredholm 6365 (MO). Georgia: Spalding, Webster 3291 (MO). Illinois: Saint Clair, Neill 8550 (MO). Indiana: Tipton, Deam 13899 (MO). Iowa: Greene, Davidse 1854 (MO). Kansas: Barber, Conrad 5973 (MO). Louisiana: 3.4 mi east/ northeast of junction of Fourchon road and LA Hwy 1, 14 July 1981, Lasseigne 6260 (MEXU). Maryland: Kent, Thompson 338 (MO). Massachusetts: Bristol, Seymour 17670 (MO). Michigan: Saint Joseph, Daniels s.n. (MO). Minnesota: Pipestone, Johson 365 (UMO). Mississippi: Harrison, 24 July 1971, Lasseigne 2844 (MO). Missouri: Barry, Steyermark 80666 (UMO). Nebraska: Antelope, Liewere 137 (UMO). New Hampshire: Cheshire, 24 July 1972, Boufford 7295 (MO). New Jersey: Atlantic, Redfield 10941 (MO). New Mexico: San Miguel, 25 August 1908, Standley 5289 (MO). New York: Essex, Crockett 7745 (MO). North Carolina: Columbus, Youngstrom 6 (MO). North Dakota: Kidder, Williams 1198 (MO). Ohio: Scioto, Demaree 10788 (MO). Oklahoma: Blaine, 24 July 1900, White 42 (MO). Pennsylvania: Erie, Shafer s.n. (MO). Rhode Island: Bristol, 29 August 1964, Seymour 22235 (MO). South Carolina: Charleston, Hill 23758 (MO). South Dakota: Clay, Johnson 5 (MO). Tennessee: Coffee, 21 August 1938, Svenson 8780 (MO). Texas: 1 mi S of Junction near the Llano River at the Texas Tech University Center, 6 October 1983, Hatch 5060 (MEXU); Collin Counth, Coit Meadow on Renner Road, 31 August 1971, Lundell 18662 (MEXU). Without locality, Lindheimer 733 (P). Virginia: Norfolk, Fernald 13878 (MO). West Virginia: Cabell, Gilbert 530 (MO). Wisconsin: Richland, 12 August 1993, Nee 43735 (MO). Wyoming: Laramie, July 1933, Nelson 995 (MO)

Key to Ungrouped Panicum Sections and Species

103

Mexico. Chiapas: Hacienda Monserrat, September 1923, Purpus 9199 (MO, US); Chismanta, Sabana Palenque, July 1939, Matuda 3722 (F, MEXU, US); junction of rio Perlas and rio Jatate, near Miramar lagoon, Sohns 1566 (US); near the 99 km post E of Villahermosa on the Villahermosa-Francisco Escárcega road, 9 August 1975, Davidse & Davidse 9396 (MO); Mun. La Trinitaria, Lagos de Montebello, 42  km northeast of La Trinitaria, 1300  m, 23 October 1971, Breedlove & Thorne 21270 (MEXU, MO); Mun. La Trinitaria, montane rain forest at Lagos de Colores, Lagos de Montebello National Park, 1370 m, 7 November 1981, Breedlove & Davidse 55039 (MO). Coahuila: 28 km north of Zaragoza, past rancho La Salada on Rio San Rodrigo, 22 September 1977, Valdés Reyna 1280 (MEXU). Jalisco: Zapopán, between Zapopán and Sapiquipán, Oliva 49 (US); Río Blanco, 1886, Palmer 510 (MEXU, MO); 2 km S of San Antonio de Padua, 10 August 1986, Santana Michel et al. 1850 (MEXU). Nuevo León: Río Ramos, 4  km south of Allende, 14 July 1983, Valdés Reyna 1548 (MEXU). Puebla: Tehuacán, March 1917, Patoni s.n. (MEXU-3704). Quintana Roo: km 9 southeast of Chumpón, 31 August 1984, Durán & Olmsted 1039 (MEXU), low flooded forest. San Luis Potosí: Lagua de la Media Luna, 980 m, Takaki s.n. (MEXU-163597). Veracruz: Orizaba, Hitchcock 6364 (US); about 8 miles north of Ciudad Alemán, 120 m, 16 August 1953, Reeder & Reeder 1996 (MEXU) Belize. Belize District, Baker’s Pine Ridge, Lundell 6938 (F, US); near Honey Camp, Meyer 110 (F); Toledo District, Near Cattle Landing, Gentle 6873 (F, G, NY) Nicaragua. Without locality, Matuda 5749 (F) Bermuda. Nonsuch Island, North Hill Cliff, Wingate G-12 (NY); shore near Frascatí, Collins 139 (NY) Cuba. Havana: Swamps west of Batabanó, León & Cazañas 5755 (NY) Matanzas: Ciénaga de Zapata, Ekman 16939 (US). Pinar del Río: Laguna Jovero to Laguna Herradura, wet land, Shafer 10927 (NY). Santa Clara: Caibarién, in swamps at Dolores, Ekman 16310 (NY); in swamps of Zapata, north of Palpití, León & Loustalot 9575 (NY); Ciénaga de Zapata, north of Bahía de Cochinos, León & Loustalot 9521 (NY) Panicum Sect. Panicum. Panicum sect. Miliaceae Stapf, Fl. Trop. Afr. 9(4): 640, 646. 1920. Type: Panicum miliaceum L., lectotype designated by A. S. Hitchcock & A. Chase, Contr. U.S. Natl. Herb. 15: 13–15. 1910 Panicum sect. Capillare (Hitchc.) Fernald, Rhodora 21(246): 110. 1919. Panicum [uncategorized] Capillaria Hitchc., N. Amer. Fl. 3(2): 200, 206. 1915. Panicum group Capillaria Hitchc. & Chase, Contr. U.S.  Natl. Herb. 15: 28, 54. 1910, nom. inval., Contr. U.S. Natl. Herb. 17: 461, 476. 1915 Panicum group Diffusa Hitchc., Contr. U.S.  Natl. Herb. 15: 29, 71. 1910, nom. inval. Panicum [uncategorized] Diffusa Hitchc., N.  Amer. Fl. 17(3): 200, 203. 1915 Plants annual or perennial, shortly rhizomatous, cespitose, with herbaceous, erect, occasionally decumbent culms; internodes hollow, occasionally solid in the basal portion, hairy to glabrous. Ligule membranous-ciliate. Blades oblong-lanceolate to lanceolate, linear-lanceolate to filiform, flat or with margins involute, hairy to glabrous. Inflorescence a terminal, exserted panicle, lax, open, less frequently contracted, few to multiflowered. Spikelets ovoid, long-ovoid to long-ellipsoid or lanceolate, glabrous or

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Taxonomic Treatment

hairy, pale or tinged with purple, closed at maturity, solitary or in pairs; lower glume (1/3–)1/2–3/4(−4/5) as long as spikelet, 3−5(−9) nerved; upper glume and lower lemma subequal, equal in length or exceeding upper anthecium in length, (5–)7–9(−15) nerved, acute to acuminate; lower palea as long as the lower lemma to reduced or absent; lower flower staminate or absent; upper anthecium ovoid to broadly ellipsoid, glabrous, not stipitate, smooth, shiny, indurate, brown to blackish at maturity, with simple or compound papillae toward the apex of lemma and palea or with simple papillae over the entire surface. Caryopsis ellipsoid to broadly ellipsoid, with punctiform, elliptic to oblong hilum, embryo 1/3 to more than ½ the length of the caryopsis. This section includes a total of 31 species in the Americas, distributed from Canada to Argentina, with ca. 28 present in Africa, India, Asia, the Pacific Islands, and Australia. Its species are commonly found in open and dry areas, in fields, and savannas. Key to the Species 1. Upper anthecium blackish at maturity; upper glume drooping at maturity, exposing the upper anthecium 2. Spikelet triflowered, with a sterile lower flower, a staminate intermediate, and a perfect upper flower; upper anthecium with the adaxial surface of the lemma toward the lower glume.............................. 42. P. quadriglume 2. Spikelet biflowered, with a sterile lower flower and a perfect upper flower; upper anthecium with the adaxial surface of the lemma toward the upper glume 3. Annual plants, with the lower glume 3/4 to 4/5 the length of the spikelet.............................................................................29. P. exiguum 3. Perennial plants, with the lower glume 1/2 to 3/4 the length of the spikelet 4. Plants 16–20 cm tall; inflorescences 3–6 cm long; spikelets 2.4–2.7 mm long. Guatemala.....................................................30. P. furvum 4. Plants 20–60  cm tall; inflorescences 7–22  cm long; spikelets (2.5–) 2.7–3.3(−3.7) mm long. South America, in Bolivia, Brazil, Paraguay, Uruguay, and Argentina...........................................41. P. peladoense 1. Upper anthecium pale to brownish, and with blackish spots at maturity, upper glume not drooping at maturity 5. Plants of aquatic environments, decumbent, with aerenchymatous culms and sheaths; culms conspicuously branched at the middle and upper nodes........... ...........................................................................20. P. aquarum 5. Plants of dry to moderately humid environments, cespitose, with culms and sheaths without aerenchyma; culms simple to sparsely branched in the middle and upper nodes 6. Inflorescences contracted, spiciform to subspiciform, with first-order branches short and spikelets appressed on the branches 7. Spikelets 2.8–3.8 mm long; lower palea as long as the upper anthecium; upper anthecium with two basal scars..................................................... ......................... 25. P. chaseae

Key to Ungrouped Panicum Sections and Species

105

7. Spikelets (4.2–)5–6  mm long; lower palea reduced to 1/2 or less the length of the upper anthecium; upper anthecium with a circular ring at base 8. Inflorescences few flowered, 3–6 cm long; lower glume 3/4 or more the length of the spikelet, separated from the upper glume by a conspicuous internode. Brazil.......................... 37. P. magnispicula 8. Inflorescences multiflowered, (5–)10–20 cm long; lower glume 1/2 or less the length of the spikelet, without a distinct internode between the lower and upper glume. Mexico...............................................................26. P. decolorans 6. Inflorescences lax, diffuse, with first-order branches divergent, spikelets scattered on the branches, rarely adpressed 9. Inflorescences drooping at maturity due to peduncle breakage 10. Plants annuals; lower palea absent to vestigial, up to 0.3 mm long.. .....................................................................23. P. capillare 10. Plants perennial; lower palea elliptic, about equal in length to the upper anthecium........................................................22. P. bergii 9. Inflorescences not drooping at maturity 12. Plants robust, up to 3 m tall; sheaths with urticant hairs................... ....................................................33. P. hirsutum 12. Smaller plants, up to 1.20 m tall; sheaths without urticant hairs 13. Perennial plants 14. Spikelets (3.3–)3.6–6.4 mm long 15. Spikelets 5–6.4 mm long; upper glume and lower lemma 3.2–4  mm longer than the upper anthecium.................................24. P. capillarioides 15. Spikelets (3.3–)3.6–4.5  mm long; upper glume and lower lemma 0.8–1.5  mm longer than the upper anthecium 16. Plants glaucous; basal leaves enrolled...................32. P. hallii 16. Plants not glaucous, basal leaves not enrolled 17. Spikelet with a stipe at its base; lower and upper glume separated by an internode 0.5  mm long; upper glume 7–9 nerved; blades linear to filiform, 0.1–0.3  cm wide................................21. P. aztecanum 17. Spikelet without a stipe at its base; lower and upper glume without a distinct internode; upper glume 9–15 nerved; blades linear-lanceolate,

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0.4–1.2  cm P. lepidulum

Taxonomic Treatment

wide..................................35.

14. Spikelets 1.8–3.1 mm long 18. Plants not glaucous; basal leaves not enrolled; blades densely hairy 19. Plants 40–90(–120) cm tall; collar densely hairy; blades erect, ascending ...........................................3 1. P. ghiesbreghtii 19. Plants (6–)15–60  cm tall; collar glabrous; blades patent.........................................27. P. diffusum 18. Plants glaucous; basal leaves enrolled; blades glabrous to sparsely hirsute 20. Spikelets (2.1–)2.7–4  mm long; upper glume and lower lemma longer than the upper anthecium; blades 4–23 × 0.1–0.5 cm......................................... ..........32. P. hallii 20. Spikelets 1.8–2.2 mm long; upper anthecium equal in length to the upper glume and lower lemma; blades 20–35(–50) × 0.5–0.8 cm.............................. ..44. P. tamaulipense 13. Annual plants 21. Upper anthecium usually obovoid, shortly stipitate, with 2 fleshy appendages at the base of the upper lemma .............. ......................................................19. P. alatum 21. Upper anthecium ovoid to ellipsoid, not stipitate, without basal appendages, with two basal scars or with a disk at the base of the upper lemma 22. Spikelets 4.5–6 mm long 23. Spikelets stipitate, with a conspicuous internode between the upper glume and lower lemma; upper anthecium lately deciduous; panicles erect............... ...............40. P. parcum 23. Spikelets not stipitate, without manifest rachilla internodes; upper anthecium early deciduous; panicles nutant........................38. P. miliaceum 22. Spikelets 1.6–4 mm long 24. Lower palea developed, 1/1 to 3/4 the length of the upper anthecium

Key to Ungrouped Panicum Sections and Species

107

25. Upper glume 5(–7) nerved; lower lemma 5 nerved; lower glume 3/4 as long as the spikelet; blades 0.2  cm wide; spikelets appressed on the branches; upper anthecium with a disk at base of the upper lemma........................28. P. ephemeroides 25. Upper glume 9–13 nerved; lower lemma 7–11 nerved; lower glume 1/3 to 1/2 as long as the spikelet; blades 0.4–1.5 cm wide; spikelets diffuse on the branches, upper anthecium with two scars at the base of the upper lemma 26. Spikelets 3.3.–3.6(−3.9) mm long; upper glume and lower lemma 1  mm longer than the upper anthecium; blades lanceolate 25–45  cm long, hirsute; inflorescences 25–40  cm long ...........................39. P. mucronulatum 26. Spikelets 2.3–3.2 mm long; upper glume and lower lemma as long as the upper anthecium or up to 0.3 mm longer; blades linear-lanceolate, 4.5–27  cm long, glabrous to sparsely papillose-pilose.....43. P. stramineum 24. Lower palea reduced, 1/2 or less the length of the upper anthecium or absent 27. Inflorescences with the main axis densely hirsute, covered with rigid tuberculate hairs; spikelets broadly ellipsoid, 3.2–3.7  ×  1.2–1.6  mm; upper anthecium smooth, with a discoid ring at its base............................ 35. P. hispidifolium 27. Inflorescence with the main axis scabrous to slightly hairy at base, without tuberculate hairs; spikelets ovoid, 1.9–2.8(−3.4)  ×  0.8–1.4  mm; upper anthecium transversely rugose to papillose over its entire surface, without a discoid ring at its base 28. Upper antechium with simple papillae on its surface and two scars at the base of the upper lemma, 0.8–1.6  mm shorter than the upper glume and lower lemma….…….34. P. hirticaule 28. Upper anthecium transversely rugose, without scars at the base of the upper lemma, ca. same length as the upper glume and lower lemma.........................45. P. venosum

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Taxonomic Treatment

19. Panicum alatum Zuloaga & Morrone Plants annual, cespitose, 13–95 cm tall, with geniculate to erect culms, branching at lower nodes; internodes 1.5–10 cm long, cylindrical, the lower ones compressed, glabrous to hispid in the upper portion, tinged with purple; nodes glabrous to shortly hairy. Sheaths 2–7  cm long, glabrous to hirsute, with caducous hairs, one margin papillose-petiolate throughout or only toward the distal portion, otherwise glabrous. Ligule membranous-ciliate, 0.7–1.8 mm long; collar glabrous. Blades lanceolate to linear-lanceolate, 2–18 × 0.3–1.7 cm, more or less erect, ascending, flat, glabrous or with the abaxial surface sparsely hairy, base rounded and acute apex, margins scaberulous, the basal ones ciliate. Peduncles up to 30 cm long, glabrous to sparsely hairy. Terminal and axillary inflorescences present, similar, panicles lax, 3.5–23 × 1.5–11 cm; main axis and branch axes glabrous, scaberulous; first-order branches alternate to subopposite, divergent; second-order branches adpressed with spikelets appressed on axes, close together; pulvini glabrous; pedicels claviform, 1–4 mm long, scaberulous, shortly hairy toward the apex to glabrous. Spikelets ellipsoid, 2.4–3(−4.5) mm × 0.7–0.9(−1.5) mm, apex long acuminate, greenish to brownish or tinged with purple; upper glume and lower lemma subequal, 0.7–0.9(−1.8) mm longer than the upper anthecium; lower glume ovate-acuminate, 1.2–1.8(−2.7) mm long, 1/2–3/4 the length of the spikelet, 5–7 nerved, the midrib scabriform; upper glume 2.2–2.7(−3.3) mm long, 7–9(–11) nerved; lower glume and upper glume separated by a short internode, 0.3  mm long; lower lemma glumiform, 2.1–2.8(−3.9) mm long, 7–9(–11) nerved; lower palea absent or reduced, when present broadly lanceolate, 0.2–0.9 mm × 0.2–0.3 mm, apex bilobed to acute, membranous, hyaline, glabrous; upper anthecium ellipsoid to obovoid at maturity, 1.5–1.8(−2.5)  ×  0.6–0.9(−1.2) mm, crustaceous, pale to dark brown at maturity, smooth, strongly papillose over entire surface or with papillae present only toward the apex of the palea, shortly stipitate, with two basal fleshy expansions 0.3–0.4  mm long, light brown at maturity; lemma 7 nerved. Caryopsis ovoid, 1–1.2(−1.7) mm × 0.6(−1) mm, whitish; hilum punctiform; embryo 1/2 the length of the caryopsis. This species is distinguished from Panicum hirticaule by having an obovoid upper anthecium, shortly stipitate, and two fleshy expansions at its base. Key to the Varieties 1. Upper anthecium strongly papillose over its entire surface, opaque....................... ...............................19a. P. alatum var. alatum 1. Upper anthecium smooth, shiny, with papillae towards the apex of the palea........ ..................................................19b. P. alatum var. minus 19a. Panicum alatum Zuloaga & Morrone var. alatum, Ann. Missouri Bot. Gard. 83(2): 224, Fig. 15. 1996. Type: Mexico. Baja California Sur: 15.5 mi S of El Arco, 27°45′N, 113°20′W, 23 October 1959, I. L. Wiggins 15160 (holotype, MO; isotypes, ARIZ 0004831, GH, MEXU 00107771, SD 00000034, SI 002791). Fig. 22 Distribution and ecology (Fig. 23). Southern USA and northern Mexico, in open, rocky places, occasionally on stream banks in humid places; grows from sea level to 1220 m; flowers and fruits between July and December.

Key to Ungrouped Panicum Sections and Species

109

Fig. 22  Panicum alatum var. alatum. (a) Habit. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d–f) Lower palea. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Caryopsis, embryo view. (j.)Caryopsis, hilum view. (a–h, from Gentry 14353, SI; i–j, from Hinton et al. 6423, US)

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Taxonomic Treatment

Fig. 23  Distribution of Panicum alatum var. alatum, P. aztecanum and P. hirticaule var. verrucosum

Panicum capillare L. var. glabrum Vasey ex Brandegee, Proc. Calif. Acad. Sci. 2(2): 211. 1889, is a nomen nudum applied to this species. Additional material examined. USA. Arizona: Phoenix, 25 September 1904, Griffiths 7317 (US). California: overflow land along Colorado River, 1 December 1905, Schellenger 3 (LA). Stateless: East of Desert Center, 16 September 1929, Jones 24493 (MO); Gila bottom, Schott 16 (MO) Mexico. Baja California Sur: Purísima, 17 February 1889, Brandegee 8 (LA, US); 12 mi. S of El Solito, 27°53′, 113°48′W, 21 October 1959, Wiggins 15135 (MEXU, MO); 11 mi. S of Millers landing, 19 October 1959, Wiggins 15097 (GH, ISC); 1 mi SW of jct. of road to Huatamote along Hwy. Mex. # 1, 365 m, 29 January 1977, Reeder & Reeder 6718 (MEXU); Cuesta de Chuenque, ca. 22 km south of Loreto on road to Puerto Escondido, 25°51′N, 111° 25′W, 10 October 1964, Carter & Ferris 4726 (LA, MEXU). Durango: Ceballos, Zona del Silencio, 1050 m, 25 October 1978, Martínez 1313 (MEXU). Guerrero: Coyuca District, Cutzamala, 8 September 1934, Hinton et al. 6423 (GH, NY, US). Michoacán: Apatzingán, 4 mi W of Apatzingsán, 1200 ft, 8 August 1941, Leavenworth & Hoogstraal 1371 (MO). Nayarit: 2 miles southeast of Las Vacas, 60–90 m, 21 September 1960, McVaugh

Key to Ungrouped Panicum Sections and Species

111

19291 (US). San Luis Potosí: 12  miles east of Ciudad Valles, 17 July 1963, McGregor et al. 840 (US); two km E of Santa Catarina along Hwy. 70, 1220 m, 23 October 1978, Reeder & Reeder 7045 (LA). Sinaloa: El Quelite cut-off to Rt. 15, 34 mi. S of Elota, 10 September 1962, Skorepa 116 (MO, WIS), pr. p.; Cerros Navachiste, about Bahía Topolobampo, 26/30 September 1954, Gentry 14353 (MEXU, SI); outskirts of Topolobampo, 9 October 1966, Gould 12110 (GH, US). Sonora: 54.5 mi. S of Hermosillo, 21 September 1974, de Wet & Harlan 4354 (MEXU, MO); 35.8 mi SE of Guaymas along Mex. 15/22 September 1974, de Wet & Harlan 4732 (MEXU); Guaymas, 17 September 1908, Hitchcock 3553 (US), 1887, Palmer 208 (LA, NY, US), 346 (US); near Guaymas, on rocky, brush-covered slopes just north of Miramar Beach, 7 October 1966, Gould 12071 (LA). Tamaulipas: between Cerro Mante and Tampico village of Granja Trinidad, 31 August 1971, Beetle et al. M-1211 (ISC, MO). Without locality, 1897, Palmer 249 (US), 251 (GH, US) 19b. Panicum alatum var. minus (Andersson) Zuloaga & Morrone, Ann. Missouri Bot. Gard. 83(2): 226. 1996. Panicum hirticaule J. Presl var. minus Andersson, Kongl. Svenska Vetensk. Akad, Handl., n.s. 1853: 135. 1855. Type: Ecuador. “In insulis Chatham et Charles” May 1852, N.  J. Andersson s.n. (holotype not known; isotype, GH; probable isotypes, MO-2098647, NY) Panicum hirticaule J. Presl var. majus Andersson, Kongl. Svenska Vetensk. Akad, Handl., n.s. 1853: 135. 1855. Panicum hirticaule var. majus Walp., Ann. Bot. Syst. 6: 952. 1861[1864], nom. illeg. hom. Type: Ecuador. Colon Archipelago: “Hab. in insula Indefatigable”, May 1853, N.  J. Andersson s.n. (holotype not known; isotype, GH) It is separated from var. alatum by its smooth, non-papillose upper anthecium over the entire surface. Distribution and ecology (Fig. 29). Variety widely distributed in the southern USA and northern Mexico, where it grows in clay or sandy soils; common along roadsides. Occasional in El Salvador, Honduras, Nicaragua, and Venezuela and also found in the Galapagos Islands, Ecuador, where it grows in rocky soils. It is found from sea level to 1860 m asl. Additional material examined. USA.  Arizona: San Bernardino Ranch, 28 August 1892, Mearns 738 (US); Tucson, 2 September 1903, Thomber 176 (LA, MO), Santa Cruz Valley near Tucson, 20 June 1881, Pringle 13940 (MO); Tucson, roadside in city, 15 September 1908, Hitchcock 3494 (US); Patagonia, Santa Cruz County, Hitchcock 3675 (US); Santa Cruz valley near Tucson, 1891, Pringle 197, 464 (US); Fourt Huachuca, Wilcox 303 (US); Yavapai Co., near Kirkland, 5 October 1930, Peebles et al. 7417 (LA). California: Bed of Fort Dry Lake, 24 miles east of Blythe, 350 ft, 30 November 1937, Grinnell & Grinnell 1082a (JEPS, LA); East of Desert Center, 16 September 1929, Jones s.n. (LA 406771). New Mexico: Gila, 29 August 1880, Greene 13107 (MO); plains of Rio Gila, 29 August 1880, Greene 258 (US); Doña Ana County, Mesilla Valley, 11 October 1904, Wooton s.n. (MO-1314437). Texas: El Paso, 23/24 August 1915, Hitchcock 13333 (US)

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Taxonomic Treatment

Mexico. Baja California: Sierra Juárez, 2 October 1966, Moran 13664 (MO). Baja California Sur: San José del Cabo, 17 October 1899, Brandegee 42 (LA, US); Llano Caquihui, Sierra de la Giganta west of Los Dolores, 24° 26′N, 110° 52′W, 17 November 1959, Wiggins 15510 (GH); 3.3 miles southwest of San Ignacio, 27° 14′N, 112° 53′W, 25 October 1959, Wiggins 15179 (LA); Sierra de la Giganta, Arroyo at northwest base of Cerro Mechudo, 24° 48′N, 110° 44′W, 2 November 1971, Moran 18878 1/2 (LA). Campeche: on W edge of Escárcega in disturbed area, 21 December 1972, Reeder & Reeder 6098 (MO). Chiapas: Chiapa de Corzo, El Chorredero 5.6  miles east of Chiapa de Corzo along Mexican highway 190, 2500 ft, 21 June 1966, Laughlin 1113 (US); Mun. Tuxtla Gutierrez, 1800 ft, 1 July 1965, Breedlove 10630 (US); about 27 miles northwest of Mexico-Guatemala border on the road to Comitán, 900 m, 23 August 1953, Reeder & Reeder 2082 (LA, MEXU). Chihuahua: Chihuahua, 4800  ft, Soderstrom 896 (USA); Meoquí, 24 August 1935, Le Sueur 032 (US); Llano de Chilicote 7  miles east of Chilicote Station, 26 September 1938, Johnston 7992 (GH, US). Coahuila: El Mesteño, Ejido 4 de March, 28 July 1976, Roig s.n. (ANSM). Guerrero: 6 km W of Colotlipa, 20 August 1978, Blanco et al. 512 (MEXU). Jalisco: Guadalajara, Río Grande de Santiago, 27  km E of Guadalajara, 5200  ft, 26 August 1941, Leavenworth & Leavenworth 1875 (MO); Puente Grande, 13 miles Se of Guadalajara, on road to Aguascalientes, 5000  ft, 17 September 1959, Soderstrom 647 (US); Tecolotlán, 1500 m, 5 October 1970, Díaz Luna 2044 (MEXU). Michoacán: Apatzingán, jungle at La Majada, 5 miles west of Apatzingán, 1000 m, 5 August 1941, Leavenworth & Leavenworth 1335 (US); Chavinda, 27 September 1946, Hernández Xolocotzi et al. X-2791 (US); ca. 5 miles north of Cotija and 22 miles south of Jiqauilpán, 6000–6200  ft, King & Soderstrom 4679 (LA, NY, US). Morelos: 2  km NW of Axochiapan, 10 August 1950, Walther 63 (US); 12 km NW of Axochiapan, 3500 ft, 10 August 1950, Lyons Jr. 59 (US), 83 (MEXU, US). Oaxaca: 12 km northeast of the city of Tehuantepec, along the Pan-American highway (routes 185 & 190), 50 m, 7 July 1958, King 391 (MEXU, US). San Luis Potosí: Palma Court, Valles, 8 August 1961, Gould 9558 (ISC, LA, MO, US). Sinaloa: about 15 miles southeast of Villa Unión, in disturbed area along the road, 23 September 1953, 90 m, Reeder & Reeder 2430 (LA, MEXU, US); vicinity of Labradas, 18 September 1925, Ferris & Mexía 5070 (MEXU); without locality, 27 August 1891, Palmer 1554 (F, NY). Sonora: 10 miles south of Hermosillo, 17 September 1908, Hitchcock 3541 (US); Hermosillo, 17 September 1908, Hitchcock 3573 (US); Guaymas, 17 September 1908, Hitchcock 3547 (US); Alamos, 16/30 September 1890, Palmer 750 (GH, US); Pinacate Region, plains northwest of Sierra Blanca, 31 33 N, 113 27 W, 30 October 1982, Ezcurra s.n. (MEXU 487793); Pinacate Junction, Pinacate Región, 7.5 km east of Los Vidrios, 31 59 N, 113 21 W, 215 m, 14 September 1986, Felger 86-339 (MEXU). Tamaulipas: in the vicinity of Ciudad Mante, Sánchez-Ken & Nieto M. 430 (MEXU). Veracruz: Puente Nacional Veracruz, Jalapa-Veracruz highway, Emiliano Zapata, 250 m, 10 August 1982, Dorantes et al. 1510 (F, MEXU) Honduras. Valle: 3 km N of Jícaro Galán along highway 1 to Tegucigalpa, 5 October 1986, Davidse & Pilz 31651 (MO, SI), savanna on heavy black clay, heavily grazed and disturbed; 3 airline km W of San Lorenzo along the road to Amapala, 5 October 1986, Davidse & Pilz 31704 (MEXU, SI).

Key to Ungrouped Panicum Sections and Species

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El Salvador. La Unión: La Unión, 13 November 1911, Hitchcock 8794 (US). Sonsonate: Acajutla, 10 August 1951, Rohweder 1648 (MO) Nicaragua. León: Puente Izapa, km 60 carretera a León, 12°15′W, 86° 44′W, 13 July 1981, Moreno 9838 (MO) Venezuela. Zulia: Guarero, Guajira, 25 December 1954, Vareschi 3745 (VEN) Ecuador. Colon Archipelago: San Cristóbal Island (Chatham Island), Wreck bay, 21 February 1905, Stewart 1293 (GH, MO, NY, SI); Wreck Bay, Chatham Island, 17 April 1932, Howell 8590 (GH); Chatham Island, May 1899, Snodgrass & Heller 548 (GH); Charles Island, May 1899, Snodgrass & Heller 454 (GH); Española, landing site on north coast, beach area and area to El Chaco, 3 July 1986, Lawesson 3143 (MO); Indefatigable Island, Academy Bay, 4 May 1932, Howell 9076 (F, GH, MO, NY); Indefatigable Island, 19 April 1923, Wheeler et  al. 122 (NY); Gardner Island, near Hood Island, 22 April 1932, Howell 8728 (GH, US); Gardner Island, May 1899, Snodgrass & Heller 648 (GH); Charles Island, Post Office Bay, 23 April 1932, Howell 8827 (GH, US); Hood Island, May 1899, Snodgrass & Heller 722 (GH); San Cristobal Island, Wreck Bay, 24 February 1964, Fournier 283 (US); Barrington Island, May 1899, Snodgrass & Heller 492 (US); Jervis Island, 6 June 1932, Howell 9757 (GH); South Seymour Island, May 1899, Snodgrass & Heller 612 (GH); North Seymour Island, 11 June 1932, Howell 9965 (GH), May 1899, Snodgrass & Heller 565 (GH) 20. Panicum aquarum Zuloaga & Morrone, Novon 1(4): 185, Fig. 1, 2. 1991. Type: Venezuela. Guárico: Dpto. Infante, Parque Nacional Aguaro-Guariquito: Congriales de la Gorra, between La Esperanza and Mesa de Cambao, ca. 9°12′-9°16′ N and 67° 48′-67°60′ W, 60 m, December 1981, F. Delascio, R. A. Montes & G.  Davidse 11206 (holotype, VEN 222482; isotypes, MO-105074, SI 002793). Fig. 24 Annual plants, 0.90–1.30 m tall; culms erect, rooting at the lower nodes and markedly branching at the upper nodes, with aerenchyma; internodes 7–19 cm long, with aerenchyma, glabrous, striate, hollow; nodes glabrous, compressed, brownish. Sheaths 8–18 cm long, slender, glabrous, shiny on the adaxial surface and with aerenchyma, the margins membranous. Ligule ca. 1.2 mm long, shortly membranous at the base and long ciliate at the apex, adaxial surface pilose; collar glabrous. Blades linear, 11–18 × 0.2–0.4 cm, the lower ones deciduous, glabrous, narrowed toward the base, the margins scaberulous, midrib conspicuous in the basal zone. Terminal inflorescences with a peduncle up to 25 cm long, scaberulous; panicles lax, pyramidal, multiflowered, 35–40 × 12–20 cm; axillary inflorescences numerous, minor, similar to the terminal inflorescence; main axis cylindrical, glabrous, pulvini glabrous, firstorder branches alternate or subopposite, divergent, branch axes triquetrous, scaberulous; second- and third-order branches alternate with scaberulous axes; spikelets congested on third-order branches; pedicels claviform, glabrous. Spikelets solitary, ellipsoid, 3–3.9 × 1–1.2 mm, glabrous, pale or tinged with purple, open at maturity; upper glume and lower lemma subequal, acuminate, exceeding the upper anthecium in length; lower glume ovate, abruptly acuminate, 1.7–2.7 mm long, 1/2 to 3/4 the length of the spikelet, (3–)5 nerved, the midrib scabriform toward the upper portion;

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Fig. 24  Panicum aquarum. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (From Blydenstein 1846, COL)

Key to Ungrouped Panicum Sections and Species

115

internode conspicuous between the lower glume and upper glume; upper glume 3–3.6 mm long, acuminate, (5–)7 nerved, not deciduous at maturity; lower lemma glumiform, 3–3.2  mm long, (5–)7 nerved; lower palea oblong-lanceolate, 1.7–2.1 × 0.6–1 mm, glabrous, hyaline; upper anthecium ellipsoid, 2.1 × 1.2–1.3 mm, crustaceous, pale, with blackish spots at maturity, with a brownish disk at the base; upper lemma 5 nerved, the margins involute; upper palea 2 nerved, with verrucose papillae toward the apex. Caryopsis broadly ellipsoid, 1.5 × 1 mm, whitish; hilum subbasal, punctiform; embryo 1/3 or less the length of the caryopsis. Distribution and ecology (Fig. 25). Species restricted to the extreme north of South America, where it is found in Venezuela, Brazil, and Colombia. It is found in flooded areas and flowers and fruits between September and May. Panicum aquarum differs from the rest of the species of sect. Panicum as it is an aquatic plant, with aerenchyma in the sheaths and deciduous blades toward the base. It is also characterized by open spikelets at maturity, with the upper glume and lower lemma (5–)7 nerved. Additional material examined. Colombia. Meta: Lago Carimagua, flat highlands, under 30 cm of water, 19 November 1963, Blydenstein 1846 (COL, G).

Fig. 25  Distribution of Panicum aquarum, P. bergii var. bergii, P. decolorans and P. ghiesbreghtii

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Brazil.  Amazonas: Mun. Humaitá, 7°31′S, 63°10′ W, campo a 15–17  km da cidade pela estrada 319 ao norte da estrada, entre igarapés Retiro e Bom Futuro, 70  m, 11 May 1980, Gemtchujnicov & Janssen 369 (IBGE, SP). Pará: Marajó, Camará River, September 1902, no collector s.n. (US-1024781); ad Tocantins flumen inter Porto Imperial et Funil, Burchell 8813 (K, W) 21. Panicum aztecanum Zuloaga & Morrone, Ann. Missouri Bot. Gard. 83(2): 229, Fig. 17. 1996. Type: Mexico. Mexico: Temascaltepec, Bejucos, barranca, 610 m, 7 October 1932, G.  B. Hinton 2010 (holotype, US 00604292; isotypes, GBH 002010, MEXU 00398114, MO-368989, NY 1770967). Fig. 26 Plants perennial, cespitose, shortly rhizomatous, not glaucous, with erect culms 50–100 cm tall, branching at the base; internodes 13–16 cm long, 1 mm in diameter, without aerenchyma, glabrous, purplish toward the upper portion; nodes glabrous, dark. Sheaths 2–10 cm long, shorter than internodes, without aerenchyma, glabrous to sparsely hairy, with delicate, silky hairs, margins membranous, glabrous, and long ciliate next to the blade junction, with whitish, silky hairs, up to 3.6 mm long. Ligule membranous-ciliate, 0.4–0.6 mm long; collar glabrous to long-hairy. Blades linear to filiform, 14–30 × 0.1–0.3 cm, involute, exceptionally flat, the basal ones not enrolled, with both sides or with the adaxial side sparsely pilose, the base narrowed, continuing imperceptibly with the sheath, apex long acute, the margins scaberulous, ciliate toward the base. Peduncles long exserted, up to 48  cm long, glabrous, with purplish tints. Inflorescences terminal, exserted, a lax, diffuse panicle, 15–18 × 8–10 cm, not deciduous at maturity; main axis glabrous, scaberulous toward the basal portion, otherwise smooth; first-order branches ascending, flexuous, naked toward the base; second-order branches with 2-3 solitary spikelets toward the apex; pulvini glabrous; pedicels 2–20 mm long. Spikelets long ellipsoid, 3.7–4.5  ×  1.5  mm, acute, glabrous, purplish, stipitate; lower glume ovate-acuminate, 2–3  mm long, 1/2-3/4 the length of the spikelet, 7-nerved, with the midrib scaberulous; upper glume 3.2–4.2 mm long, the inner surface scaberulous toward the apex, 7–9 nerved, not deciduous at maturity; lower and upper glume separated by an internode 0.5 mm long; lower lemma glumiform, 3–4 mm long, 7–9 nerved; lower palea lanceolate, 1.7–2.7 × 0.6 mm, 3/4 the length of the lower lemma, membranous, hyaline, glabrous; upper anthecium ovoid, 2.4–3 × 0.9–1.2 mm, 1.2–1.5 mm shorter than the upper glume and lower lemma, crustaceous, smooth, shiny, pale, the base circular, without scars, pale at maturity; lemma 7 nerved; upper palea with simple papillae and globose microhairs toward the apex. Caryopsis ovoid, 1.8 × 1 mm, whitish; hilum punctiform; embryo 1/2 the length of the caryopsis. Distribution and ecology (Fig. 23). Species endemic to Mexico, found in Pinus and Quercus forests, on slopes, between 600 and 1600  m  asl, flowers and fruits between August and October Species is related to Panicum lepidulum, from which it differs by having linear to filiform blades, long ciliate near the junction with the sheaths, and stipitate spikelets with a distinct internode between the lower and upper glume. Additional material examined. Mexico. Federal District: near the pyramid of Cuicuilco, Rzedowski 2047 (MEXU, US). Guerrero: 5 km NE of Tlatlauqui, Pinus

Key to Ungrouped Panicum Sections and Species

117

Fig. 26  Panicum aztecanum. (a) Plant. (b) Detail of the ligular region. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (a–b, from Hinton 2010, US; c–i, from Rzedowski 17449, US)

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Taxonomic Treatment

and Quercus forest, 14 August 1990, Calzadas 15935 (MEXU). Jalisco: 4.5 mi. north-northeast of Talpa de Allende, 1450–1500  m, 17 October 1960, Mc Vaugh 20345 (US, 2 sheets); Mun. Tecalitlán, Llano Verde, near Los Corrales, Sierra de los Corrales, 1600 m, 24 October 1963, Rzedowski 17423, 17449 (US); Mun. Tecalitlán, Barranca de San Juan de Dios, near Los Corrales, Sierra de los Corrales, 1300 m, 23 October 1963, Rzedowski 17367 (US). Sinaloa: Santa Lucía, east of Panuco, 900–1200 m, 28 August 1935, Pennell 20033 (US) 22. Panicum bergii Arechav. Plants perennial, cespitose, shortly rhizomatous, (10–)30–70(–140) cm tall; culms geniculate, erect, simple, or branching only toward the base; internodes 2–15  cm long, cylindrical, hollow, without aerenchyma, pilose, with tuberculate hairs, more abundant toward the lower portion; nodes purplish, pilose to glabrous. Sheaths striate, pale, densely to sparsely pilose, the margins membranous, ciliate. Ligule membranous at base, then long ciliate, 1–3 mm long; collar pale, with tuberculate hairs to glabrous. Blades linear-lanceolate, (3–)8–35(−60)  ×  0.2–1.2  cm, densely pilose, with long whitish hairs, to glabrescent, the base attenuate, apex acute, margins scabrous and involute to flat, midrib manifest. Inflorescences lax, diffuse, pyramidal, (4–)15–30(−40)  ×  (3–)10–25  cm, deciduous at maturity by peduncle breakage; peduncles subincluded to long exserted, glabrous to hispid; first-order branches divergent, lower ones whorled, the upper ones opposite or alternate; main axis densely hispid, with long whitish hairs, to glabrous, scabrous, triquetrous, branch axes triquetrous, hispid to scabrous, pulvini hairy to glabrous; pedicels triquetrous, scabrous, 3–20 mm long. Spikelets ovoid, 2.1–2.9 × 0.8–1.2 mm, glabrous, pale and tinged with purple, dispersed on the branches; upper glume and lower lemma subequal, 0.3 mm longer than the upper anthecium; lower glume ovate, 1–1.6 mm long, acuminate, 5-nerved, midrib scaberulous towards the apex; upper glume 2.1–2.8 mm long, not deciduous at maturity, 7–9 nerved, midrib scaberulous toward the apex; lower lemma 2–2.7 mm long, 7–9 nerved; lower palea elliptic, 1.4–2.2 × 0.6–1 mm, hyaline, glabrous, the margins finely ciliate; upper anthecium ovoid, 1.5–1.9 × 0.7–1 mm, crustaceous, smooth, glabrous, shiny, pale, brown at maturity, with two basal scars 0.1 mm long, brown at maturity; upper lemma 7 nerved, with prickle hairs at the apex; palea with simple papillae toward the apex. Caryopsis ellipsoid, 1.2–1.4 × 0.7–1 mm; hilum punctiform, embryo half the length of the caryopsis. Key to the Varieties 1. Rachis and panicle branches glabrous; sheaths and blades sparsely hairy to glabr escent..............................................................22a. P. bergii var. bergii 1. Rachis and panicle branches densely hirsute; sheaths and blades densely hispid.. .........................................................22b. P. bergii var. pilosissimum 22a. Panicum bergii Arechav. var. bergii, Anales Mus. Nac. Montevideo 1:147. 1894. Gramíneas Uruguayas: 127. 1894. Type: Uruguay. “Campos del Departamento de Montevideo, San José, Florida, Mercedes, etc.”, J. Arechavaleta s.n. (holotype not known; isotypes, W 19160024641, W 19160024883; probable isotype, US 00148174, fragment ex W). Fig. 27

Key to Ungrouped Panicum Sections and Species

119

Fig. 27  Panicum bergii var. bergii. (a) Habit. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Lower palea. (e) Upper anthecium, dorsal view. (f) Upper anthecium, ventral view. (a, from Cabrera & Zuloaga 32328, SI; b–f, from Rojas 2358, SI)

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Taxonomic Treatment

Panicum pilcomayense Hack, Bull. Herb. Boissier, ser. 2, 7(6): 449. 1907. Type: Paraguay, “Ad margines silvarum in regióne cursus inferioris fluminis Pilcomayo, flor. mens: Maj. 1906. Rojas n. 105”, T.  Rojas 105 (holotype, W 19160024416; isotypes, BAA 00002386, fragment ex G, BAA 00002387, fragment ex B, BAF 00000260, G 00009736, K 000309182, P 00740835, US 00037293, US 00139864, fragment) Panicum bergii Arechav. fo. convoluta Palacios, in Burkart, Fl. Il. Entre Rios, Colecc. Ci. Inst. Nac. Tecnol. Agropecu. 6(2): 309. 1969. Type: Argentina. Entre Ríos: Concepción del Uruguay, 16 December 1961, A. Burkart 22915 (holotype, SI 8094) Panicum burkartii Zuloaga, Bol. Soc. Argent. Bot. 17: 179, Fig.  1. 1976. Type: Argentina. Buenos Aires: Cerro Copelina, Mar del Plata, 1932, C. M. Hicken s.n. (holotype, SI-13460) Local name: “Kapi’í veve” (kapi’í = straw, veve = flying) (Zuloaga 1994) Distribution and ecology (Fig. 25). South American variety, widely distributed in Argentina, Paraguay, Uruguay, Bolivia, and southern Brazil. Less frequent in northeastern Brazil, Guyana, and Venezuela; introduced in the USA.  It inhabits fields, in open and relatively dry places. Local name: “Flying straw” (Palacios 1969; Cabrera 1970; Rosengurtt et al. 1970). This species has a marked variability in plant size, vegetative organs, and inflorescences. Larger specimens were previously characterized as P. pilcomayense; they grow profusely in northern Argentina and Paraguay, and there are isolated specimens in Venezuela, Guyana, and northern Brazil. Smaller specimens were described, for the southern limit of the species distribution, as P. burkartii. After examining abundant material of this taxon, the criterion (previously stated by Zuloaga and Morrone 1996) of considering P. bergii as the valid species to be considered is maintained. Panicum bergii is characterized, together with P. capillare L., by having a breakable inflorescence at the base at maturity, which facilitates the dispersal of the entire panicle as a diaspore. This character was cited by Palacios (1969), Nicora (1978), and Zuloaga (1989). Panicum bergii was introduced during this century in the USA. Additional material examined. USA. Alabama: Mobile, August 1891, Mohr s.n. (US). Georgia: weed around grass plots, Experiment Station, Tifton, 4 August 1938, Swallen 5393 (US). Texas: Galveston Co, just N of Dollar Point, ca. 3 mi. N of Texas City, 24 April 1974, Waller & McAden 2643 (MO); Brazos Co, roadside ditches between F.M. 2154 railroad track, 3 mi, S of College Station, 10 September 1970, Waller 2410 (MO); Galveston Co, Ave. Q between F.M. 517 and S.H. 146, Sam Leon, 17 April 1974, Waller 2620 (MO); Refugio Co, N of Tivoli, 30 April 1949, Swallen 10300 (MO) Venezuela. Apure: Llanos del Alto Apure, Jahn 197 (US). Monagas: alrededores de Jusepín, Garófalo 381 (VEN). Miranda: 4 km W of Santa Teresa, Killip & Tamayo 37046 (F, US). Portuguesa: carretera Barinas-Guanare, km 301, 14 July

Key to Ungrouped Panicum Sections and Species

121

1984, Stergios 6935 (MO, PORT, SI); Dpto. Guanare, ca. 30  km  W of Guanare, along the Guanare-Barinas highway, 8 June 1980, Davidse & Steyermark 18161 (MO, VEN) Guyana. Without locality, Schomburgk 259 (G), 456 (F), 653 (W), 656 (G, K, US), 701 (US) Peru. Loreto: Salinas de Pilluana, Huallaga, Ule 6857 (G, K). Brazil. Pernambuco: Tapera, Pickel 1403 (F, USA). Rio de Janeiro: Ilha do Gobernador, Miers 3429 (K); environs de Rio de Janeiro, Glaziou 13338 (P, K); São Pedro, 10 December 1929, Chase 10151 (F, MO, RB, US); Campos, Sampaio 2971 (US); Universidade rural, km 47, Black & Marano 51-11413 (IAN). Rio Grande do Sul: Boca do Monte, 2 February 1948, Palacios & Cuezzo 4368 (MO); Livramento, Espinilho, Araujo 289 (SI); São Gabriel, 29 July 1952, Beetle 1772 (MO). Roraima: bei der Serra do Mel, Ule 8014 (G, IAN, K, NY, US) Bolivia. La Paz: Prov. Iturralde, Luisita, sabana humeda, W del Rio Beni, Beck & Haase 9921 (K). Santa Cruz: University Farm, “El Vallecito,” Tollersey 2626 (K); Prov. Andres Ibañez, 12 km E of center of Santa Cruz, on road to Cotoca, 19 January 1989, Nee 37618 (MO) Paraguay. Alto Paraguay: Puerto Casado, December 1916, Rojas 2358 (SI). Central: Asunción, S de Villa Hayes, Rojas 2762 (US); en palmar al SW de Villa Hayes, Rosengurtt B-5623 (K). Concepción: Villa Concepción, 23 May 1876, Balansa 12 (G). Misiones: San Juan a San Ignacio, km 208, October 1951, Burkart 18643 (SI). Paraguarí: Cerro Peron, February 1875, Balansa 15 (G, P). Presidente Hayes: El Pedernal, al S de Concepción, Burkart 18397 (SI). Argentina. Buenos Aires: Balcarce, Hill Plot, INTA, Dalby 44 (NY); Arroyo Los Paretos, sobre ruta 51, 23 February 1981, Villamil 1994 (MO, SI); Punta Lara, 28 January 1978, Zuloaga & Sáenz 573 (SI), 24 March 1984, Zuloaga 1889 (MO, SI), 17 April 1987, Zuloaga et al. 3072 (SI); March del Plata, Parque Paz, Hicken 22 (SI); Los Talas, Cabrera 2484 (NY); La Plata, 14 March 1940, Cabrera 7008 (GH, M, NY); Maipú, 26 February 1947, Krapovickas 2906 (F, LIL, MO, SI); Hurlingham, Alvárez 603 (LIL, NY, SI); Quilmes, Rodríguez 99 (GH, NY); Olivos, Dawson 424 (F, NY). Chaco: Colonia Benítez, 17 January 1974, Quarín et al. 1862 (CTES, SI); Ruta 16, 4 km from Belgrano bridge, 23 February 1978, Renvoize et al. 3583 (MO, SI). Córdoba: Marcos Juárez, 25 October 1904, Stuckert 14434, 14798 (MO), 14799 (BAF, SI); Capilla del Monte, Nicora 2724 (SI); Río Primero, 26 January 1906, Stuckert 15511 (MO); Manfredi, Krapovickas 6681 (GH, SI). Corrientes: La Cruz, 7 December 1936, Parodi 12363 (MO); Río Paraná near Arroyo Sombrero, 27 November 1978, Renvoize 3640 (MO, SI); Río Miriñay y Ruta 23, Quarín 1639 (CTES, SI, US); Est. Exp. Paso de la Patria, Pérez 8 (SI); 21 km S de Loreto, Campo de Sanabria, 13 June 1974, Schinini & González 9293 (CTES, SI); Garruchos, 22 October 1954, Burkart 19692 (SI). Entre Ríos: Gualeguay, Burkart 17743 (SI); 2–3 km al N de Ceibas, Troncoso et al. 2638 (SI, US). Formosa: Fortín Pilcomayo, Krapovickas 1251 (LIL); Ruta 11 vieja, al N de Arroyo Francesa Cué, Guaglianone et al. 270 (SI), 360 (MO, SI, US); al N de San Pablo, Guaglianone et al. 595 (SI, US); Ruta 81, 10 km al W de Pirané, Guaglianone et al. 667 (MO, SI, US). Jujuy: Sierra de Zapla, Mina 9 de Octubre, 11 May 1981, Cabrera et al. 32704 (SI). La

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Taxonomic Treatment

Pampa: Anguil, Steibel 2451 (SI, US); Catriló, Fortuna 2 (US); Sierras de Lihuel Calel, Troiani & Steibel 2827 (SI); entre General Pico y Trebolares, Martínez 6077 (SI). Misiones: camino de Cerro Azul a Apóstoles, 19 October 1975, Zuloaga & Deginani 535 (SI); Escuela Pascual Gentilini, Cerro Ceferino, Zuloaga et al. 924, 3273 (SI), Cabrera et  al. 28697 (MO, SI); Apóstoles, Burkart 14455 (F, US); Loreto, Montes 14642 (F, MO), Zuloaga & Deginani 457 (SI, US); camino de Oberá a Alem, October 1975, Zuloaga & Deginani 514 (SI, US); Ruta Prov. 105, Villalonga, 13 April 1985, Zuloaga et  al. 2323 (MO, SI); de San José a Posadas, antes del Arroyo Pindapoy Chico, 16 April 1979, Zuloaga et  al. 3151 (SI); antiguo aeródromo Cataratas, October 1975, Zuloaga & Deginani 477 (SI). Santa Fe: Laguna Paiva, Hosseus 43 (CORD); Las Toscas, Quarín 110 (CTES, SI); Ruta Prov. 2, 12  km al S de Tostado, Lewis 629 (SI, US); Curupaytí, cerca de Ceres, Cabrera et al. 25988 (SI); Las Gamas, Lewis & Pire 565 (MO, SI, US). Santiago del Estero: Ruta 34, 11 km al SE de Argentina, Rotman et al. 100 (SI, US). Tucumán: barrancas del Arroyo La Falda, Venturi 7530 (F, SI); Las Cejas, Venturi 2539 (LIL, SI, US); Chañar Pozo, December 1919, Venturi 644 (GH, LIL, SI) Uruguay. Canelones: Toledo, Legrand 1455 (F); Río Santa Lucía, Gallinal et  al. PE-2362 (US). Cerro Largo: Río Negro, Gallinal et  al. PE-1290 (US). Flores: Río Yí y Arroyo Marincha, Rosengurtt B-480 (US), Rosengurtt B-681B (F). Montevideo: Montevideo, Arechavaleta 24a (US). Paysandú: Paysandú, Rosengurtt B-2228 (RB, US), 6 February 1981, Cabrera & Zuloaga 32328 (SI). Rocha: Palmar near Chafalote, 7 km NW of 19 de Abril, Bartlett 21345 (GH). San José: Kiyu, Del Puerto 685 (F); Estancia Pascual, Herter s.n. (MO-1098810). Soriano: Arroyo Grande, March 1928, Herter 544 (F, GH, M, MO, NY, US); Jackson, January 1944, Rosengurtt 5411 (MO). Tacuarembó: camino a Rivera, 32 km de Tacuarembó, 8 January 1981, Cabrera & Zuloaga 32416 (MO, SI) 22b. Panicum bergii var. pilosissimum Zuloaga, Hickenia 1(27): 151. 1978. Type: Argentina. Misiones: San José, Escuela Agrotécnica Pascual Gentilini, 13 October 1977, A. L. Cabrera, S. M. Botta, R. Kiesling, A. D. Rotman, N. M. Tur & F. O. Zuloaga 28691 (holotype, SI 008101) It is distinguished from the typical variety by having densely pilose sheaths, leaf blades, and panicles. Distribution and ecology (Fig. 34). Brazil, in Rio Grande do Sul and Argentina, occurring in the provinces of Misiones and Corrientes. It grows in fields. Additional material examined. Brazil. Rio Grande do Sul: Tupaceretan, Rambo 9845 (US) Argentina. Corrientes: Arroyo Chimiray, 12 April 1974, Krapovickas et  al. 25201 (CTES, SI); de Azara a Santo Tomé, October 1975, Zuloaga & Deginani 547 (LP). Misiones: Apóstoles, Burkart 14367 (SI); Escuela Agrotécnica Pascual Gentilini, Cerro Ceferino, 26 November 1987, Zuloaga et  al. 3271 (SI); Loreto, Montes 3478 (SI); Posadas, Establecimiento Santa Inés, Parodi 4490 (BAA), Canoniero 92 (BAA)

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23. Panicum capillare L., Sp. Pl. 1: 58. 1753. Milium capillare (L.) Moench, Methodus: 203. 1794, nom. illeg. hom., non Rottb., 1778. Chasea capillaris (L.) Nieuwl., Amer. Midl. Naturalist 2: 64. 1911. Leptoloma capillaris (L.) Smyth, Trans. Kansas Acad. Sci. 25: 86. 1913. Type: United States of America. Virginia: without locality, J.  Clayton 454 (lectotype, BM 000042600, designated by A.  S. Hitchcock, Contr. U.S.  Natl. Herb. 12: 118. 1908; isolectotype, US 00148328, fragment ex BM). Fig. 28 Panicum bobarti Lam., Encycl. 4(2): 748. 1798. Type: USA.  Virginia: without locality, Bobarti s.n. (holotype, P 00740937; isotype, US 00148183, fragment and photo ex P) Panicum capillare L. var. agreste Gatt., Tennessee Fl.: 94.1887. Type: USA. Tennessee: Ridgetop, Summer Co, 14 September 1882, A. Gattinger s.n. (lectotype, TENN, designated by A. S. Hitchcock & A. Chase, Contr. U.S. Natl. Herb. 15: 60. 1910; isolectotype, US 00148329, fragment and photo ex TENN) Panicum capillare L. var. brevifolium Vasey ex Scribn., Bull. Div. Agrostol., U.S.D.A. Div. Agrostol Bull. 5: 21. 1897, non P. brevifolium L., 1753. Panicum barbipulvinatum Nash, Mem. New York Bot. Gard. 1: 21. 1900, nom. superfl. Leptoloma barbipulvinata (Nash) Smyth, Trans. Kansas Acad. Sci. 25: 86. 1913. Milium barbipulvinatum (Nash) Lundell, Amer. Midl. Naturalist 4: 212. 1915. Panicum capillare L. subsp. barbipulvinatum (Nash) Tzvelev, Novosti Sist. Nizh. Rast.: 18. 1968. Panicum capillare L. var. barbipulvinatum (Nash) Mc Gregor, Phytologia 55(4): 256. 1984. Type: United States of America. Montana: Gallatin, Manhattan, on a shaded sand bar in the Gallatin River, 19 July 1895, C. L. Shear 436 (holotype, US 00133125) Annual plants, cespitose, 30–80 cm tall, culms erect to subgeniculate at the base, branching at the middle and upper nodes; internodes 2.4–7  cm long, cylindrical, pale and tinged with purple, pilose, with tuberculate hairs, more abundant toward the upper portion of the internode; nodes purplish, covered with whitish hairs. Sheaths larger than internodes, 2–10 cm long, striate and covered with long tuberculate hairs. Ligule membranous-ciliate, 0.9–1.5  mm long; collar brown, hairy. Blades linear-lanceolate, flat, 5–26 × 0.4–1.5 cm, slender, base subcordate, hispid, with scabrous margins, the basal ones with deciduous tuberculate hairs. Inflorescences lax, diffuse, pyramidal, 13–37 × 7–24 cm, included when young in the upper sheaths, then exserted, panicle fully deciduous at maturity by breakage of the peduncle; peduncles long exserted at maturity, up to 30 cm long, flattened, fragile, pale or tinged with purple, glabrous to sparsely hairy near the apex; spikelets scattered on the branches; main axis scabrous, triquetrous, densely hispid in the lower portion and sparsely hispid toward the panicle apex, pulvinus pilose; firstorder branches alternate, less frequently opposite or verticillate, divergent; secondorder branches divergent; axis of the branches scabrous; pedicels pilose, scabrous, 0.5–2.8 cm long. Spikelets narrowly ellipsoid to lanceolate, 2–2.8(−3.2) × 0.5–1 mm, pale and tinged with purple, apex long acute, glabrous; upper glume and lower lemma subequal, 0.8–1.2 mm longer than the upper anthecium; lower glume (0.9– )1.2–1.6 mm long, 1/2 the length of the spikelet or slightly less, 3–5 nerved, acute;

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Fig. 28  Panicum capillare. (a) Base of the plant. (b) Inflorescence. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Upper anthecium, ventral view. (From Burkart 23269, SI)

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upper glume 1.8–2.7(−3.1) mm long, not drooping at maturity, 7–9 nerved, midrib scaberulous; lower lemma 1.9–2.6(−3) mm long, glumiform; lower palea absent or vestigial, when present bilobed, up to 0.3  mm long, lower flower absent; upper anthecium ellipsoid, 1.5–1.9 × 0.5–0.8 mm, crustaceous, glabrous, glabrous, shiny, pale, brownish at maturity, without basal scars; upper lemma 7 nerved; upper palea with simple papillae toward the apex. Caryopsis broadly ellipsoid, 1.3–1.5 × 0.7–1 mm; hilum punctiform, embryo half the length of the caryopsis. Distribution and ecology (Fig. 29). Species native to North America. Naturalized in Europe and South America, being found adventitiously in South America in Brazil, Uruguay, Chile, and Argentina; it is occasional in Bermuda. It reaches up to 2200 m asl. Hitchcock (1951) and Davidse (1987) mention that in this species, dispersal occurs through the whole panicle, which is carried by the wind at maturity. The inflorescence in this species is more than half the length of the plant. Additional material examined. Canada. Nova Scotia: Windsor Junction, 31 August 1920, Fernald & Long 19754 (US). Ontario: York Co, Islington, 29 August 1946, Dore 46-137 (US). Quebec: Ottawa, 18 August 1936, Dore s.n. (US-1647912); St. Lawrence River, 3 mi. above Prescott, 25 September 1950, Dore 12887 (US); Soulange Co., Coteau du Lac, St. Lawrence River, 15 September 1959, Dore 10544 (US) USA. Arizona: near Tuba City, Coconino City, 27 September 1935, Kearney & Peebles 12868 (US). California: Phillipsville on the Redwood hwy., 20 July 1924, Heller 13864 (US). Colorado: S Platte River just W of Brighton, 4800  ft, 14 September 1941, Robbin 860 (US); Delta Co. between Delta and Austin, 28 July 1938, Goodding & Goodding 243 (US). District of Columbia: Washington, 4 September 1891, Blanchard s.n. (US-312003). Idaho: Lewiston, September 1929, Gail s.n. (US-3168275); Canyon Co, Falk’s Sore, 2,200 m, 7 July 1910, McBride 332 (US). Illinois: banks of Embarres river, 5 miles S of Charleston, Macuszek 271 (LIL). Iowa: Ames, Ball 20, 120 (US); Palo Alto Co, Freedom Twp, sec. 8, Medium Lake, 26 August 1940, Hayden 8220 (US). Kansas: NW of St. Francis, Steller & Jacobs 335 (LIL). Kentucky: Doyle Valley, W of Chaumont Road, 1 October 1949, Lix 738 (US). Maine: York Co, Ocean Park, 2 September 1931, Moldenke 6377 (US). Maryland: near Potomac, NW of Cabin John, 5 October 1904, Chase 2726 (US). Massachusetts: Harwich, 29 July 1919, Fernald & Long 17828 (US). Missouri: 2  miles NW of Holden, Steyermark 72470 (LIL). Montana: 17 mi. above Blendive, 20 July 1883, Ward s.n. (US-952984). Nebraska: North Platte, June 1896, Plank 38 (US). New Jersey: Somerset Co, Watchung, 29 August 1930, Moldenke 1372a (US). New Mexico: Animas Creek, Metcalfe 1138 (LIL). New York: Princes Bay, 6 October 1894, Kearney Jr. s.n. (US-742048). North Carolina: without locality, 1885, McCarthy s.n. (US-952946). North Dakota: Morton Co, roadside between Mandan and Bismark, Muñoz & Rollins 1006 (US). Ohio: Cincinnati, Spring Grove, 21 August 1905, Braun s.n. (US-2663130). Oregon: MacKenzie’s bridge, 1740 ft, 21 July 1927, Hitchcock 23470 (US); Roseburg, 14 July 1908, Hitchcock 2798 (US). Pennsylvania: Easton, 1894, Porter s.n.

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Fig. 29  Distribution of Panicum alatum var. minus, P. capillare, and P. capillarioides

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(US-480434). South Dakota: Belle Fourche, 4 August 1897, Griffiths 399 (US). Texas: New Braunfles, 20 June 1910, Hitchcock 5197 (US). Vermont: Passumpsic River, Lyndon, 11 September 1931, Fernald 519 (US). Virginia: Arlington, Chase s.n. (LIL). Washington: Spokane, 14 July 1908, Chase 4986 (US). Wyoming: Newcastle, 24 August 1897, Griffiths 684 (US) Bermuda. North Shore road, 31 August 1905, Brown & Britton 21 (NY, US); Paget, roadside, 21 August 1913, Collins 151 (NY, US), Paget 152 (NY) Mexico. Baja California: E of San Telmo, Rancho San José, east of San Telmo, 14 September 1930, Wiggins & Demaree 4833 (F, NY, US); banks of Río Santo Domingo, Wiggins & Demaree 4793 (US); Sierra Juárez, Rancho La Botella, Moran 16483 (US); La Bocana de Santo Tomas, on Punta Chino road, 31 32 N, 116 39 W, 20 September 1980, Moran 29255 (MEXU). Coahuila: Municipio de Villa Acuña, Serranías del Burro, Rancho El Bonito, ca. 29°01  N, 102°07  W, 1620  m, 18 September 1977, Valdés Reyna & Riskind 1230 (ANSM) Brazil. Rio Grande do Sul: Estaçao estrada do Ferro, Vacaria, Arzivenco 640 (BLA) Chile. Bio Bio: Los Angeles, Pfister 60 (LIL). Ñuble: Recinto, Castillo s.n. (US-2044754). Talca: Talca, Ruíz 6 (BAA). Valparaíso: Limache, January 1933, Garaventa 2847 (BAA, CONC). Without locality, March 1927, Gunther 5 (US) Argentina. Buenos Aires: Partido Tres Arroyos, al E de la ciudad, sobre camino que conduce a la ruta 72, 21 February 1988, Villamil 5889 (SI); Tigre, 22 September 1944, Lanfranchi 186 (SI); Daireaux, Nicora 6806 (BAA, LIL, MO); Santa Catalina, 1899, C. Spegazzini s.n. (ex herb. Parodi 2433 en BAA). C.A.B.A.: Villa Ortuzar, Parodi 1750 (BAA), 8218 (BAA, SI); Paternal, Parodi 590 (BAA). Entre Ríos: Isla Cambacuá, frente a Concepción del Uruguay, December 1957, Nicora 6511 (SI), Burkart 20264 (SI, US); vías entre Ayuí y D. Roberto, 11 March 1969, Burkart et al. 23269 (SI, US). Jujuy: alrededores de Aeropuerto El Cadillal, 14 March 1987, Zuloaga & Morrone 3060 (MO, SI), 19 January 1988, Zuloaga & Deginani 3626 (SI). La Pampa: Estación Experimental de Anguil, Legaspi s.n. (BAA 2131); Estancia El Caldén, Potrero Santa Aurelia, Rúgolo 675 (BAA, CTES); Trenel, Fortuna s.n. (LIL); Santa Rosa, Steibel 3473 (SI); E. Castex, Troiani & Steibel 4094 (SI); Caleufú, Troiani & Steibel 3902 (SI, US). Neuquén: Plottier, Ragonese s.n. (BA 14287, BAA). Río Negro: Ing. Huergo, 17 February 1948, Burkart 15943 (BAA, SI); Villa Regina, Fabris 1080 (BAA, LP); márgenes Río Negro-Allen, Hunziker 12 (BAA, LIL). San Luis: Estación Experimental Agropecuaria San LuisINTA, Villa Mercedes, Anderson 2096 (BAA) Uruguay. Montevideo: Montevideo, Felippone s.n. (SI) 24. Panicum capillarioides Vasey, Contr. U.S.  Natl. Herb. 1(2): 54. 1890. Type: USA. Texas: Cameron County, Point Isabel, G. C. Nealley 634 (holotype, US 00133129; probable isotypes, K 000674453, US 00289443). Fig. 30 Plants perennial, cespitose, shortly rhizomatous, with erect culms 30–75 cm tall, simple or sparsely branching at the middle and upper nodes, internodes cylindrical, hollow, hairy or glabrous, 1–2  mm in diameter; nodes hairy. Sheaths equal to or smaller than internodes, hirsute, hairs not urticating, green or tinged with purple, the

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Fig. 30  Panicum capillarioides. (a) Habit. (b) Ligular region. (c) Portion of inflorescence. (d) Spikelet, lateral view. (e) Spikelet, dorsal view. (f) Spikelet, ventral view. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Caryopsis, hilum view. (j) Caryopsis, embryo view. (k) Lower palea. (From Zuloaga et al. 9687, SI)

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margins ciliate. Ligule membranous-ciliate, the membranous portion 0.5 mm long, cilia 1.5  mm long; collar pilose. Blades linear-lanceolate, 12–30  ×  0.5–0.8  cm, erect, ascending, flat, rounded at base, apex attenuate, hirsute, with tuberculate hairs, to sparsely pilose, scabrous, the midrib manifest. Terminal inflorescences exserted, not drooping overall at maturity; panicles lax, diffuse, 15–30 × 10–12(−26) cm wide; peduncles subincluded to exserted, hirsute, main axis wavy, hispid toward the base to glabrous, scaberulous; first-order branches alternate to subopposite, the lower ones occasionally subverticillate, 8–18 cm long, naked toward the base, second-order branches divergent; with 1–2(–3) spikelets per branch; pulvini shortly hairy; pedicels 2–20 mm long. Spikelets solitary, long ovoid, 5–6.4 × 1–1.2 mm, pale, tinged with purple, glabrous, the upper glume and lower lemma subequal, exceeding 3.2–4  mm in length the upper anthecium; lower glume (2–)2.5–3  mm long, almost 1/2 the length of the spikelet, 5–7 nerved, acute to obtuse; upper glume 5–6  mm long, not deciduous at maturity, acute, 10–13 nerved; lower and upper glume separated by a conspicuous internode ca. 0.4 mm long; lower lemma glumiform, acute, 10–12 nerved; lower palea 1.5–2 mm long, as long as the upper anthecium, membranous, the margins scabrous; upper anthecium ovoid, 1.6–2 × 1–1.1 mm, 3.2–4 mm shorter than the upper glume and lower lemma, smooth, pale to brownish at maturity, shiny, crustaceous, with 2 basal scars, brownish at maturity, ca. 0.2 mm long, upper lemma 5–7 nerved, upper palea with verrucose papillae at its apex. Caryopsis 1.5 × 1 mm, hilum oblong, embryo 1/2 to 3/4 the length of the caryopsis. Distribution and ecology (Fig. 29). Southern USA and northern Mexico; grows in open places on sandy soils; from sea level to 1000  m  asl; flowers and fruits between April and December Panicum capillarioides is clearly distinguished from the rest of the species of section Panicum by having long ovoid spikelets, with the upper glume and lower lemma exceeding the upper anthecium by 3.2–4 mm. Additional material examined. USA.  Texas: Frio Co, Dilly, 24 June 1941, Tharp 43066 (MO); San Diego, June 1892, Halley 69 (MO); Jim Wells, “La Copita” Experimental Ranch, 15 October 1981, Hatch 4615 (MO); Kleberg Co, 2 mi. N of Riviera, of Hwy. 77, 15 October 1946, Lundell & Lundell 14683 (MO); Kenedy Co., El Toro Island, Laguna Madre, 7 April 1949, Tharp 49235 (MO); 9  miles  N of Raymondville, Gould 11456 (US); Armstrong, Swallen 1555 (US); Corpus Christi, Hitchcock 5349 (US); El Toro Island, off Laguna Madre, Tharp 49042 (MO, US); 3 miles S of Falfurrias, Swallen 10039 (US); Kingsville, Piper s.n. (US- 558794); W of Kingsville, Swallen 10254 (US); Laredo, Mally s.n. (US-1060824); Norias Division, King Ranch, Swallen 10197 (US); E of Raymondville, Swallen 10102 (US); near San Antonio, Swallen 1780 (US); San Diego, Croft 240 (US); 1 mile W of San Diego, Gould 11000 (US); Yturria Ranch, Lundell & Lundell 8719 (US); near Palacios along beach, Rogers 6609 (W); Sarita, Willacy County, Hitchcock s.n., Amer. Gr. Nat. herb. 32 (W, US) Mexico. Nuevo León: Monterrey, Hitchcock 5547 (USA). San Luis Potosí: 10 km S of Cárdenas, 1,000 m, Rzdewosky 4618 (US); in a tropical forest between Naranjo-Platinito-Salto de Agua, 400–600 m, Sohns 1430 (US). Tamaulipas: Mun.

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San Fernando, Punta de Alambre, 11 October 2007, Zuloaga et  al. 9687 (SI); Hacienda Buenavista, Wooton s.n. (US-1061806); Chamal, Swallen 1642, 1693 (US); without locality, 2 December 1930, Viereck 857 (US) 25. Panicum chaseae Roseng., B.R. Arrill. & Izag., Bol. Fac. Agron. Univ. Montevideo 103: 9, Fig. 2, 1968. Type: Uruguay. Rocha: Ruta 9, Fortaleza Santa Teresa, 3 February 1967, on roadside in sandy-clay soil, B. Rosengurtt 10804 (holotype, MVFA 0000419; isotypes, BAA, K 000309316, MVFA 0000421, MVFA 0000422, P 00740840, SI 2796, US-2946507). Fig. 31 Plants perennial, cespitose, shortly rhizomatous, 30–80  cm tall, culms erect, without aerenchyma, few noded, internodes cylindrical, 3–13 cm long, densely to sparsely hairy, with stiff tuberculate hairs; nodes purplish, hairy, with adpressed whitish hairs. Sheaths 3–12 cm long, usually smaller than the internodes, covered with tuberculate hairs, without aerenchyma. Ligule membranous-ciliate, 0.71.8 mm long; collar light brown, hairy. Blades linear-lanceolate, flat or with margins involute, erect, ascending, rigid, 925 × 0.20.6 cm, narrowed at base, with tuberculate hairs on both surfaces, more abundant on the adaxial surface, apex acuminate, the margins scaberulous, ciliate. Inflorescences contracted, 824 × 15 cm; peduncles long exserted, cylindrical, hispid, 10–31  cm long; first-order branches short, appressed to the rachis, sometimes slightly divergent; main axis glabrous, smooth, wavy, pulvini sparsely hairy to glabrous; first-order branches alternate; pedicels scaberulous, claviform, 1–5 mm long. Spikelets broadly ovoid, 2.8–3.8 × 1.1–1.5 mm, acuminate, glabrous, pale and tinged with purple, adpressed on the branches; upper glume and lower lemma subequal, equal to or exceeding the upper anthecium, up to 0.3 mm; lower glume 1.5–2.4 mm long, 1/2 to 3/4 the length of the spikelet, 5–7 nerved, midrib scaberulous, shortly pubescent on the inner surface; upper glume 2.8–3.5 mm long, not deciduous at maturity, 7–9 nerved, midrib scaberulous or not towards the apex, shortly hairy on the inner surface; lower lemma 2.7–3.5 mm long, glumiform, 9 nerved; lower palea elliptic, 1.8–2.6  ×  0.9–1.2  mm, as long as the upper anthecium, membranous, finely denticulate or not on the upper margins, apex bilobed to entire; upper anthecium ellipsoid, 2–2.6 × 1–1.3 mm, crustaceous, glabrous, smooth, shiny, pale (brown to grayish at maturity), with two small basal scars, 0.15 mm long, brown at maturity; upper lemma 7 nerved, upper palea with simple papillae at the apex. Caryopsis ellipsoid to ovoid, 1.4–1.6 × 0.8–1.1 mm; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 32). Argentina, Brazil, and Uruguay, lives in fields and flowers from October to May Species related to P. ghiesbreghtii and P. peladoense. It differs from the former in having contracted panicles, larger spikelets, and smaller blades. Panicum peladoense differs from P. chaseae by its lax panicles, with spikelets with a black upper anthecium and a deciduous upper glume at spikelet maturity. Panicum diffusum Sw., a species growing in the West Indies, differs from P. chaseae in having lanceolate blades up to 0.2 cm wide, lax panicles and minor spikelets, 2–2.5(−3) × 0.8–1 mm, with a reduced lower palea.

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Fig. 31  Panicum chaseae. (a) Plant. (b) Ligular region. (c) Ligular region, longitudinal section. (d) Spikelet, lateral view. (e) Lower glume, dorsal view. (f) Upper glume, dorsal view. (g) Inferior lemma, dorsal view. (h) Upper anthecium, ventral view. (i) Upper anthecium, dorsal view. (j) Lower palea, ventral view. (k) Upper palea, ventral view. (l) Apex of the upper palea. (m) Caryopsis, embryo view. (n) Caryopsis, hilum view. (o) Caryopsis, lateral view. (From Rosengurtt 10804, MVFA)

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Fig. 32  Distribution of Panicum chaseae, P. diffusum, P. exiguum, P. furvum, P. stramineum, and P. tamaulipense

The specimen Gallinal et al. PE-4473 differs from the others examined by having more lax inflorescences, with branches diverging from the rachis. Additional material examined. Brazil. Rio Grande do Sul: Tupaceretán, January 1935, Araujo 186 (US), Rambo 9875 (US); Belem Velho, Araujo 151 (BAA, US); Guaiba, Estaçao Experimental do UFGRS, Valls 2112 (CTES), Silva s.n. (ex BLA 2295, 2436 en SI), Kappel s.n. (ex BLA 3641 en SI); Pelotas, Araujo 527 (BAA), Barreto & Kappel s.n. (ex BLA 3372 en SI); entre Cruz Alta y Panambí, 3 November 1962, Rosengurtt & Del Puerto 8893 (BAA, SI, US); São Gabriel, Valls & Mundstock 860 (SI); alto da Morro da Gloria, Valls & Arzivenco 2026 (CTES); Porto Alegre, Morro da Gloria, Valls 1646 (CEN); Cachoeira do Sul, Km 190 da BR-290, Valls et al. 2769 (CEN); Fazenda Experimental de Criacào Bagé, Swallen 7467 (US) Argentina. Corrientes: Estancia Rincón del Ambrosio, Schwarz 9595 (LIL, LP). Entre Ríos: Nueva Escocia, Cordini 20 (BAA); Concordia, Burkart 997 (BAA); Concordia, Arroyo Gualeguaycito, 30 January 1927, Burkart 1080 (SI) Uruguay. Rivera: subida de Pena, Sierra de la Aurora, Rosengurtt B-7128 (US). Salto: San Antonio, Orihuela 92 (US), s.n. (US-1648463). Soriano: Estación Juan

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Jackson, Rosengurtt B-242 (US); without locality, Gallinal et al. PE-4473 (BAA). Tacuarembó: camino a Rivera, 32 km de Tacuarembó, 8 February 1981, Cabrera & Zuloaga 32423 (SI, US) 26. Panicum decolorans Kunth, Nov. Gen. Sp. (fourth ed.) 1: 100. 1815[1816]. Type: Mexico. Querétaro: “crescit in temperatis, apricis regni Mexicani prope Querétaro, alt. 995 hexap.”, F. W. H. H. A. von Humboldt & A. J. A. Bonpland s.n. (holotype, P 00128846; isotypes, P 00128847, US-80663, fragment and photo ex P) Plants annual, cespitose, with erect culms (15–)30–90 cm tall, geniculate or not, simple or branching at the lower nodes, few noded, without aerenchyma; internodes 6–12  cm long, cylindrical, hollow, glabrous, hirsute toward the upper portion to glabrescent, nodes shortly hairy. Sheaths 3.5–7 cm long, without aerenchyma, commonly shorter than the internodes, hirsute, with deciduous tuberculate hairs, or glabrous, margins shortly ciliate throughout or only toward the upper portion, occasionally with one margin ciliate and the remaining glabrous. Ligule membranous-ciliate, 0.6–2 mm long; collar glabrous. Blades lanceolate, 4.5–18 × 0.5–1.5 cm, flat, base rounded, apex acute, glabrous to hirsute on the abaxial surface, occasionally hirsute over the entire surface, with tuberculate hairs 1–1.5  mm long, basal margins ciliate with deciduous hairs. Inflorescences terminal, exserted, multiflowered, long-pedunculate, dense, contracted, (5–)10–20 × 0.5–1(−2) cm; peduncles wavy, up to 27 cm long, glabrous to hirsute toward the upper portion; main axis wavy, scaberulous, pulvini glabrous, first-order branches ascending, sometimes the lower one somewhat divergent, alternate, with congested spikelets on third- or fourth-order branches; pedicels claviform, scabrous, triquetrous, 1.5–4  mm long. Spikelets long ovoid, (4.2–)5–6 × 1.8–2 mm, adpressed on the branches, glabrous, pale, with purplish tints; upper glume and lower lemma subequal or upper glume slightly smaller than the lower lemma, 1–2 mm longer than the upper anthecium; stipe absent; lower glume ovate, 1.8–2.7 mm long 1/2 or less the spikelet length, without a distinct internode between the lower and upper glume, acute to obtuse, clasping the upper glume at its base, 5–7 nerved, midrib scaberulous; upper glume (3.8–)5–5.9 mm long, not deciduous at maturity, acute to acuminate, 10–13(–15) nerved, the inner surface scaberulous toward the apex; lower lemma glumiform, 10–13 nerved; lower palea reduced, 1–1.5 mm long, 1/3 or less the length of the upper anthecium, membranous, obtuse, glabrous; lower flower absent; upper anthecium ovoid, 2.7–3.8 × (1.1)–1.5 mm, glabrous, pale to brownish at maturity, smooth, shiny, crustaceous, obtuse, easily deciduous at maturity before glumes are deciduous, the base with a circular ring, brownish at maturity; lemma 7 nerved; palea with verrucose papillae toward the apex. Caryopsis ellipsoid, 2–2.5 × 1–1.5 mm; hilum oblong, embryo 1/2 to 3/4 the length of the caryopsis. Distribution and ecology (Fig. 25). A species endemic to Mexico, it grows in the states of Baja California Sur, Chihuahua, Coahuila, Guanajuato, Hidalgo, Jalisco, Michoacán, Oaxaca, Puebla, Querétaro, and San Luis Potosí and flowers between May and November; it reaches up to 1870 m asl.

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Iconography. Hitchcock & Chase, Contr. U.S. Natl. Herb. 15: Fig. 370. 1910 This species is related to P. parcum, distinguished by the characters of the inflorescence (lax and diffuse in P. parcum, 6–15 cm wide), by having a lower glume 1/2 the length of the spikelet (3/4 the length in the latter species) and by lacking a marked internode between the lower and upper glume (while P. parcum has a stipule ca. 0.6 mm long between both glumes). Additional material examined. Mexico. Chihuahua: 10  km  W of Carretas, 1675 m, 21 August 1939, Harvey 1578 (GH, US). Coahuila: Parras, 12 November 1935, Archer 3400 (US). Guanajuato: Estación El Chico, about 10  km east of Irapuato, Sohns 210 (US); 39 km al NE de León, sobre la carretera a San Felipe, 2500  m, bosque de encino, 13 August 1990, Rzedowski 49884 (ANSM); Inchamacuaro, Mun. Acambaro, 11 August 1985, Rzedowski 38973 (MEXU); 12  km al N de Salvatierra, sobre la carretera a Cortazar, 18 September 1985, Rzedowski 39002 (MEXU). Hidalgo: Mun. Jacala, 4500  ft, mountain roadside, Chase 7090 (ANSM, GH, MO, US), 7270 (GH, US). Jalisco: near Lago Chapala, 12 August 1947, Barkley et al. 7639 (F); swampy ground beside Lake Chapala, near Tuxcueca, 4400  ft, 25 August 1941, Leavenworth & Leavenworth 1839 (F, MO, US); 1 mile E of Tuxpán, on Mex 15, 1870 m, 20 October 1976, Brunken & Perino 434 (MEXU, MO). Michoacán: Chavinda, Hernández Xolocotzi et al. 2793 (US); about 13 miles east of Jiquilpan on road from Guadalajara to Morelia, 22 July 1950, Reeder et al. 1439 (MEXU). Oaxaca: 8 miles W of Huajuapan de León, on road leading to Tehuacán, Puebla, Soderstrom 445 (US); km 568 of Panamerican highway, Baldwin Jr. 14321 (US). Puebla: Tehuacán, 5500 ft, 9 August 1910, Hitchcock 6057 (US). Querétaro: Querétaro, Hitchcock 5822, 5864 (US), Agniel & Arsène 10269 (US), Agniel 10393 (US); on rocky slopes back of motel 5  miles  N of Querétaro, 8 September, Gould 11596 (US); three miles W of Querétaro, Barkley et al. 701 (MEXU, US); vicinity of Querétaro, 1912, Basile 51, 52 (US); de Ciudad de México a Querétaro, km 290, Hernández Xolocotzi et  al. 2304 (US); 6  miles southeast of Queretaro, 2100 m, 12 August 1958, Reeder & Reeder 2975 (MEXU). San Luis Potosí: Cárdenas, Hitchcock 5712 (F, GH, NY, US); Cárdenas, 19 July 1910, Hitchcock s.n., Amer. Gr. Nat. Herb. 29 (MO, US); 2 km E of Santa Catarina along hwy. 70, 1220 m, 23 October 1978, Reeder & Reeder 7042 (MEXU). Without state: Ciudad de los Tres Reyes, 1780 m, 5 September 1984, Labat 1186 (MEXU) 27. Panicum diffusum Sw., Prodr.: 23. 1788. Panicum diffusum var. genuinum Döll, Fl. Bras. 2(2): 199. 1877, nom. inval. Type: Jamaica. “Jamaica, Hispaniola.”, O.  P. Swartz s.n. (lectotype, S04-890, here designated; isolectotypes, BM 000938695, G 00099638, LD 1267065, S04-889, S06-721, S-R-3946, S-R-3947, S-R-3948, US-76920, fragment and photo) Panicum caespititium Lam., Tabl. Encycl. 1: 173. 1791. Type: “Ex Amer. merid. communic D.  Richard,” L.  C. M.  M. Richard s.n. (holotype, P; isotype, USA 00148203, fragment ex P) Panicum debile Schult. Mantissa 2: 243. 1824, nom. illeg. hom., non P. debile Desf., 1798, non Ell., 1816, nec Poir., 1816. Panicum diffusum var. debile (Schult.) Nees, Fl. Bras. Enum. Pl. 2(1): 172. 1829. Type: “Hispaniola” (holotype not known; isotype, US-974075, fragment)

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Perennial plants, cespitose and shortly rhizomatous, with erect to decumbent culms, somewhat geniculate at the base, (6–)15–60 cm tall, simple or branching at the middle nodes, internodes cylindrical, hollow, shortly hairy to glabrous, nodes short to long hairy. Sheaths striate, 2–6 cm long, glabrous, the margins membranous, shortly ciliate toward the upper portion. Ligule membranous-ciliate, reduced, 0.6–1 mm long; collar brown, glabrous. Blades linear-lanceolate, (3–)6–15 × 0.2 cm, the margins involute, flattened toward the middle portion, adaxial surface densely pilose with whitish hairs, abaxial surface glabrous or sparsely pilose, the basal margins with or without caducous tuberculate hairs, otherwise scaberulous, base narrowed, apex subulate. Inflorescences terminal, exserted, lax, 3–16  ×  1.5–8  cm; peduncles 9–15 cm long, cylindrical, glabrous; main axis wavy, scaberulous, pulvini brownish, glabrous, first-order branches alternate, rarely opposite, divergent and spaced, lower branches 3–8 cm long, naked in the lower portion, with spikelets congested toward the upper portion on second-order branches; axes of the branches and pedicels triquetrous, scabrous. Axillary inflorescences present, numerous, similar to the terminal panicle. Spikelets long ovoid, 2–2.9 mm × 0.8–1 mm, glabrous, greenish or tinged with purple, upper glume and lower lemma subequal, 0.6–0.7 mm longer than the upper anthecium; lower glume 1–1.2 mm long, 1/2 or slightly less than spikelet length, ovate, acute, 5–7(–9) nerved, with nerves anastomosed toward the apex; upper glume 2–2.8 mm long, 11–13 nerved, shortly hairy toward the apex on the inner surface; lower lemma 2–2.8  mm long. 9–11 nerved, shortly hairy toward the apex on the inner surface; lower palea ovate, reduced, 1–1.3 × 0.6–0.8 mm, 1/2–3/4 the length of the upper anthecium, hyaline, glabrous; lower flower absent; upper anthecium ellipsoid, 1.5–1.8  ×  0.6–0.8  mm, easily deciduous at maturity, pale, smooth, shiny, crustaceous, with two basal scars 0.12 mm long; upper lemma 7 nerved; upper palea with simple papillae toward the apex. Caryopsis 1.1 × 0.7 mm; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 32). This species grows in the West Indies, in the Bahamas, Cuba, Guadeloupe, Jamaica, Martinique, Puerto Rico, Dominican Republic, and St. Thomas. It reaches up to 1500 m asl and flowers throughout the year. Iconography. Hitchcock & Chase, Contr. U.S. Natl. Herb. 15: Fig. 59. 1910 In describing this species, Swartz cites “Jamaica, Hispaniola,” without citing collector or herbarium where the type material is deposited. Hitchcock and Chase (1910) state that the type is in Swartz’s herbarium, deposited in S. Several specimens from this collection were examined in S, from which specimen S04-890 was selected as lectotype, being a complete specimen and matching the protologue. Panicum diffusum is characterized by being graceful plants with conspicuously branched culms, linear-lanceolate blades, and inflorescences with spikelets adpressed on the branches and spikelets 2–2.9 mm long. Some specimens of this species are glaucous and resemble P. filipes, a North American species, distinguished by panicle type and leaf width and length and spikelet size and shape. The material cited for Brazil by Döll (1877) as P. diffusum Sw. var. subcontractum corresponds to P. exiguum Mez. In this work Döll also cites a specimen from St. Bartholomei, Forsstrom s.n. (F, S), from the West Indies and corresponding to Panicum diffusum. Panicum diffusum was also cited for Brazil, based on Döll’s work, by Beetle (1977) and Gould (1979).

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Gould (1975, 1979), Waller (1976), and Beetle (1987) erroneously indicate that the species grows in Texas, Mexico, and Brazil, without citing herbarium specimens. Additional material examined. Bahamas. New Providence, Harold Road, Britton & Brace 424 (NY, US) Cuba. Camaguey: Camaguey to Santa Yañez, Britton 2356 (NY); Sabana near Camaguey, 4 April 1912, Britton et al. 13177 (NY); Camaguey, savanna N of La Gloria, Shafer 385 (NY, US). Havana: Guanabacoa, Hitchcock 23239 (US); Arroyo Ayola, near Havana, Leon 190 (USA); Arroyo Apolo, León 923 (US); Jamanito, Ekman 1200 (NY); Santiago de las Vegas, 17 October 1904, Van Hermann 355 (NY), 2444 (NY, US), Tracy 9111 (US); near Santiago de Las Vegas, Wilson 1405 (P, US); inter Campo Florido et Loma de Coca, 18 May 1914, Ekman 995 (G, US); near Campo Florido, León 3449 (USA); Monte La Noria, León 2559 (US); Víbora, along railroad, León 2532 (US). Isla de la Juventud: Near Nueva Gerona, 12 May 1904, Curtiss 384 (G, M, P, US), 12 May 1904, Curtiss 494 (M, MO, NY, P, US); Nueva Gerona, Ekman 11732 (NY); near Madruga, Britton et  al. 649 (NY). Matanzas: Sabanilla de La Palma, León 9651 (NY). Oriente: Santiago de Cuba, in pastures N of the town, 26 June 1914, Ekman 1517 (G, NY); along Gibara carretera north of Holguín, Shafer 1437 (US); Colonia San Rafael, Manatí, León & Garteiz 5681 (US); Mir, in the savannas at the station, 22 August 1916, Ekman 7511 (G, US); Galbis, savannas near NE of the station, Ekman 7483 (US); southeast of Holguín, Shafer 1512 (NY, US). Pinar del Río: Bahía Honda, among rocks in hillside, Wilson 9443 (NY, US); San Diego de los Baños, Britton et al. 6859 (NY); near Guanajay, in grassy fields, Palmer & Riley 802 (US). Santa Clara: Sagua, Britton & Wilson 314 (NY); Santa Clara, Britton et al. 6123 (NY). Without locality, Wright 3877, 3852, 3860 (NY, US), 1540 (G, MO, P, US) Jamaica. Black river and vicinity, Britton 1380 (NY); Inverness, Lower Clarendon, 15 October 1915, Harris 12164 (MO, NY, P); Ewarton to Linstead, ditch in cut, Hitchcock 9463 (US); without locality, Macnab 184 (P) Haiti. Vicinity of Bassin Bleu, 630–1500 m, Leonard & Leonard 15090 (NY, US); vicinity of Cabaret, Baie des Moustiques, Leonard & Leonard 11875 (US), 12 January 1899, 11900 (MO, US); Départament Du Nord, vicinity of St. Michel de l’Atalaye, Leonard 7449 (NY, US); vicinity of Jean Rabel, Leonard & Leonard 12590, 13609 13828 (US); vicinity of Mole St. Nicolas, on bare rocks, between bay and north coast of Cap du Môle, Leonard & Leonard 13147 (US). Nicolas, on bare rocks, between bay and north coast of Cap du Môle, Leonard & Leonard 13147 (US); montagnes du Iron d’Eau, between Source Matelas and Source Puantes, Ekman 2151 (US); vicinity of La Vallée, Tortue Island, Leonard & Leonard 11576 (US); vicinity of Port de Paix, Leonard & Leonard 12237 (US); Mirebalais, Cook et al. 78 (US); Aux Cayes, Ekman H20 (US) Dominican Republic. Santo Domingo: Vicinity of Ciudad Trujillo, Allard 15744, 15902 (US); Azua, Valle de San Juan, 20 August 1929, Ekman 13390 (G, NY); Azua, Santo Domingo, Rose et al. 4410 (US). Santiago: Cibao Valley, Hato del Yaque, Ekman 16237 (US). Monte Cristi: Foot of El Morro, near seashore, Jiménez 3620 (US); Montecristi, Guayubín, 100 m, Abbott 1027 (NY, US). Without locality, Bertero s.n. (M). St. Thomas: Northside Bay, September 1880, Eggers 76 (G, M, MO, P, US); Buck Island, Britton & Schaffer 382 (NY, US)

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Puerto Rico. Guanajibo, near Mayagüez, 11 February 1915, Britton & Cowell 4064 (MO, NY); Quebradillas, in pockets in limestone cliff above the sea, 22 November 1913, Chase 6577 (G, MO, NY, P, US); vicinity of Quebradillas, Chase 6567 (US); vicinity of Mayagüez, near Joyuda, Chase 6308 (US) Guadalupe. Grand-terre, Richard s.n. (P); no locality, Bertero s.n. (MO-2095466, P), Duss 676 (P), 3181 (NY, US) Martinique. Without locality, 1869–1870, Hahn 1232 (P, US); Coteaux des Trois Ilets, Duss 536 (US) 28. Panicum ephemeroides Zuloaga & Morrone, Ann. Missouri Bot. Gard. 83(2): 239, Fig. 19. 1996. Panicum ephemerum Zuloaga, Morrone & Valls, Iheringia, Bot. 42: 4, Figs. 1–8, 24 and 25. 1992, nom. illeg. hom., non P. ephemerum Renvoize, 1979. Type: Brazil. Goiás: Mun. Jataí, 51 km a oeste do Rio Claro e 9 km a este do acesso a Perolândia ao longo da rodovia BR-364, 860 m, 3 April 1986, J. F. M. Valls & W. L. Werneck 9867 (holotype, CEN; isotype, SI 002801). Fig. 33 Annual, herbaceous plants, 60–70 cm tall, with erect, few noded culms; internodes cylindrical, hollow, glabrous; nodes glabrous. Sheaths herbaceous, striate and with anastomosed nerves, glabrous. Ligule membranous-ciliate, the membranous portion 0.4 mm long, cilia 1.5–2.2 mm long. Blades linear-lanceolate 20–23 × 0.2 cm, the margins involute, base rounded and apex long attenuate, margins scaberulous. Inflorescences exserted, lax, diffuse, multiflowered, ca. 30  cm long; peduncles 15–25 cm long, cylindrical, glabrous; main axis wavy, scaberulous to smooth and glabrous, with first-order branches diverging from the rachis, the lower ones alternate and the upper ones opposite to alternate, pulvini glabrous and with purplish tints, branch axes scabrous, with spikelets appressed to the secondary branches, solitary, distant and shortly pedicellate on third-order branches; pedicels claviform, scabrous. Axillary inflorescences numerous, similar to the terminal panicle. Spikelets long ellipsoid, 3.5–3.6  ×  1.3  mm, biconvex, glabrous, greenish and tinged with purple, open at maturity, upper glume and lower lemma subequal and with a long attenuate violaceous apex, exceeding 1 mm in length the upper anthecium; lower glume 2.8 mm long, 3/4 the length of the spikelet, clasping, glabrous, apex attenuate, 3(–5) nerved, the midvein scabrous toward the apex; upper glume 5(–7) nerved, the midrib scaberulous; lower lemma 5 nerved, glumiform; lower palea elliptic, 2 × 1 mm, hyaline, the margins scaberulous, otherwise glabrous; upper anthecium ellipsoid, 2.1  ×  1.1  mm, whitish, brown at maturity, crustaceous, shiny, papillose over the apex of upper palea and with a brown disk at the insertion zone with rachilla; lemma 5 nerved; palea with verrucose papillae toward the apex. Caryopsis ellipsoid, 2.4 × 1.6 mm, whitish; hilum oblong, embryo 1/3 or less the length of the caryopsis. Distribution and ecology (Fig. 34). Species known only from the typical collection in the state of Goiás, Brazil. It inhabits fields, reaching up to 860 m asl and flowers in April. It is a related species to P. stramineum; the latter differs by having its spikelets arranged toward the apex of the branches, by its spikelets 2.3–3.2 mm long, lower glume minor, 1/3 to ½ the length of the spikelet and upper glume and lower lemma 9–11(–13) nerved.

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Fig. 33  Panicum ephemeroides. (a) Habit. (b) Inflorescence. (c) Spikelet, lateral view. (d) Spikelet, ventral view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Base of lemma and upper palea. (From Valls & Werneck 9867, SI)

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Fig. 34  Distribution of Panicum bergii var. pilosissimum, P. ephemeroides, P. hispidifolium, P. lepidulum, P. magnispicula, and P. quadriglume

29. Panicum exiguum Mez, Bot. Jahrb. Syst. 56, Beibl. 125: 3. 1921. Type: Brazil. Minas Gerais: Serra de Caldas, 10 March 1876, C. W. H. Mosén 4571 (holotype, B_10_0158004; isotypes, BAA 00000898, fragment ex B, P 00740867, 00740897, S13-12871, US 00148487, fragment ex B). Fig. 35 Panicum diffusum Sw. var. subcontractum Döll, Fl. Bras. 2(2): 199. 1877. Type: Brazil. Minas Gerais: Guarda Mor, J. B. E. Pohl 2435 (lectotype, W, designated by F. O. Zuloaga & O. Morrone, Ann. Missouri Bot. Gard. 83(2): 239. 1996) Plants annual, cespitose, 10–30(−50) cm tall, culms geniculate, decumbent, rooting or not at lower nodes, then erect, densely branching; internodes compressed, hollow, hirsute, with tuberculate hairs to glabrous, 1.5–11 cm long; nodes pale to dark brown, densely hairy. Sheaths 1–4  cm long, usually smaller than the internodes, hirsute, covered with coarse tuberculate hairs, the margins ciliate. Ligule membranous-ciliate, 0.5–1.5 mm long, the membranous portion reduced, then long ciliate; collar pale, pilose. Blades oblong-lanceolate to lanceolate, acuminate, 2–6(−10) × 0.2–0.4(−0.7) cm, flat, rounded to subcordate at base, the margins scaberulous and long ciliate, cartilaginous, the rest of surface hirsute, with tuberculate

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Fig. 35  Panicum exiguum. (a) Habit. (b) Spikelet, ventral view. (c) Spikelet, dorsal view without upper glume, exposing the upper anthecium. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (From Mexía 5622, US)

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hairs to glabrescent. Inflorescences lax, diffuse, pyramidal, 2.5–8(−14) × 2–6(−8) cm, with scattered spikelets on the branches, which are divergent; peduncles subincluded to exserted, up to 20 cm long, cylindrical, hirsute to glabrescent; main axis wavy, hirsute, with long whitish hairs, to scabrous; branches alternate, rarely opposite, hirsute to scabrous, purplish, pedicels scabrous, 2–10 mm long, with or without long claviform hairs. Spikelets long ovoid, acuminate, (2.1–)2.6–3  ×  0.8–1  mm, open at maturity, glabrous, greenish and tinged with purple; upper glume and lemma subequal and exceeding 0.6–1  mm in length the upper anthecium; lower glume (1.5–)1.7–2.4  mm long, 3/4–4/5 the length of the spikelet, acuminate, (3–)5(–7) nerved, the midrib scabrous toward the apex, inner surface scabrius towards the upper portion; upper glume 2.2–2.8 mm long, 7(–9) nerved, midrib scabrous toward apex, acute to acuminate, inner surface smooth to scaberulous toward the apex, caducous at maturity, exposing the upper anthecium; lower and upper glume separated by an internode ca. 0.3 mm long; lower lemma glumiform, 2.2–2.8 mm long, 7(–9) nerved; lower palea (0.7–)1.2–1.6 × 0.5–0.7 mm, ovate, small, membranaceous, glabrous, bilobed or not at apex; lower flower absent; upper anthecium ellipsoid, 1.4–1.8 × 0.6–0.8 mm, smooth, with simple papillae clustered at the apex of palea, pale, black at maturity, deciduous before the glumes and lower lemma, with two basal scars, 0.1 mm long, brown at maturity; upper lemma 5–7 nerved; palea with simple papillae toward the apex. Caryopsis 1.1–1.4 × 0.7–1 mm wide; hilum punctiform, embryo 1/2 the length of the caryopsis. Distribution and ecology (Fig. 32). It is a species with a center of distribution in Brazil, from where it sporadically extends to Colombia, Venezuela, Bolivia, Peru, and Paraguay. It reaches up to 1000 m asl, inhabits sandy or clayey fields, and flowers and fruits between October and May. Local names: “Capim do resfriado” (Brazil, in specimen Macedo 2192); “kapi’í chu’i” (“kapi’í = straw, chu’i = small) (Zuloaga 1994) Species related to P. peladoense Henrard, from which it differs for being an annual plant, with decumbent culms branching at the base, blades with a rounded to subcordate base and spikelets with the lower glume usually 3/4–4/5 of the length of the spikelet. P. exiguum is distinguished from P. hirticaule by the upper glume being deciduous, the upper anthecium blackish, and the lower glume 3/4 to 4/5 the length of the spikelet. The specimen Swallen 4859 has a larger size than is usually found in this species, reaching 90 cm in height, with an inflorescence approximately 20 cm long. Additional material examined. Colombia. Norte de Santander: Los Estoraques, La Playa, 15 January 1974, Balick 133 (COL) Venezuela. Aragua: La Victoria, prope Colonia Tovar, Fendler 2565 (K). Distrito Federal: Yaguara, hierba de 30–40 cm de alto, en sabanas altas de colinas, November 1940, Tamayo 1469 (F, VEN). Lara: Dtto. Palavecino, dirt road between the quebrada “La Mata” and the east side of the Parque Nacional Terepaima, 15 November 1978, Burandt Jr. & Gantaume V0498 (MO). Trujillo: south of Valera, stony flat above río Matalán, 2600 m, 7 March 1940, Chase 12389 (NY, VEN)

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Taxonomic Treatment

Peru. Junín: San Ramón, 900  m, Blair 731 (K); Colonia Perené, 590  m, 22 October 1923, Hitchcock 22115 (F, GH, NY, R, US) Brazil. Ceará: Sobral, Black & Avelino 88C (IAN); Sobral, Rio Acarau, Black & Avelino 86C (IAN). Goiás: Chapada dos Veadeiros, ca. 38 km N of Veadeiros, 1,000 m, 16 March 1969, Irwin et al. 24519 (F, MO, K, NY, P, UB, US); Chapada dos Veadeiros, 4 km al S de Terezinha, 18 March 1973, Anderson 7381 (MO, NY, UB); northwest of Santa Rita do Paranahyba, 500 m, March 1925, Chase 11638, 11641 (US); near Goiania, 1,000 m, 20 March 1930, Chase 11465 (F, NY, RB, US); 20  km south of Goiás, 1000  m, Chase 11482 (US); Niquelândia, Macedo, ca. 18 km N de Niquelândia, campo limpo, 13 April 1992, Filgueiras 2282 (IBGE, SI); Santa Rita do Araguaia, 5 April 1930, Chase 11791 (US). Marãnhao: Carolina to San Antonio de Balsas, 20 March 1934, Swallen 3998 (US), 4106 (K, US); Mun. Carolina, 10–12  km a leste de Carolina na estrada para Balsas, Valls et  al. 8379 (CEN); Grajaú to Porto Franco, red clay outcrops, open ground, 8 March 1934, Swallen 3822 (IAN, K, NY, RB, US); Ilha de Balsas, región between the Rios Balsas and Parnaíba, ca. 4 km of main house of Fazenda “Morros” about 35 km S of Loreto, 350 m, 2 May 1962, Eiten & Eiten 4504 (F, K, NY, SP, US); 100 m NE of main house of Fazenda “Morros”, 35 km S of Loreto, 300 m, 3 April 1962, Eiten & Eiten 3956 (F, M, NY, SP, US). Mato Grosso do Sul: Regiao do Rio Verde, July/ September 1923, Kuhlmann 2530 (RB, SI); Corumbá, Fazenda Acurizal, Nabileque, Pott et  al. 3862, 3915 (CEN); Mun. Corumbá, Estrada do Carandazal perto do Morro do Azecte, Pott et al. 444 (CEN); Campo Grande, sand and clay, 550 m, 7 November 1930, Chase 10846 (F, GH, R, RB, US); Mun. Corumbá, Fazenda Bodoquena, Allem et al. 2233 (CEN); Corumbá, 6 April 1903, Malme 3044 (USA); Miranda, Fazenda Bodoquena, da Silva 124, 137 (MO), 1, 7 (CEN, SP); Tres Lagoas, Chase 10725 (IAN); Corumbá on Rio Paraguay, 3 March 1930, Chase 11119 (MO). Minas Gerais: Corinto, Fazenda do Diamante, 600 m, 15 April 1931, Mexía 5622 (BAA, F, G, GH, IAN, K, M, NY, P, R, US, W); Santa Terezinha, 9 March 1950, Macedo 2192 (K, MO, US), 2194 (MO, US); Serra do Cipó, km 50, 800 m, 28 March 1925, Chase 9300 (F, GH, MO, NY, US); inter Fazenda Jardim et Capivary, Regnell III 1366 (P, US). Pará: Estreito-Marabá, km l, Pinheiro & Carvalho 55, 218, 483, 634 (UB, US), 36, 74, 182, 265, 273, 297, 336, 382, 385 (US); km 2, Pinheiro & Carvalho 362, 530, 561, 512, 581, 587, 596, 603 (US); Transamazonian highway (BR-230), 6  km  W of Estreito, Rio Tocantins, 6°32′S, 47°32′W, 300  m, 28 February 1980, Plowmann et  al. 9269 (MO, SI). Paraíba: Campina Grande, 16 June 1935, Pickel 3799 (IAN, RB, US); Santa Luzia-Agua Fria, Gonçalves s.n. (SI, ex R 90480, US). Pernambuco: Arcoverde, 21 July 1971, Andrade Lima 71-6363 (F). Piauí: Gilbués, Filgueiras & Rodrigues 1332 (CEN). Rio Grande do Norte: Santa Cruz, Swallen 4859 (CEN, RB, SP). Without state and locality, Burchell 6827 (GH, K, US), Pohl 3885 (M) Bolivia. Beni: Estación Biológica de Beni, 40 km al este de San Borja, 14°50′S, 66°40′, 200  m, 15 April 1991, Killeen 2834 (MO). Santa Cruz: Buena Vista, Steinbach 7086bis (GH); Florida, El Vallecito, Lenaz s.n. (LPB); Nuflo de Chavez, San Antonio de Lomerio, 16°45′S, 61°48′W, 31 January 1985, Killeen 823 (F, MO, SI). Tarija: Avilés, 6 km SW of Chocloca, Beck 761 (K, LPB)

Key to Ungrouped Panicum Sections and Species

143

Paraguay. Amambay: Río Apa, region C.  Margarita, February 1912–1913, Hassler 10986 (G, K, MO, NY, P, US). Concepción: sabanas 8–10 km al sur del Arroyo Tagatiya-mi, alrededores del camino, 5 April 1986, Brunner 1780 (G, MO) 30. Panicum furvum Swallen, Contr. U.S.  Natl. Herb. 29(9): 416. 1950; emend. Davidse, Flora Mesoamericana 6: 306. 1994. Type: Guatemala. Huehuetenango: between Nentón and Las Palmas, via Yalisjao, Rincón Chiquite, Chiaquial, Guaxacana, in Sierra de los Cuchumatanes, 30 August 1942, 800–1,200 m asl, J.  A. Steyermark 51627 (holotype, US 00054561; isotypes, F 0046884F, US-1935080) Plants perennial, densely cespitose, with erect, few noded culms 16–20 cm tall; internodes 2–3  cm long, papillose-pilose along the upper portion, then glabrous, pale or tinged with purple; nodes brownish, pilose, with whitish hairs ca. 1.5 mm long. Sheaths 2–3.5 cm long, usually longer than the internodes, glabrous to sparsely hairy toward the basal portion, pale or tinged with purple, the margins membranous. Ligule membranous-ciliate, ca. 0.5 mm long; collar glabrous. Blades linear-lanceolate, 2–12 × 0.2–0.4 cm, erect, ascending, predominantly basal, flat or with involute margins, glabrous to sparsely hairy on the adaxial surface, base attenuate and apex acute, the lower margins long-ciliate. Inflorescences long exserted, few flowered, 3–6 × 0.5–1.3 cm; peduncles exserted, up to 9 cm long, cylindrical, slender, glabrous to sparsely hairy; main axis glabrous to sparsely hairy next to the base; pulvini glabrous; first-order branches ascending, slightly divergent, glabrous, smooth to finely scaberulous, pale and tinged with purple, the lower ones up to 3 cm long; pedicels claviform, up to 7  mm long, adpressed, glabrous. Spikelets ellipsoid, 2.4–2.7 × 0.8 mm, glabrous, pale, apex long acute, apex of the bracts violaceous; upper glume and lower lemma subequal, 0.5–0.8 mm longer than the upper anthecium; lower glume ovate-acuminate, 1.4–1.6  mm long, 1/2 or slightly more than spikelet length, 5 nerved, with scaberulous nerves toward the apex; upper glume 2.4–2.5 mm long, 7 nerved, deciduous at maturity, exhibiting the adaxial surface of the upper anthecium, the inner surface scaberulous toward the apex; lower and upper glume separated by a short rachilla internode 0.2  mm long; lower lemma glumiform, 2.4–2.6 mm long, 9 nerved, the inner surface scaberulous toward the apex; lower palea reduced, lanceolate, 1.2–1.4 × 0.5 mm, 3/4 the length of the upper anthecium, hyaline, glabrous, the apex bilobed, margins glabrous to denticulate; lower flower absent; upper anthecium ellipsoid, 1.6 × 0.7 mm, crustaceous, smooth, shiny, glabrous, blackish at maturity, with two small inconspicuous scars ca 0.1 mm long at base; lemma 7 nerved; palea with simple papillae toward the apex. Caryopsis not seen. Distribution and ecology (Fig. 32). Species known only from the type locality in Guatemala, growing on mountain slopes between 800 and 1200 m asl Swallen (1950) notes, when describing the species, that the spikelet is 1.5– 1.6 mm long. However, the analysis of typical material confirmed Davidse’s (1994) statement that the length of the spikelets varies between 2.4 and 2.7 mm in length.

144

 

Taxonomic Treatment

Panicum furvum is related, within the group of perennial species with a blackish upper anthecium and a deciduous upper glume, to P. peladoense, a South American species, the latter distinguished by plants 20–60  cm tall, with panicles 7–18 × 3–13 cm, bearing spikelets (2.5–)2.7–3.3(−3.7) mm long. Davidse (1994) indicates the relationship of P. furvum with South American species of the section and mentions that, although its presence in Guatemala is peculiar, there are no indications on the collection label of the type specimen that it is an introduced species. 31. Panicum ghiesbreghtii E. Fourn., Mexic. Pl. 2: 29. 1886; E. Fourn. ex Hemsl., Biol. Cent.-Amer., Bot. 3: 489. 1885, nom. nud. Type: Mexico. “Absque loco,” A. B. Ghiesbreght s.n. (lectotype, P 00740948, designated by A. S. Hitchcock & A. Chase, Contr. U.S. Natl. Herb. 15: 76. 1910; isolectotype, US 00148520, fragment, envelope at lower right of sheet and photo ex P) Panicum hirtivaginum Hitchc., Contr. U.S.  Natl. Herb. 12(6): 223. 1909. Type: Cuba.Without locality, 1865, C. Wright 758 (holotype, US 00037310; isotypes, K 000309160, MO-321053, MO 321055, NY 00071080, NY 00071081, NY 00071082, NY 00803744, NY 00803748, P 00740947, S04-318, US 00147883, US 00406120). Plants perennial, cespitose, shortly rhizomatous, 40–90(−120) cm tall, not glaucous, with intravaginal innovations; culms simple, erect; internodes cylindrical, 6–20  cm long, 2–3  mm in diameter, hollow, papillose-pilose, few noded; nodes densely hairy, with whitish hairs. Sheaths 5–15  cm long, lower sheaths usually smaller than the internodes, without aerenchyma, hirsute, with dense, urticant tuberculate hairs. Ligule membranous-ciliate, (0.5–)1.5–2  mm long, with long hairs beneath at the base of the blade; collar densely pilose. Blades linear-lanceolate, 16–30(−55) × (0.5–)0.8–1.4 cm, flat, erect, ascending, apex long attenuate and base rounded, the adaxial surface densely pilose, with sericeous hairs, abaxial surface hirsute, with tuberculate hairs, the lower margins ciliate, midrib manifest. Inflorescence a lax, broad, multiflowered terminal panicle, shortly exserted to partially included, 7–35 × 5–23 cm, not deciduous overall at maturity; peduncles subincluded in the leaf sheaths to exserted, to 40 cm long, glabrous to hirsute; main axis wavy, scabrous, pulvini glabrous, lower branches alternate, the subsequent ones whorled to opposite or alternate, first-order branches naked in the lower portion, spikelets paired on third-order branches toward the top; pedicels claviform, scabrous; axillary inflorescences present, similar to the terminal panicle. Spikelets long ovoid, 2.6–3.1 × 0.9–1.2 mm, glabrous, greenish, tinged with purple, open at maturity; lower glume 1.4–1.7 mm long, 1/2 or slightly less the spikelet length, acute, 5–7 nerved, midrib scaberulous toward the upper portion, the nerves anastomosed toward the apex; upper glume 2.5–3 mm long, not drooping at maturity, acute, 9–13 nerved, midrib scaberulous toward the upper portion, shortly hairy toward the apex on the inner side; lower lemma 2.5–3 mm long, 9–11 nerved, midrib scaberulous toward the upper portion, acute, shortly hairy toward the apex on the inner surface; lower palea reduced, 0.5–1.3  ×  0.5–0.8  mm, hyaline, 1/3–1/2 the length of the upper anthecium, ovate, obtuse; lower flower absent; upper anthecium ovoid,

Key to Ungrouped Panicum Sections and Species

145

1.7–2.3 × 0.8–1.1 mm, 3/4 the length of the upper glume and lower lemma, pale, smooth, shiny, glabrous, crustaceous, the apex obtuse, with two basal scars, 0.12 mm long, brown at maturity; lemma 7 nerved; palea with simple papillae and globose microhairs toward the apex. Caryopsis ellipsoid, 1.2–1.3  ×  0.7–0.8  mm, hilum oblong, embryo 1/2 the length of the caryopsis. Distribution and ecology (Fig. 25). It inhabits from the USA, the West Indies and Central America to northern South America, in Colombia and Venezuela; it grows in fields. It reaches up to 2400 m asl. Iconography. Hitchcock & Chase, Contr. U.S. Natl. Herb. 15: Fig. 65. 1910 Species is related to P. hirticaule, from which it is separated because the latter species includes annual plants, with culms usually decumbent, smaller leaves, patent, thin and with a subcordate base. Hitchcock (1927) cites this species for Bolivia with Steinbach 6979, which corresponds to P. quadriglume. This species was erroneously cited for southern South America by Hitchcock (1936), Swallen (1943), Lindeman and Stoffers (1963), Luces (1963), Palacios (1969), Adams (1972), Smith et al. (1982), and Longhi Wagner and Boldrini (1988); the material cited by these authors corresponds to P. chaseae. Additional material examined. USA.  Texas: Brownsville, 13 October 1941, Silveus 7293 (US), 19 April 1906, Tracy 9018 (US), Runyon 1551 (US); along Boca Chica road east of Brownsville, 13 October 1941, Silveus 7302 (US) Mexico. Campeche: 3 km al S de Bolonchen de Rejón, cerca de las Grutas de Xtacumbilxunan, por la carretera vía ruinas a Campeche, 25 July 1986, Cabrera 11739 (MO); Campo Experimental El Tormento, km 5 carretera Escárcega a Candelaria, 24 December 1965, Chavelas & Quero ES-109 (MEXU). Chiapas: ca 22 km SE of Teopisca along Highway 190 to Comitán, pine-oak forest, 2040 m, 11 August 1975, Davidse & Davidse 9492 (MO); grassy flats 3 km south of Comitán near old airport, 1525 m, 6 November 1981, Breedlove & Davidse 54908 (MEXU, MO); seasonal evergreen forest near new Tuxtla Airport on ridge north and above Ocozocoautla, 1000  m, 12 November 1981, Breedlove & Davidse 55442 (MO). México: Peñón, 1700 m, 8 October 1933, Hinton 4413 (MO, US); Temascaltepec, near the river, 24 August 1932, Hinton 1424 (US), 27 August 1932, Hinton 1428 (MEXU). Michoacán: about 12  miles west of Quiroga in open pine-oak forest, 7000–9000  ft, Sohns 791, 797 (US). Morelos: Oaxtepec, Matuda s.n. (MEXU-288916). Oaxaca: Oaxaca, Hitchcock 6143 (US). Puebla: Roadside along route 130, 2.7 miles south of the Puebla-Veracruz state line, 20 May 1979, Harriman & Jansen 15627 (CHAPA). Quintana Roo: Coba, in savanna, north of Lake Coba, June 1938, Lundell & Lundell 7791 (F, MEXU, MO, US). San Luis Potosí: Low ground along Río Tampaón, Chase 7480 (GH, US); without locality, 1891, Virlet 1306 (SI). Sinaloa: Rancho de la Nacha, 25 millas W de Angostura, Vera Santos 1832 (NY). Tabasco: Villahermosa, Archer 3968 (US); Fracc. Los Reyes calle Las Palmas 1 km al W de la carretera Cárdenas-Huimanguillo, 4 August 1981, Suárez 4 (MO). Tamaulipas: Chamal, Swallen 1717 (US); along route 85, ca. 4-5  miles south of Ciudad Mante, 18 February 1961, King 3791 (F, US); 10 km al NW de la

146

 

Taxonomic Treatment

desviación a Gómez Farías, 24 June 1983, Torres & Hernández 3097 (MO). Veracruz: Río Blanco, near Orizaba, 25 August 1910, Hitchcock s.n., Amer. Gr. Hb. 36 (MO, P, US); Orizaba, Hitchcock 6363, 6391 (US); environs de Veracruz, Gouin 21 (P); Córdoba, Hitchcock 6418 (US); 1 to 4 km northeast of Campo Experimental de Hule, El Palmar, Zongolica, Vera Santos 2256 (LIL, US); transecto Punta Limón a Cerro Monte de Oro, 20 June 1972, Lot et al. 1948 (MEXU); about 5 miles from El Palmar Rubber Exp. Station, growing at edge of a sugar cane field, 95 m, 28 July 1953, Reeder & Reeder 1936 (MEXU); Rancho Alegre, 7 November 1970, Chiang 191 (MEXU). Yucatán: Chichen Itzá, along roadside, Lundell & Lundell 7427 (US), Steere 1632 (US), Swalllen 2381 (US), 2404 (MEXU, MO); Uxmal, along trail near ruins, Swallen 2621 (MEXU, US); Izamal, Gaumer 2477 (F, MEXU, US); near Uxmal, 23 January 1971, Beetle M-917 (US) Belize. Corozal: between Sarteneja and Chunox, 18°17′N, 88°15′W, 10 m, 18 March 1987, Davidse & Brant 32638 (MO, SI) Honduras. Francisco Morazán: El Zamorano, 800 m, 2 July 1947, Molina 242 (F, GH, MEXU, MO), Rodríguez 694 (F), 3056 (F, US), 3220 (F, US), 27191 (F, US); vicinity of El Zamorano, Swallen 11147 (MO, US), 11150 (F, MEXU, MO, US), 11263 (US), 17 February 1947, Molina 3905 (F) El Salvador. Acajutla, near sea level, 29 November 1911, Hitchcock 8993 (SI, US) Nicaragua. Boaco: near Río Tecolostote, 7.5 km S of Hwy. 7 at El Papayal, ca. 12°11′N, 85°40′W, 60 m, 27 June 1982, Stevens 21647 (MO). Carazo: Jinotepe, 500 m, Hitchcock 8687 (US). Chinandega: Corinto, near sea level, 10 November 1911, Hitchcock 8755 (SI, US). Esteli: km 163 on Hwy. 1, ca. 11.2 km N of entrance to Estelí, ca. 13°13′N, 86°23′W, 920  m, 19 May 1981, Stevens 20198 (MEXU, MO); 16  km  N de Esteli, 900  m, 6 July 1970, Pohl & Davidse 12210 (F, MO). Managua: along new road from Hwy. 1 (N of San Jacinto entrance) to San Francisco del Carnicero, ca. 4.8  km  W of hwy. 1, 16 August 1978, Stevens 10006 (MO). Matagalpa: Matagalpa, 23 July 1969, Zelaya 2307 (MO). Rivas: San Juan del Sur, near sea level, 28 October 1911, Hitchcock 8601 (US) Costa Rica. Guanacaste: along road W of the Cañas cemetery, 70 m, 24 June 1968, Pohl & Davidse 10556 (F, MO, US). San José: San José, 1,100 m, Hitchcock 8452 (US), Jiménez 530 (US); Hacienda Taboga, Jiménez 730 (US); without locality, Pittier 2633 (US); Llano Grande de Puriscal, February 1913, Jiménez 880 (US) Panama. Ancon, 12 October 1917, Killip 4100 (US); in the vicinity of Balboa, Canal Zone, near swamp, 9 April 1911, Hitchcock 7997 (SI, US), 8014 (US); Sabanas near Chepo, 20 January 1935, Hunter & Allen 63 (MO, US); Taboga island, Gulf of Panama, 12 September 1911, Hitchcock 8065 (SI, US), 8094 (US) Bahamas. Crooked Island, Sandrail Point, 9 January 1906, Brace 4812 (NY, US) Cuba. Habana: Calvario, Finca Santa Teresa, 20 July 1911, León 2674 (P, US); cercanías de Madruga, Britton et al. 745 (NY); bancos de la bahía de Habana, León 9314 (NY). Holguín: valle del Río Matamoros, cerca de Holguín al sur, Suazer 1361 (NY). Isla de la Juventud: Santa Bárbara, Westport, Ekman 12036 (G). Matanzas: cercanías de Coliseo, León 8834 (NY). Pinar del Río: Herradura, Tracy 9116 (NY); vicinity of Herradura, Britton et al. 6349 (NY); Bahía Honda, between

Key to Ungrouped Panicum Sections and Species

147

Gerardo and Corojalito, on the railroad track, 28 December 1920, Ekman 12686 (US); weed in field on hillside, first range of hills NW of Pinar del Río, 26 November 1926, Hitchcock 23292 (US); Arroyo de las Manacas, north of Viñales,4 January 1932, León 15339 (US). Santiago de Cuba: Bayate, Ekman 2011 (G), 9667 (NY); Guaro, 20 km from Preston, 13 December 1926, Hitchcock 23397 (US); Santiago de Cuba, 1 August 1913, León 3747 (US); Santa María de Loreto, 1 October 1959, Lopez Figueiras & Acuña 341 (US); El Cristo, 200 m, scattered in savannas, Ekman 1442 (US); valley of Río Matamoros, south of Holguín, Shafer 1361 (NY). Villa Clara: Placetas del Sur, León 6429 (US); sabana de Amaro Rodrigo, León 9138 (NY) Haiti. Vicinity of Pilate, 325  m, 10 February 1926, Leonard 9607 (NY, US); vicinity of Plaisance, Departement du Nord, 400 m, 28 January 1926, Leonard 9417 (US); vicinity of Marmelade, Départament du Nord, 800  m, 20 December 1925, Leonard 8344 (US); vicinity of St. Michel de l’Atalaye, Département du Nord, 350  m, 5 December 1925, Leonard 7745 (US); vicinity of Pétion-Ville, 15 June 1920, Leonard 4901 (US); Port-au-Prince, Mont-Fleury, 23 September 1924, Ekman 1987 (US); Coffee plantation, mountain south of town, vicinity of Jean Rabel, Leonard & Leonard 13744 (US); vicinity of St. Louis du Nord, Leonard & Leonard 14338 (US); vicinity of Port au Prince, 19 May 1929, Leonard & Leonard 15754 (US); Marmelade, Camp N° 4, Nash & Taylor 1297 (NY) Jamaica. Inverness, Lower Clarendon, 22 November 1913, Harris 11691 (MO, NY, P, US), 15 October 1915, 12161 (MO, NY, P, US) Puerto Rico. Vicinity of Coamo Springs, 17 November 1913, Chase 6547 (NY); without locality, Ledru s.n. (G); Jabucoa, 8 September 1886, Sintenis 4983 (US); vicinity of Cayey, road to Guayama, 3 December 1913, Chase 6744 (US); Mayaguez, Hess 1459 (NY) Dominican Republic. Distrito Nacional: vicinity of Ciudad Trujillo, 18 December 1945, Allard 14148 (US). Santiago: Valle del Cibao, Hato del Yaque, 9 November 1930, Ekman 16129 (US) Antigua. Without locality, 14 October 1919, Hitchcock 16382 (USA), Wullschaegel 621, 622 (M); Scott’s Hill Expt. Station, 8 September 1931, Box 49 (US); NE foot of Greencastle Hill, Box 18 (US); Long Island, Box 39 (US) St. Thomas. Virgin Islands, sin colector s.n. (P) Virgin Islands. Tortola, Fishlock 206 (NY) Guadalupe. Without locality, 9 December 1939, Questel 1439 (P, US), Duss s.n. (NY, US), 1892, 3184 (MO, NY, US), 12 (P), Basse Terre, Duss 668 (P) Colombia. Huila: El Gigante, above Tolima, 500–1200 m, Lehmann 8745 (NY, US). Magdalena: Santa Marta, 1898-1901, Smith 165 (F, G, GH, K, MO, NY, P, US). Santander: Santander, Zapatoca, 22 July 1944, Fassett 25492 (COL, US). Tolima: Espinal, Lindig 102 (P) Venezuela. Aragua: en terrenos salinos de la Est. Exp. del Servicio Shell para el Agricultor, Tamayo 3949 (VEN). Distrito Federal: Ingomar, Turmerito, Luces 17 (VEN), 7 April 1940, Chase 12607 (US, VEN); entre Caracas y La Guaira, sabanas en faldas cerca de Pedro García, 23 June 1946, Burkart 17012 (SI, VEN); Hacienda Sosa, Badillo 7, 690 (VEN); La Florida, near Caracas, 12 March 1938, Alston 5298

148

 

Taxonomic Treatment

(F, NY, P, US); Los Chorros, near Caracas, 27 November 1938, Alston 5235 (F, NY). Falcón: Sierra de San Luis, arriba de Santa Marchía, 1,100 m, a lo largo de la carretera, Wingfield 5913 (MO). Guárico: carretera de Santa María de Ipire a El Socorro, 15 km antes de la última población, en desvío a Zaraza, 150 m, Zuloaga et al 4506 (SI, VEN). Lara: cerca de Sicarigua, Burkart 16870 (SI, VEN). Mérida: Brecenio Hacienda, 23 miles SE of Mérida, 6 February 1931, Reed 517 (NY, US, VEN). Miranda: Chacao, Williams 12152 (F); Los Teques, Badillo 146 (VEN). Portuguesa: Dtto. Araure, en terrenos de la fabrica de cal Guaicaipuro, Araure, Rámirez Reyes 2048 (PORT); Dtto. Guanare, lateral de vía a sector 3 de Papelón, 0–3 km de la via Guanare-Papelón, Rámirez Reyes 2428 (PORT); a 4 km al W de Guanare hacia Ciudad Barinas, 09°03′N, 69°48′W, 200 m, 4 August 1989, Zuloaga & Ortíz 4300, 4301 (MO, SI, VEN). Yaracuy: Bureche, antes de Barquisimento, 12 April 1946, Burkart 16508 (SI, VEN) Ecuador. Chacao, Miranda, 28 August 1939, Eggers 14949 (F). Manabí: El Recreo, Eggers 15419 (F, K, M, NY, P, US) 32. Panicum hallii Vasey Plants perennial, cespitose, shortly rhizomatous, with erect culms (10–)20–75 cm tall, glaucous, simple or branching at the base, few noded, without aerenchyma; internodes 1–11 cm long, cylindrical, glabrous, pale or with purplish tints; nodes pilose, covered with short whitish hairs, or glabrous. Sheaths 3–11 cm long, without aerenchyma, glabrous to hirsute, with deciduous tuberculate hairs, not urticating, margins membranous, glabrous to ciliate near the ligular region. Ligule membranous-ciliate, 0.6–1.2  mm long; collar glabrous. Blades linear-lanceolate, 4–23 × (0.1–)0.2–0.5 cm, erect, ascending, basal ones enrolled, flat or with involute margins, glaucous, glabrous to sparsely hirsute, base rounded to narrowed, imperceptibly continuing with the sheath, the apex acute, with cartilaginous margins, scaberulous, the lower ones ciliate, with papillose, whitish hairs, up to 3.5 mm long. Inflorescences terminal, exserted, lax, multiflowered, 9–31 × 3–15 cm; peduncles up to 40 cm long, cylindrical, glabrous, green or tinged with purple; main axis triquetrous, glabrous, pulvini glabrous; first-order branches alternate, divergent, glabrous, smooth; second-order branches divergent or appressed, with distant to approximate spikelets; pedicels 1–15  mm long, triquetrous, claviform, glabrous, scabrid. Spikelets long ovoid, acuminate, 2.1–4 × 0.8–1 mm, glabrous, greenish or tinged with purple; lower glume ovate, 1.2–2.4  mm long, 1/2–3/4 as long as the spikelet, apex acute to acuminate, 3–5 nerved, midrib scaberulous toward the upper portion; upper glume 2–3.5  mm long, not deciduous, apex acuminate, 7–9(–11) nerved, scaberulous on the inner surface toward the upper portion; lower and upper glume separated by a short, 0.3  mm long internode; lower lemma glumiform, 1.8–3.7 mm long, 9–11 nerved; lower palea reduced, 0.8–1.5 × 0.3–0.4 mm, 1/2–3/4 the length of the upper anthecium, membranous, hyaline, glabrous, with denticulate margins; lower flower absent; upper anthecium ovoid to ellipsoid, 1.5–2.4 × 0.7–1.2 mm, 0.8–1.5 mm shorter than the upper glume and lower lemma, crustaceous, smooth, pale, brownish at maturity, with 2 basal scars 0.2 mm long,

Key to Ungrouped Panicum Sections and Species

149

brownish at maturity; lemma 7 nerved; palea with simple papillae at the apex. Caryopsis ovoid, 1.2–1.7 × 0.6–1 mm, whitish; hilum punctiform; embryo 1/2 the length of the caryopsis. Key to the Varieties 1. Spikelets 2.1–2.8 mm long; spikelets diffuse on inflorescence branches............... .................................................32a. P. hallii subsp. filipes 1. Spikelets 2.7–4  mm long; spikelets appressed on inflorescence branc hes................................................................. 32b. P. hallii subsp. hallii 32a. Panicum hallii subsp. filipes (Scribn.) Freckmann & Lelong, Sida 20(1): 172. 2002. Panicum filipes Scribn., Contr. Herb. Franklin Marshall Coll. 1: 13. 1895. Panicum hallii var. filipes (Scribn.) F.R.  Waller, S.  W. Naturalist 19(1): 105. 1974. Type: USA. Texas: Nueces Co, growing in rich shaded ground in the upper part of the “Arroyo” at Corpus Christi, 30 ft, 31 May 1894, A. A. Heller 1809 (holotype, US-2463136; isotypes, NY 00381597, PH 00018640, US 04295490, fragment and photo) It differs from the typical subspecies in having diffuse panicles, with spikelets distant on the branches and not appressed; blades generally larger, 11–23 cm long and spikelets 2.1–2.8 mm long. Distribution and ecology (Fig. 36). The USA and Mexico. It is frequent in fields and roadsides. It flowers between March and December. Iconography. Hitchcock & Chase, Contr. U.S. Natl. Herb. 15: Fig. 60. 1910 Correll and Johnston (1970) cite P. filipes as a related species to P. hallii, from which it is distinguished by spikelet size (2–3 mm, wider panicle and larger blades in P. filipes and 3–3.7 mm spikelets in P. hallii). They also relate the species to P. pil­ comayense (= P. bergii, this one apparently introduced in Texas). There are, within the material considered of this subspecies, specimens, such as Beetle et al. 1031, Waller 2008, Hitchcock 5500, Harvey 950, that have intermediate characteristics with the typical subspecies, such as having adpressed spikelets on the ramifications, being the spikelet measurements coincident with the range of variation existing in subsp. filipes. Additional material examined. USA.  Texas: Edinburg, 18/19 June 1928, Swallen 1079 (US); Kerrville, 22 June 1910, Hitchcock 5318 (US); Robstown, 27 July 1910, Hitchcock 5395 (US); Comanche Spring, September 1849, Lindheimer 1266 (US); Denton Co, Hwy. 24 (West) within Denton City limits, 5 October 1962, Gould 10292 (US); Archer Co, 8 mi. S of Archer, 13 October 1961, Gould 9763 (US); San Patricio Co, near entrance to Lake Corpus Christi State Park, 2 October 1964, Gould 10972, 10973 (US); Frio Co, 4 mi. W of Moore on Hwy. 76, 23 May 1965, Gould 11271 (US); Kleberg Co, 28 April 1949, Swallen 10242 (USA); Rockport, 30 April 1949, Swallen 10290 (US); 2 mi. E of Eagle Pass, 31 May 1931, Swallen 1742, 1743 (US); San Antonio, 18/14 June 1910, Hitchcock 5143 (US); Corpus Christi, Hitchcock 5385 (US); Corpus Christi, 27 June 1910, Hitchcock s.n., Amer. Gr. Herb. 33 (MO, US); Brownsville, 7 August 1904, Hitchcock 220 (US);

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Fig. 36  Distribution of Panicum hallii subsp. filipes, P. hallii subsp. hallii and P. parcum

Kingsville, September 1906, Piper s.n. (US 558797); along Ennis Joslin Road by golf course, 0.9 mi. N of Padre Island Parkway, Corpus Christi, 22 July 1969, Waller 2017 (MO); Koppe Bridge Road, 100 yards E of old bridge site, Koppe Bridge on Brazos River, ca. 12 mi. SW of College Station, 7 November 1971, Waller 2419 (MO); Bexar Co, U.S. Hwy. 90, 4 mi. W of Loop 410, San Antonio, 7 June 1969, Waller 2008 (MO); Nueces Co., Ennis Joslin Road, 0.9 mi. N of Padre Island Parkway, near golf course, SE side of Corpus Christi, 11 October 1969, Waller 2193 (MO); Brazos Co, Koppe Bridge Road, E old bridge site, Waller 2413 (MO); Kendall Co, U.S 87, 17 August 1974, Waller 2978a (MO); Tarrant Co, Ft. Worth, 19 Jun, Ft. Worth, 24 June 1918, Palmer 14214 (MO); Nueces Co, U.S.  Naval Air Base, Corpus Christi, 12 June 1964, Gould 10956 (MO, US); Laredo, 1910, Olmito, 15 April 1905, Tracy 8908 (US); Gillespie Co, about 5 mi. S of Enchanted Rock, 29 June 1957, Correll & Johnston 17274 (MO); Grand Saline, 18 October 1900, Reverchon 2230 (MO); vicinity of Laredo, 22 October 1913, Rose 18052 (MO); Brazos Co, 2 mi. SW of College Station, 17 September 1957, Gould 7685 (MO); April 1931, Swallen Cameron Co, near Port Brownsville, along roadside, 9 March 1942, Lundell & Lundell 10657 (US); Brownsville, Boca Chica, low sandy ground, 10 April 1931, Swallen 1506 (US)

Key to Ungrouped Panicum Sections and Species

151

Mexico. Coahuila: 24 km E of Don Martín Dam, 335 m, 20 May 1939, Harvey 950 (US); El Berrendo, near Múzquiz, Harvey 1186 (MO, US). Nuevo León: 12 mi. S of China on the road Foward Méndez, 7 December 1960, Crutchfield & Johnston 6068 (US); Pan-American Highway from Laredo to Monterrey, near Mamulique Pass, 1500  ft, Leavenworth & Leavenworth 742 (F, MO). San Luis Potosí: El Banito, Chase 7558 (GH). Tamaulipas: El Canelo Ranch, 24 mi N of San Fernando on the Matamoros highway, 7 December 1959, Johnston 4870 (MEXU); Chamal, Swallen 1716 (US); Hacienda Buena Vista, Wooton s.n. (US 1061807); Laredo, 28 August 1971, Beetle et al. 1031 (MO) 32b. Panicum hallii Vasey subsp. hallii, Bull. Torrey Bot. Club. 11(6): 61. 1884. Type: USA. Texas: Austin, 18 May 1872, E. Hall 816 (holotype, US 00147868; isotypes, GH 00057370, K 000674450, NY 00381619) Panicum virletii E. Fourn., Mexic. Pl. 2: 29. 1886; E. Fourn. ex Hemsl., Biol. Centr.Amer., Bot. 3: 498. 1885, nom. nud. Type: Mexico. San Luis Potosí, M. Virlet 1305 (lectotype, P, designated by F. O. Zuloaga & O. Morrone, Ann. Missouri Bot. Gard. 83(2): 246. 1996; isotype, US 00323772, fragment ex P) Distribution and ecology (Fig. 36). Widely distributed in the southern USA and Mexico; grows in arid fields on rocky soils, reaching up to 1920  m  asl; flowers between March and October. Iconography. Hitchcock & Chase, Contr. U.S. Natl. Herb. 15: Fig. 62. 1910 Local name: “Sacate colorado” (in the specimen Wooton s.n.) Additional material examined. USA. Arizona: Santa Cruz Co, Patagonia, 21 September 1908, Hitchcock 3706 (US); Santa Rita Mts, Empire Ranch, 20 October 1902, Griffiths & Thornber 238 (US); 42 mi. N of Tucson, 4500 ft, 15 August 1915, Hitchcock 13255 (US); Gila Co, near road crossing of the White River about 5 mi. above junction with the Black River, 29 August 1948, Gould & Robinson 4937 (US); Pima Co, Silverbell Mts, 30 March 1946, Darrow & Haskell 3419 (US); Gila, 3500  ft, 3 September 1959, Pase 988 (US); Tombstone, 26 September 1938, Goodding 4030 (US); Rozemont, Santa Rita Mountains, 2 September 1905, Thornber s.n (MO- 2747782); Paradise, 14 September 1907, Blumer 1683 (MO, US, W); Apache Indian Reservation, Thornber s.n. (NY); P. Pearce, Thornber n.n. (NY); Arizona-Gila, 3500 ft, 8 October 1958, Pase 1048 (SI); Cochise Co, 9 mi. S of Fry, 5000 ft, 8 September 1944, Gould et al. 2409 (SI). Colorado: 25 mi. S of Pritchett, SW Baca Co, 4000 ft, 11 July 1947, Harrington 3361 (US); Las Animas Co, Carrizo Mt, SW of Utleyville, 29 June 1948, Rogers 6119 (US); Las Animas Co, S slope of Tecolote, 2 July 1947, Rogers 4491 (US). New Mexico: Doña Ana Co, near Tortugas Mts, 1270–1400 m, 30 June 1911, Standley 6441 (US); Cimarron Canyon, 21/24 August 1903, Griffiths 5004 (US); Sierra Co, 1 mi S of Hillsboro, 8 July 1937. Mann 60 (US); Guadalupe Mts, Ranger Station, 6000–7000  ft, 3/6 September 1915, Hitchcock 13517, 13539 (US); Carlsbad, Hitchcock 13445, 13490 (NY, US), Tracy 8200 (US); foothills of Organ Mountains, Barneby 2455 (NY); 20 miles S of Roswell, August 1900, Earle 302 (NY, US); Dona Ana Co, lake E of Doña Ana Mts., 6 September 1908, Standley 3983 (US); Grant. Co, Mangas Canyon, 6500  ft, 1896, Smith s.n. (US 953260); Las Cruces, 1904, Griffiths 7808 (US).

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Taxonomic Treatment

Oklahoma: Murray Co, 26 October 1938, Penn s.n. (US-1724299); Cimarron Co, 3 mi. N of Kenton, 9 July 1947, Rogers 4756 (US). Texas: Young Co., 13 mi. SE of Graham, 15 August 1965, Gould 11075a (US); Kinble Co., 5 mi. NE of junction on Mason Hwy., 26 April 1964, Gould 10929 (US); Maverik Co., about 1 mi. NE of Eagle Pass, near Hwy. junction, 24 May 1965, Gould 11302 (US); Amarillo, 12 August 1948, Hitchcock 16219 (US); Franklin Mts., vicinity El Paso, 22/25 May 1912, Chase 5900 (US); Big Spring, Howard County, Hitchcock s.n., Amer. Gr. Hb. 504 (MO, P); Kerrville, Hitchcock 5293 (US); Big Spring, 25/30 August 1915, Hitchcock 13411 (USA); Del Rio, Plank 57 (US); Laredo, 27 June 1910, Hartley Co, Channing, 19 June 1918, Palmer 14147 (MO); Alpine, 13 September 1915, Hitchcock 13597 (US); USA 377 at crossing on South LLano River, ca. 7 mi. SW of Kimble-Edwards Co. line, 7 June 1969, Waller 1994 (MO); Williamson Co., roadside S.H. 29, 15.3 mi. E of Burnet, 9 September 1969, Waller 2163 (MO); N slope Mt. Locke below McDonald Observatory, S.H. 118, 11.8 mi. NW of Davis Mountains State Park, 3 July 1972, Waller 2425 (MO); Comal Co, roadside along S.H. 46, 12 mi. W of New Braunfels, 15 October 1972, Waller 2434 (MO); roadside S.H. 29, 11 mi W of Georgetown, April 1969, Waller 1872 (MO); Uvalde Co., Garner State Park, 30 mi. N of Uvalde, 29 June 1957, Gould 7674 (MO); Culberson Co., 9 mi. E of Van Horn, 27 July 1943, Waterfall 5340 (MO); Brewster Co, 10 September 193, Tharp 43120 (MO); Coryell Co, 5 mi. W of Gatesville, 4 June 1957, Gould 7579 (MO); Brown Co., 2 mi. W of Borwnwood, 23 September 1957, Gould 7707 (MO); Handley, 23 September 1902, Reverchon 3527 (MO); Austin, 5 May 1942, Warnock s.n. (MO-1274105); Olmito, 15 April 1905, Tracy 9338 (MO); San Antonio, 20 September 1901, Bush 858 (MO); Dallas, 27 September 1900, Bush 1156 (MO); Hidalgo Co, Santa Ana National Wildlife Refuge, 14 April 1977, Solomon 2741 (MO); Amarillo, 30 May 1918, Palmer 13277 (MO); Lampasas Co., roadside U.S. 183, 11 mi. NW of Lampasas, 6 June 1969, Waller 1983 (MO); Austin, Hall 817 (MO). Utah: Beaver County, Red Knoll, east side Pine Valley, 3 September 1934, Hutchings & Stachmann n.d. (US-1648084) Mexico. Aguascalientes: 1.5 km a Real a Asientos, 22 October 1965, Atrupo 1723 (MEXU). Chihuahua: km 5 desviación a Nuevo Casas Grandes de carretera Cd. Juárez, 1600 m, 19 September 1955, Hernández & Mathus N-1911 (US); Sierra de Santa Eulalia, 2 km N of San Antonio, Harvey 1516 (US); camino de Ojinaga a Castillón, Johnston 8175 (GH); cercanías de rancho El Pino, 10 km SW de Villa Rica, Stewart 2395 (GH); Sta. Eulalia Mountains, August 1875, Pringle 376 (F, GH, MO, NY, P, US); base of Cerro Campana, 79  km  N of Chihuahua, 1620  m, 23 August 1975, Davidse & Davidse 10057 (MO). Coahuila: 5 mi S of Piedras Negras, 23 May 1965, Gould 11282 (US); Monclova, in vicinity of Chulavista Hotel, 23 May 1965, Gould 1187 (US); without locality, 1880, Palmer 1338 (US, W); 12 miles west of San Buenaventura, 1800  ft, 28 October 1964, Reeder & Reeder 3928 (MEXU); Municipio de Castaños, south of Castaños, Rancho Santa Teresa, Wynd & Muller 174 (GH, MO, NY, US); Laguna de Jaco, Johnston & Muller 1111 (GH); 20  miles  N of Saltillo, Gould 11796 (GH); 20  miles  W of Saltillo, Shreve 9821 (GH); 70  miles  W of Saltillo, Wallies 13294 (F, GH); Saltillo, 10/14 July 1910, Hitchcock 5605 (US); San Antonio de los Alamos, eastern base of the volcanic

Key to Ungrouped Panicum Sections and Species

153

Sierra de San Antonio de los Alamos, Johnston 8250 (US), 8250 a (GH); Sierra de las Cruces, eastern foothills 8 mi. north of Santa Elena Mines, 7 September 1940, Johnston & Muller 1017 (F, GH, US); Sierra del Pino, near La Noria, Johnston & Muller 467, 694 (GH), Stewart 1211 (GH); Sierra de Santa Rosa, Marsh 1278 (GH); 2 miles NW of Frontera, August 1938, Johnston 7170 (GH); 14 mi east of Paila, 400 ft, 6.9.1940, Shreve & Tinkham 9897 (US); 51 miles northwest of Múzquiz, Gould 11157 (USA); La Cuesta del Plomo on the Múzquiz-Boquillas Hwy., 28°44′N, 102°31′W, 1750–1775  m, 14 September 1972, Chiang et  al. 9201 (F, MO); 18 miles south of Sabinas, just north of El Sauz, 18 October 1957, Gould & Morrow 7906 (MO); 1 1/2 km W of Tanque La Mula which is NE of Rancho Piedra Azul, 29°09′30”N, 102°35′W, 1275  m, 24 March 1973, Johnston et  al. 10383 (MEXU, MO). Durango: eastern piedmont of Sierra Gamon about Peñón Blanco, 19/20 September 1948, 1800–1200 m, Gentry 8349 (US); 24 miles NW of La Zarca, 6200 ft, Soderstrom 819 (US); desert along Hwy. 40, 15 km SW of Pedriceña, 22 June 1964, Mick & Roe 44 (US); just to W of Mexico #40 at km 221, 26 km WSW of Torreón, 14 September 1978, Iltis et al. 114 (MEXU); 5 km al lE de La Joya, Mun. Poanas, 10 September 1984, Jimenez 7 (MEXU). Hidalgo: Mun. Jalapa, 500 ft, 1 July 1030, Chase 7256 (US); SW of Jacala, 4700 ft, Moore Jr. 3924 (GH, US). Nuevo León: Galeana, Puerto México, carretera 57, 67 km al N de Saltillo, 1600 m, 25 August 1978, Bernal & Cárdenas s.n. (MEXU 271683); orilla del Rio San Juan a 1 km del poblado San Juan, 230 m, 12 March 1986, García & Dominguez 1805 (MEXU); N of Sabinas, Hidalgo, 28 August 1971, Beetle 1061 (MO); 34.4 mi. N of Dr. Arroyo on Mex 60, 1560  m, 10 October 1976, Brunken & Perino 241 (MO); N of Monterrey, “La Escondida,” 612 m, Roybal 867 (MO, US), 883 (MO, US); cerca de Monterrey, bancos del Rio Pesqueira, Beetle 1077 (MO); Monterrey, 6/8 June 1910, Hitchcock 5537 (US). Oaxaca: Mun. Zapotitlan, 4  km al E de Zapotitlan Salinas, 25 September 1990, Sánchez-Ken et al. 328 (MEXU); 3 km al norte de la carretera a Yolotepec, Mun. de San Pedro y San Pablo, 1900  m, 23 September 1999, Paz Zambrano & Ramos Marchena 303 (MEXU). Puebla: afueras de Tehuacán, por la carretera a Esperanza, 1 September 1989, Chiang et  al. 294 (MEXU). Querétaro: 2 miles S of Querétaro, 6400 ft, Soderstrom 531 (US); 3 km al W de Las Rosas, sobre la carretera a Tequisquiapan, 2000 m, 8 September 1990, Rzedowski 50000 (ANSM, MEXU). San Luis Potosí: Valley of the Rioverde Between Rioverde and San Ciro, 850–1000  m, 12 September 1954, Sohns 1215 (US); Cárdenas, 19/20 July 1910, Hitchcock 57191/2 (US); Charcas, August 1934, Whiting 608 (US); 10 km al W de Guadalcazar, 1600 m, Rzedowski 4931 (GH, US); 8  km al W de San Lorenzo, km 62 de la Carretera San Luis Potosí a Morelos, Rzedowski 6869 (GH); Valle de Rioverde, between Rioverde and Boquilla, Sohns 1248 (F, MEXU, P, US); in the Sierra de Guadalcazar between Charco Blanco and Aguaje de García, 1100–1600 m, Sohns 1344 (US); Mun. Charcas, 6 km al E de Laguna Seca, Rzedowski 6547 (US); about 3 km east of Laguna Seca on the northwestern slopes of the Sierra de Alvarez, Sohns 1094 (MEXU, P); 10 km al S de Cárdenas, 1000  m, Rzedwosky 5174 (US); 30  miles south of San Luis Potosi, 1890 m, 12 August 1958, Reeder & Reeder 2967 (MEXU). Tamaulipas: Hacienda Buena Vista, 18 June 1919, Wooton s.n. (US-1061809); vicinity of San Miguel,

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Taxonomic Treatment

Bartlett 10591 (GH); K 678 on the Mexico City-Nuevo Laredo highway, 22  km south of Victoria, Martínez & Luyando F-2275 (US); low field by corn field, 4 mi S of Jaumave, 7 July 1949, Stanford et al. 2338 (US); vicinity of Victoria, 320 m, 1 May 1907, Palmer 554 (US); Cerro de la Tamulipeca, vicinity of San Miguel, 24 July 1930, Bartlett 10549 (US); 35 km from Victoria on the road to Casas And Soto La Marina, ca. 280 m, 3 October 1956, Martínez Martínez & Borja Luyando F-2303 (MEXU, US); 19 km from Mante on the Tampico highway, 18 September 1956, Martínez & Luyando F-1709 (US). No State: without locality, Virlet 1371 (US) 33. Panicum hirsutum Sw., Fl. Ind. Occ. 1: 173. 1797. Type: Jamaica, O. P. Swartz s.n. (holotype, S-R-3971; isotype, US 00148568, fragment and photo ex S). Fig. 37 Panicum chacoense Parodi, Darwiniana 15: 102, Fig.  8. 1969. Type: Argentina. Chaco: Isla Brasilera, 19 January 1963, A. G. Schulz 12138 (lectotype here designated, BAA 00000397; isolectotypes, BAA 00000395 BAA 00000396, CTES 0000044, G 00099650, SI-132521) Plants perennial, robust, up to 3 m tall, shortly rhizomatous; culms many noded, prostrate, without aerenchyma, with fulcrum roots at the lower not erect at the extremity, branching at the middle nodes; internodes 11–20 cm long, 0.4–0.8 cm in diameter, cylindrical, hollow, pale, with urticating tuberculate hairs, deciduous; nodes contracted, purplish, hairy. Sheaths 14–20 cm long, larger or smaller than the internodes, striate, hirsute, with deciduous tuberculate hairs, urticant, adpressed, greenish or tinged with purple, the margins membranous, glabrous or one of the margins ciliate. Ligule membranous at the base and shortly ciliate in the upper portion, 0.2–0.7 mm long, with long hairs beneath at the base of the blade; collar pilose. Blades lanceolate, 20–50 × 1.5–3.5 cm, flat or with involute margins, base subcordate to cordate, glabrous or with sparse scattered hairs, midrib manifest, margins scaberulous. Inflorescences terminal, diffuse, multiflowered, 25–45 × 5–15 cm, not deciduous at maturity; peduncles cylindrical, up to 25  cm long, glabrous; lower branches verticillate, the remaining ones opposite or alternate, appressed to the rachis to divergent, spikelets appressed, congested on the branches; main axis wavy, scabrous, pulvini slightly hairy to glabrous; pedicels 0.5–2  mm long, scabrous. Spikelets narrowly ovoid, 1.8–2.6 × 0.5–0.9 mm, deciduous, glabrous, pale, upper glume and lower lemma subequal, as long as the upper anthecium; lower glume 0.7–1.4 mm long, ovate, 1/2 the length of the spikelet or slightly less, 3-5-nerved, shortly hairy at the apex of the inner surface or entirely glabrous; upper glume 1.7–2.2 mm long, not deciduous at maturity, 7-11-nerved; lower lemma 1.7–2.2 mm long, 7–9 nerved, glumiform; lower palea lanceolate, 1.3–1.7 × 0.3–0.4 mm, 3/4 the length of upper anthecium, membranous, hyaline, the margins denticulate; upper anthecium narrowly ellipsoid, 1.2–1.6 × 0.5–0.7 mm, glabrous, smooth, shiny, pale, brown at maturity, with 2 scars at the base of lemma 0.1 mm long, brown at maturity; palea with simple papillae next to apex. Caryopsis ellipsoid, 0.8 × 0.5 mm, pale; hilum punctiform, embryo less than half the length of the caryopsis.

Key to Ungrouped Panicum Sections and Species

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Fig. 37  Panicum hirsutum. (a) Upper leaves and inflorescence. (b) Pedicels and spikelet, ventral view. (c) Spikelet, dorsal view. (d) Upper anthecium, ventral view. (From Schulz 12138, BAA)

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Taxonomic Treatment

Fig. 38  Distribution of Panicum hirsutum, P. mucronulatum and P. peladoense

Distribution and ecology (Fig. 38). A species originally described for Jamaica, it grows in the southern USA and Mexico, being found abundantly in Mesoamerica and the West Indies, having been found in much smaller proportion in South America in Colombia, Venezuela, Ecuador, Peru, Brazil, and Argentina. It lives in humid places along watercourses or on the edges of swamps, between 30 and 1800 m asl. Local names: “Cortadera,” “gamelote” (in Venezuela, according to the specimen Tamayo 1768) This species is abundant in Central America and Colombia and Venezuela, being occasional in northern Brazil. Only one collection has been recorded in Argentina, near the border with Paraguay. It differs from the rest of the species of the section for being a large plant, with wide panicles bearing small spikelets and for having sheaths with urticant hairs. Gould (1975) points out that the siliceous hairs present in the sheaths can cause skin irritation on contact with them. Panicum elatum Willd. ex Steud., Nomencl. Bot. (ed. 2) 2: 256. 1841, is a nomen nudum corresponding to this species. Additional material examined. USA.  Texas: Cameron co., vicinity of Brownsville, 1 August 1921, Ferris & Duncan 3175 (MO, US); Cameron Co.,

Key to Ungrouped Panicum Sections and Species

157

Southmost, 1941, Davis s.n. (MO-1273655); Rio Rico, near Rio Grande, about 6 miles from Mercedes, Silveus 625 (US); Brownsville, 18.4.1917, Nelson s.n. (US) Mexico. Campeche: Tuxpeña, 18 February 1932, Lundell 1355 (F, GH, MO, NY, US); a 6 km al S de Xpujil, 22 April 1982, Cabrera & Cabrera 2461 (MO, SI); 70 km al E de Escárcega, 26 March 1982, Cabrera & Cabrera 2237 (MEXU, MO). Chiapas: entrance to Cascade Mizola, 25 km S of Palenque, 91°59′W, 17°23′N, 280 m, 22 September 1988, Stevens & Martínez 25813 (MO); El Azufre, 700 m, 20 June 1983, Méndez 6220 (MO); slopes and small streams with Tropical Rain Forest along the ridges 6–12 km south of Palenque on the road to Ocosingo, 300 m, 12 October 1972, Breedlove 28837 (MEXU, MO); Mun. Ocosingo, 8  km NW of Bonampak, Lacanja-Changayab at the Río Lacanja, 350 m, 14 May 1982, Davidse et al. 20528 (MEXU, MO); tropical rain forest adjacent to small cascading river at Agua Azul, 300 m, 9 November 1981, Breedlove & Davidse 55245 (MO). Colima: Manzanillo, sea level, Hitchcock 7031 (RB, US). Oaxaca: San Antonio, 2 September 1894, Pringle 5573 (F, GH, MEXU, MO, US); District of Tuxtepec, Chiltepec and vicinity, 200  m, Martínez Calderón 568 (US); Río Verde, 7  km en línea recta al NNE de Santa María Chimalapa, 16°58′N, 94°39′W, 400 m, Vera Caletti et al. 97 (MO). Quintana Roo: 4  km NW of J.M.Morelos, along Hwy. 184, at the place where it crosses the marshes of Laguna Chichancanab, 18 May 1982, Davidse et al. 20612 (MEXU, MO); San José de la Montaña, 9 km S of Hwy. 186 on Hwy. to Tomás Garrido, 120 m, 9 May 1982, Davidse et al. 20271 (MEXU, MO); 3 km N of San Jose de la Montana, on road N to Thomas Garrido, W of Chetumal, 100 m, 9 May 1982, Davidse et al. 20246 (MEXU, MO); Coba, at edge of lake, Lundell & Lundell 7622 (US); a 6  km de la carretera Chetumal-Escarcega, sobre brecha al Ingenio Alvaro Obregón, 5 July 1980, Tellez & Cabrera 2630 (MEXU); a 16 km al Sur de San Jose, rumbo a T.  Garrido, 5 September 1980, Tellez & Rico 3414 (MEXU). San Luis Potosí: bank of Río Valles, 27 June 1910, Leavenworth 199 (GH, MO). Tabasco: Boca Cerro, Tenosique, 1 July 1939, Matuda 3577 (F, GH, US); 1.5  km  S of Villahermosa, Gilly et  al. 243 (GH); km 21 de la carretera Cárdenas-Coatzacoalcos, 17 September 1981, Magaña 427 (MEXU, MO). Veracruz: 9  km ESE of Ebano on hwy. 70, Nee & Hansen 18347 (F, MEXU); Estación Biológica San Andrés Tuxtla, 27 November 1978, 150  m, Martínez Calderón 1798 (F, MEXU, MO); Finca Sayula, Mun. Paso de Ovejas, Hernández Xolocotzi 3379 (US); near the Rio Cozalapa, about 10 miles from El Palmar Rubber Experiment Station, 100  m, 1 August 1953, Reeder & Reeder 1962 (MEXU). Yucatán: Becanche, 19 January 1956, Enriquez 347 (MEXU). Without locality, Gouin 6 (P) Guatemala. Huehuetenango: vicinity of Ixcan, Sierra de los Cuchumatanes, 24 July 1942, Steyermark 49430 (F). Izabal: Finca Murciélago, ca. 10 miles E of El Estor on Lake Izabal, 13 September 1961, Popenoe 34 (F); seashore around Punta Palma, across bay from Puerto Barrios, 22 April 1940, Steyermark 39802 (F). Petén: Parque Nacional Tikal, aguada del hotel Posada la Selva, 30 April 1970, Ortíz 1049 (F, NY); along Rio Cancuén, between mouth of Río Machaquila an bluffs above Tres Islas, 75 m, 26 April 1942, Steyermark 45998 (F); ruinas Plaza Mayor Tikal, 333 m, 16 November 1965, Molina 15281 (F, NY); 35 km E de Santa

158

 

Taxonomic Treatment

Elena, brecha El Remate-Tikal, 200 m, 9 November 1965, Molina 15426 (F, NY, US); Tikal National Park, Tikal, growing around aguada at Camp, 27 April 1959, Lundell 15918 (US) Belize. Cayo: 9.5 miles S of Georgeville, on road to Agustine, 30 May 1973, Croat 23483 (MO); vicinity of Millionario between the McCal River and Cuevas, 1900 ft, 30 May 1973, Croat 23664 (F, MO); Central Farm, Reyes 34 (US); Vaca Plateau, below Blue Hole Camp, 13 August 1980, Whitefoord 2090 (MO). Orange Walk: 1/2 mile E of San Antonio, 8 December 1976, Leino 76105 (MO). Stann Creek: Humming Bird Highway, 13 September 1954, Gentle 8380 (F, G, MO, NY, US). Toledo: Pate’s Camp, Edwards Road beyond Columbia, Gentle 7120 (F, G, NY, US); Maya Mountains, canyon along Bladen Branch from Richardson Creek to Quebrada de Oro, 16°32′N, 88°46′49″W, 100–200 m, 12 March 1987, Davidse & Brant 32401 (MO). Without locality, Smart 112 (F) Honduras. Atlántida: La Fragua, 20 m, 7 February 1928, Standley 55711 (F, US); Tela, near Lancetilla valley, 6 December 1927, Standley 53428 (F, US); vicinity of Tela, Standley 53692 (F, US). Francisco Morazán: San Antonio de Oriente, 22 July 1945, Rodríguez 3126 (F) Nicaragua. Zelaya: vicinity of Waní, including Río Ulí, ca. 13°41′N, 84°50′W, 90–100 m, 22 April 1978, Stevens 7982 (MO); prados empantanados, región entre Río Grande y Wenkalaya, 13 April 1949, Molina 2219 (F). Without department: Granade de Nicaragua, Levy 1018 (P) Costa Rica. Heredia: Finca La Selva, 100  m, 21 April 1982, Hammel 11795 (MO). Limón: near Puerto Limón, in plantations, Pittier 3634 (NY, US); margin of Bonilla Lakes, above Tunnel Camp, 13 December 1929, Dodge et al. 5610 (MO); Port Limón, near sea level, Hitchcock 8435 (US). Without locality, Tonduz 14586 (G) Panama. Bocas del Toro: 1  km NW of Bocatorito on SE side of island, 9 February 1989, Peterson & Annable 6729 (MO). Colón: 4 km E of Buena Vista, Nee 6783 (MO). Darién: Darién, September 1967, Duke 14117 (F, MO, NY); Río Sabana, Leopold 109 (MO); Santa Fe, Duke & Bristan 303 (MO). Panamá: past Chepo and Río Mamoni on road to El Llano, 25 September 1972, Tyson 6791 (MO); Agricultural Experiment Station at Matías Hernández, Pittier 6943 (US). Zona del Canal: between Frijoles and Monte Lirio, Killip 12174 (GH, US); between Gorgona and Tabernilla, Hitchcock 7964 (F, GH, K, MO, P, US); between Bohio and Frijoles, Hitchcock 8397 (USA); near Tabernilla, old field, Hitchcock 8382 (US); Pedro Miguel, Hitchcock 7957 (US); around Culebra, Pittier 3441 (US) Cuba. Cienfuegos: Soledad, 23 August 1931, Jack 6295 (NY, US). Habana: Guanimas, on beach, 22 August 1929, León 14181 (MO, US). Oriente: Los Caños, S of Guantánamo, 7 August 1913, León 3913 (US); Bayate, near roads, Ekman 6118, 6183. 10068 (US); Guaro, 20 km from Preston, 13 December 1926, Hitchcock 23396 (US); Preston, on the railroad, 13 November 1914, Ekman 3419 (NY, US). Pinar del Río: Mangas, Pueblo Nuevo, Hato del Corojal, 9 October 1923, Ekman 17578 (US). Villa Clara: Caibarién, Ekman 16312 (US). Without locality, de la Sagra 10 (P, W)

Key to Ungrouped Panicum Sections and Species

159

Haiti. Massif du Nord, 25 June 1925, Ekman 4392 (US); Plaine de L’Artibonite, Dessalines, Ekman 8534 (G). Without locality, Ekman 755 (US). Dominican Republic. Barahona: valley of Neiba, 15 August 1946, Howard 8357 (US); Barahona, Rio Yaque del Sur, 4 January 1926, Ekman 5791 (US). Duarte: valle del Cibao, Pimentel, at Río Cuaba, 23 June 1929, Ekman 13275 (US). Samaná: Sánchez, Jiménez 4309 (US) Antigua. Donovans Bridge, 1 September 1932, Box 176 (US). Santa Lucia: Mabouya Valley, 26 March 1933, Box 190 (US) Guadalupe. Entre la Riviere-Salée et St. Lucie, Duss 768 (NY); without locality, 1897, Duss 3917 (MO, NY, US), L’Herminier s.n. (G) Colombia. Bolívar: along the Caño Chacagua, N of Los Piñones, Island of Mompós, Lands of Loba, Curran 261 (US); along the Mompós-Juana Sánchez trail, Island of Mompós, Curran 250 (US); Canabetal, Pennell 3866 (GH). Cundinamarca: La Mesa, 1200 m, Triana s.n. (US-1865276). Magdalena: alrededores de Ciénaga, Romero Castañeda 8948 (MO, NY); Santa Marta, Smith 164 (COL, GH, K, MO, P, NY, US). Santander: Boca Sogamoso, Río Magdalena, 110 m, Pennell 3849 (GH, NY, US); prope Billete Blanco, Woronow et al. 4691 (US). Without state and locality, Mutis 2151 (US) Venezuela. Anzoatégui: Pariaguán, El Tigre, 16 July 1946, Oropeza s.n. (VEN). Barinas: Sistema de riego Río Boconó, Trujillo 11399 (MY). Delta Amacuro: Isla Cocuina, en sitio intervenido, Ramia 3330 (MY, VEN). Falcón: boca del Río Tocuyo, Trujillo 11492 (MY); 7 km E of Mirimire, Wingfield 5548 (MO). Guárico: Carretera Santa María de Ipire a El Socorro, 5 km antes del último poblado, 150 m, 17 August 1989, Zuloaga et al. 4507 (MO, SI, VEN); Hato Corral Viejo, Valle de la Pascua, 6 September 1955, Tamayo 4159 (VEN). Miranda: Río Grande del Tuy, above Paparo, Pittier 6332 (US). Portuguesa: Dtto. Araure, en terrenos adyacentes a la vía que une Agua Blanca con Payara, Rámirez Reyes 2051, 2054 (PORT). Táchira: Colonia Agropecuaria Los Andes, La Fría, 100 m, Medina 1246 (VEN). Trujillo: La Mesa de Esmujaque, 1800 m, Box 3703 (MY); cerca de Agua Viva, en las orillas del Río Motatán, Tamayo 1768 (US, VEN). Yaracuy: near Arenales, Río Turbio valley, 500  m, Box & Alayón 3833 (VEN); carretera entre Yaritagua y Barquimiseto, Burandt Jr. V0238 (MO, VEN); Finca “Los Apamates”, entre Urama y San Felipe, Romero 190 (MY); a orillas de la carretera entre San Felipe y Urama, Tamayo 3824 (VEN); San Felipe, orillas arenosas del Río Yaracuy, 11 April 1946, Burkart 16447 (SI). Zulia: Hacienda La Esperanza, km 14, Luces 188 (VEN); vicinity of Mene Grande, Pittier 10620 (GH, US, VEN); playas del Rio Guasare, Medina 763, 953 (VEN); San Martin, on Río del Palmar, 15 October 1922, Pittier 10534 (GH, NY, US, VEN); Distrito Perijá, 2 km W of the intersection of the Río Aricuaisa and the Maracaibo-La Fría hwy. (hwy. 6), 10  m, tall, grassy roadside plants to 2 m tall, Davidse 18851 (MO, VEN) Trinidad and Tobago. Little Tobago Island, 26 July 1914, Broadway 4895 (US); 8 miles S of Mayaro, along Mayaro- Guayaguayare road, 1 August 1970, Davidse 2510 (MO); Parrott Hill, 22 July 1913, Broadway 4586 (MO, NY, US); Manzanilla Bay, coconut groves along shore and adjacent Nariva Swamp, culms to 10 ft tall, succulent base, 19 June 1963, Soderstrom et  al. 1045 (NY, US); St. Margaretes,

160

 

Taxonomic Treatment

Mayaro, 12 June 1928, Broadway 2629 (USA); Fifth Company, Broadway 9291 (US); Manzanilla, Myers 4189 (US) Guyana. Within 30 mi of Georgetown, Rodway s.n. (US) Ecuador. Guayas: Panigón Plantation, 8  miles  S of Milagro, 50  m, 12 July 1923, Hitchcock 20580 (GH, NY, US), 20601 (US). Without state, Balao, Eggers 14538 (M) Peru. Ucayali: Canchahuayo, Vásquez et al. 7025 (MO, SI) Brazil. Pará: Lower Amazon, Estate Cacanal Grande, July 1914, Goeldi 111, 128 (US) Argentina. Chaco: Isla Brasilera, Schulz 12349 (CEN, F) 34. Panicum hirticaule J. Presl Plants annual, 25–120(−150) cm tall; culms decumbent, rooting at lower nodes to erect, branching toward the base; internodes 4–21 cm long, compressed, hirsute; nodes shortly hirsute. Sheaths 3–10 cm long, shorter than the internodes, hirsute, with tuberculate hairs, greenish to purplish, one margin ciliate, the other glabrous. Ligule membranous-ciliate, 0.9–1.5 mm long; collar hirsute. Blades lanceolate to linear-lanceolate, 6–25(−31)  ×  0.4–1.6(−2.2) cm, flat, hirsute to sparsely pilose, with whitish, tuberculate hairs, base subcordate, amplexicaul, apex subacute, the margins papillose-pilose. Terminal and axillary inflorescences present, similar, lax, 9–26(−30) × 5–18 cm; peduncles subincluded in the upper leaf sheaths to exserted, up to 36 cm long, hispid to glabrous; main axis wavy, scabrous, glabrous to sparsely hairy near the base; first-order branches alternate to subopposite, sometimes subverticillate, divergent, glabrous to sparsely hairy toward the base; second-order branches adpressed with spikelets congested; pulvini glabrous; pedicels 0.9–2.7 mm long, scaberulous. Spikelets ovoid, 1.9–2.8(−3.5) × 0.8–1 mm, glabrous, brownish to violaceous, abruptly acuminate; upper glume and lower lemma subequal, 0.8 mm longer than the upper anthecium; lower glume ovate-acuminate, 1.3–2.4 mm long, 1/2–3/4 the length of the spikelet, 3-5-nerved, midrib scaberulous to smooth; upper glume 1.8–2.9(−3.3) mm long, 7–9(–11) nerved, midrib scaberulous toward the apex to smooth; lower lemma glumiform, 1.8–2.8(−3.3) mm long, 9 nerved; lower palea lanceolate to ovate-acuminate, 0.4–0.9 × 0.2–0.4 mm, membranous, hyaline, glabrous; upper anthecium ellipsoid, 1.5–2.3 × 0.7–0.9 mm, crustaceous, glabrous, smooth, shiny, pale, with blackish tints at maturity, papillose, with simple papillae, over the entire surface, or papillae only present towards the apex of the palea, with two basal scars ca. 0.2 mm long, brown at maturity; lemma 7 nerved. Caryopsis ellipsoid, 1–1.7 × 0.7–0.8 mm, whitish; hilum punctiform, embryo half the length of the caryopsis. Key to the Varieties 1. Upper anthecium smooth, shiny, with papillae at the apex of the palea................. ................................34a. P. hirticaule var. hirticaule 1. Upper anthecium conspicuously papillose, opaque................................................ 34b. P. hirticaule var. verrucosum

Key to Ungrouped Panicum Sections and Species

161

34a. Panicum hirticaule J. Presl var. hirticaule, Reliq. Haenk. 1(4-5): 308. 1830. P. polygamun Sw. var. hirticaule (J. Presl) E. Fourn. Mexic. Pl. 2: 28.1886. Panicum capillare L. var. hirticaule (J. Presl) Gould, Madroño 10(3): 94. 1949. Type: Mexico. Guerrero: Acapulco, T.  Haenke s.n. (holotype, PR, n.v.; isotypes, B_10_0366130B, BR 0000006884147, HAL 0063369, MO-123225, US 00148571, fragment ex PR, W 0011325, W 18890236488) Panicum capillare L. var. miliaceum Vasey, Contr. U.S. Natl. Herb. 1(1): 28. 1890, non-P. miliaceum L., 1753. Panicum sonorum Beal, Grasses N. Amer. 2: 130. 1896. Panicum hirticaule J.  Presl var. miliaceum (Vasey) Beetle, Phytologia 47(5): 381. 1981. Panicum hirticaule J. Presl subsp. sonorum (Beal) Freckmann & Lelong, Sida 20(1): 172. 2002. Type: Mexico. Sonora: Lerdo, 1889, E. Palmer 947 (holotype US 00133127) Panicum hirticaule J. Presl var. glabrescens Andersson, Kongl. Svenska Vetensk. Acad, Handl., n.s., 1853: 135. 1855. Type: Ecuador. “In insula Chatham, May 1852, N. J. Andersson s.n. (lectotype here designated, BR 0000006883812) Distribution and ecology (Fig. 39). It grows from the USA, Mexico, Mesoamerica, and the Caribbean to South America, in Venezuela, Colombia, and Ecuador. It is frequent in open environments, on modified soils. It grows from sea level up to 2500 m. Local names: “Triguillo,” “zacate de año,” “zacate peludo perdis” (in Mexico); “chiri-chiri” (in Peru) Species related to P. stramineum and P. ghiesbreghtii. It differs from P. stramineum in having a reduced lower palea, brownish to purplish spikelets, densely arranged in the branches, these being appressed. Panicum ghiesbreghtii is a perennial species, with larger, linear-lanceolate blades, 16–30(−55) cm long. Spikelet size in P. hirticaule normally ranges from 1.9 to 2.8 mm in length. Some specimens from Mexico and the USA have spikelets up to 3.5 mm in length, which also have a rounded base of the upper anthecium. There are also several specimens from Peru, from the departments of Huánuco, La Libertad, and Loja (Sánchez Vega 2338, 2363, Cabanillas et al. 15, 17, Ochoa 1584, McBride 3526, López Miranda 951 and Ridoult 1919), of larger size and with spikelets of approximately 3.3–3.5 mm in length. Hitchcock (1927) cites P. hirticaule for Peru on the basis of two specimens Hitchcock 22115 and McBride 3526. The first of the above specimens corresponds to P. exiguum and the second to the previously cited specimens of P. hirticaule from Peru with larger spikelets. Swallen (1943) cites this species for Bolivia, without mentioning herbarium specimens, and this citation must correspond to P. stramineum. Larger plants from the Sonoran Desert region, previously identified as P. sonorum (one of the synonyms of this species), were cultivated for flour production by native tribes of that region. Additional material examined. USA.  Arizona: Tucson, 6 August 1903, Thornber 208 (MO); Arizona-Gila, 3500  ft, 9 September 1959, Pase 1084 (SI); Pima Co, near base of trail to top of Mt. Baboquivari, 14 September 1947, Gould 4468 (LA, MO); Pima County, 4  miles west of Havanakya, 2,500  ft, Gould &

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Taxonomic Treatment

Fig. 39  Distribution of Panicum hirticaule var. hirticaule

Haskell 3192 (GH, LA); Gila County, Pocket Creek Canyon, Sierra Ancha Mountains, 29 August 1946, Gould & Hudson 3748 (GH, LA); Cochise County, 17 miles east of Douglas, 3800 ft, 4 October 1947, Gould & Haskell 4510 (GH); Tucson, 22 August 1908, Sherff s.n. (GH); Cochise County, Rucker Canyon, Chiricahua Mountains, 5 October 1947, Gould & Haskell 4550 (US); Santa Cruz, Sycamore Canyon, 29 August 1938, Gooding & Gooding 418 (US); Santa Rita Mountains, 20 September 1902, Griffiths & Thornber 28 (US). New Mexico: Cedar Spring, Mulford 1078 (MO); 1 mile W of Hillsboro, 6000  ft, 2 October 1904, Metcalfe 1442 (GH, LA, US); Hidalgo County, 19 miles north of Rodeo, 4 October 1966, Gould 12032 (LA); Peña Blanca, Organ Mountains, 21 October 1906, Standley s.n. (US-953124). Texas: 1 mile SE of College Station, 21 April 1940, Evans s.n. (US-2434928); El Paso, 10 September 1884, Jones s.n. (LA-39220). Washington: W. Klickitat Co, bottom land near Bingen, November 1894, Suksdorf 2330 (GH, LA, MO).

Key to Ungrouped Panicum Sections and Species

163

Mexico. Baja California Sur: km 65 east of San Antonio, 6 June 1973, Beetle M-2551 (MO); 5 miles east of San Antonio, 8 January 1973, Beetle M-2646 (MO); Mesa de San Gerónimo, north from Rancho Viejo (on road from Loreto to San Javier), 23 September 1965, Carter 5016 (LA, MEXU). Chiapas: marshes along Rio Usumacinta, 25–30 km east northeast of Palenque Junction, 30 m, 12 December 1981, Breedlove 56081 (MO, NY); along road from Highway 195 to Simojovel de Allende, 2.7 mi past village of El Bosque at km 21.5, 1030 m, 16 February 1979, Croat 47699 (MO); forest at Mirador for Chicoasen Dam along road from Tuxtla Gutiérrez to the Chicoasen Dam, 850  m, 17 November 1976, Breedlove 41540 (MO). Chihuahua: N end of E side of Sierra del Cuchillo Parado, 29°37′N, 104°55′W, 1000–1300 m, 21 October 1972, Wendt et al. 9788 (MO, NY); 20 Km S of Ciudad Camargo, 1220 m, 1 July 1939, Harvey 1378 (GH, MO, US); Hacienda San Miguel, Palmer 1c (GH, NY); Nabogame, 28°30′N, 108°30′W, 1800  m, 10 November 1988, Laferriere 2725 (MO); Guasaremos, Río Mayo, 20 September 1935, Gentry 1841 (MEXU, MO, US); Sierra Madre Occidental, W of Casas Grandes, 5 miles of Hernández, 700 ft, 18 September 1960, Reeder et al. 3517 (US); 6 miles west of Piloncillo, 24 September 1938, Johnston 7866 (GH). ChihuhuaSonora: Rancho Carreteras, 1460  m, 26 August 1939, Harvey 1600 (MO, US). Colima: Alzada, 21 September 1910, Hitchcock 7089 (US); along hwy. 110, site 52, 25 July 1975, Wallace et al. 292 (MEXU, MO); Colima, July 1897, Palmer 143 (ISC, NY, US, W). Durango: 10 km N of Ceballos and 8 1/2 S of Chihuahua state line on the Torreon-Chihuahua highway, 1200  m, 26°36′N, 102°12′W, Johnston et  al. 12292 (MEXU, MO). Guerrero: south Chilpacingo, 7 December 1941, Leavenworth 966 (MO); Río Balsas, 26 August 1910, Orcutt 4197 (MO); Balsas, 1500 ft, 9 September 1910, Hitchcock 6814 (USA). Guanajuato: Irapuato, 5800 ft, 1 October 1910, Hitchcock 7424 (USA). Jalisco: 34  km west of La Barca, 27 August 1981, Beetle M-7030 (MO); ca. 5 road-miles SW of Santa Cruz de las Flores, 1550 m, 24 August 1957, McVaugh 16326 (MEXU, NY, US); Tecalitlán, al S de Tecalitlán, rumbo a Pihuamo, 12 August 1987, Ornelas U. et al 1167 (MEXU). México: Temascaltepec, Anonas, 24 July 1934, Hinton 6337 (MO, NY, US); Temascaltepec, Barranca, Ixtapán, 1000  m, Hinton 1062 (F, MEXU, NY, US). Michoacán: Chavinda, 27 September 1946. Hernández Xolocotzi et  al. x-2775 (USA); Huétamo, 1938, 240 m, Gándara 2 (US); at edge of tropical deciduous forest, La Majada, 8 May 1941, Leavenworth & Hoogstrall 1335 (GH, NY). Nayarit: 1 mile west of Mazatán, 600 m, 17 September 1960, McVaugh 19117 (NY, US); near km 31, 4–5 miles N of Compostela, 22 September 1960, McVAugusth 19333 (NY, US). Oaxaca: 2 km al SE de San Martín Toxpalán, por carretera a Teotitlán del Camino, 26 August 1980, Medrano et  al. F-1495 (MO); Los Tules Niltepec, 11 December 1985, Torres C. et  al. 7860 (MEXU, MO). Sonora: AlamoMexican boundary linear White Water, 11 September 1893, Mearns 2294 (US); El Rancho de la Nacha, 25 mi. W of La Angostura, 4300 ft, 14/20 August 1941, Vera Santos 1842 (GH, LA), 1850 (GH, LA, MEXU, MO, NY); 5 mi. E of El Rancho de la Nacha, 16 August 1941, Vera Santos 1836 (GH, US); 10 mi. S of Nogales, 16 September 1908, Hitchcock 3631 (US); summit of Cerro Prieto in the vicinity of the microwave station, 15 km E of Navojoa above the road to Alamos, 5 September 1989, Sanders

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Taxonomic Treatment

et al. 9297 (MO); 27 miles west of Hermosillo on the road to Kino Bay, 720 ft, 28 August 1941, Wiggins & Rollins 137 (GH, LA, MO, NY); Cañón de Huépari, north of Aribadi, 1310 m, 2 September 1939, Harvey 1653 (GH, MO); S of Hermosillo, 17/20 September 1908, Hitchcock 3604 (US). Tabasco: km 6.6 de Emiliano Zapata hacia Tenosique, 5 July 1981, Cowan 3398 (MEXU, MO). Veracruz: La Mancha, carretera Cardel-Nautla, 1 August 1971, Dorantes 246bis (F, GH); Baños del Carrizal, Purpus 6212 (GH, LA, MO, NY). Yucatán: Chichankarrab, Gaumer 1501 (F) Guatemala. Izabal: Los Amantes, 9 May 1919, Blake 7326A (US); vicinity of Puerto Barrios, 2 July 1922, Standley s.n. (US). Suchitepequez: South of Alotenango Farm, 7 miles south of Tiquisate along road within 3 miles of ocean, 19 June 1942, Steyermark 47766 (F); Tiquisate, 100 m, 19 June 1942, Steyermark 47859 (F, US). Zacapa: rocky slopes between San Pablo and Pepezca, 200–250  m, 8 October 1939, Steyermark 29326 (F); vicinity of Zacapa, 200 m, 6 October 1940, Standley 74383 (F) Honduras. Choluteca: 7 km by road NE of Choluteca, 50 m, 4 July 1970, Pohl & Davidse 12177 (F, ISC, MO). Comayagua: Comayagua, Estación Agrícola El Carao, 630 m, 30 July 1983, Casco 46 (MO). Cortés: San Pedro Sula, Hacienda Monte Moria, 16 April 1984, Cristoff 181 (NY). Francisco Morazán: Tegucigalpa, cerro El Berrinche, 1000  m, 6 August 1978, Castro 69 (MEXU, MO). Valle: 4 km SE of San Lorenzo, 0–5 m, 5 October 1986, Davidse & Pilz 31665 (MO) El Salvador. La Libertad: along road ca. 1 km NE of Hwy. CA-1, on road 9 km WSW of Quetzaltepeque, 18 June 1978, Pohl & Gabel 13609 (F, ISC); Río Chilana west of La Liberta, 27 July 1950, Rohweder 1497 (MO). Santa Ana: Lagunita Clara, 5 km S of Metapán, 500 m, 10 June 1970, Pohl & Davidse 11872 (F, ISC, MO); along hwy. 12, 9 km N of Texistepeque, 550 m, 25 June 1978, Pohl & Gabel 13665 (F, ISC, MO). San Salvador: Apulo, iinpastures slopes near Lake Ilopango, 18 June 1949, Williams & Molina 16751 (F) Nicaragua. Estelí: Kukamonga km 167, Portal de Belén, 1 August 1983, Moreno 21833 (ISC, MO); faldas del Chayo 13°16′N, 86°20′ W, 700–1000 m, 31 July 1983, Moreno 21790 (ISC, MO). León: along Hy. 12, 4 km S of road to Puerto Somoza, 50 m, 18 July 1970, Pohl & Davidse 12369 (F, MO). Managua: km 24 on Hwy. 12 (Carretera vieja a León), 7 km WSW of summit of Sierra de Managua, ca. 12°04′N, 86°26′W, ca. 200 m, 7 July 1977, Stevens 2672 (F, ISC, MO); Rte. 1, 12 km E of Managua, 1 August 1972, Seymour 6275 (MO); road 1, 12 km E of Managua, 31 December 1969, Seymour 2829 (GH); vicinity of Managua, Garnier 833 (US). Masaya: km 17.5 carretera Managua-Masaya, terrenos de la UCA, 12°01′N, 86°10′W, 10 June 1982, Sandino 3038 (MO). Rivas: Isla Ometepe, Volcán Concepción, Los Hatillos, camino Las Delicias, 11°33-34′N, 85°36′W, 140–350 m, 9 August 1984, Robleto 980 (MO, SI); Isla Ometepe, Volcán Maderas, cafetal Las Cuchillas, 11°28-29′N, 85°29-30′W, 200–300 m, 23 September 1984, Robleto 1269 (ISC, MO, SI); along CIA, near Lago de Nicaragua, 24 km SE of Rivas, 1 August 1971, Pohl 12674 (F, ISC, MO); San Juan del Sur, beach of Pacific Ocean, 8 July 1970, Pohl & Davidse 12228 (ISC)

Key to Ungrouped Panicum Sections and Species

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Costa Rica. Guanacaste: Playas del Coco, S end of bay, 6 July 1968, Pohl & Davidse 10653 (F, ISC, LA); upper slopes of bluffs above Pacific Ocean, S of Playas del Coco, 5 August 1966, Pohl & Calderón 10216 (ISC, MO). Puntarenas: vicinity of Cascajal, along road from Cascajal to Pigres, 30–100 m, 6 July 1949, Holm & Iltis 286 (GH). Without province: Atenas, 25 September 1911, 1165 m, Hitchcock 8517 (US) Panama. Panamá: Taboga Island, December 1923, Standley 27960 (US). Zona del Canal: Chiva Chiva trail near Miraflores Lake, 30 September 1965, Tyson 1396 (MO); Balboa, Sosa Hill, December 1923, Standley 25277 (MO, US); Contractors Hill, 28 August 1962, Dwyer 2781 (MO); Canal Zone, Balboa, November 1923, Standley 27407 (US); Balboa Heights, 4 October 1918, Killip 4326 (NY, US) Haiti. Plaine Cul-de-Sac, Port-au-Price, Damine, in fields, 25 May 1928, Ekman 9991 (G, US) Colombia. Vichada: Between Guama and Saldaña, frequent in open sandy roadsides, 300 m, 11 November 1983, Wood 4091 (COL) Venezuela. Guárico: El Socorro, 15 July 1946, Burkart 17204 (SI); Carretera Santa María de Ipire a El Socorro, 08°58′N, 65°40′W, 150  m, 17 August 1989, Zuloaga et al. 4508 (MO, SI, VEN) Ecuador. Archipiélago de Colón: Isabela Island, Volcano Alcedo, Fowler 8d (MO); Indefatigable Island, 6 mi north of Academy Bay, 10 April 1930, Svenson 236 (F, GH, LA). Guayas: Capeira, km 21, Guayaquil to Daule, 20–200  m, 15 February 1982, Dodson & Gentry 12506, 12514 (MO) Peru. Cajamarca: Prov. Cajamarca, alrededores de Choropampa, sobre la carretera de penetración Pacasmayo-Cajamarca, a la altura del km 124, anual, en bordes de campos de cultivo y laderas no cultivadas, 7 March 1981, Sánchez Vega 2363 (MO, SI, US); Prov. Chota, El Tingo a 6 km sobre la carretera CochabambaHuambos, 2000  m, 20 June 1980, suelo pedregoso arcilloso, Sánchez Vega 2338 (US); La Yunga, Sunchubamba, en ladera junto a terreno de cultivo, 23 August 1974, 1940 m, Cabanillas et al. 15 (MO, US), 17 (MO, SI, US); Cayayup, 2500 m, Ochoa 1584 (F). Huánuco: Huánuco, in cornfield, loose clumps, subdecumbent below, 28 April 1923, McBride 3526 (F, US). La Libertad: Prov. Trujillo, ChualCruz de Cascaday, 1200 m, 29 March 1953, López Miranda 951 (US); Prov. Trujillo, Simbal, 1200 m, 29 March 1953, Ridoult 1919 (NY). Piura: Prov. Talara, alrededores de Talara, 15 October 1983, Sagástegui 10912 (MO, SI); cerca a Bocapán entre Talara y Zorritos, 100 m, 19 April 1949, Ferreyra 5929 (F); 40 km west of Piura, 50  m, 29 March 1939, Stork & Horton 11361 (F, MO); 20  km  N of Lamabayaque, 100 m, 15 April 1949, Anderson 897 (LA), 899 (US) 34b. Panicum hirticaule var. verrucosum Zuloaga & Morrone, Ann. Missouri Bot. Gard. 83(2): 253. 1996. Type: Mexico. Chihuahua: between Casas Grandes and Sabinal, 4 September 1899, E. W. Nelson 6355 (holotype, US 00495307; isotype, GH) Var. verrucosum differs from the typical variety in having conspicuous papillae over the entire surface of the upper anthecium, the upper anthecium blackish at maturity, and the spikelets are larger, between 2.4 and 3.5 mm long.

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Distribution and ecology (Fig. 23). It inhabits the southern USA, in Arizona, and northern Mexico, in the states of Chihuahua, Guerrero, Jalisco, Nayarit, Sinaloa, and Sonora. It is found in fields and reaches up to 600 m asl. Local name: “Zacahuastle” (in Mexico) Additional material examined. USA.  Arizona: near Fresnal, Papago Indian Reservation, 30 September 1934, Peebles10366 (GH, US); Pima County, South Canyon, Baboquivari Mts., 15 September 1931, Gilman 37 (GH); Tucson Mountains, 25 August 1904, Griffiths 6939 1/2, pr. p. (US); 7 miles N of Nogales, USA highway 89, Ruby Road to Peña Blanca Recreation Park, 1 August 1959, Soderstrom 160 (US); north slope of Santa Rita Mountains, 13/21 September 1904, Griffiths 7194 (US) Mexico. Chihuahua: Chapingo, Hernández & Tapia 240 (US). Guerrero: Cuautepec, San Agustín Cuilutia, 10 July 1986, Herrera Castro 84 (MEXU). Jalisco: Amatitán, Barranca Santa Rosa, orilla del Río Santiago, 600 m, 4 August 1977, Villarreal 6661 (MEXU). Nayarit: El Nayar, Jesús María, 600 m, 4 August 1977, Colunga & Zizumbo 44 (MEXU). Sinaloa: alrededores del dique La Primavera, selva baja caducifolia a 10  km al sur de Culiacán, 60  m, Aviña 1248 (MEXU). Sonora: without locality, 17 September 1908, Hitchcock 3526 (USA), Wiggins 6023 (US) 35. Panicum hispidifolium Swallen, Contr. U.S. Natl. Herb. 29(9): 424. 1950. Panicum hispidum Swallen, Contr. U.S.  Natl. Herb. 29(6): 269. 1948[1949], nom. illeg. hom., non G.Forst., 1786. Type: Honduras. El Paraíso: Güinope, 1430 m, 1 January 1944, J.  V. Rodríguez 1981 (holotype, US-1869144; isotypes, EAP 61656, F 004688F, F-1166244, F-1306220). Fig. 40 Panicum flabellatum E. Fourn., Bull. Soc. Bot. France, ser. 2, 27: 293. 1880, nom. illeg. hom., non Steud., 1853. Type: Nicaragua. Managua: Isla de Ometepe, October 1869, P.  Lévy 1166 (holotype, P 00740956; isotypes, G 00099621, P 00740954, P 00740955, US 00148495, fragment ex P) Plants annual, herbaceous; culms cespitose, erect, 20–100 cm tall, geniculate or not at the lower nodes; internodes cylindrical, hollow, 4–15 cm long, 0.1–0.4 cm in diameter, hirsute, with deciduous tuberculate hairs; nodes hairy, purplish. Sheaths 4–13 cm long, smaller or larger than the internodes, densely hirsute, with long tuberculate hairs, the margins ciliate. Ligule shortly membranous at base, then long ciliate, 2–3 mm long; collar hairy, violaceous. Blades linear-lanceolate, 8–45 × 0.8–1.3 cm, flat, herbaceous, acuminate, base subcordate or rounded, hirsute, with scabrous margins, ciliate towards the base, midrib manifest. Inflorescence terminal exerted; panicles lax, diffuse, pyramidal, multiflowered, 12–50 × 10–40 cm; peduncles included in leaf sheaths to exserted, up to 40 cm long, cylindrical, hirsute; main axis wavy, hispid, covered with rigid tuberculate-based hairs, glabrous toward the upper portion of panicle, branches opposite or alternate, the lower ones occasionally subverticillate, divergent, up to 17 cm long, branch axes hispid; pedicels long, 0.5–2.5 mm long, scabrous; axillary inflorescences present, similar to the terminal one. Spikelets broadly ellipsoid, globose, open at apex, 3.2–3.7 × 1.2–1.6 mm, solitary, glabrous, greenish or tinged with purple, acute to acuminate, glumes and lower lemma with

Key to Ungrouped Panicum Sections and Species

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Fig. 40  Panicum hispidifolium. (a) Plant. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Lower palea. (e) Upper anthecium, dorsal view. (f) Upper anthecium, ventral view. (g) Caryopsis, embryo view. (h) Caryopsis, hilum view. (From Zuloaga 4363, SI)

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Taxonomic Treatment

marked veins; upper glume and lower lemma subequal to or exceeding up to 0.3 mm in length the upper anthecium; lower glume ovate, 2–2.6 mm long, 1/2-3/4 the length of the spikelet, acute, 5 nerved, the midrib scabrous, scaberulous toward the apex of the inner surface; upper glume about equal in length to the spikelet, acute, 7 nerved, the midrib scaberulous toward the apex, scaberulous at the apex of the inner surface; lower and upper glume separated by a short internode ca. 0.3 mm long; lower lemma glumiform, 7–9 nerved, acute, hairy at apex of inner surface; lower palea ovate-lanceolate, reduced, 1.6–1.8 × 0.6–0.9 mm, hyaline, the upper margins sparsely hairy to glabrous, glabrous on rest of the surface; upper anthecium broadly ovoid, 2.1–2.5 × 1.2–1.5 mm, crustaceous, glabrous, smooth, shiny, with grayish tinge at maturity and a discoid ring at base, brown at maturity; lemma 7 nerved; palea with verrucose papillae toward the apex. Caryopsis broadly ovoid, 1.6–1.8 × 1.1–1.3 mm; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 34). This species is found from Mexico and Central America, in El Salvador, Honduras, Nicaragua, Costa Rica, to Colombia and Venezuela. It grows in savannas; it is frequent along roadsides. It grows between 60 and 1430 m asl and flowers and fruits between July and December. Species related to P. hirticaule, from which it differs by having larger globose spikelets, not appressed and distant, diffuse inflorescences that reach approximately 1/2 of the total length of the plant and ligule 2–3 mm long. Swallen (1955) erroneously considers in the synonymy of P. hirticaule and P. flabellatum, whose type agrees with P. hispidifolium. Additional material examined. Mexico. Chiapas: 58  km south of Mexican Highway 190 on road to Nueva Concordia, 900 m, 11 September 1974, Breedlove 37573, 37654 (MEXU, MO). Oaxaca: near road nine miles toward Tehuantepec from Zanatepec, Morley 696 (F, GH, LA, MO) Honduras. Cortés: Ocote Arrancado, 50 km N Lago de Yajoa, 600 m, bosque tropical húmedo, November 1980, Nelson et al. 5621 (MEXU, MO). El Paraíso: Road to Yuscarán, moist places around rocks, about km 2, Swallen 11344 (US). Francisco Morazán: road between El Jicarito and El Pedregal, 800–950  m, 13 November 1948, Standley 14492 (F); in cultivated fields at El Zamorano, Muller s.n. (GH, US-2115373), Rodriguez 541 (F, MEXU), 1234 (F); vicinity of El Zamorano, Swallen 11264 (F, MO, US); beyond Las Mesas, 800 m, erect annual, scarce, 5 November 1951, Standley 29252 (F); along Río Caparrosa, above El Zamorano, 900 m, September 1948, Standley 12793 (F, GH, LA); drainage of the Río Yeguare, 2600  ft, 8 March 1948, Glassman 2145 (F); road to San Antonio, vicinity of El Zamorano, Swallen 10838 (US); vicinity of Las Mesas, ca. 5 km E of Zamorano, 3 October 1986, Davidse & Pilz 31533 (MO, SI); valle de San Pedro, 35  km  SE de Tegucigalpa, 700  m, 28.8.1983, Dueñas 2 (MO); vicinity of El Zamorano, 800–850 m, 16 November 1949, Standley 24654 (LA) El Salvador. Morazán: 2 km W of Las Delicias, 13 September 1951, 650 m, Rohweder 2507 (MO) Nicaragua. Masaya: Piedra Quemada, 1 km NW of Volcán Santiago, 350 m, 28 October 1976, Neill 1115 (MO) Costa Rica. Guanacaste: 7 km N of Bagaces along the Carretera Interamericana, 100 m, 23 October 1968, Pohl & Davidse 11291 (F, MEXU); along CIA, savanna

Key to Ungrouped Panicum Sections and Species

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pasture, 200 m, 18 October 1982, Pohl & Davidse 14172 (MO); along west side of CIA, 5 km S of La Cruz, 0.7 km S of Hacienda Inocentes road, 200 m, 10 December 1978, Pohl & Gabel 13703 (F, MO); 8 km N of Liberia, 2 August 1971, Burger & Burger 7875 (F); 5 km SE of Liberia, on the Cañas road, 1966, Harris 40, 197 (F); 21.6  km NW of Liberia, savanna, sandy soil, on the Pan American highway, 1 August 1966, Mori & Anderson 243 (F, MO); 18 km S of Liberia along the Carretera Interamericana, 100 m, 18 August 1968, Pohl & Davidse 10944 (F); without locality, Anderson 1365 (US) Colombia. Atlántico: Puerto de Colombia, 21 November 1912, Hitchcock 9927 (USA). Magdalena: Santa Marta, Smith 2152 (COL, F, G, GH, K, MO, NY, P, US) Venezuela. Anzoátegui: Las Piedritas, 16 July 1946, Burkart 17262 (SI); Pariaguán, 1 October 1939, Muller 804 (VEN); borde carretera El Tigre-Ciudad Bolívar, Ponce & Rodríguez 16 (MY); a 33  km al oeste de El Tigre, carretera a Pariaguán, 8°56′N, 64°35′W, 250 m, 17 August 1989, Zuloaga et al. 4504 (MO, SI, VEN); El Piñal, 3 km noreste de San Diego de Cabrutica, 13 August 1984, Montes 2018 (MO). Apure: Bajo Apure, Ramia & Montes 4831 (VEN). Aragua: Rancho Grande, Parque Nacional Henry Pittier, 480  m, sabanas de loma pedregosa, Montaldo & Ramia 3304 (MY). Bolívar: road to Salto Chavaripo, 88.7 km SW of Caicara del Orinoco, 7°7′N, 66°28′W, 4 September 1985, Steyermark et al. 131327 (MO); Distr. Cedeño, vicinity of Panare village of Corozal, 6 km from Maniapure toward Caicara, 6°55′N, 66°30′W, Boom & Grillo 6189 (NY). Cojedes: Galeras del Pao, August 1980, Ramia 7201 (VEN); San Carlos, sabanas entre San Carlos y Manrique, 29 September 1978, Garófalo 163 (VEN). Guárico: El Perro, 90  m, Hernández León s.n. (VEN-39936); carretera Calabozo-El Sombrero, a 7 km antes de la última población, 9°03′N, 67°3′W, 250  m, 9 August 1989, Zuloaga et  al. 4363(MO, SI, VEN); Dpto. Infante, Parque Nacional Aguaro-Guariquito, 100 m, December 1981, Delascio et  al. 11585 (MO); 12  km de Calabozo, 9 November 1971, Davidse 2950 (MO, VEN); Sabana Los Robles, cerca de El Sombrero, 14 July 1946, Burkart 17190 (SI, VEN); entre El Sombrero y Pitara Bridge, 11 September 1927, Pittier 12511 (NY, US, VEN); Estación Biológica Los Llanos, 10 km SSE de Calabozo, 60 m, Aristeguieta 4301 (VEN), 4334, 5665 (US, VEN), 5755 (VEN), November 1971, Davidse 2925 (MO); 28 km la N de Santa Rita, along highway between Chagauramas and Cabruta, 100  m, palm swamp with Mauritia flexuosa, in savanna, 21 November 1973, Davidse 4314 (MO). Monagas: without locality, Larez 605 (VEN). Portuguesa: Dtto. Ospino, alrededores de carretera y comienzos de primera bajada hacia Ospino, Rámirez Reyes 1773 (MO, PORT); Dtto. Guanare, orilla de carretera entre Las Matas y San Rafael, Rámirez Reyes 1715 (PORT). Trujillo: vicinity of Dividive, 22 November 1922, Pittier 10830 (GH, NY, US, VEN); Llanos de Trujillo, Muller 987 (VEN) 36. Panicum lepidulum Hitchc. & Chase, Contr. U.S. Natl. Herb. 15: 75, Fig. 64. 1910. Type: Mexico. Chihuahua: rocky hills near Chihuahua, 22 September 1885, C. G. Pringle 497 (holotype, US 00132937; isotypes, F 0046890F, KFTA 0000129, MEXU 00003656, NY 00381766, NY 00381767, US 00132936, W 19160022815). Fig. 41

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Taxonomic Treatment

Fig. 41  Panicum lepidulum. (a) Plant. (b) Spikelet, ventral view. (c) Spikelet, lateral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (a–g, from Davidse & Davidse 9927; MO; h–i, from Arsène 2864, MO)

Key to Ungrouped Panicum Sections and Species

171

Plants perennial, cespitose, shortly rhizomatous; culms erect, few noded, not glaucous, (15–)25–70 cm tall, without aerenchyma, simple, branching at the lower nodes, internodes 4–17 cm long, cylindrical, hirsute to glabrous, 1–2 mm in diameter, nodes hairy. Sheaths usually shorter than the internodes, without aerenchyma, hirsute with tuberculate hairs, not urticant, usually with purplish tinges, the margins ciliate. Ligule membranous-ciliate, (1–)2(–3) mm long; collar pilose. Blades linearlanceolate, 8–26 × 0.4–1.2 cm, erect, ascending, flat or with involute margins, hirsute, with tuberculate hairs, occasionally glabrous, base rounded and apex acuminate, margins scaberulous, the lower ones ciliate, midrib manifest. Inflorescences terminal, exserted, lax, diffuse, 7–20 × 4–10 cm, not deciduous at maturity; peduncles up to 30 cm long, glabrous to hirsute; main axis scabrous, hirsute toward the base to glabrous; first-order branches ascending, scattered, with second-order branches diverging, bearing 1–3 solitary spikelets; pulvini glabrous; pedicels claviform, scaberulous, 2–12  mm long. Spikelets long ovoid, (3.3–)3.6– 4.2 × 1.2–1.5 mm, acute, globose, glabrous, greenish or tinged with purple; lower glume ovate, 1.6–1.9 mm long, 1/2 or somewhat less than spikelet length, acute, 5 nerved, midrib scaberulous toward the apex; upper glume 3.3–3.6 mm long, acuminate, 9–15 nerved, midrib scaberulous toward the apex; lower lemma 3.3–3.9 mm long, acuminate, 9 nerved, midrib scaberulous toward the apex; lower palea reduced, ovate, 1.5–2 × 0.7–0.9 mm, hyaline, glabrous; lower flower absent; upper anthecium ovoid, 2.1–2.5  ×  1–1.5  mm, 0.9–1.5  mm shorter than the upper glume and lower lemma, crustaceous, glabrous, smooth, shiny, pale, with a circular disk at base, brown at maturity; lemma 7 nerved; palea with simple papillae and globose microhairs toward the apex. Caryopsis ovoid, 1.8 × 1 mm; hilum oblong, embryo 1/2 the length of the caryopsis. Distribution and ecology (Fig. 34). It is an abundant species in Mexico and occasional in Guatemala, grows in savannas or in oak forests, on often volcanic soils; it is also found on roadsides. It reaches up to 2400 m asl and flowers between March and October. Panicum lepidulum is related to P. pampinosum Hitchc. & Chase, a species from the USA, and P. parcum Hitchc. & Chase. Panicum pampinosum is distinguished by including annual plants, with inflorescences with the branches little divergent from the axis, and by having the spikelets with the lower glume 4/5 the length of the spikelet. Panicum parcum is characterized by including annual plants, with stipitate spikelets and lower glume 3/4 the length of the spikelet. Hitchcock (1951) cites this species for Utah, New Mexico, and Arizona and Waller (1976) for Arizona, without mentioning herbarium specimens. It was not possible to find specimens of P. lepidulum for the USA in the herbariums examined in that country, so these citations are considered doubtful. Additional material examined. Mexico. Aguascalientes: 8 mi. E of Calvillo, 6900 ft, 25 September 1963, Reeder & Reeder 3814 (MEXU, US); Pabellón, San Carlos, 29 October 1978, Guardado s.n. (MEXU-272159); Cerro del Picacho, 16 November 1978, González s.n. (MEXU-272160). Chihuahua: 12.8 mi. W of General Trias on Rte. 16, ca. 1780, 21 August 1979, Wagner & Solomon 4341 (MO).

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Coahuila: Sierra de la Madera, vicinity of La Cueva, Johnston 9104 (GH, MEXU). Distrito Federal: near Mexico city, Pedregal, 27 July 1910, Hitchcock 5958 (F, GH, NY, US); San Angel, 16 August 1929, Mexía 2740 (MO); Pedregal de San Angel, 1 km to the south of UNAM, 8 July 1967, Nee 183 (MEXU). Durango: Durango, 6 October 1910, Hitchcock 7591 (NY, US), August 1896, Palmer 525 (F, GH, MEXU, MO, NY, RB, US); Tobar, Palmer 533 (US); 46  miles NE of Durango on the Durango-Torreón road, 6300 ft, 30 September 1959, Soderstrom 812 (MEXU, US); 21 mi. SE of Durango, 5800 ft, 19 October 1966, Gould 12322 (USA). Guanajuato: 2–4 km east of Guanajuato, Sohns 288 (US); about 3 km NE of San Miguel, occasional in rocky soil near the top of Cerro de La Presa, 31 October 1952, Sohns 504 (US); Ojo de Agua, a small hill about one mile south of Abasolo, Sohns 253 (US); 2–4 kms east of Guanajuato, 16 October 1952, Sohns 285 (US); Santa Cruz de Juventino Rosas rumbo a Guanajuato, 2 September 1981, Beetle M-7324 (MO). Hidalgo: steep rocky volcanic outcrops at head of descent into Barranca de Metztitlán between Zoquital and Los Venados, 2000 m, Moore Jr. 4218 (GH, US); Jacala, 24 June 1939, Chase 7089 (USA). Jalisco: Paso de la Troje, near km 36, southwest of Ojuelos on road to Aguascalientes; rocky slopes on and near Cerro La Campana, McVaugh 16819 (NY, US); on rocky pastured slopes, 11 miles north of Ciudad Guzmán, Gould 9643 (US); Zapotlán, 5000  ft, 22 September 1910, Hitchcock 7117, 7238 (US); San Nicolás, 5000–6000  ft, Hitchcock 7199 (US); mountains north of Autlán, 3–5 miles above Mina San Francisco, 1600–1800 m, McVaugh 19651, 19578 (US); about 7 miles south of lagos de Moreno, 2040 m, 13 September 1953, Reeder & Reeder 2300 (MEXU, US); ca. 7 mi SE of the junction of Hwy 80 and 45 at Lagos de Moreno, 1800 m, 21 August 1975, Davidse & Davidse 9927 (MEXU, MO). México: 3  km al W de San Cristóbal Ecatepec, base de la Sierra de Guadalupe, 2400  m, 18 August 1974, Rzedowski 32145 (MEXU). Michoacán: 1.7 mile NW of Tuxpan, 1780 m, Davidse 9806 (MO); ca. 16 miles N of the junction of highways 37 and 15, Davidse 9912 (MO); vicinity of Morelia, Arséne 2417 (GH, MEXU, MO, NY, US), 2864 (MO, US), 2985 (GH, MO, US); about 16 km WSW of Morelia, km 329 on Hwy. 15 (Morelia to Quiroga), Cerro del Aguila, 2000 m, 11 September 1962, Ugent & Flores 1893 (SI); SSW of Morelia on the road to Villa Madero, 2000 m, 19 July 1963, Ugent & Flores 5801 (SI); 25 km east of Morelia on Mex 15, 2000  m, 20 October 1976, Brunken & Perino 444 (MEXU); 10 km al S de Morelia, sobre el camino directo a Atécuaro, 2300 m, 17 August 1990, Rzedowski 49933 (ANSM). Morelos: Barranca de Atzingo, 1900– 2000  m, 7 March 1972, Vázquez 3592 (MEXU). Oaxaca: Hacienda Aguilar, 1580 m, 23 July 1919, Conzatti 3603 (MEXU, US); 13.5 mi. SW of Sola de Vega along the road to Puerto Escondido, 1920 m, 14 August 1975, Davidse 9657 (MO). Puebla: Tehuacán, 5500  ft, Hitchcock 6063 (MEXU, US). Querétaro: near San Juan del Río, 18 August 1905, Rose et al. 9555 (MEXU, NY, US). San Luis Potosí: Las Canoas, 20 July 1910, Hitchcock 5756 (US); in and along open border of oak forest, 4 miles east of Ciudad de Maíz, Gould 10796 (US); near the village of San Francisco in the Sierra de San Miguelito about 25 kms. SW of San Luis Potosí, 2200–2400  m, Sohns 1066 (US); 8  km al W de San Lorenzo, km 62 carretera

Key to Ungrouped Panicum Sections and Species

173

S.L. Potosí-A. Morelos, 1800 m, Rzedowski 565 (US); W de Villa Hidalgo, 1850 m, 22 August 1954, Rzedowski 3775 (US); 3 km al W de Tierra Nueva, 7 September 1954, Rzedowski 796 (US); between Santa María del Río and Tierra Nueva, in the Sierra de Alvarez, 1700–1800  m, 7 September 1954, Sohns 1114 (US). Sonora: Valle de Teras, near La Angostura, White 3542 (GH). Zacatecas: Zacatecas, Hitchcock 7510 (US); Zacatecas, road to Huejuquilla el Alto, 1 mile west of the road-junction 18  miles south of Valparaíso on the road to Mezquitic, McVaugh 17705 (US); Chapingo, km 846 carretera Juárez, 26 km al N de Fresnillo cerca de Tropico de Cáncer, 2150 m, 12 September 1955, Hernández & Mathus 1667 (US) Guatemala. Guatemala city, 1 Dic 1911, Hitchcock 9014 (US) 37. Panicum magnispicula Zuloaga, Morrone & Valls, Iheringia, Bot., 42: 5, Figs.  9–17, 26–27. 1992. Type: Brazil. Santa Catarina: Água Doce, campo graminoso em encosta íngreme com afloramentos, 7.2  km ao sul da divisa Paraná-Santa Catarina ao longo da rodovia BR-153, 1160 m, 3 December 1987, J. F. M. M. Valls, Gomes & Silva 11521 (holotype, CEN; isotype, SI 002810). Fig. 42 Plants perennial, cespitose, shortly rhizomatous, with erect floriferous culms 20–30  cm tall, simple, without aerenchyma; internodes cylindrical, hollow, glabrous, nodes glabrous. Sheaths striate, longer than the internodes, without aerenchyma, hirsute, with whitish tuberculate hairs, upper margins long-ciliate. Ligule membranous-ciliate, reduced, ca. 0.5  mm long. Blades linear-lanceolate, 10–12 × 0.2 cm, base rounded and apex long attenuate, hirsute, with long tuberculate hairs, to hispid, margins ciliate, involute. Inflorescences terminal, exserted; panicles contracted, few flowered, 3–6 × 1 cm, with ca. 7–12 spikelets per panicle; peduncles ca. 25 cm long, cylindrical, glabrous; main axis and axis of branchlets wavy, scabrous, 3–4 first-order branches per panicle, the lower ones 3  cm long, distant and divergent from the main axis, pulvini pilose; pedicels claviform, scabrous and with long hairs near the spikelet insertion. Spikelets long ellipsoid, 5.5– 5.8 × 1.8–2 mm, solitary, adpressed on the axes, glabrous, greenish or tinged with purple, the upper glume and lower lemma subequal and exceeding the upper anthecium in length, apex attenuate; lower glume 3.5 mm long, 3/4 or more the length of the spikelet, acute, 7–9 nerved, the midrib scaberulous toward the upper portion, separated from the upper glume by a conspicuous internode 0.5 mm long; upper glume acuminate, 9 nerved, not deciduous at maturity, shortly hairy toward the apex on the inner surface; lower lemma acuminate, 9 nerved, violaceous toward the upper portion; lower palea elliptic to lanceolate, reduced, 2.5–2.8 × 1–1.2 mm, hyaline, glabrous, ciliate at margins to glabrous; absent; upper anthecium long ellipsoid, 3.4–3.6 × 1.5–1.7 mm, glabrous, crustaceous, pale, shiny, papillose, with papillae regularly distributed over the entire surface of upper palea and in lesser numbers on lemma, with a circular brown ring at the insertion zone with the rachilla; lemma 7 nerved; palea with simple papillae over the entire surface. Caryopsis ellipsoid 2.2 × 1.3 mm, pale; hilum oblong, embryo slightly less than 1/2 the length of the caryopsis.

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Taxonomic Treatment

Fig. 42  Panicum magnispicula. (a) Plant. (b) Spikelet, ventral view. (c) Spikelet, lateral view. (d) Lower glume. (e) Upper glume. (f) Lower palea. (g) Upper anthecium, ventral view. (h) Caryopsis, hilum view. (i) Caryopsis, embryo view. (From Valls et al. 11521, SI)

Key to Ungrouped Panicum Sections and Species

175

Distribution and ecology (Fig. 34). Species is only known from the state of Paraná, Brazil, known only from the type collection, where it inhabits fields in the NW of the state, at 1160 m asl. 38. Panicum miliaceum L., Sp. Pl. 1: 58. 1753. Milium paniceum Mill., Gard. Dict. (ed. 8): n 1. 1768, nom. illeg. superfl. Milium esculentum Moench, Methodus: 203. 1794, nom. illeg. superfl. Panicum milium Pers., Syn. Pl. 1: 83. 1805, nom. illeg. superfl. Leptoloma miliacea (L.) Smyth, Trans. Kansas Acad. Sci. 25: 86. 1913. Type: India, “Habitat in India” (lectotype, LINN-80.49, designated by A. S. Sherif & M. A. Siddiqi, Fl. Lybia 145: 282. 1988). Fig. 43 Annual, cespitose, herbaceous plants, 15–70 cm tall, culms decumbent, branching or not at the lower nodes, to erect, branching at the upper nodes; internodes cylindrical, hollow, hairy, 2–12 cm long; nodes dark, covered with whitish hairs. Sheaths 4–12  cm long, larger than the internodes, striate, herbaceous, densely pilose, with deciduous tuberculate hairs. Ligule 1–1.3 mm long, shortly membranous at base, then long ciliate. Blades oblong-lanceolate, 6–25 × 0.4–1.2 cm, flat, dark green, subcordate at base, with a conspicuous midrib and membranous margins, scabrous, pilose on both surfaces, the lower margins ciliate with tuberculate hairs. Inflorescences lax, multiflowered, nutant, 6–20  ×  4–11  cm, included when young in the upper leaves; peduncles cylindrical, pilose; main axis triquetrous, scabrous, undulate, with or without long whitish hairs toward the basal portion; branches alternate, rarely subopposite, divergent from the main axis and naked in the basal portion, spikelets solitary on long pedicels; pulvini glabrous, pale; pedicels scabrous, sparsely hairy, 4–9 mm long; axillary panicles similar to the terminal one. Spikelets ovoid, 4.6–5.5 × 1.5–2.1 mm, glabrous, pale and tinged with purple, upper glume and lower lemma subequal, with manifest nerves and exceeding the upper anthecium in length; lower glume 2.8–3.6 mm long, 1/2 to 3/4 the length of the spikelet, 5–7 nerved, the midrib scaberulous in the upper portion, apex attenuate; upper glume 4–5.1 mm long, 11–13(–15) nerved, the midrib scaberulous; lower lemma 4–4.8 mm long, 11–13 nerved, glumiform; lower palea ovate, reduced, 1.2– 1.6 × 0.5–0.8 mm, membranous, with a bilobed apex; lower flower absent; upper anthecium ellipsoid, 2.7–3.4 × 1.3–1.9 mm, reaching 2.3 mm when mature, smooth, glabrous, shiny, whitish, yellowish at maturity, with two basal scars ca. 0.4  mm long; lemma 7 nerved; palea with verrucose papillae next to the apex. Caryopsis 2–2.3  ×  1.4–1.8  mm, whitish; hilum punctiform, embryo ca. half the length of caryopsis. Distribution and ecology (Fig. 44). Species native to the Old World, where it has been cultivated since ancient times. Introduced in America, where it is grown for bird food, as a small cereal, or for summer grazing; it is prized for its earliness and resistance to drought (Palacios 1969). Local name: “Millet” Additional material examined. USA. California: Butte Co, on the S edge of Pentz Road, about 1 mile E of Highway 99, 10 August 1988, Ahart 6168 (MO). Florida: Pensacola, 8 August 1901, Curtiss 6867 (MO). Massachusetts: Dump Valley Road, Dorchester, 10 September 1916, Churchill 7 (MO). Mississippi: ca.

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Taxonomic Treatment

Fig. 43  Panicum miliaceum. (a) Inflorescence. (b) Spikelet, lateral view. (c) Upper anthecium, ventral view. (d) Upper anthecium, dorsal view. (From Navarro s.n., BAB)

Key to Ungrouped Panicum Sections and Species

177

Fig. 44  Distribution of Panicum miliaceum and P. venosum

4 miles N of Biloxi along highway to Hattiesburg, 24 July 1971, Lasseigne 2812 (MO). Missouri: Greenwood, 19 July 1932, Kellogg s.n. (MO 2215101); St. Louis, Lindenwood freight yard of the Frisco Railroad, 3 July 1955, Muelhenbach 667 (MO). New York: Ithaca, 21 July 1916, Metcalfe 5511 (MO). North Dakota: 18 miles S and 15 miles W of Fargo, 17 August 1971, Seiler 3825 (MO). Utah: North Park, Provo, Mengies 8023 (MO). Vermont: Chittendent Co, Flagstaff, in yard of house at Rt. 3, 66 Leisure Lane, 29 August 1983, Ricketson 677 (MO) Honduras. Francisco Morazán: El Zamorano, 800 m, Rodríguez 3512 (US) Dominican Republic. Santo Domingo: Santo Domingo, Botanical Garden, Liogier 21371 (NY) Colombia. Valle: Palmira, Estación Agrostológica, Villamizar-Jaramillo s.n. (COL-44878) French Guiana. Station des Nouragues, Bassion de l’Arataye, 31 August 1987, Feuillet 4442 (US); Chemin Hilaire-Anse de Montabo-Ile de Cayenne, 1 February 1987, de Granville 9138 (MO, US) Brazil. Distrito Federal: Plano Piloto, SQN 76, Filgueiras 1204 (IBGE). Rio Grande do Sul: São Leopoldo, Rambo 1937 (BAA). Santa Catarina: Rosario, 4 January 1964, Reitz & Klein 17008 (US). São Paulo: Pinheiros, 22 February 1934, Gehrt 31509 (US)

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Taxonomic Treatment

Argentina. Buenos Aires: Bahia Blanca, Spegazzini s.n. (BAA); Punta Lara, 24 March 1984, Zuloaga 1890 (SI); San Isidro, Pastore 849 (SI); Olivos, Dawson 713 (LP); Tigre, Lanfranchi 453 (SI), 481 (LP). Córdoba: Ascochinga, Giardelli 616 (BAA, SI); Villa María, 11 April 1938, Navarro s.n. (BAB); Villa Carlos Paz, 31 December 1935, Burkart 7197 (SI); Santa María, Alta Gracia, Alberdi 12269 (LP); Tercero Arriba, Río Tercero, 4 March 1943, Burkart 13325 (SI). Corrientes: Corrientes, Quarín 1321 (CTES). Entre Ríos: Concepción del Uruguay, 11 April 1963, Burkart 24109 (SI); Victoria, February 1934, Báez s.n. (BAA). La Pampa: Telen, Monticelli 168 (SI). La Rioja: Chilecito, Dawson 3355 (BAA). Mendoza: Godoy Cruz, Ruíz Leal 5576 (BAA, LIL); San Rafael, Dawson & Pujals 1702 (BAB); ruta 7, km 1004 al N de Santa Rosa, 26 February 1981, Nicora et al. 8361 (SI). Salta: Salta, Chachapoyas, 14 January 1989, Novara 8436 (M). Santa Fe: Rosario, Isla del Rowing, 6 January 1938, Christian s.n. (SI). Tucumán: Capital, calle lateral del Hipódromo, Türpe 3009 (LIL) 39. Panicum mucronulatum Mez, Bot. Jahrb. Syst. 56, Beibl. 125: 2. 1921. Type: Brazil. Bahia: without locality, J. Blanchet 104 (lectotype, B_10_0158002, designated by F. O. Zuloaga & O. Morrone, Ann. Missouri Bot. Gard. 83(2): 262. 1996; isolectotype, US 00139772, fragment ex B; probable isolectotype, NY). Fig. 45 Panicum virgatum L. var. pilosum Döll, Fl. Bras. 2(2): 218. 1877. Syntypes. Brazil. Bahia: Ilheus, J.  Blanchet 2961 (lectotype, BR 0000006883133, here designated; isolectotypes, G 00099728, G 00099729, US 00139771, fragment ex P). Pará: Almeirim, C. F. P. P. von Martius s.n. (n.v.) Annual, herbaceous, cespitose plants, with erect culms 60–100 cm tall, branching at the lower nodes, many noded; internodes compressed, hollow, with long whitish, adpressed hairs or with erect tuberculate hairs; nodes dark, compressed, pilose. Sheaths usually larger than the internodes, pale, covered with rigid, fragid and urticant tuberculate hairs, the margins ciliate. Ligule membranous-ciliate, 1.4–1.6 mm long, arcuate; collar densely pilose, covered with tuberculate hairs. Blades lanceolate, 25–45 × 1–1.5 cm, flat, hirsute, base subcordate, the apex acuminate to subulate, margins ciliate to scabrous, midrib conspicuous. Inflorescences terminal, exserted; panicles lax, diffuse, pyramidal, multiflowered, 25–40  ×  10–30  cm; peduncles partly subincluded in the upper leaf sheaths, hispid toward the distal portion; main axis wavy, hirsute in the basal portion, then scabrous, pulvini glabrous, first-order branches alternate, rarely opposite, divergent, axis of the branches hirsute to scabrous; pedicels triquetrous, scabrous, 2–20 mm long. Spikelets solitary, ovoid, 3.3–3.6(−3.9) × 1.3–1.6 mm, glabrous, greenish or tinged with purple, glumes and lower lemma with manifest nerves, subequal, exceeding 1 mm in length the upper anthecium; lower glume 1.5–2  mm long, 1/2 the length of the spikelet, acute, (3–)5–7 nerved, the midrib scaberulous, the nerves anastomosed toward the apex; upper glume acute, 2.8–3  mm long, 9–13 nerved; lower lemma glumiform, 3–3.3 mm long, 7–9 nerved; lower palea elliptic, 2.3–2.6 × 0.9–1.1 mm, membranous, whitish, glabrous, as long as the upper anthecium, margins and apex sparsely hairy; upper anthecium ovoid, 2.2–2.4 × 1–1.2 mm, glabrous, pale, smooth, shiny,

Key to Ungrouped Panicum Sections and Species

179

Fig. 45  Panicum mucronulatum. (a) Plant. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Lower palea. (e) Upper anthecium, dorsal view. (f) Upper anthecium, ventral view. (From Pinto 121, CEPEC)

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Taxonomic Treatment

crustaceous, with two basal scars ca. 0.2  mm long; lemma 7 nerved; palea with globose microhairs and simple papillae irregularly distributed at its apex. Caryopsis broadly ellipsoid, 1.6 × 1.1 mm; hilum punctiform, embryo half the length of the caryopsis. Distribution and ecology (Fig. 38). Species restricted to northeastern Brazil, where it grows in the states of Alagoas, Bahia, Paraiba, and Pernambuco; found along stream banks or near sugarcane fields, between 25 and 200 m asl. Local name: “Capim-de-orvalho” (Brazil) In describing Panicum virgatum var. pilosum, Döll (1877) includes two syntypes of the species from Brazil, Blanchet 2961 (from Bahia) and Martius s.n. (from Pará). Of these, the BR specimen, Blanchet 2961, was selected as the lectotype because it is a complete specimen and agrees with the protologue of the variety. Panicum mucronulatum is an annual plant, cespitose, with lanceolate, hirsute laminae, up to 45 cm long, lax, broad terminal panicles, with spikelets 3.3–3.9 mm long. Additional material examined. Brazil. Alagoas: Porto Calvo, Fazenda Porto Seguro, Campelo 2154 (CEN). Bahia: Dom Macedo Costa, Fazenda Mocambo, 5 March 1985, Noblick & Lemos 3955 (CEPEC); Mun. Santo Antonio de Jesus, BR-101 Cruz das Almas-Itabuna, km 44, Coradin et al. 3394 (CEN), 200 m; Cruz das Almas, Pinto 121 (CEPEC), 306 (IAN, US); Cachoeira, 25–50 m, coarse tufts, ner streamlet, brushy border of sugar cane field, 21 December 1924, Chase 8105 (NY, RB, US), 8105bis (US); Municipio São Sebastião do Passé, Area da Estaçao Experimental Miranda, km 62 da Rod. BR 324, Hage et  al. 1716 (CEPEC, K). Paraiba: Escola do Agronomia do Nordeste, 25 May 1953, Coelho de Moraes 677 (IAN, SP, US). Pernambuco: Recife, Sendulsky 1434A (SI, SP); Tapera, April 1928, Pickel 1393 (US); Near Vitoria, W of Recife, Davis & Andre-Lima 61068 (IBGE); without locality, Gardner 1838 (K), Glocker 208 (US) 40. Panicum parcum Hitchc. & Chase, Contr. U.S. Natl. Herb. 15: 68, Fig. 53. 1910. Panicum decolorans Kunth var. parcum (Hitchc. & Chase) Beetle, Phytologia 54(1): 4. 1983. Type: Mexico. Sinaloa: Lodiego on the Culiacán River, mountain side, not very common, 9/15 October 1891, E.  Palmer 1657 (holotype, US 00147940; isotypes, F 0046829F, ISC-v-0000577, LIL 000087, NY 00381770, P 00740972, S14-5636) Plants annual, cespitose, with erect to geniculate culms, 20–90 cm tall, simple, few noded, innovations extravaginal; internodes 5–20 cm long, 2 mm in diameter, cylindrical, glabrous to hirsute; nodes glabrous, brownish. Sheaths 10–12 cm long, usually shorter than the internodes, hirsute with tuberculate hairs, the margins ciliate. Ligule shortly membranous at base then long ciliate, 1–1.2  mm long; collar hirsute, brown. Blades linear-lanceolate, 6–45  ×  0.4–0.7  cm, flat, acute, hirsute, with deciduous tuberculate hairs ca. 1 mm long, the base narrow, margins scabrous, ciliate toward the base, midrib manifest. Inflorescences terminal, exserted, 7–30 × 6–15 cm, lax, diffuse; peduncles subincluded in the upper leaf sheaths to exserted, 10–35 cm long, hirsute in the distal portion; main axis wavy, scaberulous, glabrous to hirsute toward the base; pulvini purplish, glabrous, first-order branches

Key to Ungrouped Panicum Sections and Species

181

alternate, divergent, the lower ones 6–15 cm long, pedicels triquetrous, 5–18 mm long, scaberulous, glabrous. Axillary inflorescences present, similar to the terminal panicle. Spikelets solitary, ovoid, 4.5–6 × 1.8–2 mm, glabrous, greenish or tinged with purple, stipitate; upper glume and lower lemma subequal, 1.2–1.8 mm longer than the upper anthecium; lower glume 3–4.2 mm long, 3/4 the length of the spikelet, acute, 5–7 nerved, midrib scabrous, separated by a distinct internode from the upper glume ca. 0.6 mm long; upper glume 3.9–5.4 mm long, 9–11 nerved, midrib scabrous, shortly scabrous on the inner surface; lower lemma acute, 3.6–4.8 mm long, 9–11 nerved, scabrous on the inner surface; lower palea lanceolate, reduced, 1.2–1.8 × 0.4–0.6 mm, 3/4 the length of the spikelet, membranous, glabrous; lower flower absent; upper anthecium ovoid, 2.7–3.3 × 1.5–2 mm, crustaceous, glabrous, smooth, shiny, pale, blackish tinged at maturity; lemma 7 nerved; palea with verrucose papillae at the apex. Caryopsis ovoid, 2.3–2.5 × 1.4–1.8 mm, whitish; hilum punctiform; embryo half the length of the caryopsis. Distribution and ecology (Fig. 36). It is found from Mexico to Costa Rica, in open rocky terrain or pine forests, between sea level and 2070  m elevation, and flowers between July and October. Iconography. Hitchcock & Chase, Contr. U.S. Natl. Herb. 15: Fig. 53. 1910 Panicum parcum differs from P. decolorans in having lax panicles with branches divergent and stipitate spikelets, with the lower and upper glumes separated by a distinct internode. Additional material examined. Mexico. Chiapas: 9 km southwest of Cintalapa, 8 October 1968, Gould 12757 (GH, US); 14 km E of Ocozocuatla, in open grassy field, 6 October 1968, Gould 12729 (MO, US); near junction at Las Cruces on road to La Ciénaga, 853 m, 18 September 1981, Breedlove 52871 (MO); small forested stream and adjacent savanna 2 km SE of Arriaga, 100 m, 27 August 1974, Breedlove 36875 (MEXU, MO); 9 km north of Tuxtla Gutierrez, along road to El Sumidero, 2500 ft, 27 October 1965, Breedlove & Raven 13492 (F, US), 13833 (F, MEXU, US); 10 km east of Arriaga on Mexican Highway 200, 31 October 1981, Breedlove & Davidse 54261 (MEXU, MO); western side of Tuxtla Gutierrez, 13 September 1965, Gould 11676 (US). Chihuahua: Río Negro, 28 August 1937, Le Sueur 173 (US). Colima: Alzada, 21 September 1910, Hitchcock 7081 (RB, US). Durango: 14 mi SE of Durango, about 600  ft, 16 October 1966, Gould 12288 (US). Guanajuato: km 20 N de Silao, 16 September 1946, Hernández Xolocotzi et al. X-2468 (US). Guerrero: Balsas, 9 September 1910, Hitchcock 6782 (F, MO, NY, US), 6808, 6810, 6811 (US); Coyuca, Santa Barbara, 29 September 1934, Hinton 6686 (GH, MO, NY, US); along road from Almoloya-Teloloapan-Iguala, 5900– 9000 ft, 10 October 1953, Sohns 960 (US); Cañón de La Mano, entre Los Amates y El Naranjo, 10  km al N de Iguala por el ferrocarril, 900–1000  m, 12 September 1986, Catalán & Terán 92 (MO). Jalisco: about 13  miles northwest of León, Guanajuato in a grassy area with scattered thorny shrubs, 2070 m, Reeder & Reeder 2288 (MEXU, US); steep rounded hills ca. 8 miles southwest of La Resolana, road to La Huerta, McVaugh 21107 (US). Michoacán: West-facing slopes of Cerro de Carboneras, ca. 22  km  S of Uruapan, 3300–3700  ft, King & Soderstrom 4831 (MEXU, NY, US); on steep slopes of barrancas on road from Uruapan to Nueva

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Taxonomic Treatment

Italia, Sohns 834, 836 (US); along road from Tsitzio-Tiquicheo-HuetamoAltamirano, Sohns 905, 922, 945 (US). Morelos: Xochiltepec, 24 September 1938, Lyonnet 2639 (MEXU, MO, US). Nayarit: Nayar, 8 km al NW de San Juan Peyotán, en los límites de Nayarit y Durango, 22°30′N, 104°30′W, 600  m, 21 September 1989, Flores. & Tenorio 1197 (MEXU, MO); Mun. Nayar, 3.9 km al NE de Jesús María, camino a Huejuquilla, 22°16′N, 104°30′W, 530 m, selva baja caducifolia, 15 September 1989, Flores et al. 1053 (MEXU, MO); Mun. Nayar, 7 km al NW de San Juan Peyotan, brecha a Huazamota, 22 26 N, 104 29 W, 600 m, 21 September 1989, Tenorio & Flores 16264 (MEXU). Oaxaca: 11 mi. W of Zanatepec, 150  m, 28 August 1953, Reeder & Reeder 2152 (MEXU, US); 40 miles NE of Juchitan, in moist opening in brush-forest area, Gould 11669 (US); 151 km SW of Oaxaca, on highway 190, 1100 m, 15 October 1976, Brunken & Perino 368 (MO); 10 miles W of Tehuantepec on Mex 190, 240 m, 14 October 1976, Brunken & Perino 360 (MO); near km post 184 on Hwy. 190, ca. 40 miles NW of Tehuantepec, 710 m, 13 August 1975, Davidse 9587 (MO); about 42 miles west of Tehuantepec on grass-covered slopes, 760 m, Reeder & Reeder 2164 (MEXU, US). Puebla: Mesa del Pochote 3 km al E de El Salado, bosque de Quercus, 1310 m, 20 October 1985, Guízar 1850 (ANSM, CHAPA). Sinaloa: Cofradía, vicinity of Culiacán, 31 October 1904, Brandegee 4 (US); Las Mesas, Sierra Surotato, ca. 3000  ft, 15 September 1941, Gentry 6667 (GH, MEXU, MO, NY), Harvey 6667 (MO). Temascaltepec, Bejucos, Hinton 1996 (GH, NY), 2000 (NY, US). Without state: Alzada, 4 October 1910, Orcutt 4687 (US) Guatemala. Chiquimula: grassy plains and low slopes around Chiquimula, 400 m, 20 October 1939, Steyermark 30064 (F, US). Jutiapa: Región of El Tablón, northeast of Jutiapa, 850–900 m, 31 October 1940, Standley 75925 (F, US). vicinity of Jutiapa, 850  m, 24 October 1940, Standley 75102a (F, US). Zacapa: near La Fragua, 200–500 m, 14 October 1940, Standley 74761 (F, US); rocky hills in vicinity of Santa Rosalía, 2 mi. south of Zacapa, 200 m, 7 October 1939, Steyermark 29310 (F); lower slopes of Sierra de las Minas, along trail above Río Hondo, 250– 900 m, 11 October 1939, Steyermark 29053 (F) Honduras. El Paraíso: Road to Yuscarán, edge of brushy places, 800  m, 5 November 1951, Swallen 11371 (MO, US); road to Danlí, not far from Río Choluteca, Swallen 11250 (US). Francisco Morazán: región of Las Mesas, pine hills around the Zamorano valley, rocky pine woods, 800–900 m, 2 November 1951, Swallen 10765 (MO, US), 2 November 1954, 11254 (F, MEXU, MO, US); San Antonio del Oriente, 1200 m, Swallen 10909 (US) Nicaragua. León: along Hwy. 12, 2 km SE of Jct. with Puerto Somoza road, 40 m, 8 July 1971, ca. 12 km E of Puerto Somoza, Pohl 12707 (F, MO, NY); El Velero, at mouth of Estero San José, ca. 10 km by road S of Hwy. 32, 12°08′N, 86°45′W, 9 September 1984, Stevens 23096 (MEXU, MO, SI). Managua: Inst. Pedagogico de Varones, Vernier 1505 (GH) Costa Rica. Guanacaste: South end of Playas del Coco, 14 November 1968, Pohl & Davidse 11437 (F, US)

Key to Ungrouped Panicum Sections and Species

183

41. Panicum peladoense Henrard, Blumea 4(3): 504. 1941. Type: Paraguay. Paraguarí: Cerro Pelado, 3 April 1883, B. Balansa 4357 (holotype, L 0044832; isotypes, G 00030394, G 00030395, G 000097241, K 000307787, MO-2114103, L 0044831, L 0044833, P 00371645, P 00371646, P 00371647, US 00147949, US 00147947, US 00147948). Fig. 46 Panicum campestre Nees, Fl. Bras. Enum. Pl. 2(1): 197. 1829, nom. illeg. hom., non P. campestre Nees ex Trin., 1826. Panicum. cayennense Lam. var. campestre (Nees) Pilg., Bot. Jahrb. Syst. 30(1): 132. 1901. Type: Brazil. São Paulo: between Taubaté and Pindamonhangaba, C. F. P. P. von Martius s.n. (holotype, M; isotypes, US 00148207, fragment and photo ex M, US 00406127, fragment ex M) Panicum bergii Arechav. var. leiophyllum Hack. & Lindm., Kongl. Svenska Vetensk. Acad. Handl., n.s., 34(6): 10, table 4b, 1900. Type: Brazil. Rio Grande do Sul: Cachoeira, 11 February 1893, C.  A. Lindman 1185 (holotype, S12-2548; isotypes, B_10_0249080, P 00740773, US 00323770, W 19160024882) Plants perennial, cespitose, shortly rhizomatous, 20–60 cm tall; culms erect, few noded; internodes 3.5–10.5 cm long, cylindrical, hollow, densely hispid to glabrescent; nodes purplish and with whitish hairs densely to sparsely arranged. Sheaths 3.5–8 cm long, usually smaller than internodes, pale, densely pilose, with adpressed tuberculate hairs to glabrescent, the margins ciliate. Ligule shortly membranous then long ciliate, 0.5–1.7 mm long; collar light brown, pilose. Blades lanceolate to linear-lanceolate, 7–22 × 0.2–0.4 cm, erect, ascending, flat, the base narrow, continuing imperceptibly with the sheath, densely pilose, with tuberculate hairs on both surfaces or only on the abaxial surface to glabrous, margins scaberulous and long ciliate, cartilaginous. Inflorescences lax, diffuse, pyramidal, 7–18 cm × 3–13 cm; peduncles subincluded in the upper leaf sheaths to exserted, up to 30 cm long, cylindrical, glabrous; main axis cylindrical, scabrous, with or without long sparse hairs, first-order branches alternate or opposite, sometimes subverticillate toward the base, axes of the branches scabrous to pilose, pulvini glabrous; pedicels claviform, scabrous, 2–20  mm long. Spikelets long ovoid, acuminate, (2.5–)2.7– 3.3(−3.7) × 1–1.2 mm, solitary, glabrous, pale and tinged with purple; lower glume ovate, acute, (1.5–)1.7–2.1(−2.4) mm long, 1/2 to 3/4 the length of the spikelet, 5–7 nerved, the midrib scaberulous toward the upper portion; upper glume (2.3−)2.6−3.1(−3.4) mm long, 7–9 nerved, deciduous at spikelet maturity and exposing the upper anthecium, the inner surface scaberulous toward the apex; lower and upper glume separated by an internode ca. 0.3 mm long; lower lemma glumiform, 2.5–3(–3.4) mm long, 7–9 nerved; lower palea elliptic, 1.7–2.2 × 0.6–0.9 mm, membranous, hyaline, the margins finely hairy to glabrous, apex bilobed to entire; upper anthecium ellipsoid, (1.7–)1.9–2.1(−2.3) × 0.8–1.1 mm, 1–1.2 mm shorter than the upper glume and lower lemma, crustaceous, glabrous, glabrous, smooth, shiny, pale, black at maturity, with two basal scars ca 0.1 mm long; lemma 7 nerved; palea with simple papillae toward the apex. Caryopsis 1.2–1.3 × 0.6–0.9 mm; hilum punctiform, embryo 1/3 the length of the caryopsis.

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Taxonomic Treatment

Fig. 46  Panicum peladoense. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Mature upper anthecium, ventral view. (i) Caryopsis, embryo view. (j) Caryopsis, hilum view. (From Zuloaga & Deginani 137, SI)

Key to Ungrouped Panicum Sections and Species

185

Distribution and ecology (Fig. 38). It inhabits fields in Argentina, Bolivia, Brazil, Paraguay, and Uruguay, reaches up to 1200 m asl, and flowers from October to March. Local name: “Kapi’í tî” (kapi’í = straw, tî = clear) (Zuloaga 1994) Henrard (1941) mentions, in the original description of the species, that it has glabrescent or sparsely pilose sheaths, nodes, and blades. The character of the pilosity has been found to be variable, ranging from densely pilose to glabrescent specimens. This species was described by Nees in 1829 as P. campestre, this name being invalidated by the existence of an earlier homonym described by Trinius in 1826, the latter species belonging to section Rudgeana (Zuloaga et  al. 2018). Henrard (1941) erroneously states that P. campestre described by Nees in 1829 corresponds to P. ghiesbreghtii. A related species to P. peladoense is P. exiguum Mez, the latter species being distinguished by being an annual plant, with culms abundantly branched and decumbent at the base, with the lower glume 3/4 or more the length of the spikelet. Panicum peladoense is separated from P. chaseae because the latter taxon includes plants with contracted panicles, with broadly ellipsoid spikelets and brownish upper anthecium at maturity, without a deciduous upper glume. Additional material examined. Brazil: Distrito Federal, Reserva Ecologica do IBGE, Filgueiras 1060 (SI, SP); Area da FAL, 5 May 1991, Azevedo & Filgueiras 950 (MO). Goiás: between Viannápolis and Ponta Funda, open clay, sandy cerrado, Chase 11280 (US). Mato Grosso: Santa Rita do Araguaya, on Rio Araguaya, Chase 11858 (US). Mato Grosso do Sul: Mun. Dourados, some 20 km E of Itaúm, on road to Dourados, Pedersen 11109 (K, MO); Campo Grande, 7 February 1930, Chase 10780, 10786 (US), Fazenda Experimental de Criaçao, 27 March 1946, Swallen 9308 (US); Ponta Porá, Swallen 9427 (US); Tres Lagoas, 300 m, 4 February 1930, Chase 10723 (NY, RB, US); between Campo Grande and Dourados, 400–500  m, 14-17 February 1930, Chase 10886 (F, US). Minas Gerais: Farias, Serra da Bocaina, Chase 10534 (US); 30 km SW of Campanha, along highways 267 and 381 to São Paulo, 850 m, 25 February 1976, Davidse et al. 10618 (MO, SP); Serra do Cipó, 110  km NE of Belo Horizonte, W of Vaccaria, 800  m, 28 March 1925, Chase 9279 (F, GH, MO, NY, US); Poços de Caldas, 1100–1500 m, Chase 10665 (US); 13  km E of Lavras along highway 265 to Barbacena, 970  m, 27 February 1976, Davidse et  al. 10712 (MO, SP); Serra do Mantiqueira, between Sitio and Dr. Sá Fortes, 1200 m, 1 March 1925, Chase 8696 (F, GH, K, MO, NY, US); between Sucupira and Omega, S of Uberlandia, 700–900 m, leaning, sandy clay campo, 14 March 1930, Chase 11207 (US). Pará: Lagoa Aura Preta, Drouet 1953 (F). Paraná: Campo Morào, Swallen 8964 (US); Mun. Arapoti, Campo Cerrado by Jaguariaíva boundary, 17 January 1965, Smith et al. 14726 (K, NY, P, RB, US); Jaguariaíva, Swallen 8682 (US); Ponta Grossa, Estaçao Experimental, Swallen 8311 (US); Ponta Grossa, Campo, Fazenda de Criaçao, Swallen 8771 (US); Mun. Chapada Santo Antonio, Jaguariaíva, Dombrowski 11096 (K); Rodovia BR-277, prox. ao Rio Capivari, Mun. Palmeira, Hatschbach 43533 (MO). Rio Grande do Sul: Morro Santa Teresa, Porto Alegre, Clayton 4366 (K); Don Pedrito,

186

 

Taxonomic Treatment

15 April 1946, Swallen 9097 (US); Estaçao Experimental Agronomica do UFRGS, 9 February 1972, Valls 1929 (CEN, SI); Alto do Morro da Gloria, Porto Alegre, Valls & Arzivenco 2033 (CEN), Valls 1645 (CEN); Morro Santa Teresa, Porto Alegre, 11 September 1949, Rambo 43391 (LIL, US), 43692 (US); Vacaria, Swallen 8203 (US); Cachoeira, Exp. I. Regnell, Lindmann A-1187 (US); Pelotas, 22 April 1946, Swallen 9156 (US). Santa Catarina: 17 km NE of the S. Catarina-RGS border, BR-116, 11 March 1976, Davidse et al. 11145 (MO, NY, SP). São Paulo: Casa Branca, 700 m, sandy clay campo, 16 January 1930, Chase 10589 (K, US); Ypiranga, Chase 10697 (US); Jundiahy, Brade 8144 (IAN, R); Fazenda Campininha, 10 km NEE of Padua Sales, 22°11′S, 47°10′W, 15 July 1960, Eiten et al. 2116 (F, G, NY, US); São Paulo, Pickel 5154, 5795 (US); Mun. Itapetininga, Sorocava to Itapetininga, 4 February 1965, Clayton 4507 (K, NY, US); Mun. Itapetininga, Faz. Sta. Luzia do Campo Largo, SSE da cidade do Itapetininga, Machado de Campos 209 (NY, US); Botucatu, Rubiao Junior, 12 March 1980, Gemtohujnicov s.n. (ex SP-179335 en SI). Without state: without locality, Riedel 2155 (K), Burchell 6794-2 (K), Sellow s.n. (K) Bolivia. Santa Cruz: Florida, 10 km E of Samaipata, Renvoize et al. 4044 (K); Cordillera, 50 km S of Santa Cruz, Renvoize et al. 3933 (K) Paraguay. Amambay: in altiplanitie Sierra de Amambay, Hassler 12099 (G). Concepción: San Salvador, Rojas 2784 (BAA, US). Guairá: praderas de Itaigú, pres de Villa Rica, 17 December 1876, Balansa 13 (BAF, G, K, P, US) Paraguarí: Cerro Peró, prés de Paraguarí, 29 October 1876, Balansa 14 (G, K, P, US) Argentina. Córdoba: Dpto. Calamuchita, Yacanto Calamuchita, “La Tirelire”, Tirel 156 (P); 7 km W de Sta. Rosa de Calamuchita, Quarín 2909 (CTES, SI, US). Without locality, 9 January 1919, Castellanos 43 (F, SI, US). Corrientes: Dpto. Capital, Ruta 12, Riachuelo, Eskuche 2538- 5 (SI). Rincón de Luna, Pedersen 5812 (BAA, GH, K, P, US); Dpto. Empedrado, Estancia Las Tres Marías, Pedersen 1882 (P, US); El Sombrerito, Estacion Experimental del INTA, Quarín et al. 1969 (CTES, SI); Río Paraná near Arroyo Sombrero, 27 February 1978, Renvoize et al. 3637 (K, MO, NY, P, SI, US); Dpto. Gral. Paz, Lomas Vallejos, Ibarrola 3481 (LIL); Dpto. Ituzaingo, 12 km E de Ruta Nac. 12, camino a San Carlos, Krapovickas et al. 17861 (CTES); Itambé, Bertoni 5532, 5534 (LIL); Estancia El Plata, Meyer 6329 (LIL); Dpto. Mburucuyá, Estancia Santa Teresa, Pedersen 5482 (GH, K, MO, NY, P), Quarín 2866 (SI); Dpto. Saladas, Estancia Rincón del Ambrosio, Schwarz 9492 (LIL); Dpto. San Miguel, 21 km S de Loreto, Schinini et al. 8190 (CTES); Dpto. San Roque, M.  F. Mansilla, Ibarrola 2889 (LIL, NY, P); Dpto. Santo Tomé, Garruchos, Estancia Garruchos, Pedersen 5457 (GH, K, NY); Estancia Rincón de las Mercedes, Pedersen 5428 (NY, P). Jujuy: Dpto. Dr. Manuel Belgrano, Sierra de Zapla, Mina 9 de Octubre, Zuloaga & Deginani 137, 3447 (SI); Dpto. Santa Bárbara, Abra de los Morteros, Cabrera et  al. 22167 (LP). Misiones: Dpto. Apóstoles, Apóstoles, Martínez Crovetto 8978 (BAA). Dpto. Capital, Posadas, Establecimiento Santa Inés, 4 February 1922, Parodi 4512 (US), 4513 (BAA, US); Dpto. Candelaria, Loreto, Montes 11238 (LP); Concepción de la Sierra, 28 January 1926, Parodi 6935 (BAA, US); Dpto. San Ignacio, Colonia Corpus, Schwarz 6499 (LIL, US); Gobernador Roca, Schwarz 6484 (BAA, LIL, MO, US); Campo San Juan, 21 November 1984, Fontana F94-56 (SI)

Key to Ungrouped Panicum Sections and Species

187

Uruguay. Rivera: Estancia Ripol, ruta 5 próximo a ruta 29, Del Puerto 1527 (P). Tacuarembó: Tacuarembó, Gruta de los Cuervos, 8 February 1981, Cabrera & Zuloaga 32385 (SI), 32388 (MO, SI, US) 42. Panicum quadriglume (Döll) Hitchc., Contr. U.S. Natl. Herb. 24(8): 460. 1927. Panicum cayennense Lam. var. quadriglume Döll, Fl. Bras. 2 (2): 220. 1877. Panicum bergii Arechav. var. quadriglume (Döll) Henrard, Meded. Rijks-Herb. 40: 52. 1921. Type: Brazil. Minas Gerais: Caldas, 18 March 1847, A. F. Regnell III-1406 (holotype, W 19040010627; isotypes, BR 0000006882907, P 00371640, S05-9165, S13-12770, S-R-3943, US 00148345, fragment ex BR, US 00133135). Fig. 47 Plants perennial, cespitose, 15–80 cm tall; culms erect, few noded, branching at the middle nodes, internodes 4–10 cm long, cylindrical, covered with tuberculate hairs to glabrescent; nodes pale, covered with long whitish hairs. Sheaths smaller than internodes, 3–9 cm long, densely to sparsely hirsute, with tuberculate hairs, the margins ciliate. Ligule 0.5–1 mm long, base shortly membranous, then ciliate; collar pale, glabrous to hairy. Blades linear-lanceolate, 6–26  ×  0.3–0.6  cm, erect, ascending, rigid, flat, the base narrow and the apex acuminate, densely to sparsely hirsute on both surfaces with deciduous tuberculate hairs, the margins scabrous and ciliate, with deciduous tuberculate hairs. Inflorescences terminal, exserted, lax, diffuse, pyramidal, 7–20  ×  3–13  cm, when young included in the upper sheaths; peduncles subincluded in the upper leaf sheaths to exserted, 8–20 cm long, cylindrical, glabrous; main axis glabrous, triangular and scabrous, first-order branches alternate, rarely subopposite or verticillate toward the top, diverging from the rachis, scabrous; pulvini glabrous; pedicels scabrous, greenish to purplish, 3–13 mm long; axillary panicles present, similar to the terminal inflorescence. Spikelets solitary, long-ovoid, acuminate, 2.5–4.2 × 1–1.4 mm, glabrous, greenish or tinged with purple, triflowered, with two lower lemmas, one naked and the remaining without its corresponding palea, and upper anthecium with its dorsal surface disposed toward the lower glume; lower glume 1.2–2.3 mm long, 1/2 the length of the spikelet, acute to obtuse, 5-9-nerved, the midrib scaberulous toward the upper portion; upper glume 1.9–3.6 mm long, deciduous, exposing the upper anthecium, acute to obtuse, 7–9 nerved, the midrib scaberulous; lower and upper glume separated by a short internode ca. 0.2 mm long; lower lemma I 2.2–3.7 mm long, acuminate, 5–9 nerved; lower palea I absent; lower lemma II 2.2–3.8  mm long, acuminate, 7–9 nerved; lower palea II elliptic, 1.5–2.3 × 0.7–0.9 mm, as long as the upper anthecium, membranous, hyaline, the apex bilobed, shortly ciliate or not towards the upper margins; upper anthecium narrowly ovoid, 1.6–2.3 × 0.8–1.2 mm, 0.3–0.9 mm shorter than the spikelet, with its dorsal surface toward the lower glume, crustaceous, glabrous, smooth, shiny, pale, black at maturity, with two basal scars ca. 0.1 mm long, brown at maturity; lemma 7 nerved; palea with simple papillae next to the apex. Caryopsis broadly ellipsoid, 1.6 × 1–1.1 mm, pale; hilum punctiform, embryo half or more the length of the caryopsis.

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Taxonomic Treatment

Fig. 47  Panicum quadriglume. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (From Steinbach 6979, USA)

Key to Ungrouped Panicum Sections and Species

189

Distribution and ecology (Fig. 34). South American species, found from Peru and Brazil to Bolivia, Paraguay, and Argentina. It inhabits fields, reaches up to 2600 m asl and flowers and fruits between January and May. Local names: “Kapi’í Cayena” (kapi’í = straw); “Cayena grass” (Zuloaga 1994) Henrard (1941) mentions that P. quadriglume would be a teratological species, due to the fact that it has 4 “glumes,” of which he indicates that the fourth corresponds to a glume III and not to the lemma of the second flower. However, the analysis of abundant material of the species rules out Henrard’s hypothesis, thus coinciding with what was previously stated by Palacios (1968). This peculiar spikelet structure is practically unique within the material examined so far in the genus Panicum, a similar structure having occasionally been found in a specimen of Ocellochloa craterifera (Sohns) Zuloaga & Morrone (= P. crateriferum Sohns) (Zuloaga and Sendulsky 1988; Sede et al. 2009). In this species, as in P. peladoense and P. exiguum, the upper anthecium is black and deciduous independently of the glumes at spikelet maturity. Panicum bergii Arechav. fo. quadriglume Henrard, Meded Rijks-Herb. 40: 52. 1921, is a nomen nudum corresponding to this species. Panicum eccentricos Hitchc. & Chase ex Rojas, Revista Jard. Bot. Mus. Hist. Nat. Paraguay 2: 164. 1930, is a nomen nudum corresponding to this species. Additional material examined. Peru. Cajamarca: valley of Río Tabaconas, 05°19′ S, 79°14′W, 1700 m, 11 April 1947, Fosberg 27768 (US). Cuzco: bajando a Chavares, 14 April 1966, Vargas 17225 (US). Huancavelica: entre Quintabamba y Virgen Pampa, 1200 m, 11 April 1964, Tovar 4585 (US). San Martín: Alto Río Huallaga, 360–900 m, December 1929, Williams 5800 (F, GH, NY, US). Brazil. Goiás: Goyandira, 800 m, campo, 26 March 1930, Chase 11560 (RB, US); Mun. Presidente Kennedy, 12  km west of village of Presidente Kennedy, 3°25′S, 48°37′W, 1 February 1980, Plowman et al. 8226 (MO, NY); 58 km NW of Rio dos Bois, 500 m, 1 April 1930, Chase 11678 (US . Marãnhao: Grajahú to Porto Franco, 8 March 1934, Swallen 3844 (IAN, K, RB, US). Mato Grosso: between Rondonopolis and São Lourenco, 16°20′S, 54°30′W, 12 April 1930, Chase 11924 (US), 11990 (IAN, RB, US); Corrego Feio, Hatschbach 23576 (K, MO). Mato Grosso do Sul: Miranda, Fazenda Bodoquena, da Silva 144 (MO); Mun. Ponta Porá, Ponta Alto, 12 February 1983, Hatschbach 46144 (K); vicinity of Dourados, 22°S, 54°40′W, 400 m, 18-21 February 1930, Chase 10976 (F, GH, MO, NY, US); Aquidauana, 200 m, Chase 11049 (RB, US); Campo Grande, Chase 10866 (R, US); Maracajú, Swallen 9390 (US); Mun. Bela Vista, Logo ao norte da ponte sobre o Rio Machorras, cerca de 10 km ao N de Bela Vista, Valls et al. 7664 (CEN); Rio Poguba, São Lourenço, Rondón 2585 (RB). Minas Gerais: Caldas, Mosén 4469 (P); Belho Horizonte, Barreiros, Kuhlmann s.n. (RB); Belho Horizonte, Black 51-11086 (US); Buritis, near Rio São Francisco, 1 January 1930, Chase 10450 (K, US, W); Serra do Curral, SE of Belo Horizonte, trail to Pico, 1000 m, 26 March 1925, Chase 9089 (F, GH, MO, NY, US); Corinto, Fazenda do Diamante, Mexía 5543a (GH, NY, US); Antonio Justiniano, 17 km S of Oliveira, 900 m, 16 March 1925, Chase 8897 (F, GH, MO, NY, RB, US, W); Uberaba, Chase 12115 (US); Lavras, Chase 88021/2

190

 

Taxonomic Treatment

(US); Belo Horizonte, pequeno morro adiante do Lago Pampulha, Black 51-11686 (IAN, US); without locality, Weddell 2531 (P). Wiithout state and locality, Riedel s.n. (P), Burchell 6769 (K) Bolivia. Beni: near Reyes, 300  m, April 1947, Cutler 9084 (F, US). La Paz: Apolo, Williams 1010 (NY); Nor Yungas, December 1917, Buchtien 4174 (GH, MO, NY, US); Yungas, 1890, Bang 493 (F, G, K, M, US); Inquisivi, climb between the Río Khara Bridge, ca. 3–4 km Licoma Pampa, 16°48′S, 67°14′W, 2600 m, 12 March 1989, Lewis 35344 (MO, SI). Santa Cruz: Ñuflo de Chávez, Comunidad Puesto Nuevo, 60 km of Concepción 16°25′S, 62°00′W, 450–750 m, 21 February 1986, Killeen 1828 (F, SI); Buena Vista, Steinbach 2204 (LIL), 5299 (G, GH, NY), 5639 (G), 6979 (BAA, F, G, GH, K, MO, US, W), 7076bis (BAA, F, G, GH, K, LIL, MO, NY, US); Ñuflo de Chavez, Estancia El Recreo, 2  km  N of Concepción, 16°02′S, 62°08′W, 21 March 1987, Killeen 2397 (F, MO, SI); Ñuflo de Chavez, Estancia Las Madres, on road to Monte Verde, 16°00′S, 62°00′W, 500  m, 13 February 1986, Killeen 1770 (F, MO, SI); Andres Ibañez, 1 km E of Int. airport Viru Viru, 16 km N of Santa Cruz, 17°40′S, 63°10′W, 400 m, 1 January 1986, Killeen 1563 (F, MO); Ñuflo de Chavez, Estancia Las Madres, 15 km N of Concepción, 16°02′S, 62°00′W, 500 m, 6 May 1985, Killeen 891 (F, MO, SI); Ñuflo de Chavez, Pueblo de Concepción, laguna west of town, 16°05′S, 62°00′W, 500  m, 25 March1986, Killeen 1848 (F, MO, SI); Ñuflo de Chavez, 2 km S of Concepción, on road to Lomerio, 16°10′S, 62°00′W, 480 m, 14 February 1985, Killeen 856 (F, MO) Paraguay. Amambay: Pedro Juan Caballero, 21 March 1945, Rojas 12741 (LIL, MO), 12819 (BAA); Cabecera Aquidabán, Sierra de Amambay, Rojas 6654 (BAA); in regióne calcarea cursus superioris fluminis Apa, Hassler 10967 (BAF, G, K, NY, P, US), 11682 (BAF, G, K, NY, P). Concepción: entre el Río Apa y el Río Aquidabán, Fiebrig 5102 (F, G, GH, M), Centurión, Fiebrig 5095 (G, K); San Salvador, Rojas 2745 (BAA, US). Cordillera: Fortín Lopez, Hassler 1942 (BAA, G, K, NY, P); región del lago Ipacaray, Hassler 12494 (BAA, F, GH, K, MO, NY, US). Paraguarí: cerca de Sapucay, Hassler 13022 (US) Argentina. Misiones: Dpto. Montecarlo, Colonia Montecarlo, 20 January 1949, Schwindt 1154 (LIL, MO, US) 43. Panicum stramineum Hitchc. & Chase, Contr. U.S. Natl. Herb. 15: 67, Fig. 50. 1910. Panicum capillare L. var. (Hitchc. & Chase) Gould, Madroño 10(3): 94. 1949. Panicum hirticaule J.  Presl var. stramineum (Hitchc. & Chase) Beetle, Phytologia 47(5): 383. 1981. Panicum hirticaule J.  Presl subsp. stramineum (Hitchc. & Chase) Frekmann & Lelong, Sida 20(1): 172. 2002. Type: Mexico. Sonora: Guaymas, November 1887, E.  Palmer 206 (holotype, US 00148035; isotypes, BM 000938692, F 0044440F, K 000309147, NDG 06750, NY 00381773, NY 00381774, NY 00381775, SI 002819, US 00148034, US-823636, US-823638, W 19160019514). Fig. 48 Panicum caatingense Renvoize, Kew Bull. 37(2): 325, Fig. 5. 1982. Type: Brazil. Bahia: 41 km N of Senhor do Bonfim on the BA 130 highway to Juazeiro, 50 m, 26 February 1974, R.  M. Harley et  al. 16384 (holotype, CEPEC; isotypes, K 000003787, MO-105087, NY 01546733, P 02610957, RB, S05-9123, US 00288689, WISv0254200WIS).

Key to Ungrouped Panicum Sections and Species

191

Fig. 48  Panicum stramineum. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (h) Caryopsis, embryo view. (i) Caryopsis, hilum view. (From Palmer 206, US)

192

 

Taxonomic Treatment

Plants annual, cespitose, with erect culms 10–70(–125) cm tall, geniculate and branching at the lower nodes; internodes 2–15 cm long, cylindrical, glabrous; nodes short to long hairy. Sheaths 2–9 cm long, glabrous to hirsute, with papillose hairs, green to purplish, the margins membranous, glabrous, or with one margin ciliate. Ligule membranous-ciliate, 0.9–2.4 mm long, with or without long hairs beneath at the base of the blade; collar glabrous to hispid. Blades linear-lanceolate, 4.5–30(−42)  ×  0.3–1.3(−2) cm, the basal ones smaller, erect, ascending, slightly divergent, flat, glabrous to sparsely papillose-pilose on both surfaces, the base rounded base and apex attenuate, margins scaberulous, glabrous to papillose pilose toward the base. Terminal and axillary inflorescences present, similar, lax, 2.5–22(−38) × (1–)4–10(−20) cm; peduncles up to 24 cm long, cylindrical, glabrous; main axis wavy, scaberulous; first-order branches alternate to subopposite, divergent, naked toward the base; second-order branches scaberulous, divergent, with spikelets scattered arranged; pulvini glabrous to shortly hairy; pedicels claviform, 0.7–2.5 mm long, scaberulous, with whitish hairs toward the apex. Spikelets ovoid, 2.3–3.2 × 0.8–1.4 mm, turgid, globose, abruptly acuminate, pale; upper glume and lower lemma subequal, as long as or exceeding 0.4 mm the upper anthecium; lower glume ovate-acuminate, 0.7–1.2(−1.5) mm long, 1/3–1/2 or slightly more the length of the spikelet, obtuse to acute, 5–7 nerved, with the midrib scabrous toward the apex; upper glume 1.8–3.1 mm long, 9–11(–13) nerved; lower lemma glumiform, 1.7–3 mm long, 9–11 nerved; lower palea ovate-lanceolate, (1.4–)2.3–2.7 × 0.5–0.9 mm, equal to or longer than the upper anthecium, membranous, hyaline, glabrous, upper margins ciliate to glabrous; upper anthecium ellipsoid, 1.5–2.5 × 0.7–1.3 mm, crustaceous, smooth, shiny, pale and grayish tinged at maturity, with two basal scars 0.2 mm long; lemma 7 nerved; palea with simple papillae toward the apex, sometimes distributed over the entire surface. Caryopsis ovoid, 1.2–1.6  ×  0.6–0.9  mm, pale; hilum punctiform; embryo half the length of the caryopsis. Distribution and ecology (Fig. 32). Species present in the USA and Mexico and in South America from Venezuela to Argentina. It grows in fields or in open sites along roadsides or railroad tracks. It is also frequent in deciduous forests or flooded sites, occasionally leaning on vegetation; it lives from sea level to 1100 m elevation. Local name: “Capim-lanudo” (in Brazil) Panicum caatingense, a species synonymized with P. hirticaule by Zuloaga (1989) and Killeen (1990) should be considered in the synonymy of P. stramineum. In P. stramineum the lower glume varies between 1/3 and 1/2 the length of the upper glume and lower lemma. This species is related to P. hirticaule, from which it differs by having globose, turgid spikelets, more or less abruptly acuminate and with the lower palea developed, reaching the same length as the lower lemma. The plants usually have erect leaves, little divergent from the stem. Additional material examined. USA.  Arizona: Pima Co, 18 August 1937, Proctor 2571 (US); under Mezquite bushes, White’s Mills, 5 September 1867, Palmer 270 (MO); sandy plains near Mexican boundary, 29 August 1894, Pringle s.n. (MO-2752212); Casa Grande, Silveus 3461 (US); Tucson, Pima Co, near

Key to Ungrouped Panicum Sections and Species

193

Souther Pacific Hospital, 1 September 1934, Higgins 6990 (US); 1/2 mi. NW Cielito, 24 September 1938, Hers & Mendez 4000 (US); Tucson, 18 September 1930, Abrams 12902 (US); Pima Co., 17 October 1945, Goodding 192-45 (US) Mexico. Chiapas: Mun. Arriaga, deciduous forest, at Río Las Arenas, 15  km northwest of Arriaga, 130 m, 27 August 1974, Breedlove 36779-A (MO). Guerrero: Lagunillas, 1 August 1898, Langlassé 263 (US). Michoacán: Tancitaro, road to Hda. California, 1000 ft, 12 August 1941, Leavenworth & Hoogstraal 1453 (GH, MO, NY, US); Río San Antonio near junction with Río Tepalcatepec, Rancho El Capire, 3 August 1940, Leavenworth 442a (US). Nayarit: between Concepción and Acaponeta, 29 July 1897, Rose 1889 (MEXU, NY, US); between Rosario and Acoponeta, 28 July 1897, Rose 1883 (US); 26.8 mi. S of Sinaloa state line in Nayarit, 24 July 1975, Dunn et  al. 21855 (MO, NY). Nuevo León: 12 mi S of Monterrey, 14 July 1933, Mueller 404 (US). Oaxaca: Ixtepec, August 1935, Fisher 35314(F, MO, NY, US); along the Hwy. to Tehuantepec (route 185), along the Hwy. to Ixtepec, 1 km NW of Juchitán, 100 m, 26 July 1958, King 771 (MEXU, US). Sinaloa: between Rosario and Acaponeta, Rose 1878 (GH, US); vicinity of Labradas, 18 September 1925, Ferris & Mexía 5069 (MO, US); Culiacán, 27 August/15 September 1891, Palmer 1538 (F, US). Sonora: Orozi, near Río Yaqui, 7 September 1935, Pennell 20221 (MO, US); Guaymas, 1887, Palmer 168a (US); 3.3 miles south of Oasis de los Chinos, 57.8 miles south of Santa Ana, 21 September 1974, de Wet & Harlan 4353 (MEXU) Colombia. Cundinamarca: Alto Magdalena province, Nilo municipality, Hacienda La Guaira, Pagüey river, 350 m, 16 October 2004, Giraldo-Cañas 3773 (COL) Venezuela. Apure: between Balza and Manati canos, Mantecal-Quintero road, 7°45′N, 69°40′W, 5 August 1989, Zuloaga et  al. 4340 (MO, PORT, SI, VEN); between Achaguas and San Fernando, near Bethel, 7°48′N, 67°55′W, 6 August 1989, Zuloaga et al. 4350 (MY, PORT, SI, VEN); Mantecal, 10June 1971, Ramia 4444, 4452 (VEN); Mantecal, 17 July 1975, Ramia & Montes 5345 (US, VEN). Carabobo: Valencia, ca. 4 km E of Valencia on the highway between Valencia and Maracay, 8 June 1978, Burandt Jr. & Wingfield V0242 (VEN). Falcón: Hato Uvento, Wingfield 6840 (MO); Paraguaná, 2 km E of Sta. Ana, 4 January 1980, Wingfield 7361 (CORO, MY, VEN); Paraguará, Pueblo Nuevo, January 1939, Tamayo 1000 (VEN). Guárico: S de Calabozo, 22 July 1976, Castillo 583 (VEN); La Rubiera near Calabozo, Grisol 36 (VEN). Zulia: Perijá, Las Piedras-San José-Las Laras road, at Km 9 E of the junction with Perijá road, 10 September 1977, Bunting 5447 (VEN); Miranda, road between El Mecocal (km 19 E of the Puente Sobre el Lago, of the Marchacibo-Coro road) and El Consejo, at km 6 SE of El Mecocal, 24 September 1977, Bunting 5568 (VEN). Without state: Hato Becerra, 2 km S of the Orituco River, 11 July 1961, Blydenstein 148 (VEN) Peru. Tumbes: along the road to Tutumo, about 20 km from Tumbes, 26 April 1969, Simpson & Schunke V. 829 (F, US) Bolivia. Beni: Prov. Gral. Ballivian, Espíritu, in the zone of influence of the Yacuma River, 200 m, 12 March 1990, Beck 15176 (SI); Trinidad-Beni, 200 m, July 1944, Cárdenas 3533 (US), 3534 (GH, LA). Chuquisaca: Isirenda, 26 January

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Taxonomic Treatment

1992, Saravia Toledo & Nelson Joaquin 10412 (SI). Santa Cruz: Cordillera, 400 m, Renvoize & Cope 3912 (K, US); Chiquitos, Est. San Ignacio, 22 km N of San José de Chiquitos, 17°35′S, 60°45′W, 320  m, 1 February 1986, Killeen 1711 (F, SI); Chiquitos, San José de Chiquitos, 17°47′S, 60°45′W, 320  m, 30 January 1986, Killeen 1702 (F, SI). No department: pampas near Lake Rogagua, 1100  m, 14 November 1921, Cárdenas 1647 (NY) Brazil: Bahia: Iaçú, Faz. Suíbra, Tanque da Pindoba, 18 km leste da cidade, 15 March 1985, Noblick 3728 (CEPEC); 90  km ao N do Senhor do Bonfim, Mina Caraíba, Castellanos 25909 (R); 4 km north of Senhor do Bonfim, disturbed ground by roadside, 550 m, 40°11′W, 10°27′S, 24 February 1974, Harley et al. 16294 (MO, R, RB, US). Ceará: Iguatú, Swallen 4396 (GH, RB, US). Marãnhao: Mun. Loreto, Ilha de Balsas, region between the Balsas and Parnaiba rivers, 26  km south of Loreto, 200 m, 11 February 1970, Eiten & Eiten 10579 (US); Perizes, 6 July 1954, Black et  al. 54-16594 (US). Mato Grosso do Sul: Fazenda Bodoquena, Seçao Guaicurus, Allem et al. 2145 (MO); base do Cerro do Urucum, 23 km S Corumbá, 22 November 1977, Allem & Vieira 1479 (MO, SI). Piauí: Fazenda Nacional to Picos, low ground near small stream, 4 April 1934, Swallen 4192 (RB, US). Pará: Ilha do Marajó, Faz. Gavinho, Goeldi 261 (NY) Paraguay. Alto Paraguay: Colonia San Lázaro, near Río Apa, January 1931, Rojas 5476a (US). Boquerón: Tinpunké, near Río Pilcomayo, 28 October 1985, Mereles 715 (SI); between Estancias Santa Ramona and San Alberto, Puesto Sastre, 9 March 1949, Ramírez 51 (SI); Parque Nacional Tte. Enciso National Park, 21°10′S, 61°35′W, 10 January 1984, Hahn 1911 (MO, SI); W of Estancia La Patria, 7 December 1992, Nicora et al. 9737 (SI); Ruta Trans-Chaco, 21°26′S, 61°25′W, 6 March 1979, Schinini & Bordas 16368 (SI). Concepción: San Salvador, 4 March 1917, Rojas 11027, 11047 (US). Presidente Hayes: Ruta Concepción-Pozo Colorado, 28 km from the bridge, 18 February 1990, Palacios 1856 (MO) Argentina. Catamarca: Santa Lucía, 2 April 1950, Brizuela 982 (GH, NY). Formosa: Dpto. Matacos, Ingeniero Juárez, 14 January 1957, Burkart 20275 (SI, US). Jujuy: Dpto. El Carmen, Los Lapachos, 21 March 1984, Ahumada & Castellón 5023 (SI); Dpto. El Carmen, Ruta 34, 6  km from Pampa Blanca, 2 February 1975, Zuloaga & Deginani 412 (SI); Dpto. Ledesma, Est. Exp. Yuto, 12 November 1975, Otero 48 (SI); Campamento Caimancito de YPF, entrance, 7 December 1986, Zuloaga et al. 2515 (SI). Dpto. San Pedro, Chaguaral, Finca Jure, 18 April 1983, Ahumada & Castellón 4829 (SI); Dpto. Dr. Manuel Belgrano, Ruta nac. 66, near Aeropuerto El Cadillal, 20 January 1988, Zuloaga & Deginani 3702 (SI). La Rioja: Dpto. Gobernador Gordilllo, Gobernador Gordillo, 14 March 1942, Nicora s.n. (SI-18478); Dpto. Famatina, Famatina, 17 February 1941, Burkart 12262 (SI); Dpto. Gral. San Martín, Estancia La Diana, ca. 800 m, March 1907, Kneucker 730 (GH, ISC, LA, MO, NY, SI, W). Salta: Dpto. Anta, Salta Forestal, sector I, 320  m, 19 February 1987, Saravia Toledo 1305 (SI); Dpto. Anta, Salta Forestal, 11 km NNW, 320 m, 11 April 1986, Saravia Toledo 1243 (SI); Dpto. Anta, San Javier, 8 km S of J.V. González, ruta 16, 350 m, 18 February 1987, Saravia Toledo 1301 (SI); Dpto. Campo Santo, Güemes, 3 April 1945, O’Donell 2629 (NY, SI); Dpto. Gral. José de San Martín, near Dragones, Ruta 81, April 1975, Cabrera et al. 26542 (SI)

Key to Ungrouped Panicum Sections and Species

195

44. Panicum tamaulipense F.R. Waller & Morden, Syst. Bot. 8(2): 221. 1983. Type: Mexico. Tamaulipas: Sierra de Tamaulipas area, between La Chona and Río Santa Olaya, low deciduous forest, 50 m, 26 September 1956, F. Martínez Martínez & G.  Borja Luyando F-2161 (holotype, TEX 00370129; isotype, US 00148051) Plants perennial, cespitose, 50-80(−120) cm tall, culms simple, glaucous, with intravaginal innovations, erect, few noded, internodes 3–10  cm long, cylindrical, hollow, 1–2 mm in diameter at base, nodes hairy with short hairs. Sheaths 8–12 cm long, equal to or larger than the internodes, striate, hirsute, tinged with purple, with adpressed tuberculate hairs, one margin ciliate, the other glabrous. Ligule membranous-ciliate 0.5–1.2  mm long; collar hairy to glabrous. Blades linear-lanceolate, 20–35(−50) × 0.5–0.8 cm, erect, ascending, rigid, flat, slightly curled at base, glaucous, glabrous, base narrow, apex acute, the margins slightly involute, scaberulous, midrib conspicuous. Inflorescences exserted, 15–30 × 6–15 cm, lower branches up to 18 cm long; peduncles up to 50 cm long, glabrous, smooth, pale or tinged with purple; main axis scabrous, glabrous, slightly undulate; pulvini glabrous, first-order branches alternate, divergent, pedicels claviform, 4–15 mm long, divergent, flexuous, glabrous, scaberulous. Spikelets ovoid, 1.8–2.2 × 0.7–1 mm, glabrous, pale or tinged with purple, apex abruptly acuminate, nerves greenish; upper glume and lower lemma subequal, as long as the upper anthecium; lower glume ovate-acuminate, 1–1.2 mm long, reaching 1/2 the length of the spikelet, 5 nerved; upper glume 1.8–1.9 mm long, acuminate, 7–13 nerved; lower lemma glumiform, 1.8 mm long, 7–9 nerved; lower palea lanceolate, 1–1.2 × 0.3 mm, 3/4 the length of the upper anthecium, membranous, hyaline, glabrous, the apex finely denticulate; upper anthecium ovoid, 1.5–2 × 0.6–1 mm, crustaceous, glabrous, smooth, shiny, with two basal scars ca. 0.2 mm long, brown at maturity; lemma 7 nerved; palea papillose toward the apex. Caryopsis ovoid, 1 × 0.5 mm, whitish; hilum punctiform; embryo half the length of the caryopsis. Distribution and ecology (Fig. 32). Species endemic to Mexico, where it is found in fields in the departments of San Luis Potosí and Tamaulipas, reaches up to 350 m asl, and flowers and fruits between May and December. Species related to P. hallii subsp. filipes, from which it differs in having smaller spikelets, 1.8–2.2 mm long and blades 20–35(−50) × 0.5–0.8 cm. Additional material examined. Mexico. San Luis Potosí: El Banito, 100 ft, 22 July 1939, Chase 7558 (US); Mun. El Pujal, Chase 7492 (US); 2  miles  S of Huichihoayan, Mc Gregor et  al. 870 (US); Ciudad Valles, 5 May 1962, Beetle M-718 (US); Valles, 800 ft, 17 July 1933, Fischer 3313 (US); in a tropical forest along Mexico Highway 55 to Xilitla, Sohns 1431 (US). Tamaulipas: El Mante to Limón, 3 May 1931, Swallen 1618 (US); 64 km from Soto La Marina on the road to Casas and Victoria, 350 m, 4 October 1956, Martínez Martínez & Borja Luyando F-2364 (US); 35 km from Victoria on the road to Casas and Soto La Marina, 280m, 3 October 1956, Martínez Martínez & Borja Luyando F-2333 (US); 5 mi N of San Rafael toward Soto la Marina, 800 ft, 11 December 1959, Crutchfield & Johnston 4956 (MEXU)

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Taxonomic Treatment

45. Panicum venosum Swallen, Contr. U.S. Natl. Herb. 29(9): 415. 1950. Urochloa venosa (Swallen) Morrone & Zuloaga, Darwiniana 32(1-4): 74. 1993. Type: Mexico. Michoacán: near irrigation ditch 1 mile SW of Apatzingán, 5 August 1940, W. C. Leavenworth 481 (holotype, US 00148083; isotype, MO-123241) Annual plants, ca. 40  cm tall; culms erect to geniculate; internodes glabrous, 4–6 cm long; nodes hairy. Sheaths 3–7 cm long, glabrous, with one margin ciliate, the remaining glabrous. Ligule membranous-ciliate, 1  mm long; collar glabrous. Blades lanceolate, 4–9 × 0.5–0.9 cm, base subcordate and apex attenuate, glabrous, the margins cartilaginous, smooth, the lower ones long-petiolate, then glabrous. Inflorescences lax, 12–14  ×  4–5  cm; main axis scaberulous, first-order branches divergent, alternate to opposite, naked toward the base; axes of branches triquetrous, glabrous, scaberulous, ending in a developed spikelet; second-order branches present; pulvini glabrous; pedicels triquetrous, scaberulous; spikelets solitary or in pairs. Spikelets ovoid, 2.4–2.6 × 1–1.4 mm, glabrous, pale green, upper glume and lower lemma subequal, with manifest nerves; lower glume ovate-acuminate, 1.3– 1.8 mm long, ½ or slightly more the length of the spikelet, membranous, glabrous, clasping, 5–7 nerved; upper glume 2.5  mm long, membranous, glabrous, 9–11 nerved; lower lemma glumiform, 9 nerved; lower palea reduced, ½ or less the length of the lower lemma, hyaline; lower flower absent; upper anthecium ellipsoid, 1.9  ×  1  mm, biconvex, shortly apiculate, indurate, glabrous, pale, transversely rugose, with simple papillae on its surface; upper lemma 5 nerved, upper palea 2 nerved, transversely rugose, the margins involute, smooth and shiny. Caryopsis ovoid, 1.5 × 0.7 mm; hilum punctiform, embryo half the length of the caryopsis. Distribution and ecology (Fig. 44). Species endemic to Mexico, only known from the type collection in the state of Michoacán Panicum Sect. Repentia Stapf, Fl. Trop. Afr. 9: 640, 648. 1920. Type species. Panicum repens L. Panicum sect. Coloratae Stapf, Fl. Trop. Afr. 9: 641, 648. 1920 Plants perennial, occasionally annual, when perennial with rhizomes manifest, rigid, short to elongate; culms erect, internodes hollow, without aerenchyma. Ligule membranous-ciliate. Blades oblong-lanceolate to lanceolate, flat to involute, hairy to glabrous. Inflorescence a terminal panicle, exerted, lax to contracted. Spikelets long ovoid to ellipsoid, glabrous, not open at maturity, pale or tinged with purple; lower glume ¼ to 1/3(exceptionally 1/2) the length of the spikelet, nerveless to 1–5(−7) nerved, upper glume and lower lemma 9–11 nerved; lower palea present, hyaline, glabrous; lower flower staminate or absent; upper anthecium ellipsoid to ovoid or broadly ovoid, glabrous, pale, not stipitate, smooth, shiny, indurate. Caryopsis ellipsoid; hilum oblong, embryo less than half to half the length of the caryopsis. Key to the Species 1. Plants cespitose, shortly rhizomatous; lower glume up to ½ of spikelet length, 1−5(−7) nerved..........................................................................46. P. coloratum

Key to Ungrouped Panicum Sections and Species

197

1. Plants with longt rhizomes; lower glume 1/5 to ½ the length of the spikelet, 0(−3) nerved. 2. Spikelet (3–)3.2–4 mm long..................................................48. P. pedersenii 2. Spikelet 2.4–2.7(−2.9) mm long. 3. Lower glume reduced, 1/5 to 1/3 the length of the spikelet, the apex usually truncate, occasionally obtuse or acute.......................................................... ............................49. P. repens 3. Lower glume 1/3 to ½ the length of the spikelet, apex acute........................ .............................................................47. P. gouinii 46. Panicum coloratum L., Mant. Pl. 1: 30. 1767. Type: Egypt. “Habitat in Cairi,” Forsskal s.n. (lectotype, LINN-80.46, designated by F. O. Zuloaga, Fl. Guianas, ser. A, 8: 384. 1990) Plants perennial, cespitose, shortly rhizomatous; culms erect to decumbent, 20–95 cm tall; nodes glabrous, occasionally pubescent. Sheaths hispid to glabrous. Ligule membranous-ciliate. Blades lanceolate to linear-lanceolate, (4–)11– 25 × (0.3–)0.5–1.1 cm, glabrous or hairy, cordate to rounded at base, the apex acute to acuminate. Inflorescence an exserted terminal panicle, 5–28(−37) cm long, with ascending to adpressed or divergent first-order branches; spikelets solitary on claviform pedicels, glabrous. Spikelets ovoid to ellipsoid, 2–3  ×  1–1.1  mm, glabrous, upper glume and lower lemma subequal; lower glume 0.5–1 mm long, ¼ to 1/2 the length of the spikelet, membranous, 1–5(−7) nerved, glabrous, acute; upper glume 7–9 nerved; lower lemma glumiform, 7–9 nerved, acute; lower palea present, lower flower staminate; upper anthecium ovoid, 1.5–2.6 × 0.7–0.8 mm, indurate, smooth, shiny, glabrous. Caryopsis not seen. Distribution and ecology (Fig. 50). Species widely distributed in the Old World, occasionally introduced in America, found in the USA and Venezuela and grows in open and humid environments. Additional material examined. USA. Georgia: Spalding, Webster 3246 (MO). Texas: Lampasas, Hatch 5847 (MO) Venezuela. Lara: Finca Monte Carmelo, El Tocuyo, 5 April 1978, Burandt Jr. V0120 (MO) 47. Panicum gouinii E. Fourn., Mexic. Pl. 2: 28. 1886. Panicum gouinii E. Fourn. ex Hesml., Biol. Cent.-Amer. Bot. 3: 489. 1885, nom. nud. Type: Mexico. Veracruz: Veracruz, 4 July 1867, F. M. G. Gouin 4 (holotype, P 00371642; isotype, BAA 00002340, fragment ex P). Fig. 49i-q Panicum gouinii E. Fourn. var. pumilum E. Fourn., Mexic. Pl. 2: 28. 1886. Panicum gouinii E. Fourn. var. pumilum E. Fourn. ex Hesml., Biol. Cent.-Amer., Bot. 3: 489. 1885, nom. nud. Type: Mexico. Veracruz, 15 April 1850, M.  Virlet 1300 (lectotype, P 00740949, here designated; isolectotypes, P 00740950, US 00148539, fragment ex P)

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Taxonomic Treatment

Fig. 49  Panicum repens. (a) Spikelet, lateral view. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Lower glume. (e) Upper glume. (f) Lower lemma. (g) Lower palea. (h) Upper anthecium, ventral view. Panicum gouinii. (i) Spikelet, lateral view. (j) Spikelet, ventral view. (k) Spikelet, dorsal view. (l) Lower glume. (m) Upper glume. (n) Lower palea. (o) Upper anthecium, ventral view. (p) Caryopsis, hilum view. (q) Caryopsis, embryo view. (a–h, from Burkart & Gamerro 21661, SI; i–q, from Zuloaga & Sáenz 574, SI)

Key to Ungrouped Panicum Sections and Species

199

Panicum repens L. var. confertum Vasey, Bull. Torrey Bot. Club 13(2): 25. 1886. Panicum halophilum Nash, Bull. Torrey Bot. Club 28(2): 87. 1901. Type: USA. Mississippi. Hancock Co, in sandy beach of Gulf, St. Louis, 13 September 1883, A. B. Langlois 49 (holotype, US 00147993) Perennial plants, 0.10–0.50  m tall, with elongated rhizomes, cataphylls hairy cataphylls, culms, erect, simple, internodes cylindrical, glabrous, 2–6  cm long; nodes glabrous or hairy, dark. Sheaths smaller or larger than the internodes, 3–6 cm long, aphyllous at the base of the cane, distichous, the margins ciliate toward the upper portion or glabrous. Ligule shortly membranous at base then ciliate, reduced, 0.4–0.8  mm long; collar light brown, glabrous. Blades lanceolate to linear, 4.5– 13  ×  0.2–0.4  cm, base narrow, the margins involute and finely scabrous, adaxial surface pilose toward the base or glabrous. Inflorescence terminal, exserted; panicle contracted, few flowered, 5.4–10 × 1–6 cm; main axis triquetrous, scabrous, pulvini glabrous, first-order branches alternate, adpressed, spikelets congested on the branches, pedicels scabrous, 1–4.5 mm long. Spikelets ovoid, 2.4–2.9 × 0.9–1.1 mm, glabrous, pale or tinged with purple; lower glume ovate, acute, 1–1.6 mm long, 1/3 to 1/2 the length of the spikelet, 1–7 nerved; upper glume 2.2–2.7 mm long, covering or not the apex of the upper anthecium, 9–11 nerved; lower lemma 2.2–2.7 mm long, acute, 9–11 nerved; lower palea elliptic, 2–2.4 × 0.7–0.9 mm, membranous, glabrous; lower flower staminate, anthers orange, 1.5 mm long; upper anthecium ovoid, 1.9–2.1 × 0.8–1 mm, pale, brown at maturity, glabrous, smooth, shiny, indurate, lemma 7 nerved. Caryopsis ellipsoid, 1.5 × 0.8 mm, brownish and with black spots; hilum oblong, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 50). It is found in the southern USA, Mexico, and Cuba and in South America, in Brazil, Paraguay, Uruguay, and Argentina. It grows in coastal dunes or on the edges of rivers or streams, on sandy soils. Of the two syntypes from Mexico cited by Fournier in describing Panicum gouinii var. pumilum, Liebmann 450 (syntype, C, US-80681, photo and fragment) and Virlet 1300, the latter specimen from P is selected as lectotype, as it corresponds to the protologue and is a complete specimen. This species has been synonymized with P. repens (Davidse 1994). However, it differs from the latter in that the latter species includes larger plants, up to 1 m tall and spikelets with the lower glume up to 1/3 of the spikelet length, 1–3(−5) nerved. Additional material examined. Mexico. Veracruz: Veracruz, Muller 2177 (P, US), 31 August 1910, Hitchcock 6551 (US), 23 April 1894, Pringle 5569 (MEXU, US), Schiede 895 (US); orilla este de la Laguna Verde, 23 June 1972, Vazquez Yañes 635 (MEXU); Coatzacoalcos, isthmus of Tehuantepec, 8 March 1895, 8 March 1895, Smith 913 (MEXU, US); camino entre Veracruz y Antón Lizardo, 8 km al este de Boca del Río, 20 April 1974, Koch 7461 (MEXU, US); Paso de Doña Juana, 18 July 1981, Moreno Casasola et al. 175 (MEXU) Costa Rica. Limón: Guapiles, Hitchcock s.n., Amer. Grass Herb. 40 (F) Cuba. Habana: Vedado, Ekman 702 (G); Vibora, 14 May 1921, Ekman 12819 (G, NY, US); Santos Suarez, Ekman 16801 (G, NY, US); Lomas de Camoa, at a

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Taxonomic Treatment

Fig. 50  Distribution of Panicum coloratum, P. gouinii, P. pedersenii, and P. repens

small laguna, 29 September 1923, Ekman 17554 (NY, US); Arroyo Apolo, León 296, 563 (US); Castillo de Atarés, near the gashouse, Ekman 13301 (NY) Brazil. Pará: Marajó, Miranda 3238 (G). Paraná: Areial, Mun. São Jose dos Pinhaes, Hatschbach 47654 (K); Serra do Mar, Banhado, Dusén 14574 (K). Rio Grande do Sul: Mun. Bagé, Hulha Negra, Allem & Vieira 1819 (CEN); Mun. Herval do Sul, 15 km E Herval do Sul, Allem & Vieira 1844 (CEN); Montenegro, Agua Exp. Station, Clayton 4401 (K, SP); Osorio, Lagoua Itapena, Kappel- Froner s.n. (SP); Morro da Gloria, Porto Alegre, Orth s.n. (SP 51157); Cachoeira in uliginosis et palustribus camporum, Lindman A1209 (P); Corral Alto, Trinta 1163 (SI). Santa Catarina: Balneario Camboriú, Hatschbach 46653 (K); Florianópolis, Ilha de Santa Catarina, March 1981, Zuloaga et al. 879 (SI, US); Sombrío, 10 m, Reitz 2015 (G, NY). São Paulo: water font in city of Cananeia, Clayton & Eiten 4687 (K, NY, SP) Argentina. Buenos Aires: Ensenada, Punta Lara, 28 January 1978, Zuloaga & Sáenz 574 (SI), Zuloaga 2994 (SI). General Madariaga, Martínez Crovetto 2532 (BAB). Isla Maciel, Clos 701 (BAB). Ituzaingó, Martinez Crovetto s.n. (BAB); Isla Santiago, 7 March 1942, Cabrera 7453 (SI). Pipinas, Clos 1883 (BAB). Rio Salado

Key to Ungrouped Panicum Sections and Species

201

y La Aurora, Vervoorst s.n. (BAB). Corrientes: Concepción, Paraje Rincon de Luna, entre Batel y Batelito, Estancia Los Tres Rincones, 2 February 1963, Cano & Cámara Hernandez 685 (SI); Manuel Derqui, 8 February 1976, Quarín 3324 (CTES); Estancia Santa Teresa, Pedersen 1418 (NY). Entre Rios: Concordia, Clos 245 (BAB); Holt, Martinez Crovetto s.n. (BAB), Milano 5100 (BAB); Dpto. Colón, Villa San José, Arroyo Perucho Verne, 7 April 1968, Burkart et al. 26865 (SI, US) Uruguay. Montevideo: Carrasco, Herter 432 (F, NY). San José: Barra de Santa Lucia, Osten 4512 (G) 48. Panicum pedersenii Zuloaga, Hickenia 1(27): 148. 1978. Type: Argentina. Corrientes. Dpto. Mburucuyá, Estancia Santa Teresa, 6 March 1961, T. M. Pedersen 5868 (holotype, BAA 00000392; isotypes, BAA 00000393, CTES 000045). Fig. 51 Perennial plants 0.70–1  m tall, strongly rhizomatous, rhizomes covered with hairy cataphylls; culms erect; internodes 5.5–12 cm long, cylindrical, hollow, glabrous, pale; nodes glabrous, dark, compressed. Sheaths 5–11 cm long, usually larger than internodes, pale or tinged with purple, pilose to glabrescent, margins membranous, ciliate to glabrous toward the upper portion. Ligule membranous at the base, then long ciliate, 1.7–2.4 mm long; collar pale, pilose to glabrous. Blades lanceolate, 11–27 × 0.4–0.7 cm, flat or with slightly involute margins, the adaxial surface pilose with adpressed hairs, basal margins ciliate, otherwise scabrous, midrib manifest. Inflorescence a lax, diffuse, terminal panicle, 8–23 × 4–15 cm wide; main axis scabrous, triquetrous, pulvini glabrous, first-order branches; pedicels 1.5–7  mm long, scabrous. Spikelets lanceolate, (3–)3.2–4  ×  0.9–1.4  mm, glabrous, upper glume and lower lemma subequal, exceeding the upper anthecium in length; lower glume ovate, 1.3–1.7 mm long, up to 1/3 the length of the spikelet, glabrous, 5–7 nerved, obtuse; upper glume 2.9–3.8 mm long, 9–11 nerved; lower lemma glumiform, 3–3.7 mm long; lower palea elliptic, 3–3.6 × 0.8–1.1 mm, membranous, glabrous; staminate flower present, three anthers, 2.1–2.3 mm long; upper anthecium ellipsoid, 2.5–2.9 × 0.9–1.1 mm, glabrous, smooth, shiny, pale, palea with papillae at apex; anthers orange, 1.8–2 mm long, stigmas plumose, purplish. Caryopsis not seen. Distribution and ecology (Fig. 50). Species of restricted distribution, found in the province of Corrientes in Argentina, southern Brazil, and Paraguay. It grows in low fields or edges of lagoons and flowers from December to March. Additional material examined. Brazil, Mato Grosso: Posto XV, 14 February 1970, Hatschbach 23525 (MO) Paraguay. Central: Aregua, 10 January 1875, Balansa 17 (G, K, P); Paraguaria centralis, in regione lacus Ypacaray, November 1913, Hassler 11451 (F, G, K); Capital, San Lorenzo, Arroyo Aratiry, 4 March 1984, Schinini 23924 (CTES, G). Cordillera: Cordillera de Altos, Hassler 3652 (G, K, NY, P, W) Argentina. Corrientes: Dpto. Ituzaingó: Ituzaingó, 7 December 1973, Quarín et al. 1766 (CTES); Laguna Soto, Schwarz 9144 (LIL), 18 January 1950, Schwarz 9192 (LIL, MO); Ruta Prov. 17, 20  km  SW de San Miguel, 30 March 1974, Krapovickas et al. 24632 (CTES, US), 24669 (CTES); Carambola, Estancia “Fortín del Iberá, 30 December 1982, Pedersen 13473 (MO)

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Fig. 51  Panicum pedersenii. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, ventral view. (From Pedersen 5868, BAA)

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49. Panicum repens L., Sp. Pl. ed. 2, 1: 87. 1762. Type: “Habitat in Hispania, inde missum a Claud. Alstraemer.”, C. Alstroemer s.n. (lectotype, LINN-80.47, designated by A.  S. Hitchcock & A.  Chase, Contr. U.S.  Natl. Herb. 15: 85. 1910). Fig. 49a-h Panicum littorale C. Mohr ex Vasey, Bot. Gaz. 4(1): 106. 1879. Type: USA. Alabama: Mobile Co, Mobile, July 1877, C. Mohr s.n. (holotype, USA 00132949; isotypes, NY 00546631, NY 00546632) Plants perennial, strongly rhizomatous, culms erect, 0.50–1 m tall, many noded, simple, branching or not at the lower nodes, basal sheaths persistent, distichous; internodes 2–12 cm long, cylindrical, glabrous; nodes glabrous to sparsely hairy, brown. Sheaths usually smaller than the internodes, 2–8 cm long, papillose-pilose, with deciduous tuberculate hairs, or only pilose toward the upper portion. Ligule membranous-ciliate, 0.4–0.6  mm long; collar glabrous to sparsely pilose, pale. Blades lanceolate, 9–20 × 0.2–0.4 cm, the margins involute, base rounded, dense to sparsely pilose on the basal portion of the abaxial surface, lower margins ciliate with tuberculate hairs, otherwise scabrous. Inflorescence terminal exerted; panicles lax or more commonly contracted, 12–24  ×  2–6  cm; main axis wavy, scabrous, pulvini glabrous, first-order branches alternate, branch axes and pedicels triquetrous, scabrous, pedicels 2–10 mm long. Spikelets long ovoid, 2.4–2.6 × 0.8–1 mm, glabrous, pale; lower glume 0.8  mm long, truncate, (0–)1–3(−5) nerved; upper glume 2.2–2.4 mm long, acute, 9–11 nerved; lower lemma glumiform, 9–11 nerved; lower palea elliptic, 1.9–2.1 × 0.7–0.9 mm, membranous, glabrous; lower flower staminate, anthers purplish, 1–1.3  mm long; upper anthecium long-ovoid, 1.9– 2.1 × 0.7 mm, glabrous, smooth, shiny, indurate, pale. Caryopsis not seen. Distribution and ecology (Fig. 50). It is a cosmopolitan species, distributed in the tropics and subtropics throughout the world. Originally from Europe, it has been introduced in America as a fixer of dunes; it lives in sandy soils. Local names: “Torpedo grass”, “torpedo grass”, “pensrepens” Holm et al. (1977) report that P. repens is an important herbicide-resistant weed in crops. They indicate that caryopsis formation is rare in spikelets of this species and mention that, when present, the caryopsis is pale. Additional material examined. Belize. Toledo: in wet area, Mason’s land, Wilson Road, 12 October 1949, Gentle 6863 (F, US) Bahamas. New Providence, weedy area about Caribbean Nursery near roundabout on Village Road, 22 May 1980, Correll & Popenoe 51405 (MO) Cuba. Havana: Santos Suarez, near Habana, in ditches, 21 June 1923, Ekman s.n., Amer Gr. Herb. 701 (MO) Brazil: Alagoas: Porto Calvo, Faz. Mucaita, Campelo 2177 (CEN); Matriz de Camaragibe, Fazenda Santa Lucia, Campelo 2158 (CEN). Mato Grosso: Mun. Poconé, Km 19 MT3 (Transpantaneira), Allem & Vieira 1686 (CEN). Pará: Belem, IAN, Black 56-18910 (IAN, UB); Rio Paracavari, Fazenda Vitoria, Black & Engelhard 50-89761/2 (IAN). Rio Grande do Sul: São Leopoldo, Dutra 588 (IAN). São Paulo: Praia Grande, Florida, Sendulsky 1078 (SP); Cananeia, Ilha do

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Cardoso, Sendulsky 1299 (SP); Agua Funda, Bordo 1 (SP); Reserva do Instituto de Botanica de São Paulo, Parque do Estado, Rosa 3697 (SI, SP), 21 February 1976, Davidse & D’Arcy 10478 (K, MO, SP) Argentina. Corrientes: Dpto. Paso de los Libres: Laguna Mansa, 30 October 1973, Quarín 1657 (CTES, G); Krapovickas & Cristóbal 21720 (CTES); Gobernador Virasoro, Estancia Las Marías, Barrett s.n. (BAB). Entre Ríos: Santos Rubio s.n. (SI); Dpto. Concordia: Concordia, Colonia Yeruá, 12 April 1960, Burkart & Gamerro 21661 (SI) Panicum Sect. Rudgeana (Hitchc.) Zuloaga, Ann. Missouri Bot. Gard. 74(3): 470. 1987. Panicum [no rank] Rudgeana Hitchc., N. Amer. Fl. 17(3): 200, 205. 1915. Type species: Panicum rudgei Roem. & Schult Plants annual or perennial, cespitose, shortly rhizomatous, with erect to geniculate culms, rooting at lower nodes, internodes cylindrical to compressed, solid to hollow, hirsute to glabrescent; nodes densely hairy to glabrous. Sheaths hirsute, with or without urticant tuberculate hairs to glabrous. Ligule membranous-ciliate. Blades lanceolate to linear-lanceolate, hispid to hirsute or glabrous, flat or with involute margins, base rounded to subcordate. Inflorescence an exserted, lax, open terminal panicle or terminal and axillary panicles numerous, forming a cluster in the upper third of the plant. Spikelets ovoid, obovoid to ellipsoid, open at maturity, solitary or in pairs, hairy to glabrous, pale to purplish; lower glume 1/2 to 3/4 the length of the spikelet, 5–9(−11) nerved; upper glume and lower lemma somewhat larger than the upper anthecium, 7–9(−11) nerved, rachilla conspicuous or not between glumes; lower palea elliptic or absent; lower  flower staminate or absent; upper anthecium ellipsoid, broadly ellipsoid to ovoid, glabrous, stipitate at base, smooth, shiny, indurate, palea with verrucose papillae at the apex. Caryopsis ovoid to broadly ovoid; hilum punctiform, embryo less than half the length of the caryopsis. It includes five species. The species of section Rudgeana inhabit open, dry, and sunny places; they are frequent in savannas of Central and South America, from sea level up to 1500 m. The species of this section are characterized by being cespitose, shortly rhizomatous plants, with a membranous-ciliate ligule and lanceolate to linear-lanceolate blades, flat or with involute margins. The inflorescences are lax and open, and there may be only terminal panicles, in P. cervicatum and P. ligulare, or terminal and axillary panicles, forming a set, in P. campestre, P. cayennense, and P. rudgei. The spikelets vary from hairy to glabrous, have a lower glume ½ to ¾ of the spikelet length, a subequal upper glume and lower lemma and a conspicuous stipe is present at the base of the upper anthecium; the stipe consists of two parts: a membranous portion toward the palea and another hardened, smooth, shiny portion toward the upper anthecium lemma. The membranous portion is reduced in P. campestre and P. cayennense (Fig. 2D), is larger in P. rudgei and even more conspicuous in P. cervicatum and P. ligulare; in the latter species, it extends into one or two wings covering the base of the upper anthecium (Fig. 2B).

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Key to the Species 1. Inflorescences terminal, lax; axillary panicles usually absent; spikelets 4.4–9 × 1.2–2.5 mm; stipe prominent, ca. 1 mm long 2. Stipe glabrous; sheaths papillose-pilose, with urticating hairs; spikelets 4.4– 5.7  mm long................................................................................53 P. ligulare 2. Stipe pilose; sheaths glabrous to pilose, without urticating hairs; spikelets 7–9  mm long..................................................................................52. P. cervicatum 1. Inflorescences terminal and axillary present, forming a cluster in the upper third of the plant; spikelets 2.1–3.5 × 1.1–1.3 mm; stipe 0.5 mm long or smaller 3. Spikelets 3–3.5  mm long; sheaths densely hairy, with stiff, urticating hairs ...........................................................................................54. P. rudgei 3. Spikelets 2.1–2.8 mm long; sheaths without urticating hairs. 4. Plants perennial; spikelets ovoid, sparsely hairy, 2.6–2.8 mm long.............. .......................................................................50. P. campestre 4. Plants annual; spikelets obovoid, glabrous, 2.1–2.6 mm long...................... .............................................................51. P. cayennense 50. Panicum campestre Nees ex Trin,, Gram. Panic.: 197. 1826. Type: Brazil, without locality, F. Sellow s.n. (holotype, LE-TRIN-0613.01). Fig. 52 Panicum rotundum Hitchc. & Chase, Contr. U.S.  Natl. Herb. 15: 139, Fig.  134. 1910. Type: Brazil. Minas Gerais: without locality, 1845, J.  F. Widgren s.n. (holotype, US 00148001; isotypes, BAA 00002405, fragment, BR 0000006883805, K 000309183, P 00740839, S14-5805) Perennial plants, usually with rigid adventitious roots, culms 30–75 cm tall, erect to geniculate, rooting or not at the lower nodes, branching basally; internodes cylindrical to compressed, 3–10  cm long, hirsute, with adpressed to glabrescent rigid hairs; nodes densely hairy, with long whitish hairs. Sheaths 3–7 cm long, usually shorter than the internodes, the basal ones longer, hirsute, covered with tuberculate, not urticant hairs, margins ciliate. Ligule membranous-ciliate, 1.5–2.5  mm long, with hairs beneath at the base of blade; collar pale to brownish. Blades linear-lanceolate, 7–25 × 0.5–0.7 cm, densely hirsute on both surfaces, flat or with involute margins, acuminate, base rounded to subcordate, margins scabrous and ciliate, midrib manifest. Inflorescence a lax terminal panicle and axillary panicles present, forming an oblong compound inflorescence; main axis sparsely hirsute, at least towards the basal portion, scabrous, flexuous, first-order branches alternate, divaricate, scabrous; pulvini pilose; pedicels long, scabrous. Spikelets ovoid, 2.6– 2.8 × 1–1.2 mm, sparsely pilose, pale to purplish toward the apex or purplish; lower glume 1.8–2.2 mm long, ½ to ¾ the length of the spikelet, acuminate to subulate, with long stiff hairs toward the apex, 5–9 nerved, midrib scabrous; upper glume and lower lemma subequal, 2.6–2.7  mm long, acuminate, 7–9 nerved, sparsely hairy

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Taxonomic Treatment

Fig. 52  Panicum campestre. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e–f) Lower palea. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Caryopsis, embryo view. (j) Caryopsis, hilum view. (From Sendulsky 637, SP)

on the inner surface and with or without long sparse hairs on the outer side; lower palea elliptic, 1.8–2  mm long to absent; upper anthecium broadly ellipsoid, 1.8–2 × 1 mm, pale, lemma 7 nerved, palea with compound papillae on its apex, stipe ca. 0.2 mm long. Caryopsis 1.2–1.3 × 1 mm; hilum punctiform, embryo less than half the length of caryopsis.

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Fig. 53  Distribution of Panicum campestre and P. cayennense

Distribution and ecology (Fig. 53). Species endemic to Brazil, where it grows from Pará and Bahia to the state of Paraná, in fields or cerrado on sandy or clay soils, from sea level to 1500 m asl, and flowers between December and May. Additional material examined. Brazil: Bahia: Serra Geral de Caetite, 9.5 km S of Caetite on road to Brejinho da Ametistas, Harley 21319 (CEPEC). Federal District: Brasília, Bacia do Rio Sao Bartolomeu, Equipe IBGE 2919, 3046, 6188, 6503 (IBGE); Planaltina, CPAC- Chapada entre Sobradinho e CPAC a direita da Rodovia, Almeida 355 (IBGE); BR-020 CPAC Planaltina, Almeida 1036 (IBGE); Recor, Filgueiras 997 (IBGE); Recor, Vizinhancas das instalacoes (herbarium), Equipe IBGE 1467 (IBGE); Reserva Ecologica do IBGE, Equipe IBGE 5946, 6937 (IBGE); University of Brasilia, 23 February 1965, Clayton 4795 (MO, NY, US); 4842 (MO, NY, US); E of Lagoa Paranoa, 10 December 1965, Irwin et al. 11181 (F, GH, MO, NY); 15  km  S of Brasilia, 28 August 1964, Irwin & Soderstrom 5700 (US); Sobradinho, Clayton 4875 (NY); Goiás: 6-7 km E of Alto Paraiso, 7 March 1973, Anderson 6515 (MO, NY); 26  km NE of Catalao, 23 January 1970, Irwin et al. 25210 (F, MO, NY, US); 75 km N of Corumbá de Goiás, 22 January 1968, Irwin et al. 19000 (F, GH, MO, NY, NY, US); 14 km S of Niquelandia, Irwin et al. 34386, 34387 (F, NY, US); 16 km N of São Joao da Aliança, Dawson 14442 (US);

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between Viannapolis and Ponta Funda, 17 March 1930, Chase 11315 (US); Serra do Rio Preto, 14  km E of Cabeceiras, 16 November 1965, Irwin et  al. 10354 (US); Corumbá de Goiás, Macedo 4482 (BAA, US). Luziania, ao longo da Rodovia BR-40, Filgueiras & Soejarto 238 (IBGE). Mato Grosso:1 km NE of Garapé, 1 October. 1964, Irwin & Soderstrom s.n. (US-2642542). Mato Grosso do Sul: Campo Grande, Chase 10790 (GH, RB, US), 4 July 1946, Swallen 9598 (US). Minas Gerais: Barbacena, Serra Mantiqueira, Chase 8645 (F, GH, NY, RB, US); Corinto, Fazenda Diamante, 8 April 1931, Mexía 5567 (F, GH, MO, NY, R, US); 11 km N of Gouveia, Anderson et al. 35584 (MO, NY); Faria, Serra da Bocaina, 6 January 1930, Chase 10533 (F, NY); 10  km  W of Barao da Cocais, Irwin et  al. 28827 (F, MO, NY, US); 7 km W of Campanha, 26 February 1976, Davidse et al. 10651 (MO, NY); 33 km NE of Francisco Sa, Irwin et al. 23071 (F, MO, NY, SP, US); 35  km  SW of Gouveia, Anderson et  al. 35148 (F, MO, NY); 9  km NE of Estiva, 25 February 1976, Davidse et  al. 10544 (MO); Ouro Preto, Chase 9354 (F, NY, US); 2 km S of Itacolumy, Irwin et al. 29364 (F, MO); Poços de Caldas, 18 January 1930, Chase 10637 (US); Piloes, Macedo 4876 (NY, US); lower slopes of Sierra da Piedade, Irwin et al. 28730 (NY); between Sucupira and Omega, Chase 11198 (USA); Oliveira, Chase 8856 (US); Lavras, Chase 8815 (US); Maia 18 (RB); Jardim, Widgren 908 (US); Lagoa Santa, Chase 8995 (US); Hermilo Alves, 14 February 1962, Duarte 6352 (MO, SI, US); Parque Municipal de Belho Horizonte, Occhioni s.n. (RB 44096); Passa Quatro, Sampaio 6249 (R). Pará: Marchajó ate Natal, Schwacke 62 (R); Fazenda Conceiçao, Rio Aura, Black 54-16109 (R). Paraná: Jaguariahiva, Dusén 16393 (F, GH, NY, US); 10074 (US); Swallen 8678 (US); 2 km W of Rio Itarare and road PR-11, Davidse et al. 11375 (MO, NY). Rio de Janeiro: Monte Serrat, Serra da Itatiaia, Chase 8358 (F, GH, MO, NY, US); Resende, Hoehne & Gert 17583 (GH, US); Campos, Serra de Itatiaia, Damazio s.n. (RB 203141). São Paulo: 8 km N of Avaré, 5 February 1965, Clayton 4526 (BAA, GH, MO, NY, SP, US); Campinas, Novaes 1269 (US); Ytu, Rusell 186 (US); Mandaqui, Usteri 9820 (SP, US); 10 km S of São Paulo, Parque do Estado, Sendulsky 208, 311 (SP, US), 637, 716, 725 (SP), Fonseca 13 (MO, NY, SP, US); 3 km from Cajurú, Sendulsky 126 (SP, US); 16 km NNE of Padua Sales, Eiten 1669 (NY); São José dos Campos, Lofgren 212 (RB) 51. Panicum cayennense Lam., Tabl. Encycl. 1: 173. 1791. Panicum cayennense Lam. var. patulum Döll, Fl. Bras. 2(2): 220. 1877., nom. illeg. superfl. Panicum rigiophyllum Döll, Fl. Bras. 2(2): 220. 1877, nom. inval. Type: French Guiana. Cayenne: Cayenne, D. Stoupy s.n. (holotype, P-LAM; isotypes, BAA 00003682, fragment ex P, BAA 00003683, fragment ex P-LAM, US 00148343, fragment ex P-LAM). Fig. 54 Panicum sessilicaule Desv., Mém. Soc. Agric. Angers 1: 199. 1831. Type. [United States of America]. “Habitat in Carolina,” without collector, s.n. (holotype, P 00740936; isotypes, BAA 00002410, fragment ex P, US 00139990, fragment ex P)

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Fig. 54  Panicum cayennense. (a) Plant. (b) Ligular region. (c) Spikelet, lateral view. (d) Spikelet, dorsal view. (e) Spikelet, ventral view. (f) Spikelet, ventral view without the lower glume. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Lower palea. (i) Caryopsis, hilum view. (k) Caryopsis, embryo view. (l) Upper palea, lodicules and gynoecium. (From Zuloaga et al. 4348, SI)

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Taxonomic Treatment

Panicum pedunculare Willd. ex Steud., Syn. Pl. Glumac. 1: 77. 1855[1853]. Type: “P. cayennense Nees. Agr. Bras. 195. Brazil.” (holotype not known, syntype, US 00139852, fragment) Panicum cayennense Lam. var. curtatum Döll, Fl. Bras. 2 (2): 220. 1877. Type: “extra fines in via inter Cayenne et Baduel (Yelski, inter plantas a cl.? Rostafinski benigne mecum communicatas).", 1866, C. von Jelski s.n. (holotype, W, n.v.; isotype, US 00148344, fragment ex KR) Annual cespitose plants, with erect or decumbent culms, usually branched at lower and middle nodes, 1.10  m tall, culms often zigzag, few noded; internodes compressed, hispid to glabrous, hollow; nodes dark, hairy. Sheaths 2–8 cm long, shorter or longer than the internodes, papillose-pilose, with dense tuberculate hairs, margins ciliate. Ligule membranous-ciliate, 0.8–1.6 mm long; collar pilose, pale. Blades linear-lanceolate, 5–28 × 0.4–1 cm, flat, acuminate, narrowed at base, hispid to glabrous, the margins ciliate, scabrous, midrib conspicuous. Terminal and axillary inflorescences present at the upper nodes, forming an elongate inflorescence 5–30 cm long, reaching about 2/3 of the total plant length; panicles with main axis wavy, flexuous, hispid towards the base, scabrous on the rest of the surface, pulvini pilose, the branches alternate to opposite, sometimes pseudoverticillate, divergent, branch axes flexuous, scabrous. Spikelets obovoid, 2.1–2.6 × 1.1–1.3 mm, globose, glabrous, pale or tinged with purple; lower glume 1.2–1.8 mm long, more than 1/2 the length of the spikelet, acuminate, hairy on the inner surface, 5 nerved, midrib scabrous toward the apex; upper glume 2.2–2.5 mm long, acute, 7 nerved; lower lemma 2–2.4 mm long, 7 nerved; lower palea elliptic, 1.6–1.9 × 0.6–1.1 mm, membranous, glabrous; lower flower absent; upper anthecium broadly ovoid, 1.5– 1.8 × 0.9–1.2 mm, pale, smooth, glabrous, shiny, shortly stipitate at base. Caryopsis broadly ovoid, 0.9  ×  0.7  mm, pale; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 53). It grows from Mesoamerica and the West Indies to South America, where it is found in Venezuela, Colombia, the Guianas, Bolivia, and Brazil. It is frequent in fields on sandy or clay soils. 0–1500 m asl and flowers all-year round. Panicum cayennense is related to P. campestre Nees ex Trin, a species that differs in being an annual plant, with larger, hairy spikelets; in P. cayennense the culms are short, branched and with numerous axillary inflorescences. Panicum floribundum A. Rich. ex Lam., Encycl. 4: 742. 1798, is a nomen nudum corresponding to this species. Additional material examined. Mexico. Campeche: about 43  miles SW of Mamantel in an area of tropical vegetation, 200 ft, 21 December 1972, Reeder & Reeder 6111 (MEXU, MO). Chiapas: Escuintla, 3 July 1937, Matuda 1799 (F, MEXU, MO, NY, US); Savannah near airport of San Quintín, along the Río Jataté, 1200 ft, 24 February 1965, Breedlove 9183 (MO); roadside pastures and savannas west of Catazaja on road to Villahermosa, 100 m, 10 November 1981, Breedlove & Davidse 55250 (MO). Oaxaca: Tuxtepec, 16 April 1968, Martínez Calderón 1676 (MEXU, MO, NY); Santiago de Jocotepec, Vera Santos 3372 (MO, US); along the

Key to Ungrouped Panicum Sections and Species

211

highway to Tehuantepec (route 185), km 17 carretera Las Limas-L. Bonita, 6 July 1964, Chavelas & Pérez 116 (MEXU). Tabasco: 21 km W of Cardenas, Conrad & Conrad 2959 (MO); km 15 de la desviación de Huimanguillo hacia Francisco Rueda, 12 July 1980, Cowan 3119 (MEXU, MO); km 55.8 de la desviaciónde Huimanguillo hacia Fco. Rueda, 40 m, 31 October 1979, Cowan 2590 (MEXU). Veracruz: Vicinity of Rio Tonto, 6 km W of Campo Experimental de Hule, Vera Santos 2275 (NY, US) Guatemala. Izabal: S of Rio Dulce, at Shell Station, Le Doux et al. 106 (NY); at Shell Station and vicinity just S of Río Dulce, 75 m, 8 August 1975, Le Doux et  al. 2125 (MEXU, MO). Petén: sabanas y bosques secundario de Santa Rita, 20 km al sur de Santa Elena, 10 November 1965, Molina 15533 (MO, US). Belize. El Cayo: Mountain Pine Ridge, San Agustin, July/August 1936, Lundell 6585 (F, NY, US) Honduras. Atlántida: Vicinity of Tela, Standley 54701 (US) Costa Rica. Alajuela: Grecia, Hacienda La Argentina, 26 July 1933, Valerio 601 (F, US), 605 (F). Puntarenas: 0.5  km  S of Buenos Aires, Pohl & Davidse 10761 (F); between San Antonio and Boruca, Pohl & Davidse 10979 (F); Buenos Aires, open grassland aroun a pon at airport, 385, 8 June 1978, Pohl & Gabel 13871(MO). San José: Buenos Aires, Tonduz 3685 (US); Valerio 1062 (F) Panama. Chiriquí: 5  miles  S of Boquete, McDaniel 6807 (MO); vicinity of David, 7 October 1911, Hitchcock 8372 (F, MO, NY, US). Panamá: near Arraijan, 21 July 1938, Woodson, Jr. 1402 (MO, NY, US) Cuba. Isla de la Juventud: near Nueva Gerona, 1 January 1904, Curtiss 267 (MO, US); vicinity of San Pedro, 15 February 1916, Britton et al. 14455 (MO, US); Isla de Pinos, Taylor 34 (MO, US). Oriente: Cayo del Rey, 25 October 1912, Ekman 10028 (US). Pinar del Río: Herradura, 15 August 1907, Tracy 9073 (US), 9093 (MO), 26 August 1910, Britton et al. 6520 (US); Sierra de Cabra on Guane road, 9 September 1910, Britton et al. 7275 (US); Laguna jovero and vicinity, Shafer 10510 (US) Jamaica. Halliss Savanna, Upper Clarendon, 30 November 1915, Harris 12226 (MO) Dominican Republic. Distrito Nacional: Sierra Prieta, Villa Mella, 12 September 1970, Liogier 17408 (US). La Vega: Vicinity of Pedra Blanca, 14 October 1967, Allard 16060, 16067 (US); Cordillera Central, Sabana de la March, El Valle, 14 Aur 1930, Ekman 15694 (US) French Guiana. Cayenne, Leprieur s.n. (MO, US); route de Rachombeau, Hoock s.n. (NY) Colombia. Cauca: Buenos Aires, Lehmann 5268, 5269 (US). Córdoba: Ayapel, Hacienda Simba, Fernández 11 (MO). Guainía: inmediately S of Casuarito, lajas along the Rio Orinoco, 5°40′ N, 67°37′W, 22 June 1984, Davidse & Miller 26378 (MO, SI). Magdalena valley, savanna, 11 September 1924, Allen 659 (MO). Guaviare: municipio San José del Guaviare, inspección La Fuga, sitio Filo de Hambre, Giraldo-Cañas 2603 (COAH, HUA). Meta: de Puerto Gaitán a Planadas, 1 km de la ruta principal, 280 m, 14 June 1989, Zuloaga 3978 (COL, SI); 10 km al E de Cumaral hacia Paratebueno, bosque de galería, 520 m, 18 June 1989, Zuloaga

212

 

Taxonomic Treatment

4093 (COL, SI). Vichada: región guayanesa, municipio Puerto Carreño, afloramientos rocosos del tipo “lajas” entre Punta de Lajas y el cerro El Bita, ribera del río Orinoco, 40–100  m, 4-5 January 2004, Giraldo-Cañas 3645, 3649 (COL). Without department: Magdalena valley, savanna, without date, Allen 659 (MO). Without department, locality or date, Mutis 5339, 5378, 5498, 6110 (US) Venezuela. Amazonas: Dept. Atures, Rio Orinoco, alrededores de Siquita, 19 July 1969, Bunting et  al. 3637 (MY); Puerto Ayacucho, Williams 13085 (F, US, VEN); near Capuana, Davidse & Huber 16811 (MO); dptos. Atures y Atabapo, alrededores de Santa Bárbara del Orinoco, Steyermark et al. 117119 (MO, VEN); 23 km NE of Puerto Ayacucho, Davidse & Huber 15430 (MO, VEN); Dpto. Atures, 3–6 km N of Sanariapo along the road to Puerto Ayacucho, Davidse et al. 16770 (MO). Anzoátegui: Pariaguaán, 1 October 1939, Muller s.n. (US); Rio Mapire, afluente norte del Rio Orinoco medio, Estacion Musinacio, Rosales & Valles 143 (MO); cabeceras del Rio Guaraguara, mesa de Guaripa, Luces 64 (VEN); Los Canos, Rio Caris, Pittier 14479 (VEN); Río Laisme, en la cienaga del Morichal, Luces 96 (VEN). Apure: Payarita, entre San Fernando y Achaguas, Ramia & Montes 4730 (VEN); en sabanas entre Achaguas y San Fernando, cerca de Bethel, 6 August 1989, Zuloaga et  al. 4348 (SI). Barinas: Sur de Pedraza, Ramia 1807 (VEN); costa del Río Suripa, Ramia 1757 (VEN). Delta Amacuro: Isla La Tórtola, Ramia 2167 (VEN). Guárico: 28 km N of Santa Rita, Davidse 4319 (MO, PORT). Monagas: 3 km E of Jusepín, Davidse et al 4548 (MO, PORT); camino a Cazorla, entrada al Hato Becerra, Blydenstein 204 (VEN) Suriname. In distr. Pará, Kappler 1495 (MO) Brazil. Amapá. Rio Pedreira, Fróes & Black 17322 (US); Macapá, Fazendinha, Black & Lobato 50-9659 (US). Amazonas: 2 km S of Labrea, Prance et al. 8177 (F, GH, MO, NY); km 27 of road Humaitá-Porto Velho, Prance et  al. 3517 (MO). Bahia: Col. Valença, Pinto 1021 (US). Goiás: 2 km SW of Araguaina, Eiten 10154 (US). Mato Grosso: 20 km S of Garapú, Irwin & Soderstrom 6485 (US). Mato Grosso do Sul: 100  km  W of Coxim, Bommer 54 (NY, US); Paiaguás, Fazenda Alvorada, Allem & Vieira 1001 (MO); Xavantina-Cachimbo road, W of km 229, Philcox et al. 3631 (NY, RB). Marãnhao: Barra do Corda to Grajahú, Swallen 3674 (RB, SP, US). Pará: Ilha de Marajó, Fazenda Gavinho, Goeldi 245 (F, US); Oriximina, Cachoeria Porteira, Davidson et al. 10692 (MO); Belém, Archer 7587 (US); Cuminá, Kuhlmann 1701 (US). Rondonia: 2–4 km E of Abuna, Prance et al. 8600 (MO, NY, R); 2–4 km E of Mutum Paraná, Prance et al. 8831 (F, MO) Bolivia. La Paz: Prov. Iturralde, Luisita, 13°05′S, 67°15′W, 180  m, Beck & Haase 9976 (SI). Pando: Provincia Federico Román, Río Orthon, 14 September 1985, Moraes 590 (SI). Santa Cruz: Buena Vista, Steinbach 6935 (BAA, F, GH, LIL, MO, NY, US); Ichilo, Montero to Puerto Grether, 25 km SE of Puerto Grether, 200 m, Renvoize et al. 3954 (K, MO); Ñuflo de Chavez, San Antonio de Lomerio, 16°45′S, 61°48′W, 500  m, 30 January 1985, Killeen 797 (SI); Ñuflo de Chavez, Rancho Corralitos and granite mountain near Zapoco, 90 km SW of Concepción, 16°30′S, 61°40′W, 450 m, 23 November 1985, Killeen 1486 (SI)

Key to Ungrouped Panicum Sections and Species

213

Fig. 55  Panicum cervicatum. (a) Spikelet, ventral view. (b) Spikelet, dorsal view. (c) Spikelet, lateral view. (d) Lower lemma, dorsal view. (e) Lower lemma, ventral view, enclosing the lower palea. (f) Upper anthecium, dorsal view. (g) Upper anthecium, ventral view. (From Morrone & Pensiero 529, SI)

52. Panicum cervicatum Chase, J. Wash. Acad. Sci. 32(6): 164, Fig. 10, 1942. Type: Brazil. Mato Grosso do Sul: Tres Lagoas, 4 February 1930, A.  Chase 10737 (holotype, US 00133136; isotypes, BAA 00000384, G 00099651, RB 00538803, SP 003133, US 00289442, WISv025420WIS). Fig. 55 Panicum olyroides Kunth var. denudatum Döll, in C. Martius, Fl. Bras. 2(2): 230. 1877. Type: Brazil: “in Brasilia occidentali,” Tamberlik s.n. (holotype not known; isotypes, W 0023590, W 0023591)

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Taxonomic Treatment

Panicum vinaceum Swallen, Fieldiana, Bot. 28 (1): 27. 1951. Type: Venezuela. Bolívar: Gran Sabana, between Kun and waterfall at Rue-Meru, south of Mount Roraima, 1,065  m, 2 October 1944, J.  A. Steyermark 59173 (holotype, US 00153814; isotype, F 0046933F) Plants perennial, 40–100  cm tall; culms erect, simple, occasionally branched; internodes 7–23 cm long, cylindrical, glabrous to sparsely hairy below nodes; nodes densely hairy to glabrous. Sheaths 7–13 cm long, lower sheaths larger than internodes, pale, densely hirsute to glabrous, the margins ciliate. Ligule membranousciliate, 0.6–2  mm long. Blades lanceolate, 16–36  ×  0.8–1.6  cm, rigid, base subcordate, apex acuminate, flat or with involute margins when dry, hispid to strigose or glabrous on both surfaces, margins scabrous, long ciliate, with tuberculate hairs, midrib manifest. Inflorescence lax, exserted, diffuse, terminal panicle, 25–60  ×  12–35  cm, spikelets in pairs; main axis scabrous, striate, first-order branches alternate or opposite, scabrous, pulvini hairy, pale; axillary inflorescences occasionally present; pedicels scabrous, 2–20  mm long. Spikelets ellipsoid, 7–9 × 2.1–2.5 mm, glabrous, pale to purplish; lower glume 3.5–3.8 mm long, acuminate, 7–11 nerved, midrib scabrous toward the apex; upper glume 6.7–8.4 mm long, sparsely pilose to glabrous, long pilose toward the base, 7–11 nerved, midrib scabrous; lower lemma glumiform, 6.2–7.3 mm long, pilose toward the base, purplish, 7–9 nerved; lower palea elliptic to obovate, 4–5.8 × 1.3–2.2 mm, membranous, the margins pilose; lower flower absent; upper anthecium ovoid to broadly ellipsoid, 4–4.5  ×  1.8–2.1  mm, 2.5  mm wide at maturity, brown; stipe ca. 1  mm long, pilose, rachilla pilose below the stipe. Caryopsis 2.8–3.2 × 1.5–2.2 mm. Distribution and ecology (Fig. 56). Species restricted to Venezuela, Bolivia, Brazil, and Paraguay, where it grows in sandy or sandy-clay savannas, in fields or cerrados, between 400 and 1300 m asl, and flowers between December and October. This species has a disjunct distribution, growing in savannas in Venezuela, Bolivia, and Amazonas states (corresponding to the material originally described as P. vinaceum) and in fields in Bolivia and Brazil. Additional material examined. Venezuela. Amazonas: Atures, alrededores de Puerto Ayacucho, 15 km al N, en ambos lados de la carretera hacia El Burro, 5 47′N, 67 30′W, 80 m, Huber 862, 1351, 2132 (MO, VEN), 961 (VEN); carretera Puerto Ayacucho-El Burro, Guinand 13 (VEN); Dpto. Atures, sabanas y bosques en la región de Rincones de Chacorro, 30 km al N de Puerto Ayacucho, 5 km al NE de Galipero, Huber 5268 (VEN); Isla del Ratón, 1 June 1940, Williams 13221 (F, US, VEN); Dpto. Atures, a 200 m al N de Puente sobre el río Galípero, a unos 30 km al N de Puerto Ayacucho, 5°50′N 67°31′W, 24 June 1984, 24 June 1984, Guánchez & Mercado 2813 (MO, VEN); Dpto. Atures, sabanas de Galipero, alrededores de Puerto Ayacucho, Garófalo 480 (VEN). Bolívar: La Gran Sabana, al S de El Dorado a lo largo del camino a Santa Elena, 890 m, 5 December 1973, Davidse 4929 (F, MO, NY, US); Sabanas de Santa Elena, suelo arenoso, 25 February 1946, Tamayo 2964 (MO, NY, USA, VEN); Distr. Roscío, sabanas en el valle del río Kukenan inferior, en la región de Campo Alegre, approx. 14 km al SW de San Ignacio de Yuruaní, 4°55′N, 61°12′W, 870  m, 23 June 1983, Huber & Alarcón 7587 (MO,

Key to Ungrouped Panicum Sections and Species

215

Fig. 56  Distribution of Panicum cervicatum, P. ligulare and P. rudgei

VEN); Distr. Roscío, sabanas y arbustales sobre glacis arenoso y vertiente rocosa approx. 7 km al N de Santa Elena de Uairén, approx. a la altura de la entrada al Hato La Divina Pastora, 4°41′N, 61°03′W, 850  m, Huber & Alarcón 7807 (MO, NY, VEN); Gran Sabana, Hato Divina Pastora, Tamayo 2907 (VEN); La Gran Sabana, alrededores de la quebrada de Jaspe, Garófalo et  al. 1230 (VEN); Distr. Roscío, sabana abierta en arbustal sobre banco de arena, en la región del río Yuruaní inferior, approx. 5 km al W de San Ignacio de Yuruaní, 5°00′N, 61°11′W, 850 m, 12 June 1985, Huber 10446 (MO, NY, VEN); 8 km al norte de Santa Elena de Uairén, Gran Sabana, 4°38′N, 61°03′W, 900 m, 14 August 1989, Zuloaga et al. 4444 (MO, SI) Brazil. Bahia: Road to Posse, 225 km SW of Barreiras, 12 April 1966, Irwin et  al. 14657 (MO, NY, SP, US); Espigão Mestre, 100  km WSW of Barreiras, Anderson et al. 36654 (F, MO, NY, US); Rio Roda Velha and highway BR-020, 8 April 1976, Davidse et  al. 12084 (MO, NY). Distrito Federal: Universidade de Brasília, Clayton 4809 (SP), 4839 (K, NY, SP); 20 km E of Brasília, 14 October 1965, Irwin et al. 9213 (F, MO, NY, US); Brasília, Belém 1970 (CEPEC); 15 km E of Brasília, 30 August 1964, Irwin & Soderstrom 5711 (F, MO, NY, US); 1 km W of Sobradinho, 5 December 1965, Irwin et al. 11077 (MO, NY, US); Brasília, entre UNY y Parque Flor, 18 April 1963, Pires et al. 9176 (F, SP, US); Brasília, Lago Sul,

216

 

Taxonomic Treatment

November 1988, Zuloaga 3837 (SI). Goiás: Rio da Prata, 6 km S of Posse, 7 April 1966, Irwin et al. 14509 (US); 1 km W of Veadeiros, Irwin et al. 12574 (F, MO, NY, US); 3  km  N of Cristalina, 2 March 1966, Irwin et  al. 13268 (F, MO, NY, US); 16 km SW of Goiás-Bahia border, 8 April 1976, Davidse et al. 12192 (MO, NY); between Jatahy and Rio Araguaia, Chase 11736 (US); vicinity of Annápolis, 25 March 1930, Chase 11519 (US); 38 km N of São Joao da Aliança, Dawson 14354 (US); 40 km W of Rio Verde, 2 April 1930, Chase 11713 (US); W of Santa Rita do Araguaia, 5 Abr 1930, Chase 11863 (US); between Viannápolis and Ponta Funda, Chase 11281 (US). Maranhão: Barra do Corda to Grajahú, 1 March 1934, Swallen 3648 (RB, SP, US). Mato Grosso: Rodovia Cuiabá-Santarem, Lemes 4125 (RB); Rondonópolis, Rio Paguba, Rondón 2566 (RB, USA); Diamantina, Weddell 3081 (USA). Mato Grosso do Sul: Xavantina-Cachimbo roada, 85 km from Xavantina, 2 June 1966, Hunt & Ferreira 5739 (NY, SP, US); NW of São Lourenço, Chase 11959 (US). Minas Gerais: Lagoa Santa, 14 February 1864, Warming s.n. (US); 26 km NE of Patrocinio, 29 January 1970, Irwin et al. 25582 (F, NY, SP); Serra do Cipó, 110 km NE of Belho Horizonte, Chase 9138 (F, GH, MO, NY, US); between Sucupira and Omega, S of Uberlandia, Chase 11167 (US); 3–4  km de Prata, Sendulsky 18 (SP), 37 (SP, US); Frutal, Valls 649 (US); Caldas, Regnell III 1369 (US); Pratinha, Dorsett 189b (US). Pará: Serra do Cachimbo, BR-163 CuiabáSantarém, km 823, 7 November 1977, Prance et  al. 24993 (MO, NY); Serra do Cachimbo, 15 December 1956, Pires et al. 6302 (BAA, US). Rondonia: Vilhena, 2 January 1979, Silva & Pinheiro 4101 (MO, NY). São Paulo: Moji-Guaçu, Mattos 12255 (SP); Cajurú, Sendulsky 169 (SP); Botucatú, Gottsberger 950-95B (SP); Casa Branca, Chase 10951 (US); Cabaceiras, Pickel 5887 (US); de Santa Rita a São Simao, Sendulsky 148 (US); 4 km SS de Paraguaçu Paulista, Clayton 4598 (K, SP, US); campo de Itirapina, Black 51-11072 (BAA, US) Bolivia. Santa Cruz: Santiago de Chiquitos, San Micerato, Cárdenas 4506 (US); Chiquitos, cerca de El Carmen, February 1950, Cárdenas 4503 (US) Paraguay. Amambay: Parque Nacional Cerro Corá, camino a Colonia Naranjaí, próximo a la naciente del Arroyo Aquidabán, 2 May 1992, Morrone & Pensiero 529 (SI) 53. Panicum ligulare Nees ex Trin, Gram. Panic.: 206. 1826. Type: [Brazil]. “V. sp. imperfectum Brasil (N. ab Esenb.),” C.  F. P. von  Martius 3800 (lectotype, LE-792.01, designated by F. O. Zuloaga, 1987: 475). Fig. 57 Panicum ligulare Nees, Fl. Bras. Enum. Pl. 2(1): 196. 1829, nom. illeg. hom. Type: Brazil. Pará: Habitat in campis prope Almeirim, C. F. P. von Martius 3800 [lectotype, M, designated by F. O. Zuloaga, Ann. Missouri Bot. Gard. 74(3): 475. 1987] Plants cespitose, 1.30–2  m tall, with thick adventitious roots and cataphylls woolly, culms erect, many noded; internodes 8–24 cm long, solid to hollow, hairy to glabrous; nodes brownish, hairy to glabrous. Sheaths 8–23  cm long, greenish to purplish, papillose-pilose, with deciduous urticating hairs, the margins ciliate. Ligule membranous-ciliate, 1.8–3.2 mm long, with long hairs beneath at the base of blade, collar pale, densely villous. Blades linear-lanceolate, 30–55 × 0.9–1.9 cm,

Key to Ungrouped Panicum Sections and Species

217

Fig. 57  Panicum ligulare. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e–f) Lower palea. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Upper anthecium, lateral view. (From Irwin 14904, US)

218

 

Taxonomic Treatment

flat, base subcordate, apex acuminate, densely pilose on both surfaces to glabrescent, the margins ciliate, scabrous, flat to involute, midrib manifest. Inflorescence terminal panicle, lax, diffuse, multiflowered, 47–65 × 15–30 cm; main axis striate, hairy toward the base, otherwise scabrous, first-order branches divergent, alternate to opposite, sometimes verticillate at base of inflorescence, pulvini hairy, pale to brownish; axillary inflorescences usually absent; pedicels claviform, 2–20  mm long, scabrous. Spikelets ellipsoid, 4.4–5.7  ×  1.2–1.6  mm, glabrous, greenish to purplish; lower glume 2.9–3.8 mm long, ½ to ¾ the length of the spikelet, subulate, shortly hairy toward the apex of inner surface, 5–7 nerved, midrib scabrous; upper glume 7–9 nerved; lower lemma glumiform, 4.1–4.9  mm long, acuminate, 5–7 nerved; lower palea elliptic, 3–3.3 × 0.9–1.5 mm, glabrous, membranous, whitish, the margins with or without short hairs; lower flower absent; upper anthecium ovoid, 2.5–3.2 × 1.1–1.5 mm, pale, palea with compound papillae at the apex; stipe conspicuous, glabrous, with two wings ca. 0.8–1.1 mm long, hardened portion 0.4– 0.7 mm long. Caryopsis ovoid, 2.4 × 1.3 mm. Local name: “Capim elefante” (Brazil) Distribution and ecology (Fig. 56). Species restricted to Brazil, where it grows from the states of Maranhão and Bahia to Mato Grosso, in fields, between 500 and 1100 m asl and flowers from March to October. Additional material examined. Brazil: Bahia: 150  km  SW of Barreiras, 15 April 1966, Irwin 14904 (F, MO, US); entrance to Lençois on BR-242, km 8, at land exit, Zuloaga et al. 9031 (SI). Distrito Federal: Barragem do Paranoá, closed, 20 May 1987, Filgueiras 1261 (IBGE, SI); Chapada da Contagem, ca. 20 km NE of Brasília, 16 August 1964, Irwin & Soderstrom 5166 (US), 28 October 1965, Irwin et  al. 9653 (F, MO, NY, US). Goiás: Chapada dos Veadeiros, Alto Paraíso, ca. 10 km between Alto Paraíso e Colinas, 5 September 1994, Filgueiras & Fonseca 2992 (IBGE, SI); 20 km N of Cristalina, Serra dos Cristais, 7 March 1966, Irwin et al. 13699 (F, GH, MO, NY, US), 13700 (F, MO, NY, US); 35 km NE of Catalao, 20 March 1968, Irwin et al. 21525 (F, US); Serra Dourada, Glaziou 22525 (US); vicinity of Goiás, 20 March 1930, Chase 11460 (F, GH, NY); 26–31  km  S of Goiania, 10 April 1976, Davidse et al. 12278 (MO). Maranhão: Carolina to San Antonio de Balsas, 20 March 1934, Swallen 4094 (US); Serra do Penitente, Miranda 128 (RB). Mato Grosso: between Rondonópolis and São Lourenço, Chase 11987 (US); Rio Turvo, 210 km N of Xavantina, 27 May 1966, Irwin et al. 16122 (F, MO, NY, US); Serra Azul, 77 km from Barra do Garças, Hunt 6075 (NY, US); Serra do Roncador, 86  km  N of Xavantina, 1 June 1966, Irwin et  al. 16386 (F, NY, US); Xavantina-Cachimbo road, 215 km from Xavantina, 27 May 1966, Hunt & Ferreira Ramos 5606 (NY, US), 8 km NE of Base Camp, 12°54′S, 51°52′W, Ratter et al. 2090 (NY, RB); Campos Novos, Kuhlmann 1745 (RB) 54. Panicum rudgei Roem. & Schult, Syst. Veg. [Sprengel] 2: 244. 1817. Panicum scoparium Rudge, Pl. Guian. 1: 21, table 29. 1805, nom. illeg. hom., non Lam., 1798. Panicum cayennense Lam. var. divaricatum Döll, Fl. Bras. 2 (2): 220. 1877, nom. inval., based on P. scoparium Rudge. Type: French Guiana. Without locality, J. Martin s.n. (holotype, BM 000835112; isotypes, BR 000000688738, USA 00139980, fragment and photo ex BM). Fig. 58

Key to Ungrouped Panicum Sections and Species

219

Fig. 58  Panicum rudgei. (a) Plant. (b) Ligular region. (c) Spikelet, lateral view. (d) Spikelet, ventral view. (e) Spikelet, dorsal view. (f) Spikelet, ventral view without the lower glume. (g) Upper anthecium, dorsal view and lower lemma. (h) Lower palea, lodicules and stamens. (i) Upper anthecium, dorsal view. (j) Upper anthecium, ventral view. (k) Upper palea, lodicules and caryopsis. (l) Caryopsis, hilum view. (m) Caryopsis embryo view. (From Abbott 16824, SI)

220

 

Taxonomic Treatment

Panicum pilosissimum Roth, Syst. Veg. (ed. 15bis) 2: 458. 1817. Type: Guyana. “Roth nov. plant Spec. Ms..... In Essequebo”, C.  H. Mertens s.n. (holotype, B_10_0248954; isotype, US 00139868, fragment ex B) Panicum rudgei Roem. var. brasiliense Raddi, Agrostogr. Bras.: 48. 1823. Type: [Brazil]. Rio de Janeiro. “.... in viciniis fluminis inhumirim, in locis silvosis et herbosis.”, G. Raddi s.n. (holotype, PI, n.v.; isotype, US 00139945, fragment ex PI) Panicum dasytrichum Spreng, Syst. Veg. [Sprengel] 1: 317. 1825. Type: “Panicum dasytrichum Spr. hirsutum Willd. herb. Hoffmansegg.”, J. C. von Hoffmannsegg s.n. (US 00148404, fragment ex B) Panicum rhigiophyllum Steud, Syn. Pl. Glumac. 1: 76. 1855[1853]. Type: Brazil. Bahia: “P. rigens Salzm. Hrbr. Bahia.”, P. Salzmann s.n. (holotype, P 00740790; isotypes, G 00099811, HAL 0133146, MPU 024595, MPU 024596, MPU 024597, MVFA 0000450, P 00740791, US 00147995, W 18890239844) Plants perennial, shortly rhizomatous, cataphylls hairy, culms decumbent, geniculate to erect, 0.30–1.30 m tall, often zigzag, branching at the upper nodes; internodes 5–15  cm long, stiff, hollow, cylindrical, densely to sparsely hairy; nodes hairy, covered with adpressed whitish hairs to glabrous. Sheaths 4–13  cm long, densely papillose-pilose with stiff, urticating hairs, margins ciliate. Ligule membranous-ciliate, 1.5–2 mm long, with long hairs beneath at the base of blades; collar pale, densely to sparsely hairy. Blades linear-lanceolate, 20–40 × 0.6–1.1 cm, acuminate, base straight or narrowed, flat or with margins involute, densely hirsute to glabrescent, the basal margins ciliate or not, otherwise scabrous, midrib conspicuous. Inflorescences terminal and axillary at the upper nodes of plant forming an elongate inflorescence cluster, comprising 1/3 or more of plant length; panicles lax, diffuse, 25–50 × 10–20 cm; main axis wavy, scabrous to hairy, pulvini long hairy or glabrous, pale to brownish, first-order branches alternate and divaricate, flexuous, branch axes scabrous to hairy; pedicels flexuous, hairy and scabrous. Spikelets ovoid, 3–3.5 × 1–1.2 mm, solitary, acuminate, greenish and tinged with purple to violaceous, sparsely hirsute with irregularly distributed rigid hairs; rachilla conspicuous between glumes, lower lemma and upper anthecium; lower glume 2–2.7 mm long, 2/3 of spikelet length, acuminate, with stiff hairs toward the upper portion to entirely covered with hairs, 3–5 nerved, midrib scabrous; upper glume 2.7–3 mm long, acuminate, hairy on the inner surface, 7–9 nerved, midrib scabrous; lower lemma 2.5–2.9  mm long, acute, with long hairs or glabrous; lower palea elliptic, 1.8–2.3  ×  0.5–1.1  mm, membranous, the margins shortly hairy; lower flower staminate; upper anthecium ellipsoid, 1.8–2.2  ×  0.8–1.1  mm, indurate, smooth, shiny, glabrous, palea with verrucose papillae toward the apex; stipe conspicuous, the membranous portion ca. 0.4 mm long, hardened portion ca. 0.5 mm long. Caryopsis 1.5 × 1 mm wide. Local names: “Penabo” (Guahiba language, Vichada), “ta” (Makuna language, Vaupés) (Zuloaga and Giraldo Cañas 2013) Distribution and ecology (Fig. 56). Species widely distributed from Mesoamerica and the West Indies to Bolivia and Brazil, between sea level and

Key to Ungrouped Panicum Sections and Species

221

1000 m asl. It inhabits open, dry places, in savannas, fields or enclosed areas, usually on sandy soils. Additional material examined. Mexico. Tabasco: km 35 de la desviación de Huimanguillo hacia Francisco Rueda, 16 November 1979, Cowan 2677 (MEXU, MO); Achotal, Balancan, 9 May 1939, Matuda 3087 (F, GH, MEXU, US); 5 km de la carretera que va a Mecatepec por el lado de la sabana de Francisco Rueda, 22 February 1982, Magana & Zamudio 746 (MEXU) Guatemala. Izabal: Montaña del Mico, 6 mi. S of Izabal, Steyermark 38581 (F); Santa Cruz, N of Lago Izabal, Steyermark 39673 (F)Belize. Cabbage Hall, 20 March 1967, Dwyer et al. 454 (F, MO); Swasey Brach, Monkey River, 12 January 1942, Gentle 3862 (F, GH, MO, NY, US); Machaca, Gentle 6893, 6923 (F, NY, US). Stann Creek: In pineland, Eve’s Pine Ridge, 5 January 1955, Gentle 8510 (MO); in pine ridge, Commerce Bight Pine Ridge, 16 March 1954, Gentle 8156 (MEXU). Toledo: Savanna between the foothills of the Maya Mountains at Chun Bank and the Southern Highway, 88°41-42  W, 16° 29-32 N, 70 m, 14 March 1987, Davidse & Brant 32456 (MO); in broken pine ridge, 12 miles, San Antonio-Punta Gorda Road, 10 March 1949, Gentle 6686 (MEXU) Honduras. Gracias a Dios: orilla del Río Dursuna, 21 February 1979, Nelson & Vargas 5021 (MO); orilla del riachuelo Aras Tinni, 35 km SO Puerto Lempira, 21 February 1979, Nelson & Vargas 4987 (MO) Nicaragua. Zelaya: entre Siuna y Limbaikán, 17 March 1971, Seymour 4977 (F, MO, NY) Costa Rica. Alajuela: Buenos Aires, January/February 1943, León 1184 (US), Tonduz 3679, 4875 (US); Los Palmares, Pittier 10588 (US). Puntarenas: Buenos Aires, 24 February 1972, Molina 27395 (F, MO, US); east of CIA, road to Buenos Aires, 31 December 1974, Pohl et al. 13116 (F, MO); Algo de los Mogos, camino a Rincón de Osa, 14 February 1986, Zamora et al. 1184 (MO); sabanas de Buenos Aires, cerca del río Sebror, 30 November 1983, Sánchez et al. 346 (MO). San José: vicinity of El General, Skutch 3065 (GH, MO, NY, US), Pittier 12064 (US); savanna pond SW of Buenos Aires, 400 m, January 1983, Gómez 19690 (MEXU, MO) Panama. Jaboga, 31 October 1917, Killip 4163 (US); Canal Zone, near Fort Randolph, 28 December 1923, Standley 28598 (MO, US), Perlas Archipelago, San José Island, Johnston 324 (GH, US) Jamaica. Clarendon Parish: in secondary tropical forest on the Mason River Field Station, ca. 3 miles NW of Kellits, Pruski 1560 (NY); Clarendon, Bull Head, Hart 1510 (NY); Bunker’s Hill Savanna, 26 September 1912, Harris 11170 (NY, US); Halliss Savanna, Upper Clarendon, 30 November 1915, Harris 12235 (MO, NY, US); James Hill, Upper Clarendon, 31 December 1917, Harriss 12845 (NY, US); Mason River Savanna, 275  miles NW of Kellits, 28 March 1965, Proctor 26301 (NY, US) Colombia. Amazonas: Corregimiento de Araracuara, trocha al Rio Yari, 18 April 1986, Galeano et  al. 904 (COL); Corregimiento de Araracuara, Aguirre Galviz 877 (COL). Antioquia: Morro Pan de Azúcar, Orozco et  al. 767 (COL). Boyacá: Muzo, 1000–1400  m, December 1859, Lindig 1067 (COL). Casanare: sobre los llanos Aguasclaras, April 1963, Saravia 2685 (COL); Rio Casanare,

222

 

Taxonomic Treatment

Esmeralda, 19 October 1938, Cuatrecasas & García Barriga 3840 (COL). Guainía: Corregimiento de San Felipe, Rio Negro, 28 September 1971, Pabón et  al. 224 (COL); caserío de Karanacoa, en el Rio Guainía, 12 October 1977, Espina et al. 179 (COL); Río Guainía, Puerto Colombia, Schultes et  al. 17936 (US). Guaviare: Municipio San José del Guaviare, serranía La Lindosa, afloramientos rocosos de Ciudad de Piedra, 250 m, November 1995, Giraldo-Cañas 2513 (COAH, COL, SI); municipio San José del Guavire, trocha Nuevo Tolima, en cercanías al Batallón “José Joaquín París”, 250  m, March 1996, Giraldo-Cañas 2576 (COAH, COL, HUA, SI); municipio San José del Guaviare, Puerta de Orión, 23 March 1997, López 2338 (COAH, COL). Huila: ca. 8 km W of Palermo, towards Santa Marchía, 700 m, 16 March 1985, Wood 4752 (COL). Meta: 15 km E de San Martín, al S del Caño Camoa, 11 September 1963, Blydenstein 1658 (COL, MO); Hato Horizontes, Blydenstein 969 (COL); 20 km SE of Villavicencio, 17 March 1939, Killip 34319 (COL); 7  km E de Cumaral, al S del Río Guacavia, Blydenstein 1570 (COL); 1/4 km SW de San Martín, Herrmann 11196 (COL); El Mico airstrip, before Rio Guejar, Philipson et al. 1329 (COL); 43 km NEE of Puerto López along road to Puerto Gaitán, Davidse 5106 (COL, MO); 50 km al E de Puerto López, 300 m, 14 June 1989, Zuloaga 3977 (COL, SI); Laguna Carimagua, al S de Orocue, Hilton s.n. (COL); Puerto López, Hacienda San Cayetano, Blydenstein 1706 (COL); Villavicencio, Apiai, Humbert 27815 (COL); Llanos de San Martín, La Serranía, Hoya del Río Ariari, Jaramillo et al. 1047 (COL); carretera de Bella Vista a Pinalito, Echeverry 2304 (COL); 73 km W of Las Gaviotas, Davidse 5390 (MO). Norte de Santander: La Motilona, hoya del Río de Oro, 20 May 1965, García Barriga 18723 (NY, US). Santander: entre Puerto Wilches y Puerto Santos, km 16, Killip & Smith 14859 (F, GH, MO, NY, US); Campo Capote, 6° 38′N, 73° 55′W, Nee & Mori 3783 (COL); Bajo Magdalena, Barrancabermeja, 3 January 1962, Uribe 3967 (COL). Tolima: El Convento, W of San Lorenzo, Pennell 3509 (F, GH, MO, NY, US). Vaupés: Alto Vaupés, alrededores de Miraflores, 2 February 1944, Gutiérrez & Schultes 731 (COL); Río Piraparana, Caño Teenema, 5 September 1952, Schultes & Cabrera 17192 (COL); Cerro de Circasia, 10 October 1939, Cuatrecasas 7201 (COL, US). Vichada: Gualandayas, 100  km E de Gaviotas, 30 December 1983, Wood 4192 (COL); Caño Urimica, Cabrera 2006 (COL); Territorio Faunístico El Tuparro, entre los Ríos Tomo y Tuparro, El Tapon, 27 April 1971, Daniel & Vasquez 64 (COL); 25 km E of Curamibo along unimproved dirt road between Las Gaviotas and Santa Rita, 28 December 1973, Davidse 5325 (COL, MO); sabanas entre el campamento y el Río Muco, 13 December 1971, Cabrera 1511 (COL); 10 km W of Las Gaviotas, 30 December 1973, Davidse 5367 (COL, MO, NY) Venezuela. Amazonas: Cerro Duida, Maguire 29424, 29060 (NY); Serranía Parú, 16 February 1951, Cowan 31486 (NY, US); 20 km S of Puerto Ayacucho, 2 November 1971, Davidse 2841 (MO); 5 km NE of San Carlos de Río Negro, 20 November 1977, Liesner 3703 (MO); 25 km S of Samariapo, 29 June 1975, Gentry & Berry 14600 (MO); Yavita, Williams 13879 (F, US); pie del Cerro Huachacamari, Huber 4990 (MO); El Manguito, 1 km N of Caño Caname, 10 May 1979, Davidse et al. 17482 (MO); alrededores de Canaripo, Huber 1981 (MO); 8 km S de Puerto Ayacucho, 13 April 1978, Davidse & Huber 14916 (MO). Anzoátegui: vicinity of

Key to Ungrouped Panicum Sections and Species

223

Santo Tomé, Chase 12841 (GH, US). Apure: end of the Galerías de Cinaruco, 28 February 1978, Davidse & González 14667 (MO); near the Rio Meta at Fundo El Algarrobo, 16 February 1978, Davidse & González 14217 (MO). Barinas: 16 km SW of the Merida intersection just outside of Barinas, 18 November 1971, Davidse 3182 (MO, NY). Bolívar: Sabanas de Santa Teresa, Tamayo 2808 (MO); Gran Sabana, S of Mt. Roraima, Steyermark 59429 (F, US); 0,5 km NE of Urimán, Steyermark & Wurdack 22 (F, NY, US). Monagas: E de Maturín, ca. caserío La Pica, Aristeguieta 4048 (F, MO, NY); Laguna Mosu, 12 km N de Capirito, Trujillo 14194 (F). Zulia: 60  km NW of Santa Barbara-San Carlos del Zulia, near Campamento El Rosario, de Bruijn 1473 (MO, NY, US) Trinidad and Tobago. O’Mearey Savanna, 18 June 1963, Soderstrom 1010 (US), Britton & Hazen 1563 (NY, US); Pitch Lake, 7 December 1912, Hitchcock 10083 (MO, NY, US); St. Joseph, 23 December 1912, Hitchcock 10181 (US); Piarco Savanna, S of Dabadie, Britton 688 (NY) Guyana. Tumatumari, Gleason 40 (GH); Waini River, April 1923, de la Cruz 3634 (F, GH, MO, US); Waramuri Mission, Horuka River, October 1922, de la Cruz 2576 (F, GH, MO, NY, US); Bartica, Hills Estate, 10 December 1919, Hitchcock 17191 (F, MO, NY, US); Kaieteur Falls, Potaro River, October 1923, de la Cruz 4478 (F, GH, MO, NY, US) Suriname. Zandery, Samuels 233 (GH, NY, US); Kwatta, 21 June 1944, Maguire 23912 (F, GH, MO, NY, US); Sanderijl, 14 November 1934, Archer 2745 (US) French Guiana. Vicinity of Cayenne, 31 May 1921, Broadway 348 (GH, NY, US); 10  km from St. Laurent, on road to Cayenne, 20 December 1954, Cowan 38943 (US) Peru. Loreto: Nauta, Río Marañón, 11 October 1980, Gentry et al. 29965 (MO); vía Nauta-Iquitos, 30 June 1979, Díaz & Jaramillo 1270 (MO) Brazil. Amapá: Araguarí River, 20  minutes downriver from Porto Platón, 16 September 1961, Pires et al. 50973 (F, GH, NY, US); Porto Platón, Silva 2782 (NY, RB). Amazonas: Cucuí, Rio Negro, Nascimento et al. 194 (MO); km 20 on road from Humaitá to Labrea, 27 November 1966, Prance et al. 3386 (F, MO, NY, US); Fortaleza Savanna, Rio Puciari, Prance 13791 (F, GH, NY). Bahia: Salvador, Chase 7883 (F, GH, MO, NY); 35 km E of Eunápolis, Harley 17285 (CEPEC, MO, NY); Marchaú, Belém & Pinheiro 2121 (CEPEC), 18 May 1985, Zuloaga et  al. 2464 (RB, SI, US); road BR-418, 16 km del cruce con BA-001, Mori et al. 9674 (CEPEC, NY). Mato Grosso: Serra Azul, 75 km S of Xavantina, 17 June 1966, Irwin et al. 17302 (F, MO, NY, US); 270  km  N of Xavantina, Ratter 2069 (NY); Serra do Roncador, 60 km N of Xavantina, 24 May 1966, Irwin et al. 15961 (F, MO, NY, US). Mato Grosso do Sul: Tres Lagoas, 4 February 1930, Chase 10745 (US). Pará: Santarém, March 1934, Swallen 3721 (US); Soure, Ilha do Marajó, 21 June 1934, Swallen 4974 (US); Acará, Thomé Assú, Mexía 5921, 5975 (F, GH, MO, NY, US); 73  km NE of Castanhal, 6 April 1980, Davidse et  al. 17939 (MO, NY); 17 km SE of Vigia, along road Pa-140, 30 March 1980, Davidse et al. 17610 (MO, NY). Pernambuco: vicinity of Recife, Chase 7675 (F, US), Poazeves, Pickel 3137 (US). Rio de Janeiro: Silvestre, Holway et al. 1116 (US); Merity, 20 km N of Rio de Janeiro, Chase 8465 (US). Rondonia: Porto Velho, Black & Cordeiro 52-15348

224

 

Taxonomic Treatment

(US). Roraima: Igarapé Agua Boa, Rio Mucajai, 22 January 1967, Prance et al. 4035 (MO); Sierra de Sururucu, 14 February 1969, Prance et  al. 9899 (F, MO); Boca da Mata, base de Serra Tepequem, 10 February 1967, Prance et  al. 4274 (MO). São Paulo: São Simão, Kuhlmann 4110 (US); 7 km de São José dos Campos, 4 October 1961, Eiten & Mimura 3351 (MO, US) Bolivia. Beni: 15  km  W de Guayamerín, camino a Riberalta, 12 April 1979, Krapovickas & Schinini 35068 (CTES, US). La Paz: San Carlos, Buchtien 32 (GH, MO, NY, US); Apolo, Williams 1020 (NY); San Antonio, December 1907, Buchtien 1159 (US). Santa Cruz: Noel Kempff Mercado National Park, 1–3 km E of Los Fierros encampment, 18 May 1995, Abbott 16824 (MO, SI)

Ungrouped Species 55. Panicum bartlettii Swallen, Publ. Carnegie Inst. Wash. 436: 346. 1934. Type: Guatemala. Petén: Uaxactum, 23 March 1931, H. H. H. Bartlett 12245 (holotype, US 00133098; isotype, MICH 1108732) Perennial plants, with decumbent, rooting and branching culms at the lower nodes, stoloniferous, then erect, with simple to branched culms at middle nodes, up to 100 cm tall, internodes 6–11 cm long, cylindrical, hollow, rigid, glabrous, nodes compressed, brownish, glabrous. Sheaths striate, commonly smaller than the internodes, glabrous or sparsely hairy, one margin short to long ciliate, the other glabrous. Ligule membranous, reduced, glabrous, auricles short, pilose to glabrous; collar brown, glabrous. Blades ovate-lanceolate, flat, herbaceous, 8–20 × 1–2.9 cm, glabrous or pilose with long, adpressed hairs on both surfaces, base subcordate and asymmetrical, pseudopetiolate, pseudopetiole brown, sparsely pilose with long hairs; apex acuminate, margins scabrous, the basal ones ciliate or not. Inflorescence terminal exerted, peduncle ca. 20 cm long, cylindrical, glabrous; panicle lax, diffuse, multiflowered, 15–30 × 12–30 cm, branches alternate, divergent and ascending, spikelets solitary and scattered on the branches; main axis cylindrical, glabrous, pulvini glabrous, branches and pedicels smooth, scabrous, pedicels long. Spikelets ellipsoid, acuminate, greenish, glabrous, 1.8–2 × 0.8–1 mm, with diminutive vesicles over the entire surface; upper glume and lower lemma subequal, exceeding the upper anthecium in length; lower glume ovate, ca. 0.9 mm long, 1/2 or slightly less than spikelet length, acute, 1 3 nerved, the nerves anastomosed toward the apex; upper glume and lower lemma (3–)5 nerved, scaberulous on the outer surface, shiny on the inner surface, lower glume separated by an internode from the upper glume, rachilla conspicuous between the upper glume and upper anthecium; lower palea absent, lower flower absent; upper anthecium ellipsoid, glabrous, 1.3–1.5  ×  0.6– 0.8  mm, smooth, shiny, whitish, with bicellular microhairs. Caryopsis obovoid, whitish; hilum punctiform, embryo 1/3 the length of the caryopsis.

Ungrouped Species

225

Fig. 59  Distribution of Panicum bartlettii and P. brevifolium

Distribution and ecology (Fig. 59). Species is restricted to Mexico, Belize, and Guatemala, where it grows on the margins of jungles, 0–700  m asl, and flowers between February and April. Panicum bartlettii is morphologically related to P. trichanthum Nees, a species widely distributed in America, the latter being distinguished by having reduced lower glume, 0.3–0.5 mm long, reaching ¼ or less the length of the spikelet, nerveless and with a truncate or obtuse apex; in addition, this species has a developed lower palea. Both species are phylogenetically related (Zuloaga et al. 2018) and have a still uncertain position within the tribe Paniceae. Additional material examined. Mexico. Campeche: Hacienda San Pablo, near Champoton, Collins 37 (US); Tuxpeña, 14 January 1932, Lundell 1181 (MO, US); 65 km south of Conhuas, in the Calakmul Regional Center, 17 March 1983, Cabrera et al. 4447 (MEXU, MO). Chiapas: near the junction of the Rio Perlas and Rio Jatataté at San Quintín and near Laguna Miramar, 200m, 17 March 1955, Sohns 1602 (MEXU, MO, P, US), 1651 (MEXU, US), 1714 (MEXU, US); at COFOLASA camp 24 km SE of Crucero Corozal, road to Boca Lacantum, 220 m, 25 February

226

 

Taxonomic Treatment

1985, Martínez 11359 (MO). Hidalgo: steep slopes at Hidalgo-San Luis Potosí border, km 343-344 on highway below Chapulhuacán, Moore Jr. 2885 (US). Oaxaca: Las Palmas, San José Chiltepec, on road to Tuxtepec, Schultes & Reko 565 (US). Quintana Roo: Coba, Lundell & Lundell 7657 (US); 6–10 km NE of Felipe Carrillo Puerto, on road to Vigía Chico, 3 March. 1989, Cabrera & Villanueva 16374 (MEXU); 3 km N of San José de la Montaña, on road N of Thomás Garrido, W of Chetumal, S of Hwy. 186, 9 May 1982, Davidse et al. 20245 (MEXU, MO); 8 km N of Unión, 110 km SW of Chetumal by road, 7 May 1982, Davidse et al. 20156 (MEXU, MO); 8 km south of San José, towards Tomás Garrido, 17 February 1981, Cabrera 1249 (MEXU); 15  km E of Ucum, by road to Ingenio Alvaro Obregón, 5 March 1980, Téllez 1678 (MEXU); 20–22 km W of Puerto Morelos, and 3–5 km W of Vallarta, 5 May 1982, Davidse et al. 20108 (MEXU, MO); 2 km N of Estero Franco, Chetumal-La Unión road, 9 March 1982, Cabrera et  al. 2100 (MEXU, MO); 12 km south of Ingenio Alvaro Obregón, 11 April 1980, Téllez 1982 (MEXU, MO); 9 km N of La Unión, 4 March 1980, Téllez 1671 (MEXU, MO); 14 km NE of Cobá, 3 February 1980, Téllez 1413 (MEXU, MO); 72 km south of Ucum, on the Chetumal-La Unión road, 24 April 1982, Cabrera & Cabrera 2494 (MEXU, MO). San Luis Potosí: mountains along the gravel road to Jalpan ca. 7–8 miles northeast of Xilitla, 27 March 1961, King 4346 (F, MO, NY); south of Ciudad Valles, road to Tamazunchale, km 409 on road to Mexico City, Beetle M-547 (US); Motzorongo, February 1892, Smith 585 (MO, US), 632 (US); Lejem, Mun. San Antonio, 16 April 1979, Alcorn 2766 (MEXU). Tabasco: W-W highway near Zapate Bobal, 12 March 1976, Calzada & Arellano 2230 (MO). Veracruz: vicinity of Río Tonto, 6 km west of Campo Experimental de Hule El Palmar, Zongolica, Vera Santos 2826 (US) Guatemala: Izabal: 1  km south of San Felipe, 13 March 1970, Harmon & Fuentes 2075 (MO). Petén: 6  km from Tikal village, north side, Tikal National Park, 21 April 1971, Ortíz 1693 (F, US); Belén, open field, behind school, 15 May 1986, Walker 1414 (MO); Tikal National Park, Tikal, Contreras 628 (G, US); Sayaxche, bordering airport, Lundell 17770 (US); La Libertad and vicinity, 20 February 1934, Aguilar 378 (MO, US); Tikal National Park, on Remate road south of Tikal, Lundell 15668 (G, US); along Rio Cancuen, between La Concordia and El Cambio, 23 April 1942, Steyermark 45889 (MO); Sayaxche, around airport, Lundell 17920 (US) Belize. Corozal: Tiger savanna, ca. 12 airline km W of Little Belize, 18°11′N, 88°17′W, 17 March 1987, Davidse & Brant 32539 (MEXU, MO, SI). El Cayo: River Bluffs, Bartlett 11452 (US); EL Cayo, Bartlett 12016 (US); Broken ridge, Gentle 9647 (US); Cayo Road, Dover Farm, Reyes 17 (US); vicinity of old lumber camp at Grano de Oro, 2 June 1973, Croat 23332 (F, MO); Little Coquericot, Belize river, 27 March 1933, Lundell 3868 (US), 3876 (F, US); Gracie Rock, Sibun river, 10 April 1935, Gentle 1601 (MO, US); ca. 3 mi. S of Grano de Oro on road between Millionario and L.  Flor, disturbed forest, 2 June 1973, Croat 23387 (MO); Vaca Plateau, above Blue Hole Camp, 14 August 1980, Whitefoord 2111 (MO). Orange Walk: road between Lamanai and San Felipe, 88°39′W, 17°46′N, 19 March 1987, Davidse & Brant 32704 (MEXU, MO). Toledo: Cero, near hill, Gentle 6971 (F, G, US); Union Camp, 13 May 1979, Whitefoord 1699 (MO)

Ungrouped Species

227

56. Panicum brevifolium L., Sp. Pl. 1: 59. 1753. Type: “Habitat in India” (LINN80.64, lectotype designated by W. D. Clayton & S. A. Renvoize, Gramineae (Part 3), Fl. Trop. E. Africa: 496. 1982; isolectotype, US 01164892) Panicum litigosum Steud, Syn. Pl. Glumac. 1: 89. 1855[1854]. Type: Brazil. Bahia: “Bahia, Brazil”, P. Salzmann s.n. (holotype, P 00740837; isotypes, G 00099654, K 000003647, K 000003648, K 000003649, MO-316357, MPU 024434, MPU 024435, P 00740786, P 00740787, US 00132947, US 00132948, W 18890239826) Annual plants, with decumbent culms, geniculate and rooting at the lower nodes, then erect, 0.20–1 m tall; internodes hollow, cylindrical, glabrous; nodes glabrous. Sheaths ciliate to glabrous, the margins hairy. Ligule membranous, shortly ciliate in the apical portion, ca. 1 mm long. Blades ovate-lanceolate, 4–9 × 1–2.3(−3.5) cm, the base amplexicaul, cordate, apex acuminate, flat, pilose to glabrous, margins scaberulous, the basal ones ciliate. Inflorescence a lax, diffuse, terminal panicle, (2.5– )4.5–16 × (2–)4–10 cm; main axis scabrous, pilose to glabrous, first-order branches divergent, flexuous; pedicels claviform, scaberulous. Spikelets ovoid, 1.4– 1.6(−2) × 0.7–0.9 mm, pilose; lower glume 4/5 or as long as the spikelet, hyaline, 1(−3) nerved, pilose to pubescent, acute, rachilla manifest between the lower and upper glume; upper glume and lower lemma subequal, membranous, 5-nerved; lower palea elliptic, 1.2–1.8 mm long, hyaline, glabrous; lower flower absent; upper anthecium ovoid, indurate, smooth, shiny. Caryopsis not seen. Distribution and ecology (Fig. 59). Species is widely distributed in the tropics of the Old World, being found in Africa and Asia. It has been introduced in America, where it grows in Brazil, in forest margins. This species is placed in a clade of non-Kranz species (Zuloaga et al. 2018), in which it is related to Panicum hirtum Lam. and P. glandulopaniculatum Renvoize, the latter growing in Africa. It is characterized by including annual plants, with ovate-lanceolate laminae, flat, hairy spikelets, with the lower glume 4/5 or as long as the spikelet. Additional material examined. Brazil, Bahia: Olivenca-Maruim highway, between kms 7–10, 19 May 1985, Zuloaga et al. 2473 (MO, RB, SI, US); without locality, Luetzelburg 6002 (NY) 57. Panicum haenkeanum J. Presl, Reliq. Haenk. 1: 304. 1830. Type: Mexico: “Hab. in Mexico,” without locality, T.  Haenke s.n. (holotype, PR, n.v.; isotype, US 00148559, fragment and photo ex PR). Fig. 60 Panicum costaricense Hackel, Oesterr. Bot. Z. 51: 428. 1901. Type: Costa Rica. Inter Buenos Aires et Térraba, 1891, H.  Pittier 3636 (lectotype, BR 0000006882815, designated by A. S. Hitchcock & A. Chase, Contr. U.S. Natl. Herb. 15: 134. 1910; isolectotypes, G 00227837, US 00148384, fragment ex W) Perennial plants, with decumbent culms, rooting and branching at the lower nodes, then erect, supporting on vegetation, up to 50 cm tall; internodes 3–13 cm long, cylindrical, hollow, papillose-pilose with short tuberculate hairs; nodes dark, compressed, glabrous. Sheaths 2–6 cm long, shorter than internodes, striate, hirsute,

228

 

Taxonomic Treatment

Fig. 60  Panicum haenkeanum. (a) Plant. (b) Ligular region. (c) Portion of the inflorescence. (d) Lower glume. (e) Spikelet, ventral view without the lower glume. (f) Spikelet, dorsal view. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Upper lemma, lodicules and gynoecium. (From Zuloaga et al. 4368, SI)

Ungrouped Species

229

with caducous tuberculate hairs, one margin shortly ciliate, the remaining membranous. Ligule membranous-ciliate, 0.5–0.7 mm long, with long hairs beneath at the base of the blade; collar densely pilose with whitish hairs. Blades lanceolate, 5–14 × 0.4–1.2 cm, flat, slightly asymmetrical and narrowed at base, shortly pseudopetiolate, apex attenuate, the abaxial surface sparsely hairy with adpressed hairs, abaxial surface shortly hirsute, midrib manifest on the abaxial surface, hispid toward the base; pseudopetiole light brown, hairy. Inflorescence terminal, exerted, peduncle cylindrical, 6–7 cm long, hirsute; panicle lax, oblong, 8–20 × 5–13 cm; main axis wavy, scabrous to hispid, pulvini densely pilose, first-order branches alternate to occasionally subopposite, ascending, spikelets in pairs, one short and the remaining long pedicellate, pedicels slender 2–15  mm long. Spikelets long ellipsoid, 2.3– 2.6 × 0.7–0.8 mm, sparsely pilose, with whitish hairs toward the margins of glumes and lower lemma; lower glume 1.2–1.4  mm long, 1/2 or more the length of the spikelet, ovate, acuminate, 3 nerved, separated from the upper glume by a conspicuous internode; upper glume and lower lemma subequal, acuminate, 5 nerved, upper glume covering the lower lemma with margins; lower palea reduced, hyaline, glabrous; lower flower absent; upper anthecium long ellipsoid, 1.5–1.8 × 0.6–0.8 mm, smooth, shiny, indurate, whitish, apiculate, with prickle hairs, stomata, and bicellular microhairs toward the apex, also with simple papillae toward the upper portion. Caryopsis not seen. Distribution and ecology (Fig. 61). It is present from Mesoamerica and in South America from Colombia and Venezuela to Bolivia and Brazil; it grows in forest margins, between sea level and 800 m asl. Panicum haenkeanum is a species of uncertain position within the tribe Paspaleae, having a basic chromosome number of x = 10 and being anatomically a non-Kranz taxon. The species is characterized as including perennial plants, with decumbent, rooting and branching culms at the lower nodes, a lax terminal panicle, hairy spikelets, with the lower glume ½ the length of the spikelet, 3 nerved, upper glume and lower lemma 5 nerved, reduced lower palea, lower flower  absent, and apiculate upper anthecium, with bicellular microhairs and simple papillae on its surface. Additional material examined. Belize. Toledo: lower slopes of Richardson Peak, Maya Mountains, directly N of the junction of Richardsoncreek and Bladen Branch, 16°34′N, 88°46′W, 300–620 m, 4 March 1987, Davidse & Brant 31985 (MO) Honduras. Gracias a Dios: bank of the Río Mocorón, village of Mocorón, 21 February 1979, Nelson & Vargas 5077, 5110 (MO, SI) Nicaragua. North Caribbean Coast: Comarca del Cabo, Waspan, 13 March 1971, Seymour 4665 (MO). Zelaya: at ca. km 47 on road from Puerto Cabezas to Rosita, ca. 5.3 km W of Río Wawa ferry, ca. 14°06′N, 83°35′W, 1 May 1978, Stevens 8561 (MO); along road between El Empalme and Limbaika, ca. 1.5  km  SE of Palmera, ca. 13°35′N, 84°20′W, 60 m, scandent in understory in dense evergreen forest, 24 February 1979, Stevens 12849 (MO) Costa Rica. Entre le Convento et Buenos Aires, 250 m, February 1891, Pittier 3661 (US); plaines du Cordoncillal, Pittier 3640 (US); buissons a Boruca, December 1891, Pittier 4626 (US, W). Guanacaste: along road to Hacienda Inocentes, 3 km

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Taxonomic Treatment

Fig. 61  Distribution of Panicum haenkeanum and P. hirtum

E of CIA, 200 m, scrambling in brush near a stream in open savanna, 10 December 1978, Pohl & Gabel 13689 (F, MO); 2 km E of Liberia, 24 October 1968, Pohl & Davidse 11330 (F); 1  km  W of Las Animas, 200  m, 4 December 1968, Pohl & Davidse 11537 (MEXU). Puntarenas: 1 km SE of Buenos Aires intersection, along the CIA, scrambling in brush, culms to 3 m long, 16 December 1978, Pohl & Gabel 13728 (F, MO). San José: vicinity of El General, January 1939, Skutch 4053 (MO) Panama. Chiriquí: vicinity of San Felix, December 1911, Pittier 5247 (US). Coclé: ca. 2 mi W of Nata, rocky slopes of low hill between Mata Palo & El Corteza, 27 December 1970, McDaniel & Tyson 14702, 14709 (MO). Herrera: 3 km W of Las Minas on road to El Toro, 23 January 1981, Systma & D’Arcy 3150 (MO); 4 mi S of Los Pozos, in deep shade on hillside, 17 January 1966, Tyson 2681 (MO). Panama: ca. 10 km SW of San Carlos along the Inter-American Hwy., 18 November 1975, Davidse & D’Arcy 10132 (MO); summit of Cerro Pelado, 1 km N of Gamboa, secondary tropical moist forest, 30 November 1973, Nee 8485 (MO); hills above Campana, 23 December 1938, Allen 1317 (F, MO); vicinity of Aguarubia, dry savannas, Killip 4282 (US); Sabinas, 2 miles east of Juan Díaz, Cornman 638 (US). Canal Zone: Ancon Hill, Canal Zone, Killip 4183 (US), Piper 52656 (US); Canal Zone, Pedro Miguel, 27 January 1918, Killip 4309 (US); Canal Zone, between

Ungrouped Species

231

Corozal and Ancon, December 1911, Pittier 2169, 2636 (US); Canal Zone, between Panama and Corozal, Hitchcock 9206 (US); Canal Zone, near Frijoles, Cornman 712 (US); Culebra, Canal Zone, Hitchcock 9168 (F, US); Pecora, Killip 4237 (US); San Jose Island, Perlas Archipielago, edge of forest, trailing, decumbent, Johnston 1108 (P, US); savannas 3 miles north of Orange River, Killip 4256 (US); vicinity of Bella Vista, Piper 5260 (US); Fort Clayton, 12 January 1966, Blum 2076 (MO) Colombia. Meta: Puerto Gaitán-Planadas road, 4 km from the central road, 26 March 1971, Pinto 1548 (COL) Venezuela. Apure: Distr. Pedro Camejo, bank of the rio Meta, ca. 19 airline km WSW of Paraquita, 6°14′N, 68°05′W, edge of savanna and gallery forest, 12 February 1978, Davidse & González 13989 (MO, VEN). Bolívar: Hato la Vergareña, Cerro Moro, Garófalo et  al. 450 (VEN); 10  km  SW of rio Aro along hwy. 19 between Caicara and Ciudad Bolívar, forest and Trachypogon-Curatella-Byrsonima savanna, in partial shade along edge of savanna and forest, 24 November 1973, Davidse 4477 (MO, PORT, VEN). Carabobo: carretera a Bejumo, Box 3889 (MY) 3937 (MY, VEN); La Mona, near Chirgua, 700 m, 1 January 1939, Alston 5944 (F, NY, P). Portuguesa: between Aparición and Ospino, 26 December 1925, Pittier 12011 (NY, US, VEN). Sucre: 7 km E of the Mochima hwy. intersection along hwy 9 between Cumaná and Puerto La Cruz, 16 December 1973, Davidse 5031 (MO, VEN); 5 km SW of Santa Fe along hwy. between Puerto La Cruz and Cumaná, 15 December 1973, Davidse 5008 (F, MO, VEN), growing in brushy areas in forest; between Guarita and Los Altos, in humid and shady places, Tamayo 2134 (VEN); in fields of Los Altos, Tamayo 2151 (VEN); Distr. Sucre, 8 km directly S of Santa Fe by road, 64° 24′W, 10°16′N, 230 m, rocky hill savanna with Trachypogon, Byrsonima and Curatella, 19 November 1981, Davidse & González 19046 (MO, US, VEN); between Barcelona and Cumaná, secondary road south of Santa Fe, 9 August 1989, Zuloaga et al. 4368 (SI) Brazil. Amazonas: Humaitá, Campo para norte da estrada 319, km 657, Januaryssen & Gemtchujnicov 373 (CEN, IBGE, SI, SP). Maranhão: Carolina, 18 March 1934, Swallen 3889 (CEN, RB, SP, US); Imperatriz, fazenda Victoria, Pires & Black 1710a (IAN), 6 August 1949, Pires & Black 1698a (IAN, US). Mato Grosso: Serra do Roncador, ca. 60 km N of Xavantina, 25 May 1966, Irwin et al. 16060 (MO, UB, US); Buraçao, Linha Telegrafica, Kuhlmann 1786 (SI); ca. 20 km S of Xavantina, 12 June 1966, Irwin et al. 17039 (MO, NY, US). Pará: Bahia do Alto Tocantins, Pires 1936 (IAN); Bom Jardim-Itaituba, Swallen 6940 (US). Rondônia: Rio Guaporé, en direçao a Serra Conceiçao, Black & Cordeiro 52-14754 (IAN); Rio São Miguel, afluente do Guaporé, Serra do Limoeiro, 20 June 1952, Black & Cordeiro 52-15178 (IAN, US) Bolivia. Santa Cruz: Ñuflo de Chavez, Estancia Los Cruces, 45  km NE of Concepción, August 1985, Killeen 1094 (F, SI); Ñuflo de Chavez, ca. 40 km south of Asención dos Guarayos, Hopkins et al. 214 (NY) 58. Panicum hirtum Lam., Encycl. 4: 741. 1798. Type: French Guiana. Cayenne: Cayenne, J. Martin s.n. (holotype, P-LAM 00740884; isotypes, G 00099620, P 00740885, P 00740886, US 00148570, fragment ex P-LAM). Fig. 62

232

 

Taxonomic Treatment

Fig. 62  Panicum hirtum. (a) Plant. (b) Ligular region. (c) Inflorescence axis with glands. (d) Spikelet, dorsal view. (e) Spikelet, ventral view. (f) Lower palea. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (i) Upper anthecium, lateral view. (j) Caryopsis, embryo view. (k) Caryopsis, hilum view. (From Strudwick et al. 4389, MO)

Ungrouped Species

233

Panicum axipilium Steud. Syn. Pl. Glumac. 1: 79. 1855[1853]. Type: French Guiana. “Guiana No. 15” (holotype, P 00731504; isotype, US 00148163, fragment ex P) Annual plants, culms densely branching and rooting at the lower nodes, then erect, (8–)15–30(−60) cm tall; internodes purplish, sparsely hispid, cylindrical, hollow, 1.5–7 cm long; nodes hairy. Sheaths shorter than internodes, 1–4 cm long, striate, papillose-pilose, more so  toward the upper portion, one margin ciliate, the remaining membranous. Ligule reduced, membranous, ca. 0.3 mm long, with hairs beneath at the base of blade. Blades ovate-lanceolate, 1.5–6  ×  0.5–1.5(−2) cm, shortly pseudopetiolate, flat, cordate and amplexicaul at base, apex acuminate, the adaxial surface sparsely papillose-pilose, abaxial surface almost entirely glabrous, basal margins long ciliate, otherwise scabrous. Inflorescences terminal, exserted or partially included in the upper leaves, peduncle up to 15 cm long, cylindrical, hispid to glabrous; panicles 2–8 × 2–3.5 cm, ovate, open, multiflowered; main axis hirsute to glabrous, wavy, pulvini brownish, glabrous, first-order branches ascending, at an angle of 45°, smooth, cylindrical, with conspicuous yellow glands 0.3–0.5 mm long; pedicels slender, glabrous. Spikelets in pairs, ovoid to ellipsoid, 1.2–1.5  ×  0.6– 0.7  mm, obliquely inserted into the pedicels and inclined downward, greenish, sparsely to densely hispid, in the latter case with long whitish hairs covering bracts; lower glume ca. 1.4 mm long, nearly as long as the spikelet, hyaline, ovate, 3 nerved; upper glume ovate, as long as or slightly shorter than the spikelet, 3–5 nerved, the apex slightly incurved; lower lemma as long as the spikelet, ovate, 3–5 nerved, apiculate; lower palea ovate-lanceolate, 1–1.2 × 0.4–0.5 mm, acute, hyaline, whitish, the margins ciliate; lower flower staminate, stamens 3; upper anthecium broadly ellipsoid, smaller than the upper glume and lower lemma, 0.9–1.1  ×  0.6  mm, planoconvex, shortly stipitate at base, whitish, indurate, smooth, shiny, with conspicuous bicellular microhairs and simple papillae on its surface; lemma 3 nerved, anthers 0.6 mm long. Caryopsis broadly ellipsoid, brownish, plano-convex, 0.7 × 0.5 mm; hilum punctiform, embryo less than 1/2 the length of the caryopsis. Distribution and ecology (Fig. 61). It is found in jungles and savanna edges at low altitudes from Mesoamerica to northern South America, occurring in Colombia, Venezuela, Ecuador, Guyana, Suriname, French Guiana, and northern Brazil, between sea level and 1000 m asl. It has been introduced in tropical Africa. Panicum hirtum is characterized by having glands in the inflorescence axes, spikelets arranged obliquely on the pedicels, with the lower glume as long as the spikelet and stipitate upper anthecium, smaller than the upper glume and lower lemma and with bicellular microhairs and simple papillae on its surface. It is a nonKranz species, unrelated to the genus Panicum s. str. In a recently published phylogenetic study (Zuloaga et al. 2018), the species is placed in an “incertae sedis” clade of the Paniceae, in which it appears related to other species possessing glands on the inflorescence, such as P. glandulopaniculatum Renvoize and P. heterostachyum Hack, African species; it is also related to P. brevifolium, the latter being differentiated by lacking glands in the inflorescence axes, spikelets not arranged obliquely on the pedicels and upper anthecium not stipitate. This clade also includes species of the genera Sacciolepis and Trichanthecium Zuloaga & Morrone.

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Taxonomic Treatment

Additional material examined. Belize. Stann Creek: Silk Grass Road, 13 January 1954, Gentle 8099 (F, G, MEXU); Toledo: Swasey Branch, Monkey River, 20 January 1942, Gentle 3883 (F) Nicaragua. Costa Caribe Norte: Waspan, along bank of Coco River, alt. 10–100 m, 19 January 1970, Atwood & Seymour 3546 (F) Colombia. Magdalena: Santa Marta, Smith 2261 (NY). Meta: Río Meta, Nueva Antioquia, forest, 28 October 1938, Cuatrecasas 4207 (COL, F), alrededores de Villavicencio, Molina & Barkley 53 (COL, US), January 1856, Triana 2179 (P); Villavicencio, Triana s.n. (COL 2270), 2179 (US). Vichada: Río Orinoco, Puerto Carreño, 23-24 October 1938, Cuatrecasas 4085 (F, US). Without locality, Lindig 1057 (P) Venezuela. Amazonas: 12.5  km  S of Puerto Ayacucho, 1 November 1971, Davidse 2806 (MO, VEN); Dpto Río Negro, near Cerro de la Neblina Base Camp which is on Río Mawarinuma, 140 m, 25 November 1984, Liesner 17268 (MO, NY, VEN); Dpto. Río Negro, Neblina Base Camp on the Río Mawarinuma, 0°50′N, 66°10′W, 4 July 1984, Davidse & Miller 26918 (MO, NY, VEN). Anzoátegui: San Joaquín, Lasser & Vareschi 4260 (VEN); Fondo El Samán, entre Quebrada El Onoto y el Río Cabrutica, 11 September 1984, Montes 2580 (MO). Barinas: Barinas, Angulo 1083 (VEN). Bolívar: Island at junction of Río Paramichi and Río Paragua, 4°12′N, 63°05′W, 500 m, Steyermark 90559 (NY, US, VEN); raudales de Maihia a lo largo del Río Paragua, 500  m, Steyermark 90508 (NY, US, VEN); San FélixCastillos de Guayana, Aristeguieta 5811 (MY, NY, PORT, VEN); Isla La Berraca near mouth of Rio Nichare in Río Caura, 250 m, 6°04′N, 6°02′W, 11 August 1985, Horner et al. 203 (MO). Carabobo: Upper Chirgua valley, Bejuma road, Box 3891 (VEN). Cojedes: Dtto. Pao, carretera entre Tinaco y El Baúl, Hato Paraima, Garófalo 171 (VEN). Guárico: Mesa de El Sombrero, Woronow s.n. (US), Pittier 12231 (G, NY, US, VEN); at intersection of Rio Orituco and road from Calabozo to Cazorla, 90 m, 4 November 1973, Davidse 3713, 3724 (MO, VEN); Parque Nac. Aguaro- Guariquito, Quebrada El Cascón, afluente del Río San Bartolo, 100  m, Delascio et al. 10053 (VEN). Lara: El Altar, Smith V5562 (MO); El Altar, al W de La Miel, Smith V5563 (VEN). Portuguesa: Dtto. Ospino, lateral de vía entre campamento autopista J.A.Páez y vía Ospino-La Aparición, 200 m, Ramirez Reyes 2402 (PORT). Sucre: Distrito Sucre, 8 km S directly S of Santa Fe by road, 19 November 1981, Davidse & González 19047 (F, MO, NY, US, VEN) Trinidad and Tobago. Piarco Savanna, S of Arauca, among shrubs in swampy land, 28 December 1912, Hitchcock 10363 (RB, US); without locality, Fendler 928 (P); six miles N of Arima, shady bank, 25 December 1912, Hitchcock 10310 (G, MO, NY, P, US, W); Dibe valley, 15 April 1920, Britton & Coker 1754 (NY, US); St. Joseph, hills near savanna, 23 December 1912, Hitchcock 10177 (MO, US); Port of Spain, St. George road, Hitchcock 10320 (US); Aripo road, near 1/2 mile post, on banks, 31 December 1926, Broadway 6450 (MO, US) Guyana. Moka Creek flood on Lethem road, 5 September 1963, Goodland 605 (US); basin of Rupununi River, near mouth of Charwair Creek, 1-4 November 1937, Smith 2398 (F, MO, NY, US); Waipa village, South Pakaraima Mountains, Maguire et al. 46249 (NY); St. Ignatius, Harrison 1339 (NY, US); Rupununi, Sand Creek,

Ungrouped Species

235

Wilson-Browne WB-61 (NY); Rupununi River, Karanambo, 2 September 1988, Maas et al. 7199 (US) Suriname. 9 km N of Lucie Rivier and 12 km W of Oost Rivier, 275 m, 16 July 1963, Maguire et al. 54208 (NY, P, SI, US), Irwin et al. 54530 (F, NY, P, US); 2 km below confluence of Oost Rivier, 6 July 1963, Maguire et al. 53979 (F, NY, P, US) French Guiana. Cayenne, 1859, Sagot 1338 (G, P, US); Petit Mitaraka, 2 km N of Mitaraka, Sastre 1639 (P, RB); Roche Koutou-Bassin du Haut Marouini, 17 August 1987, de Granville et  al. 9385, 9396 (MO, US); Haut Approuague, Cros Montagne, 26 July 1958, Hoock 302 (US). Without locality, Leblond s.n. (P), Leprieur s.n. (G, R) Ecuador: Oro: between Portovelo, gold mine near Zaruma, and El Tambo 600– 1000 m, 30 August 1923, Hitchcock 21253 (US); between Portovelo and El Tambo, 2 September 1923, Hitchcock 21320 (US) Brazil. Amazonas: Río Uaupés, 15 October 1947, Pires 677 (IAN, NY, US); Rio Negro, Uaupés, Mun. Amazonas, Rodrigues & Coêlho 4749 (NY). Ceará: Jaguarate, Cachorinha, Spruce 2687 (G, P). Maranhão: Santa Bárbara, SSE of Loreto, on shore of Rio Parnaíba, 7°30′S, 45°3′W, 200 m, 25 May 1962, Eiten & Eiten 4709 (F, NY, SP, US); BR 316, Caxias-Codó, km 98, Coradin et  al. 2659 (CEN); Carolina, Black & Pires 50-9734 (IAN); Ilha dos Botes, duas leguas abaixo de Carolina, Pires & Black 1951 (IAN); Ilha dos Botes, Rio Tocantins, Pires & Black 1991 (IAN, US); proximo a Tasso Fragaso, Miranda 130 (RB). Pará: Cacaval Grande, María Francisca, 9 July 1952, Black 52-15586 (IAN, US); Marajó Island, Río Caracara, June 1914, Goeldi 98 (NY, US); within 2  km downstream of Sete Varas, airstrip on banks of Rio Curua, borders of forest flooded at high water, Strudwick et al. 4389 (F, MO, NY, US); Cacanal Grande, June 1914, Goeldi 123 (US); Marajó Island, Estate Gavinho, Goeldi 223 (F, NY, US, W). Piauí: Mun. Buriti dos Lopes, Palmeira, Sucre 9236 (SI); ad Oeiras, Gardner 2457 (G, US, W). Roraima: Serra do Mel, Surumú, Ule 8027 (IAN, US); Bôa Vista, Black 51- 12507 (IAN, NY); Rio Branco, Sao Marcos, Black 51-13323 (IAN); Rio Branco, Fazenda Capela, Black 51-13176 (IAN, P); Bôa Vista, Black s.n. (IAN) 59. Panicum millegrana Poir., Encycl., Suppl. 4: 278. 1816. Type: French Guiana, Cayenne, J. Martin s.n. (holotype not known; syntypes, LE-TRIN-0830.01, P). Fig. 63 Panicum hirsutum Lam., Encycl. 4: 741. 1798, nom. illeg. hom., non Swartz, 1797. Type: French Guiana. Cayenne. Cayenne, J.  B. Leblond s.n. (holotype, P 00740762; isotypes, P 00740848, US 00148569, fragment ex P) Panicum patentissimum Desv. ex Poir., Encycl., Suppl. 4: 283. 1816. Type: Dominican Republic. Santo Domingo. Santo Domingo, A. N. Desvaux s.n. (holotype, P 00740970; isotype, US 00139845, fragment and photo ex P) Panicum rugulosum Trin, Gram. Panic.: 195. 1826. Type: Brazil. “in umbrosis”, January 1825, G. H. von Langsdorff s.n. (holotype, LE-TRIN-0934.02; isotypes, BR 000000883461, USA 00138947, fragment and photo ex LE; probable isotypes, NY 00414043, P 00740761)

236

 

Taxonomic Treatment

Fig. 63  Panicum millegrana. (a) Upper portion of culm and inflorescence. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, lateral view. (g) Upper anthecium, ventral view. (a, from Schinini 8445, CTES; b–g, from Martin s.n., LE)

Ungrouped Species

237

Panicum lasianthum Trin, Sp. Gram. 3(21): Table 245. 1830. Panicum rugulosum var. lasianthum (Trin.) Hack., Ergebn. Bot. Exp. Südbras. 1: 11. 1906. Type: Brazil. Without locality, G. H. von Langsdorff s.n. (holotype, LE-TRIN-0779.01; isotypes, MVFA 0000437, fragment ex US, US 00148227, fragment ex LE). Panicum dispersum Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér 6, Sci. Math., Seconde Pt. Sci. Nat. 3, 1(2-3): 282. 1834, Type: Brazil. Bahia: Ilheos, 1831, L.  Riedel s.n. (holotype, LE-TRIN-0679.01; isotypes, MVFA 0000423, fragment ex US, US 00148435, fragment ex LE, possible isotype, NY 00381737) Panicum rugulosum Trin. var. subvelutinum Döll, Fl. Bras. 2(2): 259. 1877. Panicum sellowii Nees var. subvelutinum (Döll) Henrard, Fl. Suriname 1(1): 387. 1943. Type: Suriname. Onoribo, H.  R. Wullshlaegel 1612 (holotype, BR 0000006884123) Panicum subglobosum Hack, Bull. Herb. Boissier, ser. 2, 4(3): 274. 1904. Type: Paraguay. Cordillera. Prope Cordillera de Altos, 1885-1895, É. Hassler 640b (holotype, W 19160022667; isotypes, BAA 00002416, fragment ex G, G 00009729, G 00030305, NY 00414058, NY 00414059, US 00140032) Panicum rugulosum Trin. fo. effusum Hack., Repert. Spec. Nov. Regni Veg. 6(2126): 343. 1909. Type: Paraguay. Caaguazú. In viciniis Caaguazú, É. Hassler 9107 (lectotype, W 19160024240, here designated; isolectotypes, G 00004997, G 00004998, G 0000499, G 00005000, G 00009728, MO-1330597, W 19060001551, W 19060001552) Panicum multinodosum Swallen, Phytologia 14(2): 83. 1966. Type: Brazil. Pará. Bragança, 27 June 1934, J. R. Swallen 5037 (holotype, US 00132991) Perennial plants, rhizomatous, with culms decumbent, rooting and branching at the lower nodes, supporting and climbing on the surrounding vegetation, then erect, 0.70–1.50 m tall; internodes 3–20 cm long, cylindrical, subwoody in the basal portion, hairy to glabrous; nodes purplish, hairy to glabrous. Sheaths 1.8–15 cm long, shorter than the internodes, papillose-pilose with deciduous tuberculate hairs to glabrous, margins membranous, ciliate; collar glabrous to densely pilose. Ligule reduced, 0.2–0.4 mm long, membranous and shortly laciniate in the upper portion, with hairs beneath at base of the blade; collar densely pilose, with whitish hairs, to glabrous. Blades ovate-lanceolate, 6–14 × 1–3.8 cm, densely pilose, with strigose pilosity, to glabrous, base subcordate and slightly asymmetrical, adaxial surface with tuberculate hairs toward the basal portion, margins scaberulous, the lower ones ciliate with deciduous tuberculate hairs. Inflorescence lax, diffuse, pyramidal, multiflowered terminal panicle, 10–40 × 6–30 cm; peduncle up to 6 cm long; main axis scabrous, pulvini hairy to glabrous, brownish, first-order branches opposite or alternate, pedicels slender, 1.2–7  mm long, divergent from the branches. Spikelets obovoid, 1.6–2.5 × 0.9–1.2, open at maturity, hirsute to shortly hairy or glabrous, greenish, brownish at maturity; lower glume 1-1.7 mm long, 1/3 to 1/2 or slightly more the length of the spikelet, 1(−3)-nerved, acute to attenuate; upper glume 1.7– 2.2 mm long, not completely covering the upper anthecium, obtuse, 5 nerved, with a scabrous midrib in its upper third; lower lemma as long as the spikelet, 5 nerved; lower palea ovate-lanceolate, reduced, 1–1.5  ×  0.4–0.6  mm, membranous,

238

 

Taxonomic Treatment

glabrous; lower flower absent; upper anthecium ovoid, 1.6–2.2 × 0.9–1.2 mm, glabrous, indurate, brown, transversely rugose and with bicellular microhairs over its entire surface, apex acute and incurved. Caryopsis obovoid, 1.1–1.3 × 0.8–1 mm; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 64). Species widely distributed from Mexico, Mesoamerica, and the Caribbean, in South America from Colombia and Venezuela to Paraguay and Argentina. It is frequent in jungle edges and roadsides or in cultivated fields. It grows between sea level and 2000 m asl. Local names: “Paja peluda” (Venezuela), “yacuapi-li” (in specimen Montes 186); “kapi’í mbokaja’í” (kapi’í = straw, mbokaja = palm, i = small) (Zuloaga 1994) Panicum millegrana is a species related to P. sellowii Nees (Zuloaga 1979; Zuloaga et al. 2018), both taxa sharing obovoid, globose spikelets with a reduced lower palea and transversely rugose, brownish upper anthecium and with bicellular microhairs. Panicum sellowii is distinguished by including plants with lanceolate blades, 6–13  ×  0.7–1.4  cm, few flowered panicles, 6–22  ×  2.5–11, with naked branches at the base and congested spikelets at the upper end of the branches. In describing P. rugulosum Trin. fo. effusum, Hackel cites two syntypes as belonging to this form: Hassler 9107 and 9107a. Of these, Hassler 9107 from W

Fig. 64  Distribution of Panicum millegrana

Ungrouped Species

239

was selected as the lectotype of the form, being a complete specimen and corresponding to the protologue. Panicum pilosum Rupr. Ex Galeotti, Bull. Acad. Roy. Sci. Bruxelles 9: 239. 1842 and Panicum sellowii Nees var. longevaginatum Rupr., Bull. Acad. Roy. Sci. Bruxelles 9: 239. 1842, are nomen nudum corresponding to this species. Additional material examined. Mexico. Chiapas: along road to Pueblo Nuevo Solistahuacán, 5  miles south of Jitotol, Mun. Jitotol, 19 August 1965, Breedlove 11920 (F, US); above Finca Cuxtepec, Mun. Angel Albino Corzo, 1380  m, 11 August 1981, Breedlove 52108 (MO); Mun. of Cintalapa, 23 km west of Las Cruces along road to La Mina Microwave Station, 870 m, 30 October 1981, Breedlove & Davidse 54125 (MO); near Córdoba, open clay soil bank, 26 August 1910, Hitchcock s.n. (F, MO); 14 km north of Tuxtla Gutiérrez on road to Cañón Sumidero, 1150 m, 10 October 1976, Brunken & Perino 327 (MEXU). Veracruz: 3  km  SSW of Zongolica along gravel road to Texhuacán, 18°39′N, 97°00′W, 1350 m, 8 February 1984, Nee & Taylor 29458 (F). Without locality, Schiede s.n. (P) Guatemala. Guatemala: along Hwy. 9, at edge of Guatemala (city), weedy field, cinder soil, 1500  m, 26 August 1988, Stevens & Martínez 25218 (MO). Huehuetenango: between San Andrés and San Marcos, Sierra de los Cuchumatanes, 1300–1500 m, 3 September 1942, Steyermark 51834 (F). Quetzaltenango: lower south-facing slopes of Volcán Santa María, between Finca Pirineos and Los Positos, between Santa María de Jesús and Calahuaché, 1300–1500  m, 8 January 1940, Steyermark 33708 (F). San Marcos: between Finca El Porvenir to “Número 6”, lower south-facing slopes of Volcán Tajumulco, 1300–1500  m, 7 March 1940, Steyermark 37138 (F). Santa Rosa: Aguacaliente, Kellerman 7743 (F) Honduras. Morazán: along Quebrada El Gallo above El Jicarito, 900–1000 m, 12 August 1949, Standley 22483 (F); El Zamorano, Standley 22020 (F); along Río Caparrose, above El Zamorano, 900 m, September 1948, Standley 12634 (F); along road from El Zamorano toward San Antonio de Oriente, 900 m, 2 September 1949, Standley 23356 (F); aguas abajo de la Quebrada El Gallo al norte de Zamorano, drainage of the Río Yeguare, 1000 m, 12 August 1949, Molina 2591 (F) Colombia. Cundinamarca: Mun. Pandi, vereda El Caracol, 29 July 1981, Díaz Piedrahita 3292 (COL). Magdalena: Rincón Hondo, Magdalena Valley, Allen 249 (MO); Sierra Nevada de Santa Marta, at the edge of forest between Quebrada Botella and finca Los Arroyitos, 1700 m, 27 September 1972, Kirkbride Jr. 2224 (MO) Venezuela. Amazonas: Dpto. Río Negro, Neblina massif, Canyon Grande, along the Río Mawarinuma between the mouth of the canyon and the first major fork of the river, ca. 7 airline km ENE of Puerto Chimo, July 1984, Davidse & Miller 27303 (MO, VEN). Barinas: 79 km from Barinas along road to San Cristóbal, bank of Bumbun River, 16 July 1964, Breteler 4039 (NY, US). Bolívar: San Pedro de las Dos Bocas, west bank of river, forest along river, adjacent savanna and slopes, 200 m, 24 July 1978, Liesner & González 5479 (MO, VEN); woods in vicinity of “Los Patos”, 2 km SE of Los Patos, N of río Hacha about 15 km, 30 km S of El Manteco, 1,200 m, Steyermark 86911 (VEN); 21 km E of río Caura along hwy. 19 between Caicara and Ciudad Bolívar, savanna with clumps of large trees, 24

240

 

Taxonomic Treatment

November 1973, Davidse 4467 (MO, PORT); 14 km al sureste de Upata, hacia Villa Lola, 200 m, 16 August 1989, Zuloaga et al. 4493 (MO, SI, VEN) Guyana. Wanama River, Northwest District, May 1923, de la Cruz 4045 (F, MO, NY, US); Orealla, Jenmann 197 (US), 6676 (NY) Suriname. Carolina, 13 December 1934, Archer 2892 (US); 9 km N of Lucie, 12 km W of Oost River, Maguire et al. 54237 (NY); near Auca, km 1, JodensavanneMapanekreek area, July 1961, Boerboom 8769 (NY, US); near Biegieston, Marowjine River, Lanjouw & Lindeman 1955 (NY) French Guiana. Matabon, Cayenne, 18 July 1921, Broadway 638 (NY, US); St. Madelaine Rd., Cayenne, Broadway 11 July 1921, 733 (NY), 818 (NY, US); Moulin Vidal, Cayenne, Hoock 119 (P, US); Montagnes de la Trinité, Inselberg NW, de Granville et  al. 6153 (MO, NY, US); route de Saut Sabbat, environs de Morpio, Hoock 129 (NY, P) Brazil. Amapá: 4 km a Norte-Nordeste da ponte do Lago Curiau em direçao a Rio Pedreira, Valls 11678 (CEN). Amazonas: Desaperta, Parana do Ramos, 18 March 1924, Kuhlmann 1667 (SI, US); Falls of Madeira, Rusby 233 (US). Bahia: Maracas, Rodovia Maracas-Contendas do Sincora (BA 026), Região de Mata de Cipó, 14 February 1979, Silva et al. 223 (CEPEC, K, MO). Ceará: Parna, proximo a Cachoeira do Cafundo, Fernandez 4225 (IBGE); Linha telegrafica da Serra Grande, Lofgren 295 (R); Baturité to Guaramirango, 30 April 1934, Swallen 4416 (NY, R, US). Distrito Federal: Sobradinho, De Paula 1766 (IBGE). Goiás: Chapada dos Veadeiros, 10 km S of Alto do Paraiso, 21 March 1969, Irwin et al. 24846 (US); 75 km N of Corumbá de Goiás on road to Niquelandia, Irwin et al. 19032 (NY, US); near Goiabeira, between Annápolis and Goiás, 1000 m, 20 March 1930, Chase 11398 (US). Maranhão: Caxias to Barra do Corda, Swallen 3520 (P, US), 3486 (US); Barra do Corda to Grajahú, Swallen 3622 1/2 (R, US); São Luis to São José, Swallen 3461, 3462 (US). Mato Grosso: 75 km de Barra do Garças, Hunt 5999 (US); Campos Novos, Kuhlmann 2690 (RB); entre Parecís e S. Antonio, Linha Telegrafica, estrategicas de Mato Grosso ao Amazonas, Kuhlmann 1755 (SI); Rio Papagaio, Kuhlmann 2689 (RB); Fazenda São Geraldo, Allem & Vieira 1988 (CEN). Mato Grosso do Sul: Mun. Mondo Novo, Porto Renato, Hatschbach 48993 (K); Mun. Ponta Porá, Ponto Alto, Hatschbach 46148 (G, K, NY, US); Campo Grande, Chase 10810 (RB, US); between Campo Grande and Dourados, 400–450 m, Chase 10916 (US); vicinity of Dourados, Chase 10979 (US); Corumbá on Rio Paraguay, Chase 11122 (US); Fazenda De Caranba, Mun. Dourados, Estrada Maracaju 35 km N from Dourados, 700 m, Mizoguchi 572 (MO). Minas Gerais: Serra do Cipó, Chase 9125 (US); Serra do Espinhaço, 48 km W of Montes Claros, road to Agua Bôa, Irwin et al. 23982 (US); Serra do Curral, Southeast of Belho Horizonte, 900–1000 m, Chase 8943 (F, MO, NY, US); Serra do Cipo, Skvortzov 304 (UB); Serra do Itabirito, ca. 50 km SE of Belho Horizonte, 1,500 m, Irwin et al. 19740 (MO). Pará: Aramarai, Rio Tapajós, Swallen 3221 (K, R, SP, US); Marajó Island, near Soure, Goeldi 295 (F, MO, NY, SP, UB); Mun. Marapanim, Campo de Matapiquara, ca. 1  km east of Matipaquara, 73  km NNE of Castanhal by road, Davidse et  al. 17897 (MO, NY, US); Marajó, Schwacke 507 (RB); Mosqueira, Swallen 4891 (SP); Obidós, Swallen 5077 (RB), Black et  al. 57-19425 (IAN);

Ungrouped Species

241

Campina do Mangaba, Martins Pinheiro, Coradin 80 (IAN, MO); Belem, Pires & Black 355, 409 (IAN); Bolterra, beira de capoeira, Black 47-1866 (IAN); Belem, Rio Guajara, Black 54-16161 (IAN); camino de Jubum, para Cordeixa, Ilha do Marajo, Black 48-3536 (IAN); Santarém, Spruce 11 (G, P), 12 (NY, P), 677 (P), Swallen 3255 (IAN, R, RB, SP), 6914 (US); Mun. Tucuruí, Breu Branco, km 40 south of Reprêsa Tucuruí along Hwy. BR 422, Plowmann et al. 9697 (MO). Paraná: Mun. S.  Mateus do Sul, Rio Potinga, Hatschbach 13802 (K); Mun. Palmeira, Corrego da Anta, Hatschbach & Pedersen 35890 (K, NY); Mun. Rio Branco do SulItaperucu, Dombrowski 2455 (K); Curitiba, Capado da Imbuia, Dombrowski 2368 (K); Bocaiuva do Sul, Clayton 4263 (K, SP, US); Curitiba, 900 m, Jonsson 16 a (F), Dusén 16a (G, MO); Paraguai, Baixo Parana, proximo ao Rio Acarai, Sendulsky 1869 (SI); Mun. Iratí, Riozinho da Cima, 9 km south of Irati, 800 m, Smith & Klein 14951 (US); Mun. Uniào da Vitoria, Rio Espingarda, Hatschbach 4399 (US); Serrinha, Dusén 3759 (IAN, R); Tamanduá, Dusén 7688 (MO, US); 2  km E of Cascavel along Highway BR-277 to Curitiba, 670 m, Davidse et al. 11261 (MO, NY); Tres Barras, 760 m, Dusén 17606 (MO); 8 km NE of the Paraná-Santa Catarina border at the Rio Negro, 820 m, Davidse et al. 11020 (F, MO); Mun. Palmeira, Rio Caniu, 8 January 1980, Hatschbach 42679 (MO). Pernambuco: Tapera, Pickel 2275, 2280 (USA); Dois Irmaos, vicinity of Recife, 10–20 m, Chase 7729 (F, MO, NY). Rio Grande do Sul: Caxias do Sul, Vila Oliva, Valls 261 (CEN); Caracol, Rambo 28796 (US); Cruz Alta, Lindman 1477 (US); Caxias do Sul- Conceiçao, Wasum et al. 5471 (MO, US); Morro Santana, Mattos 3587 (SP); Santa Rita, prope Farroupilha, Rambo 45722 (SI); Vacaria, Arzivenco s. n., BLA 12003 (SI); Paso do Inferno, cerca de Canela, Brescia & Marchesi 4068 (P); Carazinho, BR 285, km 231, Valls 2562 (CEN, CTES); Farroupilha, Rambo 40254 (US); Sapucaia, Rambo 42068 (MO); Mun. de Montenegro, S. Salvador, Orth 2717 (MO). Rio de Janeiro: Itatiaia, Holway 1824 1/2 (US); Petrópolis, Goes & Dionisio 122, Otavio s.n. (RB); Petrópolis, Vale do Bo Bonsucesso, Sucre 2192, Braga 48 (SI); just S of Petrópolis, open roadside cliff in montane forest, 530 m, Davidse et al. 11415 (MO); Corcovado, 465 m, Chase 9741 (US); subida antiga para Petrópolis, Sucre 8578 (RB); Without locality, Widgren 1213 (US). Roraima: Rio Mucajai, Black 51-13354 (IAN); Bôa Vista, Magalhaes 3 (IAN); Mun. Bonfim, margem da estrada para o Cantá, 2°50′N, 60°35′W, Silva 608 (MO). Santa Catarina: Brusque, moista shady banks Azambuja, Smith & Reitz 6038 (R, SI); Mun. Lajes, N of Rio Caveiras, Smith & Klein 16093 (US); Mun. Pôrto Uniào, between Fazenda Frei Rogério and Rio Pintado, southeast of Pôrto Uniào, Smith & Klein 10774 (NY, R); Mun. Xanxeré: Pinheiral, 9 km east of Xanxeré, Smith & Klein 11832 (NY, R); Mun. Xanxeré, 8 km north of Abelardo Luz, Smith & Klein 11467 (NY); cercanías de Florianópolis, Zuloaga et al. 1880 (MO, SI, US); Tijucas, Reis 32 (US); 19 km SW of Lages along Highway BR-116 to Porto Alegre, 960  m, Davidse et  al. 11132 (MO, NY). São Paulo: Anhembi, Fazenda Barreiro Rico, Bokermann s n. (SP); 50 km de Avare, na rodovia Avare-São Manuel, Mattos 14535 (SP); Campinas, Estrada Souza-Pedeira, Vera Carnielo 6762 (RB); Capital, Pickel 4640 (US); Cotia, 34 km SW of Center of City of São Paulo, Sendulsky 661 (SI); Horto Florestal, Pickel 4566 (SP); Iguapé, Morro das Pedras, Brade 9163 (IAN, SI); Itapira, Hoehne s. n. (SP, US); Paraguaçu Paulista, 6 km N

242

 

Taxonomic Treatment

of City of Paraguaçu Paulista, Clayton 4561 (SP); Prata, Holway 1717 (US); Parque do Estado, Sendulsky 565 (US); 34 km SE ciudad, Sendulsky 668 (US); São Paulo, Reserva Biologica, Parque Estadual das Fontes do Ipiranga, Sendulsky 1477 (SI); 1 km W of Rio San Mateus, Clayton 4578 (K, SP); Tremembe, Holway 1571, 1615 (US); Villa Augusta, Holway 1594 (US); Mun. de São Carlos, Fazenda Canchim, ca. 8 km NE of São Carlos, Eiten et al. 3157A (MO) Bolivia. Chuquisaca: Hernando Siles, 12  km E de Monteagudo, 1200  m, 11 March 1981, Renvoize & Cope 3897 (K, LPB, MO). Santa Cruz: Santa Cruz, Ñuflo de Chavez, Estancia San Sebastián, 25  km  S of Concepción, 22 February 1987, Killeen 2320 (F, MO, SI); Ñuflo de Chavez, Concepción, Estancia San Josesito, 16°10′S, 62°00′W, 1 June 1985, Killeen 966 (F, MO, SI); Santa Cruz, Velasco, 5 km E of Santa Rosa de la Roca, 15°40′S, 61°30′W, 390 m, 27 January 1986, Killeen 1675 (MO) Paraguay. Alto Paraná: Alto Paraná, Irala, Montes 11057 (US); Puerto Bertoni, Bertoni 5849 (G, W). Amambay: in silvis in regione cursus superioris fluminis Apa, Hassler 8332 (G), 11444 (G, K, P, US). Caaguazú: bords de L’Uirangua, au NE de Caaguazú, Balansa 47 (P); Ruta 2, km 120, 8 February 1989, Zardini & Velázquez 10516 (MO). Central: Lago Ipacaray, Fiebrig 561 (G, P); Asunción, Balansa 48 (G, K, P); prope Sapucay, Hassler 12462 (NY); Asunción, Monte del Jardín Botánico, Burkart 18380 (SI, US); Lago Ipacaray, Hassler 11462 (P), 11589a (MO), 12455 (F, G, MO, NY, US); Tarumandy, Schinini 6195 (CTES, G); environs of Remanto Castillo, along the Paraguay river, about 20 km south of Asunción, 16 January 1974, Conrad 2255 (MO). Concepción: in regióne cursus superioris fluminis Apa, Hassler 8332 (G, MO, NY, P), 11444 (F, G, K, NY, P, US), 11444a (G); Río Apa, Hassler 11644 (P). Cordillera: Salto Piraretá del río Piribebuy, Vázquez Avila 195 (SI, US); San Bernardino, Marchesi 4383 (US); Emboscada, Itá Trompo, 23 February 1987, Bordas & Schmeda 4253 (G, NY, US); Eusebio Ayala, Schinini 2474 (CTES, G); Tobatí, “Ybytú Silla” mesa, southern area, 300  m, Zardini & Velázquez 26697 (MO); Caacupé, Rosengurtt B-5794 (F, K, MO); prope Cordillera de Altos, Hassler 640 (G, P). Guairá: Carobeni Nuevo, camino a Itapé, Rosengurtt B-5897 (MO, P); Villa Rica, Joergensen 4085 (F, LIL, MO, NY, SI); prés de Villa Rica, Balansa 48a (K, P), 48b (G); Cordillera de Ybytyruzú, road MelgarejoAntena, 6 km N of Antena, 25°45′S, 56°15′W, Zardini 13449 (MO); without locality, Balansa 2941, 2942 (P). Misiones: Estancia La Soledad, 2  km de Santiago, Schinini & Vanni 26047 (CTES, G); San Ignacio, Rosengurtt B- 5772 (P); 10 km N de Ayolas, Schinini & Vanni 25990 (CTES.  K). Paraguarí: Caapucú, Barreritos, Anderson 1156 (US); Idaparay- Pirayú, Schinini 4285 (CTES, F, G, K); Cerro Hu, prés de Paraguarí, Balansa 4355 (P); Piraretá-Piribebuy, Schinini 4393 (CTES, G); Palacios Mountain, 25°25′S, 57°10′W, Zardini & Velázquez 13193 (MO); Parque Nacional Ybycu’í, Campo cerrado on Arroyo Mina basin, 5 km N of administration area, Zardini 11843A, 11852 (MO); Cerro Mbatoví, Zardini & Soria 3894 (MO). San Pedro: Alto Paraguay, Primavera, Woolston G.29, G.62 (K, NY); Azucarera Tebicuary, Pavetti & Rojas 9714 (US) Argentina. Corrientes: Concepcion, Tabay, Arbo et al. 976 (CTES, SI); 8 km ruta 17, camino a Concepción, Paso Crucecita, 30 March 1975, Schinini et al. 11044

Ungrouped Species

243

(CTES, SI); Empedrado, Estancia Las Tres Marías, 24 February 1977, Pedersen 11679 (CTES, G, SI); San Martin, Cerro Nazareno, 14 February 1979, Schinini et al. 17098 (CTES, SI); Yapeyú, costa del Río Uruguay, 12 February 1979, Schinini et  al. 16978 (CTES, SI); Santo Tomé, Garruchos, Cabaña San Juan Bautista, 12 April 1974, Krapovickas et al. 25132 (CTES, SI); Dpto. San Martín, Yapeyú, orillas del Río Uruguay, 9 February 1978, Cabrera & Sáenz 29041 (SI, US); Dpto. San Martín, La Cruz, Tres Cerros, 13 April 1976, Cabrera et  al. 27694 (MO, SI); Estancia Santa Teresa, Pedersen 2591 (CTES, MO, P, US); Dpto. Mburucuyá, Estancia San Juan, Pedersen 4818 (G, P, US); Dpto. San Miguel, Loreto, 8 March 1974, Schinini 8445 (CTES). Misiones: La Mina San Juan, Montes 186 (F); Apóstoles, de Azara a Concepción de la Sierra, 10 km de Azara, Zuloaga et al. 1899 (MO, SI, US); Dpto. Cainguás, Ruta 14 pasando Campo Grande, Zuloaga et al. 661 (SI, US); Candelaria, Santa Ana, camino a las Ruinas Jesuíticas, Romanczuk et al. 507 (SI); Santa Ana, camino al Balneario Municipal, 11 April 1985, Zuloaga et al. 2219 (MO, SI); Iguazú, Arroyo Tupicuá, 27 January 1983, Guaglianone et al. 1039 (MO, SI); Iguazú, puente sobre el Arroyo Uruguay, Guaglianone et al. 977 (MO, SI); Posadas, Ekman 648 (US); San Ignacio, camino al Peñón del Teyucuaré, 11 April 1985, Zuloaga et al. 2267 (MO, SI); San Ignacio, Montes 15319 (F, SI); San Pedro, Montecarlo, Montes 15390 (SI); Gobernador Roca, Schwarz 1979 (G); San Ignacio, Puerto Viejo, Scharwz 2285 (G, US); San José, Bertoni 2710 (G); San Ignacio, Montes 1933 (CTES, F, G, US); San Ignacio, Montes 14889 (MO) 60. Panicum sellowii Nees, Fl. Bras. Enum. Pl. 2(1): 153. 1829. Type: Brazil. Without locality, F.  Sellow s.n. (holotype, B_10_0383435; isotypes, BR 0000006883799, HAL 0082593, L 0819922, MVFA 0000882, fragment ex LE, MVFA 0000883, fragment ex B, US 00139991, fragment ex LE). Fig. 65 Panicum beyrichii Kunth, Révis. Gram. 2: 231, table 27. 1830. Panicum rugulosum Trin. var. glabrescens Döll, Fl. Bras. 2(2): 259. 1877. Type: “Panicum sellowii Nees ab Esenb. in herb. Beyrich. Crescit in nemoribus prope novum Friburgum Brasiliae”. Brazil. Rio de Janeiro, Nova Friburgo, December 1822, H. C. Beyrich s.n. (holotype, B_10_0366161; isotype, .US 00148179, fragment ex B) Panicum valenzuelanum A. Rich., Hist. Fis. Cuba, Bot. 11: 304. 1850. Type: Cuba. “Crescit in locis montosis partis occidentalis insulae Cubae Vuelta de abajo dictae, Valenzuela s.n.”, J: M.  Valenzuela s.n. ex R. de la Sagra 6 (holotype, P 00740965; isotypes, P 00740966, P 00740967. P 00740968, US 01231580, fragment and photo ex P) Panicum probandum Steud, Syn. Pl. Glumac. 1: 76. 1855[1853]. Panicum puberulum Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 3, 1(2-3): 277. 1834, nom. illeg. hom., no (Link) Kunth, 1829. Type: Brazil, without locality, no collector s.n. (type, USA) Panicum rugulosum Trin. var. hirtiglume Griseb., Cat. Pl. Cub: 233. 1866. Type: Cuba: without locality, C.  Wright 3455 (holotype, W 18890123384; isotypes, BM 000938702, G 00099731, G 00099803, GH 00135364, GH 00135407, K 000309177, MO-321035, NY 00803745, S04-319, S04-320, US 00148002, US 00139949, fragment ex GOET)

244

 

Taxonomic Treatment

Fig. 65  Panicum sellowii. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Detail of a unicellular macrohair. (f) Lower palea. (g) Upper anthecium, dorsal view. (h) Upper anthecium, ventral view. (a, from Schinini 9356, CTES; b–h, from Sellow s.n., B)

Ungrouped Species

245

Panicum rugulosum Trin. var. condensatum Hack, Repert Spec. Nov. Regni Veg. 6(21-26): 344. 1909. Type: Paraguay. Caaguazú: in dumetis pr. Caaguazú, É . Hassler 9051a (holotype, G 00009721) Plants perennial, shortly rhizomatous, with culms leaning on the vegetation, up to 3 m long, rooting and branching at lower nodes, then erect, 40–80 cm tall; internodes 3–12 cm long, cylindrical, rigid, hairy to glabrous; nodes hairy to glabrous. Sheaths 2–6 cm long, smaller than the internodes, papillose-pilose, with deciduous tuberculate hairs to glabrous, the margins ciliate. Ligule 1.5–2.5 mm long, membranous and shortly ciliate at apex; collar shortly pilose. Blades lanceolate, 6–13 × 0.7– 1.4  cm, flat, base subcordate and slightly asymmetrical, shortly pilose on both surfaces or with the abaxial surface glabrous, basal margins ciliate, with deciduous tuberculate hairs, otherwise scabrous. Inflorescence a terminal ovate panicle, lax, 6–22 × 3–11 cm; main axis puberulose to scabrous, pulvini pilose, brownish, lower branches opposite, the rest alternate, divergent, branch axes and pedicels shortly pilose to scabrous, spikelets congested on short pedicels at the top of the branches. Spikelets obovoid, 1.6–2.4 × 1–1.2 mm, pilose, with short, stiff hairs, to glabrous; lower glume 0.8–1.8 mm long, ovate-lanceolate, acute, 1–3 nerved; upper glume covering or not the apex of the upper anthecium, obtuse, 5 nerved; lower lemma as long as the spikelet, glumiform, 5 nerved; lower palea reduced, ovate, 1–1.5  ×  0.4  mm, hyaline, glabrous; lower flower absent; upper anthecium long ovoid, plano-convex, 1.8–2 × 0.9–1.1 mm, brown, slightly rugose transversely and with bicellular microhairs and simple papillae all over its surface, apex acute, incurved. Caryopsis broadly ellipsoid, 1.3  ×  0.8  mm, light brown; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 66). It is found from southern Mexico, Mesoamerica, and the West Indies to Argentina; it has been introduced in Asia and Africa. It frequently inhabits forest edges, in humid and shady places where it supports the vegetation, 200–1900 m asl. Local names: “Kapi’í kokito” (kapi’í = straw, kokito = coquito, Zuloaga 1994); “guardasireno,” “ilusión golondrina,” “ilusión de hojita ancha,” “paja ilusión” (Colombia) Species related to P. millegrana (see observations under the latter species); both have obovoid spikelets, with variable pilosity, usually greater in P. sellowii and less in P. millegrana, with a reduced lower palea, lower flower absent, and upper anthecium transversely rugose, the  upper anthecium  with bicellular microhairs on its surface. Additional material examined. Mexico. Chiapas: Mun. de Ixtapa, near Ixtapa, 915 m, 1 November 1981, Breedlove & Davidse 54319 (MO); 18–20 km north of Ocozocoautla along road to Mal Paso, 800 m, 27 September 1972, Breedlove 28117 (MO); 3–5  km above Jaltenango along road to Finca Prusia, 900  m, 11 October 1974, Breedlove 38598 (MEXU). Veracruz: ca. 4 km SSW of Xalapa, 10 October 1978, Iltis et al. 946 (F); 6 km (by road) ESE of Ixhuacán de los Reyes, 19°20′, 97°04′W, 1500 m,, 30 October 1981, Nee 22491 (MO); about 27 miles north of El

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Taxonomic Treatment

Fig. 66  Distribution of Panicum sellowii and P. trichanthum

Palmar Rubber Experiment Station, on the road to Cordoba, 550  m, 15 August 1953, Reeder & Reeder 1979 (MEXU) Guatemala. Alta Verapaz: lowland forest in valley, 2 1/2  miles west of Cubilgüitz, 250–300 m, 28 February 1942, Steyermark 44244 (F). Chimaltenango: San Martín, 14 October 1938, Johnston 1614 (F). Izabal: near Quiriguá, hilly pine forest, 26 April 1939, Standley 72464 (F) Belize. El Cayo: Mountain Pine Ridge, San Agustín, Lundel 6557, 6734 (F). Stann Creek: in high ridge, on hillside, 19 miles Section, Stann Creek Valley, 23 September 1957, Gentle 9339 (F) El Salvador. Morazán: Cerro Cacahuatique, Finca San Pedro, 1050  m, 13 September, 1951, Rohweder 1996 (MO) Honduras. Distrito Central: Llamapa, s.w. of Talanga, Rancho Santa Regina de Archaga, 1400  m, 6 January 1978, Pohl & Gabel 13521 (F, MO). Francisco Morazán: camino a San Antonio, Rodríguez 1392 (F); Tanque, Rodríguez 853 (F); Campus of Escuela Agrícola at El Zamorano, Quebrada El Gallo, 800 m, 29 July 1970, Pohl & Davidse 12460 (F, MO); road to El Zuyatillo, north of El Zamorano, 800 m, 29 July 1970, Molina & Molina 25789 (F, MO); near El Zamorano, dense

Ungrouped Species

247

shade among stream, 15 October 1951, Swallen 10783 (MO). Olancho: Montaña Los Zapotes, 10 km NO de Campamento, 1057 m, 25 August 1979, Soto 53 (MO) Nicaragua. Zelaya: Bilwaskarma, valley of Río Coco, 20 m, 10 July 1970, Pohl & Davidse 12257 (F, MO) Costa Rica. Cartago: canyon ot the Río Reventazón behind the Instituto Interamericana de Ciencias Agrícolas near Turrialba, 600–700 m, 6 August 1968, Pohl & Davidse 10831 (F). Puntarenas: Las Alturas along the Río Cotón, 8°56′N, 82°50′W, 1340 m, 28 August 1983, Davidse 24106 (MO); foothills of the Cordillera de Talamanca, lower montane forest in the area of Sitio Cotón (Cotonsito), along the road to Sitio Coto Brus, 1500–1700 m, 4 September. 1983, Davidse 24541 (MO); Cordillera de Talamanca, forested slopes along the Río Burú, upper slopes of Cerro Burú, ca. 9°01′N, 82°52′W, 1600 m, 19 August 1983, Davidse et al. 23765 (MO). San José: in shade of remnant forest near Puriscal, 1200 m, 21 July 1971, Burger & Burger 7630 (F); ca. 1 km E of Frailes, near city water tank, 1700 m, forest margiin, 6 September 1968, Pohl & Davidse 11073 (F); 1 km S of Rivas, 6 km NE of San Isidro del General, 725 m, 14 July 1966, Pohl & Calderón 10049 (MEXU) Panama. Chiriquí: Foothills, vicinity of El Boquete, Hitchcock 8299 (SI). Coclé: along road between El Cope and Bajo Grande, ca. 7  km  W of El Cope, 1000 ft, 28 July 1978, Hammel 4189 (MEXU, MO). Veraguas: 18 km W of Las Minas, N slope of Cerro Alto Higo, 2400–3000 ft, in cut over field, 6 August 1978, Hammel 4246 (MO) Cuba. Pinar del Río: Sierra de Cabra, on Guane road, bank of stream, September 1910, Britton et  al. 7201, 7284 (NY); Pinar del Río, thickets, Shafer 561 (NY); Arroyo Sumidero, dry hillside, Shafer & León 13682A (NY); Las Ánimas, wayside in open thickets, León 12607 (NY). Without locality, Wright 3855 (K, NY) Colombia. Antioquia: Tablaza, Caldas, Gartner 1600 (COL). Atlántico: entre Baranoa y Galapa, 80–140 m, December 1960, Dugand 5395 (COL), 4 November 1961, Dugand 5877 (COL); Barranquilla, barrio Altos del Prado, 50–70  m, 15 January 1962, Dugand 5983 (COL); Barranquilla, Elías 560 (RB). Caquetá: Morelia, Vereda de Santander, Quebrada of Río Bodoquero, Soderstrom 1405 (K). Cauca: Río Ortega, Pennell & Killip 7253 (US); El Tambo, Chisquío, von Sneidern 529 (NY); de El Estrecho a Balboa, 18 km de El Estrecho, 1280 m, 27 June 1989, Zuloaga 4244 (COL, SI). Cesar: entre La Paz y Manaure, finca de San Antonio, 1950 m, 19 November 1985, Torres 2912 (COL). Cundinamarca: at the junction of the Río Murca and Quebrada El Puente, Wood 4947 (K); Vereda La Libertad, San Francisco, Wood 3555 (K); Mun. Nilo, Inspección de Pueblo Nuevo, Finca La Turena, Torres et al. 750 (COL); Pandi, Vereda El Caracol, Díaz Piedrahita et al. 3292 (COL); Chisquío, Finca Los Derrumbos, Asplund 10846 (G, US); Mun. San Bernardo, Vereda Portones, Díaz Piedrahita et al. 3272 (COL). Magdalena: Santa Marta, Smith 2146 (COL, F, G, NY, P, US), 168 (COL, NY, P, US). Meta: Los Llanos, Villavicencio, Cuatrecasas & García Barriga 1980 (COL, US), Pennell 1598 (US); Restrepo, Salinas, subida al cerro, selva nublada secundaria, 600–700 m, 12 June 1989, Zuloaga 3940 (COL, SI); carretera de Villavicencio a Bogotá, 6 km de Villavicencio, 600  m, Zuloaga 4020 (COL, SI). Santander: municipio San

248

 

Taxonomic Treatment

Vicente de Chucurí, marden izquierda de la quebrada de El Medio, 1270–1460 m, 13 November 1985, Torres 2774 (COL) Venezuela. Aragua: carretera Maracay-Choroní, Tamayo 1638 (PORT); vicinity of Valencia, Pittier 9054 (G); mountains N of Maracay, Box 3952 (VEN). Bolívar: island on sand with low grassy mats at junction of rio Paramichi and rio Paragua, Steyermark 90562 (NY, VEN); Hato Divina Pastora, Tamayo 2919 (VEN); 1–10 km NW of Upata, on road to S. Felix, Steyermark 57679 (F, NY, VEN). Distrito Federal: Cerro Naiguatá, Loma de las Delicias, entre Quebrada de Basenilla y Quebrada Guayoyo, 9–12 km SW de hacienda Cocuizas, Steyermark 97479 (VEN); Parque Nacional El Ávila, selva nublada en Loma del Caño, 1600 m, Manara s.n. (VEN113044); Cordillera de la Costa, frente a Caracas, vía Mariperez-El Papelón, 1700  m, Morillo & Manara 1427, 1432 (VEN); vicinity of IVIC headquarters, west-NW of Caracas, 7 February 1973, Croat 21305 (MO, VEN). Falcón: Sierra de San Luis, entre Curimagua y San Luis, 1300–1400 m, Steyermark 99139 (VEN). Lara: Dtto. Moran, carretera Guarico-Villa Nueva, approx. 5 km desde Villa Nueva desde la carretera, Burandt Jr. V0326 (VEN). Miranda: La Providencia, 1000 m, en las cabeceras de la quebrada de Maitana, Pittier 13734 (VEN); Valle de Sartenejas, Univ. Simón Bolívar, 1200–1380  m, Gomez & Praderio 474 (VEN). Monagas: entre La Sabana de Las Piedras y Cerro Negro, NW of Caripe, Steyermark 61825 (F, NY). Portuguesa: en terrenos de la UNELLEZ, Mesa Alta de Cavacas, Ramírez Reyes 1399 (PORT). Sucre: BellaVista, carretera Cumaná-Puerto La Cruz, Farinas 176 (VEN). Táchira: montaña de Guafitas, just W and N of El Pinal, 250-300 m, Steyermark et al. 119536 (NY, VEN); Dtto. Córdoba, Cerro La Camiri, just S of the town of Río Negro, 6 November 1982, Davidse & González 21544 (VEN) Trinidad and Tobago. St. Joseph Savanna, above St. Augustine, Soderstrom 108 (K, US); Erin Savanna, south of Point Fortin, 24 June 1963, Soderstrom 1125 (K, NY, US); St. Ames, 17 Sep 1921, Broadway 5909 (MO) Guyana. Kopinang village, 600 m, 5 April 1988, Hahn et al. 4240 (US) Ecuador. El Oro: Zaruma, climbing in thicket, 1100 m, Asplund 15816 (NY) Brazil. Amapá: Macapá, Porto Platón, Mattos 10000 (SP). Bahia: 1 km S of Rio de Contas on side road to W of the road to Livramento do Brumado, 980 m, 15 January 1974, Harley et al. 15068 (CEPEC, MO, P); 12 km N of Rio de Contas, on the road to M. Grosso, 1200 m, 17 January 1974, Harley et al. 15202 (CEPEC, NY, P); 3 km N of the town of Rio de Contas, 21 January 1974, Harley et al. 15376 (CEPEC, NY, P); Estrada Jeque-Maracas, 2da. Parada, Vianna 1468 (R); 16 km NW of Lagoinha, on side road to Minas do Mimoso, 4 March 1974, Harley et al. 16640 (CEPEC, NY, P); Serra do Sincorá, Lagoa Encantada, 19 km NE of Ibicoara near Brejao, Harley et al. 15804 (P); Seabra, Lagoa da Bôa Vista, Pinto 132/80 (US). Ceará: Campo Sales to Crato, 13 April 1934, Swallen 4313 (IAN, R, US). Distrito Federal: Parque Nacional da Gama, Clayton 4961 (K, SP); Riberão da Contagem, ca. 25  km  N of Brasília, 14 December 1965, Irwin et  al. 11299 (MO, NY, US); Chapada da Contagem, Irwin et  al. 19413 (NY, UB); Brasília, Bacia do Rio S.  Bartolomeu, Equipe IBGE 4509, 5093, 6145 (IBGE); 25  km  SW of Brasília, Irwin et al. 13061 (F, NY, UB); Chapada da Contagem, 20 km NE of Brasília, Irwin et al. 9645 (F); Corrego Alogado, Parque Municipal do Gama, 20 km SW of Brasília,

Ungrouped Species

249

Irwin et  al. 14095 (F, NY, UB); Corrego Forquilha, Pereira 157 (IBGE). Goiás: 26 km NE of Catalao, Irwin et al. 25208 (F, NY); Corumbá de Goiás, Serra dos Pirineus, 10 km NW Cocalzinho, Burman & Filgueiras 416 (IBGE), 421 (IBGE); 12 km S of Corumbá de Goiás, Irwin et al. 10849 (F, NY); Planalto do Brasil, Serra do Caiapó, ca. 45 km S of Caiaponia on road to Jataí, Irwin & Soderstrom 7494 (SI); Luziania, Heringer 16119, 17760 (IBGE); 15  km ao sul da cidade Luziania, Heringer 17558 (IBGE, UB); Santo. Antonio do Descoberto, Filgueiras 698 (IBGE); Serra Dourada, 30 km E of Goiás Velho, Irwin et al. 11871 (NY, P, SP, UB); Serra dos Pirineus, 20 km E of Pirenópolis, Irwin et al. 34361 (NY, SP). Maranhão: Caxias to Barra do Corda, Swallen 3608 (RB, SP). Mato Grosso: entre Parecis e San Antonio, Kuhlmann 1755 (US). Minas Gerais: ca. 9  km NE of Estiva and 21 km SW of the intersection of highways 381 and 459, Davidse & D’Arcy 10541 (SP); Caldas, Regnell III-1359 (K, R); 20 km E of Diamantina, Irwin et al. 28045 (F, NY, P); 48 km W of Montes Claros, road to Agua Boa, Irwin et al. 23892 (F, NY, P, SP, UB); Passa Quatro, Sampaio 5995 (SI); Arará, Macedo 4240 (CEN); Poços de Caldas, Leitao Filho et al. 10 (CEN); Caapuera of Lavras, Black 844B (UB); Pedra Azul, estrada Almenara, Klinger 16860 (RB); 15 km N of São Joao da Chapada, Irwin et al. 28150 (F, NY, UB); Viçosa, Kuhlmann 1947, 1949 (RB); without locality, Langsdorff s.n. (K). Pará: Serra do Cachimbo, Pires s.n. (IAN 91973); Santarém, Obidós, Spruce 13 (P). Paraná: Rio Pasa Dois, Hatschbach 15980 (K, P); entre Icaraima e Casa Branca, Hatschbach 15839 (K); Mun. Xambre, Perola, Hatschbach 15842 (K); 41 km E of Guarapuava along Highway BR- 277 to Curitiba, 270 m, Davidse et al. 11323 (NY); Mun. Cianorte, Fazenda Lagoa, Hatschbach 14269 (F, K); Curitiba, Capao de Imbuia, Clayton 4204 (SP). Rio Grande do Sul: 13 km de Carazinho a Sta. Barbara, Quarín 1158 (CTES, SI); Morro Sapucaia, Sapucaia do Sul, Fernandes 109 (SI); Tenente Portela, Reserva Florestal do Turvo, Valls et al. 2583 (CEN). Rio de Janeiro: Macieiras, Serra de Itatiaia, Chase 8317 (NY, RB, US), Petrópolis, Sucre 2518 (SI); Petrópolis, Vale Bon-sucesso, Sucre 2196 (RB); São Gonçalo, Luetzelburg 26703A (F); Teresópolis, Freire de Carvalho 619 (RB). Santa Catarina: Mun. Dionísio Cerqueira, Pinheiral and ruderal near Dioníosio Cerqueira, 800  m, Smith & Reitz 9661 (NY, R); Azambuja, Brusque, Reitz 5927 (NY). São Paulo: 5  km E of main road on Araxa new track, about 8  km  SE of Uberlandia, Porto Ferreira, Goodland 358 (NY); Butantán, Hoehne s.n. (SP); Casa Verde, Pickel s.n. (SP 42423); near village of Parana-Piacaba, Skvortzov 283 (UB); Fazenda Campininha, N of Rio Moji-Guacu, Eiten & Eiten 2676 (K, NY, SP, UB); 9  km east of Sete Barras, shore of Rio de Ribeira, Clayton & Eiten 4675 (K); Descalvado, Fazenda Graciosa, Sendulsky 298 (SP); from Descalvado along road to São Carlos, Fazenda Graciosa, Sendulsky 727 (SI, SP); Instituto da Botanica, Parque do Estado, Sendulsky 575, 1101 (SP); Jundiai, Usteri s.n. (SP 9850); Porto Ferreira, Sendulsky 154 (SI, SP); Serra da Cantareira, Hauff 86 (SP) Bolivia. Cochabamba: Chapare, Steinbach 9095 (NY). La Paz: Murillo, Valle del Zongo, Renvoize & Cope 4253 (K,  LPB); San Carlos, Buchtien 53 (G, NY); Bella Vista, 26 December 1923, Hitchcock 22752 (NY, US); Hacienda Casana, Tipuani, 1,400 m, 5 January 1923, Buchtien 7114 (MO). Santa Cruz: Chiquitos, Serranía de Chiquitos, 5 km S of San José, Killeen 1732 (F, SI)

250

 

Taxonomic Treatment

Paraguay. Alto Paraná: Reserva Itabó, Itaipú Binacional 963 (MO). Caaguazú: in vicinis Caaguazú, Balansa 52 (G, P). Central: Villa Elisa, Pedersen 5904 (K, P), Balansa 51 (G, K, P); Lago Ipacaray, Hassler 12462 (G, GH, K). Concepción: Zwischen Rio Apa und Rio Aquidaban, Fiebrig 5373 (G, K). Cordillera: Cerro Caacupé, 8 km del pueblo, Arenas 1226 (F, K); Cordillera de Altos, Cerro de Tobatí, 6 March 1984, Schinini 23973 (SI) Argentina. Corrientes: 42 km E de Ituzaingó, costa del río Paraná, 31 March 1976, Quarín et al. 3373 (CTES); Dpto. San Miguel, 12 km NE de San Miguel, Estancia Curuzú Laurel, 14 June 1974, Schinini 9356 (CTES). Misiones: Gral. Manuel Belgrano, de J. M. de Rosas a Bernardo de Irigoyen, 4 May 1984, Zuloaga et al. 2119 (SI); Oberá, entre Alem y Obera, 12 km de Oberá, 2 May 1984, Zuloaga et al. 1976 (SI); entre Oberá y Campo Viera, Arroyo Yasa, Zuloaga et al. 1984 (SI); San Ignacio, Salto Tabay, 20 February 1971, Burkart 28260 (SI); San Pedro, 24 February 1984, Pedersen 13675 (SI); Teyucuaré, July 1945, Martinez Crovetto 3519, 3524 (SI); Loreto, Montes 434 (NY), 1788 (G); Yabebiry, Montes 752 (LIL, NY) 61. Panicum trichanthum Nees, Fl. Bras. Enum. Pl. 2(1): 210. 1829. Type: Mexico. Without locality, F. W. H. H. A. von Humboldt s.n. (lectotype, B_10_0521660, designated by A.  S. Hitchcock & A.  Chase, Contr. U.S.  Natl. Herb. 15: 131. 1910). Fig. 67 Milium microspermum Lag., Gen. Sp. Pl.: 2. 1816. Panicum microspermum (Lag.) E.  Fourn., Mexic. Pl. 2: 22. 1886. Sporobolus microspermus (Lag.) Hitchc., J. Wash. Acad. Sci. 23(10): 453. 1933. Type: Mexico. “"Habitat in Nova Hispania. Semina communicavit perill. D. Sesse”, D. Sessé s.n. (holotype, MA; isotype, US 00433438, fragment) Panicum guayaquilense Steud, Syn. Pl. Glumac. 1: 85. 1855[1854]. Type: Ecuador. Guayas: Guayaquil, April 1846, W.  Jameson 560 (holotype, P 00740816; isotypes, BAA 00002347, fragment ex P, BM 000938714, G 00099788, G 00099789, K 000309301, K 000309302, USA 00148555, fragment ex K, W 18890088459) Panicum trichanthum Nees var. modestum Döll, Fl. Bras. 2(2): 248. 1877. Type: Brazil. Ad Tocantins fluvium inter Porto Imperial et Funil, W. J. Burchell 8791 (holotype, BR; isotypes, K 000309235, USA 00140065, fragment ex BR, W 18860009172) Panicum granuliferum Kunth var. longifolium Kuntze, Revis. Gen. Pl. 3(3): 361. 1898. Type: Bolivia. Velasco, 200 m, July 1892, C. E. O. O. Kuntze s.n. (lectotype, NY 00381747, here designated; isolectotype, NY 003811746). Brazil. Mato Grosso. Without locality, C. E. O. Kuntze s.n. (syntypes, NY 00381748, NY 00381749) Plants perennial, shortly rhizomatous, 0.50–1.30 m tall; culms creeping, rooting and branching at the lower nodes, then erect, rigid, branching at the upper nodes and leaning on the vegetation; internodes 2–10  cm long, cylindrical, hollow, pale or tinged with purple, glabrous; nodes compressed, purplish, glabrous. Sheaths striate, 1.7–8.5  cm long, usually smaller than the internodes, glabrous, margins

Ungrouped Species

251

Fig. 67  Panicum trichanthum. (a) Upper portion of culm and inflorescence. (b) Spikelet, ventral view. (c) Spikelet, dorsal view. (d) Spikelet, lateral view. (e) Upper anthecium, ventral view. (f) Caryopsis, hilum view. (g) Caryopsis, embryo view. (From Zuloaga 2217, SI)

252

 

Taxonomic Treatment

membranous, short to long hairy toward the upper portion. Ligule membranous, reduced, 0.3–0.5 mm long, with long whitish hairs beneath at the base of the blade; collar brown, glabrous to sparsely hairy. Blades lanceolate, flat, 1.6–12.3 × 0.2–2 cm, subcordate to cordate, shortly pseudopetiolate, pseudopetiole brown, blade glabrous to sparsely pilose on both sides or only on the adaxial surface, the margins scabrous and sparsely pilose. Inflorescence terminal, exerted, peduncle cylindrical, glabrous, 15 cm long; panicles lax, diffuse, multiflowered, 10–30 × 6–26 cm, when young included in the upper leaves; main axis glabrous, smooth, pulvini hairy to glabrous, brownish, first-order branches alternate, rare opposite, divergent and distant, spikelets solitary, scattered on long glabrous, smooth, wavy pedicels, 1–15  mm long. Spikelets ellipsoid, 1–1.9 × 0.6–0.9 mm, glabrous, dark green, glumes and lower lemma shiny, with minute vesicles over the entire surface; lower glume 0.3–0.5 mm long, 1/4 or less the spikelet length, nerveless, obtuse to truncate, glabrous; upper glume 1.3–1.6 mm long, 5 nerved, slightly nerved; lower lemma 1.4–1.6 mm long, (3–)5 nerved, glumiform; lower palea ovate-lanceolate, 0.8–1 × 0.2 mm, glabrous, hyaline; lower flower absent; upper anthecium ellipsoid, 1.3–1.6  ×  0.5–0.8  mm, smooth, lustrous and pale, brown at maturity, with bicellular microhairs; lemma 5 nerved, the glumes often fallen first and the upper anthecium remains. Caryopsis obovoid, 1 × 0.7–0.8 mm, whitish; hilum obovate, embryo half the length of the caryopsis. Distribution and ecology (Fig. 66). A widespread species in the Americas from Mexico, Mesoamerica, and the Caribbean to Argentina. It lives in humid and shady places, on the edges of streams or in the undergrowth, on the edges of jungles or forests; 0–1200 m asl In flower between November and May. Local names: “Kituka watla” (in Miskito), “zacate sereno” (Mexico). “takuari ñu” (takuari = tacuarilla, ñu = field) (Zuloaga 1994) Species related to P. bartlettii (see remarks under the latter species). In describing Panicum granuliferum Kunth var. longifolium, Kuntze cites syntypes from Bolivia, Velasco, and Brazil, Mato Grosso. Of these, NY 00381747 is designated as the lectotype of the species, as it is a complete specimen corresponding to the protologue. Additional material examined. Mexico. Campeche: Éjido Pach-wits, 32 km S of Xmaben on the dirt road from Hopelchén to Xpujil, 19°00′N, 89°20′W, 200 m, 12 Mar 1990, Sanders et  al. 9687 (MO). Chiapas: Mun. of Ocozocoautla de Espinosa, large marsh and adjacent slopes, 13 km south of Ocozocoautla, 900 m, 29 October 1981, Breedlove & Davidse 54051 (MO); 5 km NW of Tumbo, roadside, 17°06′N, 91°39′W, 620 m, 26 September 1988, Stevens & Martínez 25887 (MO); en Arroyo Agua Azul, a 3 km al W de Frontera Corozal, 200 m, 1 November 1985, Martínez 14651 (MO); Río Lacantum, on riverbank, 9 February 1964, Lundell 17857 (MEXU). Oaxaca: Chiltepec, Tuxtepec, 23 August 1965, Martínez Calderón 221 (MEXU). San Luis Potosí: Camp area at El Salto, 1 September 1972, Gallian & Leake 1067 (MO); valley of Río Tampaon, 20 July 1939, Chase 7518 (MO); Tamasopo, 1200  ft, 24 August 1937, Fisher s.n. (MO). Tabasco: 1  km al N de Monte Grande, orilla de camino, junto canal de agua, 13 March 1983, Fernández

Ungrouped Species

253

et al. 1485 (MO). Veracruz: south edge of Tesechoacán, W side of Río Tesechoacán, 18°07′N, 95°39′W, 50  m, 6 December 1981, Nee 23829 (MO); Tlaliscoyán, 24 November 1967, Martínez Calderón 1565 (MEXU, MO); Estación Biológica San Andrés Tuxtla, 7 November 1968, Martínez Calderón 1786 (MEXU, MO); Veracruz, Gouin 19 (P); km 6 carretera Misantla Naolinco, 520 m, 30 January 1976, Avendaño et al. 112 (MEXU) Honduras. Atlántida: Brushy roadside 9 km E of Tela, scrambling in brush, 26 June 1970, Pohl & Davidse 12078 (MO). Cortés: 5 kms de San Pedro Sula, Río Arenales, 40 m, 18 December 1950, Molina 3831 (MEXU, MO). Gracias a Dios: Ahuas Bila, 200 km S.O. de Puerto Lempira, orilla del río Wankí, Coco a Segovia, bosque húmedo tropical, 100 m, 5 May 1985, Nelson & Cruz 9452 (MO) El Salvador. Vicinity of San Salvador, wet soil along stream, 650–850 m, March 1922, Standley 23298 (MO) Costa Rica. Cartago: Instituto, Turrialba, 16 November 1950, 600  m, León 2243 (MO). Guanacaste: small lagunas near San Bernardo, ca. 14  km n.n.e of Bagaces, 300 m, 21 February 1969, Pohl & Davidse 11727 (MEXU, MO). Heredia: Finca La Selva, the OTS Field Station on the Río Puerto Viejo just E of its junction with the Río Sarapiquí, 100  m, 21 April 1982, Hammel 11794 (MO). Limón: Cahuita, abundant on roadsides and in banana grove, 14 June 1973, Pohl 12914 (MO). Puntarenas: northeastern slopes of Fila de Cal, between San Vito de Coto Brus and Ciudad Neily, 8°41′N, 82°56′W, 500-620  m, 12 July 1985, Grayum & Hammel 5641 (MEXU, MO). Without locality, Tonduz 8600 (P) Panama. Bocas del Toro: Vicinity of Chiriquí Lagoon, 31 October 1940, von Wedel 1449 (MO); Old Bank Island, vicinity of Chiriquí Lagoon, 5 February 1941, von Wedel 1998 (MO); approximately 3.5 km S of Tiger Key on the mainland, 20 February 1989, Peterson & Annable 6943 (MO). Darién: vicinity of Cana, second growth forest and open pastures at old town of Cana, 23 June 1959, Stern et al. 497 (MO). Panamá: Río Tapia, wet forest, common, 7 December 1924, Standley 26154 (MO). Los Santos: Guayabo, a place several miles west of Tonosí, 1 March 1963, Stern et  al. 1894 (MO). Veraguas: Cerro de Tute, 8 August 1963, Dwyer 4292 (MO). Zona del Canal: Barro Colorado Island, sand deposit at end of Fuertes Cove, 1 May 1968, Croat 5254 (MO); Culebra, 26 August 1911, Hitchcock s.n., Amer. Gr. Hb. 209 (MO) Cuba. Habana: Vento, thickets on rocky slopes, 11 January 1905, Curtiss 598 (MO, P). Las Villas, Corralillo; near Río Amimao, vicinity of Soledad, 17 December 1941, Gonzales 414 (MO, P). Without locality, de la Sagra 8 (P), Wright 753 (G, MO) Puerto Rico. Río Blanco, June 1881, Eggers 461 (G) Colombia. Antioquia: Puerto Bernio, Archer 1462 (COL); carretera LiborinaSabanalarga, 3 km de Liborina, 910 m, 8 March 1986, Callejas et al. 2197 (MO). Boyacá: entre San Luis del Gaceno y Santa Marchía, 19 June 1989, Zuloaga & Londoño 4110 (MO). Caquetá: Florencia, río Hacha, Echeverry 1648 (COL). Chocó: carretera San José del Palmar-Novita, alrededores del Campamento de Curundo, 23 August 1976, Forero et  al. 2366 (COL, MO). Cundinamarca: La Esperanza, García Barriga 3202 (COL). Magdalena: Santa Marta, Smith 2151 (COL, MO). Meta: margen del río Negrito, entre Villavicencio y Puerto López,

254

 

Taxonomic Treatment

Cuatrecasas & García Barriga 3546 (COL). Norte de Santander: Cúcuta y El Rosario, Carvajalino & Díaz 42 (COL). Quindío: without locality, 1600 m, January 1853, Triana s.n. (COL-2338). Risaralda: along road between Pueblo Rico and Istmina, 14 km NW of Pueblo Rico, 5°3′N, 76°00′W, 780 m, 22 February 1990, Croat 70846 (MO). Santander: Badillo, Río Magdalena, 16 January 1918, Pennell 3918 (MO). Valle del Cauca: Hacienda El Medio, carretera Panamericana entre La Paila y Zarzal, 950  m, 29 February 1992, Silverstone 6512 (MO); Hacienda El Guachal, Vereda de Cauca Seco, por la recta Cali-Palmira, 880 m, 11 May 1988, Ramos & Paz 914 (MO) Venezuela. Aragua: between San Mateo and Villa de Cura, Alston 6053 (F, P); prope Colonia Tovar, Fendler 1643 (MO). Distrito Federal: Hacienda Sosa, Badillo 67 (VEN). Falcón: Sierra de San Luis, Quiragua, 900 m, 17 January 1978, Ruiz 3081 (MO). Lara: Paso del río Bucare, 16 January 1928, Pittier 12617 (MO, VEN); Distrito Jiménez, Parque Nacional Yacambú, Quebrada Negra from the base of the tunnel to the junction with the Río Yacambú, 9°41′N, 69°30′W, 620 m, 24 October 1982, Davidse & González 20995 (MO). Mérida: entre La Vega y San Juan, valle del Chama, 29 January 1928, Pittier 12763 (MO, VEN). Miranda: Distr. Páez, Fila de Guayas between rio Guayas and rio Guapo, ca. 15 km directly SW of El Guapo, 66°04′W, 10°04′N, 400–700  m, 6 June 1977, Davidse & González 13666 (MO, VEN). Monagas: Maturín to Quiri Quiri, Chase 12578 (US, VEN). Sucre: Distr. Sucre, El Guayabito, along the rio Guayabo at its juncture with the rio Zumbador, 20-22 November 1981, Davidse & González 19266 (MO, VEN). Táchira: Río Umuquema, carretera hacia La Fría, Medina 1243 (VEN). Trujillo: La Cristalina, 1900-2000 m, Tamayo 1862 (VEN). Yaracuy: entre la Ciénaga y Carbonero, 30 December 1928, Pittier 13069 (MO, VEN); selva de El Rodeo, Aristeguieta & Foldats 1355 (VEN) Suriname. Paramaribo, near Agric. Experim. Station, shadowy roadside, 19 July 1933, Lanjouw 42 (MO) French Guiana. Haut Itany, Hoock 133 (P) Ecuador. Guayas: Las Américas, 28 September 1952, Faberling & Wibom 326 (MO); Tenguel, 27 August 1946, Giler 1773 (MO). Los Ríos: Hacienda Clementina on Río Pita, Asplund 5512 (P). Without department: Südl. Naranjapata, 550 m, 28 November 1933, Schimpff 532 (MO) Peru. Cuzco: La Convención Province, Río Manguriari, 12°47′S, 72°40′W, 750  m, 2 February 1991, Nuñez & Ortíz 12809 (MO). Loreto: Yarinacocha, Pucallpa, Aldave & Fernández 5634 (US); Maynas, Dpto. Iquitos, Uspa-Cano across Río Itaya from Iquitos, 30 September 1975, McDaniel & Rimachi 20250 (MO); Sintica, on Río Ucayali, Prov. Requena, 5°35′S, 74°15′W, 9 November 1947, Fosberg 29052 (MO); above Pongo de Manseriche, mouth of Río Santiago, sandy riverbank, 200 m, 29 November 1931, Mexía 6190 (MO); Iquitos, west edge of city, 31 June 1972, Croat 17446 (MO); Río Paranapura above Yurimaguas, 10 July 1972, Croat 17954 (MO). Madre de Dios: Parque Nac. del Manú, Cocha Cashu Station, 11°50′S, 71°25′W, 350  m, 6 August 1984, Foster 9779 (MO, SI). San Martín: Quebrada de Huaquisha, margen derecha del río Huallaga, 400–450 m, 1 July 1974, Schunke Vigo 7096 (MO)

Ungrouped Species

255

Brazil. Amazonas: S. Gabriel, Pires 818 (IAN); São Pedro, Kuhlmann 72 (RB). Goiás: 16 km SW of the Goias-Bahia border along highway BR-020, 8 April 1976, Davidse et al. 12194 (MO, SP); Serra do Caiapó, ca. 30 km S of Caiaponia, road to Jataí, 950 m, 29 June 1966, Irwin et al. 17992 (MO). Mato Grosso: 26 km N of Xavantina on Xavantina-Cachimbo road, near João Loepz bridge, Ratter & Ramos 240 (P); Miranda, retiro de Tição de Fogo, 18 km da séde da Fazenda Bodoquena, no rumo N-NE, 100–200 m, 10 June 1973, da Silva 61 (MO); Miranda a zero km da séde da Fazenda Bodoquena, 13 June 1973, da Silva 114 (MO). Mato Grosso do Sul: Corumbá, Vali & Pott 150 (CEN). Minas Gerais: Anna Florencia, east of Ponte Nova, 400  m, Chase 9471 (MO). Pará: Cacaval Grande, Canal Fernando Costa, Black 52-15439 (IAN); Estate “Cacanal Grande”, 10 July 1914, Goeldi 121, 122 (MO). Rio Vermelho, regiäo do Tocantins, Froes 27045 (IAN). Paraná: 9 km S of Jardinópolis along the road to Medianeira, 300 m, 13 March 1976, Davidse et al. 11221 (MO, SI, SP); Fazenda Mururé, 15 May 1969, Hatschbach 21499 (MO). Rio Grande do Sul: Mun. Butiá, Polo Carboquímico, Abruzzi 611 (CEN); vicinity of Monte Negro, tropical forest margin, 3 July 1952, Beetle 1486 (MO). Rondonia: Ouro Preto, entre Vila de Rondonia e Porto Velho, Valls 1188 (CEN); Rio Yata, Silva 365 (IAN) Bolivia. Beni: ca. 5 km SW of Riberalta, 11°02′S, 66°07′W, 220 m, 3 June 1982, Solomon 7919 (MO, SI). Chuquisaca: Hernando Siles, 12 km E de Monteagudo, 1200 m, 11 March 1981, Renvoize & Cope 3898 (K, LPB, MO), 3890 (K, MO). La Paz: Polo-Polo a Coroico, Buchtien 445 (BAF), 19 December 1921, 3612 (MO); Prov. Nor Yungas, carretera 3, between Coroico and Caranavi, 6 November 1976, Davidson 4824 (MO); NNW de Coroico, Suapi, Beck 8344 (K), Solomon 14188, 8930 (MO), Buchtien 446 (K). Santa Cruz: 1 km arriba de Comarapa, Steinbach 242 (F); Buena Vista, Steinbach 1213 (F); 30  km de Sta. Cruz de la Sierra, 26 October 1980, Zuloaga et al. 1412 (MO, SI); Prov. Velasco, San Juancito, 27 km al N de San Ignacio, 400  m, 1 May 1986, Seidel & Beck 355 (MO, SI); Prov. Santiesteban, 12 km NE of crossroads at Montero, along highway from Montero to Colonia Okinawa 1, 17°17′S, 63°09′W, 275 m, 22 October 1990, Nee 39458 (MO); Prov. Ichilo, Parque Nacional Amboró, ca. 15 km SE up the Río Pitasama from the Río Surutú, 17°44′S, 63°40′W, 700 m, 31 August 1985, Solomon & Urcullo 14188 (MO); Prov. Andres Ibañez, 1 km E of Int. Airport Viru Viru, 16 km N of Santa Cruz, 17°40′S, 63°10′W, 400 m, 1 January 1986, Killeen 1556 (MO); Santa Cruz, Chiquitos, 5 km N of El Porton on road t o Entre Ríos, 18°10′S, 60°05′W, 300 m, 12 October 1987, Killeen 2773 (MO); Ñuflo de Chavez, Jinca, an Ayoreo community 90  km  SE of Concepción, 16°33′S, 61°40′W, 450  m, 21 July 1985, Killeen 1053 (MO); Ñuflo de Chavez, Concepción, 16°10′S, 62°00′W, 500  m, 6 August 1985, Killeen 1091 (MO); Cordillera, Tatarenda, 30 km S of Rio Grande on road to Camiri, 19°08′S, 63°15′W, 700 m, 10 October 1985, Killeen 1259 (MO). Tarija: Ipa, 30 km N de Villa Montes, camino a Santa Cruz, 63°25′W, 21°S, 12 April 1977, Krapovickas & Schinini 31194 (CTES, MO, SI); ruta Tarija- Villa Montes, Cañadas, 156 km al E de Tarija, 980 m, Krapovickas et al. 19192 (CTES, P); Villa Montes, Quebrada de Tampinta, 28 May 1971, Krapovickas et al. 19389 (CTES, MO); Prov. Arce, 29.2 km S of Emboroza-Sidras road on road to Bermejo, 12.7 km S of Naranjo Agrio, 600 m, 21 April 1983, Solomon 9881 (MO)

256

 

Taxonomic Treatment

Paraguay. Alto Paraguay: Parque Nacional Defensores del Chaco, entra la casa forestal de Aguadulce y Aguarica, 21 October 1980, Fernández Casas 4366 (G); Mayor Pedro Lagerenza, 60°45′W, 20°05′S, 11 April 1978, Schinini & Bordas 15188 (CTES, MO). Boquerón: Cerro León, 2 June 1984, Billiet & Jadin 3281 (MO); 12 km E de Pratts Gill, 23 October 1986, Molas & Vera 1382 (MO); Col. Fernheim, 10  km  S de Filadelfia, 13 September 1990, Vanni et  al. 2047 (CTES, MO). Canindeyú: Colonia Fortuna, 7 km de Curuguaty, 7 May 1974, Arenas 668 (MO). Central: Luque, 20 March 1875, Balansa 46 (G, P), 46a (G); in regióne lacus Ypacaraí, Hassler 12450 (G, MO), 12477 (G), 12579 (G, MO, NY); Asunción, 16 August 1874, Balansa 46 (P); Tavarory, entrance to Tavarory savanna, 25°30′S, 57°30′W, 25 March 1992, Zardini & Aquino 31413 (MO); border Tavarory-Acosta NU, 25°30′S, 57°30′W, 18 May 1992, Zardini & Aquino 31692 (MO). Concepción: Zwischen Río Apa und Río Aquidabán, Sanja-Mocotí, Fiebrig 5176 (G); Arroyo Tagatiya-mí, 57°32′W, 22°37′S, 140  m, 4 April 1986, Brunner 1719 (G, MO). Cordillera: Cordillera de Altos, Hassler 640a (G, P); Co. Tobatí, 8 March 1984, Schinini 24028 (CTES, G, SI); confluence of Río Paraguay and Río Salado, inundated savanna on clay 25°09′S, 57°30′W, 3 March 1990, Zardini & Velázquez 19873 (MO), 21 July 1990, Zardini & Velázquez 22272 (MO). Guairá: Cordillera de Ybyturuzú, road to Cantera Jhú, 2 km S to Coronel Oviedo, 25° 48′S, 56° 20′W, 27 September 1989, Zardini & Velásquez 14578 (MO); Cordillera de Ybyturuzú, road Melgarejo-Antena, 5 km S of Melgarejo, forest on mountain slopes, 25° 55′S, 56° 15′W, 28 May 1989, Zardini & Velásquez 12203 (MO), km 15, 14 July 1989, Zardini & Velásquez 13546, 13554 (MO); a 6 km de Melgarejo hacia Cerro Acatí, bosque ribereño del Arroyo Tacuara, 17 June 1989, Aguayo 421 (MO); Cordillera de Ybytyruzú, Melgarejo-Cerro Acati, 8 km S of Melgarejo, 25 48′S, 56 16′W, 10 July 1992, Zardini & Tilleria 32673 (MO). Paraguarí: Forets de Guarapí, 5 April 1883, Balansa 4356 (G); Macizo Acahay, in forest on Eastern Peak, 25°54′S, 57°09′W, 500 m, 30 June 1988, Zardini & Cuevas 5328 (MO); Acahay Massif, forest on western slopes of southeastern peak, 25°54′S, 57°09′W, 400 m, 11 June 1989, Zardini 12943 (MO); Acahay Massif, easternmost Peak, western part, 25°52′S, 57°08′W, 13 January 1992, Zardini & Tilleria 29780 (MO); Acahay Massif, gallery forest along creek on easternmost peak, 25°54′S, 57°09′W, 350  m, 11 June 1989, Zardini & Velázquez 12704 (MO). Presidente Hayes: Pilcomayo River, Morong 1571 (G, MO); in regióne cursus inferioris fluminis Pilcomayo, May 1906, Rojas 90 (G). San Pedro: Alto Paraguay, Primavera, Woolston G.36, G.258 (NY, SI) Argentina. Chaco. Dpto. 1° de Mayo, Colonia Benitez, Quarín et  al. 1869 (CTES, SI). Dpto. DoNovemberan, 13 km N de La Verde, Estancia Dos Tranqueras, 21 September 1982, Schinini 22718 (CTES, LIL, MO). Dpto. San Fernando, Fontana, Meyer 321 (LIL); Resistencia, Venturi 8255 (SI), Parodi 8307 (CTES). Corrientes: Dpto. Capital, Escuela de Agricultura, Schinini 8598 (CTES, SI); Perichón, costa del río Paraná, 7 April 1974, Schinini 8715 (CTES, MO); Puente Pesoa, Schinini 8694 (CTES, SI); Riachuelo, Schinini & Martinez Crovetto 12911 (CTES). Dpto. Empedrado, Río Empedrado y ruta 12, 31 August 1973, Quarín & Schinini 1215 (CTES, MO, SI); Estancia Las Tres Marías, 10 January 1952, Pedersen 1430 (MO). Dpto. Esquina, Guayquiraró, Quarín et al. 2156 (CTES, LIL, SI); Dpto. Gral. Paz. 19 km S de Caa-Catí, Ahumada 2352 (CTES); Dpto. Ituzaingó,

Ungrouped Species

257

11 km N de San Carlos, Krapovickas et al. 24955 (CTES, SI); Dpto. Mburucuyá, Estancia Santa Teresa, 15 April 1976, Cabrera et  al. 27752 (MO, SI); Estancia Santa María, 4 May 1952, Pedersen 1710 (CTES, MO); Dpto. Saladas, Río Santa Lucía, Paso Naranjito, Krapovickas et  al. 24552 (CTES, SI); Dpto. San Cosme, entre Costa Toledo y Puerto González, Krapovickas & Cristobal 11895 (CTES, SI); Dpto. San Luis del Palmar, 10 kmSE de San Luis del Palmar, ruta 6, Quarín & Tressens 1337 (CTES, SI); Dpto. San Martin, Torrent, Ibarrola 173 (LP), 1719 (LIL). Entre Rios: Dpto. Concordia, Concordia, Parque Rivadavia, Burkart et al. 23252 (SI); Dpto. Feliciano, Paso Yunque, 18 May 1964, Burkart et al. 25275 (SI), 25276 (MO); Dpto. La Paz. Isla Curuzú-Chalí, Burkart et  al. 26815 (SI); Paso Telégrafo, sobre el río Guayquiraró, Burkart & Bacigalupo 21068 (SI). Formosa: Dpto. Bermejo, Puerto Bermejo, Castellanos s.n. (LIL 325867); Dpto. Formosa, Estancia Monteagudo, Guaglianone et al. 335 (SI); Dpto. Pirané, Casco Cué, 31 January 1946, Morel 764 (LIL, MO); Dpto. Pilcomayo, Laguna Vera, Morel 5005 (LIL); sur de Riacho Negro a 3 km, 20 July 1947, Morel 3502 (MO). Jujuy: Dpto. Ledesma, Abra de Cañas, Vervoorst et al. 4524 (SI); Calilegua, toma del río Zora, Cabrera et al. 30379 (SI, US); Yuto, El Bananal, 21 September 1976, Cabrera et al. 28055 (MO, SI); Dpto. San Pedro, Ingenio La Esperanza, Bartlett 20313 (LP, SI); San Pedro de Jujuy, Herrera 504 (GH); Dpto. Santa Bárbara, Laguna La Brea, Fries 81 (US); Laguna San Miguel, Cabrera et al. 26322 (SI, US); camino a Valle Grande, 26 April 1975, Cabrera et al. 26418 (MO). Misiones: Dpto. Candelaria, Santa Ana, camino al Balneario Municipal, 11 April 1985, Zuloaga et al. 2217 (MO, SI); Dpto. Capital, Posadas, Burkart 14203 (SI); Dpto. Eldorado, Arroyo Piray-Guazú y ruta 12, 29 November 1987, Zuloaga et  al. 3291 (MO, SI); Dpto. Iguazú, Cataratas, Hunziker 891 (SI), Krapovickas 2480 (LIL, LP); Dpto. Lib. Gral. San Martin, Puerto Leoni, Cabrera, Corte y Gebhard 86 (LP); Dpto. Montecarlo, Montecarlo, Montes 15381 (SI); Dpto. San Ignacio, Campo Viera, Montes 15373 (MO, SI), 15374 (MO); Dpto. San Javier, Arroyo Itacaruaré, camino viejo, 1 May 1984, Zuloaga et al. 1952 (MO, SI); San Javier, 31 January 1926, Parodi 7059 (MO); Dpto. San Pedro. entre San Pedro y Puerto Piray, km 60, 3 May 1984, Zuloaga et al. 2074 (MO, SI). Salta: Dpto. Gral. Jose de San Martin, 16  km  W de General Ballivián, Krapovickas & Schinini 30922 (CTES); Dpto. Orán, Orán viejo, Burkart 20311 (SI); Tabacal, 12 August 1923, Parodi 5150 (MO); de río Pescado a Orán, Cabrera et al. 26537 (SI). Santa Fe: Dpto. Gral. Obligado, Villa Ana, Quarín 1186 (CTES, SI); Dpto. San Javier, entre Romang y Alejandra, Prado 260 (SI); Dpto. Vera, al E de Guampita, Lewis 526 (SI) 62. Panicum trichoides Sw., Prodr.: 24. 1788. Type: Jamaica. Without locality, O. P. Swartz s.n. (lectotype, S06-642, specimen at left, designated by A. S. Hitchcock, Contr. U.S.  Natl. Herb. 12: 140. 1908; isolectotypes, LD 1266465, LINNHS118-64-1, S06-641, S06-814, S06-815, S06-816, S-R-4016, SBT 10758, USA 00140069, fragment and photos). Fig. 68 Panicum capillaceum Lam., Tabl. Encycl. 1: 173. 1791. Panicum filamentosum Pers., Syn. Pl. 1: 83. 1805, nom. inval. Panicum micranthum Salzm. ex Döll, Fl. Bras. 2 (2): 249. 1877, pro syn. nom. nud. Type: Puerto Rico: Without locality,

258

 

Taxonomic Treatment

Fig. 68  Panicum trichoides. (a) Plant. (b) Spikelet, lateral view. (c) Spikelet, ventral view. (d) Spikelet, dorsal view. (e) Lower palea. (f) Upper anthecium, ventral view. (g) Upper anthecium, dorsal view. (h) Caryopsis, hilum view. (i) Caryopsis, embryo view. (From Davidse et  al. 18474, MO)

Ungrouped Species

259

A. P. Ledru s.n. (holotype, P-LAM 00740821; isotypes, BAA 00002311, fragment ex P-LAM, P, US 00148326, fragment ex P-LAM) Panicum acutifolium Willd. ex Spreng, Syst. Veg. [Sprengel] 1: 320. 1825[1824], nom. inval. Type: Venezuela. Cumanacoa, F.  W. H.  H. A. von Humboldt s.n. (holotype, B-W 18772-01-0; isotype, US 00148111, fragment ex B) Panicum capillaceum Lam. var. strictius Döll, Fl. Bras. 2 (2): 249. 1877. Syntypes: Brazil. Piauí: without locality, January 1840, G. Gardner 3509 (K 000309228, K 000309227, US 00148327, fragment ex K) Plants annual, stoloniferous, culms decumbent and rooting at the lower nodes, then erect, 10–60  cm tall, branching at the upper nodes; internodes hairy, nodes dark, densely hairy. Sheaths shorter than the internodes, densely to sparsely papillose-pilose, more so toward the upper portion, the margins ciliate. Ligule membranous, reduced, 0.3–0.5 mm long, shortly ciliate in the upper portion; collar pilose. Blades ovate-lanceolate, 2–7 × 0.4–2 cm, flat, glabrous to sparsely pubescent, papillose-pilose toward the base, this asymmetrical, cordate, and slightly amplexicaul. Inflorescence exserted or partially included in the upper blades, peduncle up to 5 cm long; panicle terminal, lax, diffuse, 5–20 cm long; main axis, branches and pedicels glabrous, occasionally hirsute, first-order branches ascending, pedicels long, slender. Spikelets obovoid to ellipsoid, 0.9–1.4 × 0.5–0.7 mm, greenish or tinged with purple, hairy; lower glume 0.5–1.1  mm long, acute, 1–3 nerved; upper glume slightly shorter than the spikelet, elliptic, obtuse to acute, 3–5 nerved; lower lemma as long as the spikelet, elliptic, 3–5 nerved, acute; lower palea reduced, 0.6–0.9 mm long, hyaline, glabrous; lower flower absent; upper anthecium ovoid, 0.9–1.3 × 0.5– 0.6 mm, pale, shiny, finely and transversely rugose and with bicellular microhairs, acute. Caryopsis broadly ellipsoid, 0.8 × 0.5 mm; hilum punctiform, embryo less than half the length of the caryopsis. Distribution and ecology (Fig. 69). Grows from the USA (cited for Texas), Mexico, Mesoamerica, and the West Indies to Brazil, Bolivia, and Argentina; also introduced in Asia and Africa. It frequently inhabits disturbed sites and is common in humid environments on forest edges; 0–1100 m asl. Local names: “Cocullito” (in specimen Steyermark et  al. 100143); “Marsh grass” (in Guyana) Panicum trichoides was described by Swartz on the basis of a specimen that is a mixture of what this author considered P. trichoides and a portion of P. trichanthum; he adds to the original description the citation of a plate found in Sloane’s work, which corresponds to P. trichoides. Hitchcock (1908), in studying the species described by Swartz for America, analyzes the type of the species, indicating that there are two different materials: one that agrees with what Grisebach (1864) calls P. brevifolium and the other similar to the type of P. trichanthum; he mentions that it is evident that Swartz considers his species similar to P. brevifolium and variable enough to include the two existing plants in the specimen; he concludes that P. trichoides Sw. should be considered valid, applied to the American species usually described as P. brevifolium and take the plant found in the type, from S, to the left as the type of the same.

260

 

Taxonomic Treatment

Fig. 69  Distribution of Panicum trichoides

This species was included (Hitchcock and Chase 1910) in the informal Trichoidia group, characterized by including annual plants, with ovate to oblong-lanceolate laminae, diffuse panicles, spikelets less than 1.4  mm long, minutely papillose. Pilger (1940) gives this group section rank; later, Zuloaga (1981) places the species in section Ovalifolia Stapf. Panicum trichoides is morphologically related to P. millegrana and P. sellowii, with which it shares having hairy spikelets, with the lower glume 1/3 to ½ the spikelet length, reduced lower palea and absent lower flower, and upper anthecium slighty rugose transversely. The latter two species are distinguished by including perennial plants, with spikelets 1.6–2.5 × 0.9–1.2 mm, open at maturity, with the upper anthecium markedly rugose transversely. Additional material examined. Mexico. Baja California: Margin of arroyo 2 mi. S of El Valle Perdido along road to La Junta, 23°41 N, 110°06 W, 7 November 1959, Wiggins 15359 (MEXU). Campeche: Tuxpeña, 26 November 1931, Lundell 989 (MEXU). Chiapas: Mun. Cintalapa, 23 km west of Las Cruces along road to La Mina Microwave Station, 870 m, 30 October 1981, Breedlove & Davidse 54080 (MO); Mun. Villa Corzo, 56 km south of Mexican Highway 190 near junction to Jericó on road to Nueva Concordia, 700 m, 2 November 1981, Breedlove & Davidse

Ungrouped Species

261

54401 (MO); Mun. Ocozocoautla de Espinosa, 13  km south of Ocozocoautla, 900 m, 29 October 1981, Breedlove & Davidse 54047 (MO); Mun. of Chiapa de Corzo, above El Chorreadero, 800 m, 29 October 1981, Breedlove & Davidse 54030 (MO); Mun. Amatenango de la Frontera, along Río Cuilco between Nuevo Amatenango and Frontera Comalapa, 1100 m, 20 November 1976, Breedlove 41585 (MEXU, MO); 13 km E of Ocozocoautla on Rte. 190, then N on road to Aguacera, 630–700 m, 2 October 1984, Huft et al. 2285 (MO); along road from Huixtla to Motozintla de Mendoza, 4.8  km  N of Huixtla, 200  m, 13 February 1979, Croat 47521 (MO); ca. 14 miles S of Escuintla along gravel road to Finca Tres de Mayo, ca. 4 miles NE of El Triunfo, 400 m, 21 August 1977, Croat 43887 (MO); Mun. Ocozocoautla, El Aguacero, canyon of the Río La Venta, 540–600 m, 19 November 1984, Davidse et al. 30063 (MO); en los alrededores de la zona arqueológica de Palenque, 2 August 1984, Cabrera & Cabrera 7002 (MEXU, MO); Mun. Mazatán, El Roble, 16 October 1984, Ventura & López 503 (MO); Mun. Cintalapa, 23 km west of Las Cruces along road to La Mina Microwave Station, 870 m, 19 September 1981, Breedlove 52930 (MO). Colima: 2 km all E de Agua de la Virgen, brecha a Las Conchas, 200 m, 17 October 1984, Santana & Cervantes 752 (MEXU); Colima, 22 October 1910, Orcutt 4515 (MEXU). Guerrero: Puerto Marqués, terreno frente a la gasolinería Revolcadero, 15 September 1981, López-Forment 1417 (MEXU); Copacabana, 15 September 1968, Boege 848 (MEXU). Jalisco: near Guadalajara, Hitchcock s.n., Amer. Gr. Hb. 70 (MO, P); barranca near Guadalajara, 23 September 1891, Pringle 3828 (MEXU, MO; P, W); Rancho El Nogal, km 80 brecha Tamazula a Manuel M. Dieguez, 19 November 1973, Díaz Luna 4550 (MEXU). Michoacán: cerca de Rancho Viejo, camino Tierras Blancas-Bastán del Cobre, 610 m, 6 October 1981, Núñez & Silva 3196 (MEXU, MO); west facing sloopes of Sierra Madre del Sur, ca. 32  km north of Playa Azul, 25 October 1961, King & Soderstrom 4927 (MEXU). Nayarit: Mun. Nayar, 8 km al NW de San Juan Peyotán, en los límites de Nayarit y Durango, camino a San Juan Peyotán, 22°30′N, 104°30′W, 600  m, 21 September 1989, Flores & Tenorio 1180 (MO); a 10 km al S de Ahuacatlán, camino a Amatlán de Cañas, 21°00′N, 104°30′W, 19 October 1986, Téllez 9906 (MO); María Madre, Tres Marías Island, 22 October 1925, Ferris 5605 (MO); Ojos de Agua, near Yxtlán del Río, 1100 m, 23 September 1926, Mexía 747 (MO). Oaxaca: a 2.5 km al N de Candelaria Loxichia, camino Pochutla-Oaxaca, 600 m, 24 October 1982, Martínez et al. 2413 (MEXU, MO); 2 km al S de Valle Nacional, carretera Tuxtepec-Ixtlán, 11 March 1983, Tenorio et al. 3378 (MEXU); a 2 km al N de Putla de Guerrero, 28 October 1980, Téllez 3911 (MEXU, MO); 67 km al S de Tecomavaca, 7–10 km al S de Cuicatlán, 17°41 N, 96°56 W, 16 September 1991, Salinas 6163 (MEXU); orilla E de la Presa de Temascal, 100  m al N de la cortina, 60  m, 23 November 1986, Mendoza et al. 2720 (MEXU); Cerro Guiengola, 10 km al NW de Tehuantepec, 1100  m, 27 September 1984, Torres & Cabrera 6322 (MEXU). Puebla: Yancuictlalpán, Cuetzálan, 17 October 1980, Basurto & Patrón 177 (MEXU); El Jengibre, Mun. de Jalpán, 200 m, 1 November 1966, Cruz Cisneros 1490 (IPN). Quintana Roo: 5  km al norte de la carretera transversal, sobre el camino a la zona Arqueológica de San Gervasio, Isla de Cozumel, 25 November 1985, Cabrera & Cabrera 9877 (MEXU); Tres Reyes, 12 November 1980, Cabrera

262

 

Taxonomic Treatment

& Cortez 273 (MEXU); a 15 km al norte de Cobá, rumbo a Nuevo Xcan, 19 October 1980, Téllez et al. 3811 (MEXU). San Luis Potosí: in a tropical forest along Mexico highway 55 to Xilitla, 23 September 1954, Sohns 1448 (MEXU). Sinaloa: Las Mesas, Sierra Surotato, 15 September 1941, Harvey 6610 (MO). Tabasco: km. 21 de la carretera Cárdenas-Coatzacoalcos, 11 August 1981, Suárez 24 (MO); Grutas de Ocona near Teapa, 20–100 m, 12 November 1984, Davidse et al. 29501 (MO). Veracruz: Catemaco, 5 km al este del Pen. de Moreno, 3 October 1986, Ortíz 1093 (MO); Salto de Eyipantla, 5 km (by air) S of San Andrés Tuxtla, 150 m, 18°23′N, 95°12′W, 4 December 1981, Nee 23649 (MO); Mun. Amatlán de Los Reyes, 3 km antes de la Pesca, 18 November 1990, Oliva & Ramírez 882 (MEXU); in a banana plantation at Pueblo Nuevo, about 4  miles from El Palmar Rubber Experiment Station, 12 August 1953, Reeder & Reeder 1976 (MEXU); Ceiba Blanca, 60 m, 25 October 1973, Dorantes et  al. 2263 (MEXU). Yucatán: San Anselmo, Gaumer 895, 2027 (P); 22 km al NE de la ciudad de Mérida, en la Zona Arqueológica de Dzibilchaltum, 24 November 1986, Cabrera & Cabrera 12881 (MO) Guatemala. St. Thomas, 29 May 1909, Deam 6041 (MO). Chontales: without locality, Friedrichsthal 859 (W). Izabal: vicinity of Quiriguá, 15 May 1922, Standley 24577 (MO). Petén: Tikal National Park, Ortiz 228 (MO, US). Santa Rosa: Santa Rosa, November 1892, Heyde & Lux 4299 (MO). Suchitepequez: 3 km east of Río Mora on CA-2, 30 July 1970, Harmon & Dwyer 3394 (MO) Belize. Cayo: Along Sibun river near crossing of Humming Bird highway, Gentry 8450 (SI); in high ridge, base of hill, 45 Miles Section, Humming Bird Highway, 22 February 1956, Gentle 9037 (MO); along Sibun River, just west of Hummingbird Highway, 200–300  ft, 21 June 1973, Croat 24867 (MO). Stann Creek: Monkey River, Southern Highway, sandy open bar, 25 March 1970, McDaniel 12933 (MO). Toledo: Mira Mar Hill, Edwards Road beyond Columbia, 30 December 1947, Gentle 6351 (MO); Maya Mountains, canyon along Bladen Branch from Richardson Creek to Quebrada de Oro, 16°32′N, 88°46′W, 100–200 m, 12 March 1987, Davidse & Brant 32392 (MO); Solomon Camp, vicinity of the junction of Richardson Creek and Bladen Branch, foothills of the Maya Mountains, 16°32′N, 88°45′W, 12 March 1987, Davidse & Brant 32042 (MEXU, MO) Honduras. Atlántida: Lancetilla, 5 km SW de Tela, Canabes 86 (SI); orilla del Río Cangrejal, 10–20 km S de La Ceiba, 3 May 1980, Nelson 5467 (MO); bank of Danto river, slopes of Mt. Cangrejal, 1000 ft, 2 August 1938, Yuncker et al. 8777 (MO); 20 km E of La Ceiba, along Hy. to Jutiapa, weedy pasture along a large river, 30 m, 12 December 1977, Pohl & Gabel 13322 (MO). Choluteca: caserío 10 km al N de Jícaro Galán, 30  m, 2 December 1975, Nelson et  al. 3186 (MO). Colón: Trujillo Colón, 20 April 1984, Ramos 190 (MO); carretera Trujillo-Bonito Orientall, 12 March 1983, Delgado 133 (MO). Copán: Vivistorio, 13 December 1976, Villeda 30 (MO). Cortés: 5 km W of Choloma, on road to Jutosa, along a small stream, 100 m, 24 December 1977, Pohl & Gabel 13382 (MO); a lo largo de quebrada El Encanto, Montaña La Cumbre, 200 m, 5 December 1950, Molina 3591 (MO). El Paraíso: Finca “La Blonda”, Planes Cifuentes, 82 km NE de Danlí, 26 December 1981, Escobar 162 (MO). Francisco Morazán: Rio de la Orilla, southeast of El Zamorano, 750–800 m, Standley 24175 (P); orilla Quebrada La Orejona, 28 October

Ungrouped Species

263

1980, Borjas 94 (MO). Gracias a Dios: orillas del río Sicre o Sikri, 24 January 1976, Vargas et al. 401 (MO). Islas de la Bahía: camino entre Roatán y Sandy Bay, 20 March 1978, Nelson & Romero 4470 (MO). Olancho: orillas de la quebrada El Salto, Campamento, 800 m, 4 September 1982, Pineda 16 (MO); faldas del cerro El Boquerón, 20  km NE de Juticalpa, 700  m, 4 September 1983, Flores 49 (MO). Valle: Amapalal, Isla de Zacate Grande, 21 October 1978, Zelaya 229 (MO) El Salvador. San Miguel: Laguna de Jocotal, Volcán de San Miguel, 60 m, sin colector s.n. (MO-2325861). San Salvador: vicinity of Tonacatepeque, 30 December 1921, Standley 19527 (MO); vicinity of San Salvador, 30 March 1922, Standley 23187 (MO). Sonsonate: Finca Cuyancuya, 380 m, Rohweder 1327 (MO) Nicaragua. Boaco: 4 km al S de Boaquito, “San Antonio,” 12°26′N, 85°44′W, 200 m, 21 October 1982, Moreno 18081 (MO); 2 km al N de Boaquito, camino a Santa Lucía, 12°28′N, 85°44′W, 300  m, 21 October 1982, Moreno 18117 (MO); 1  km al S del empalme Juigalpa-Camoapa, Río Quizaltepe, 140  m, 28 October 1983, Grijalva & Aranda 3199 (MO). Carazo: 2  km al N de Jinotepe, 11°51′, 86°13′W, 560 m, 31 August 1981, Moreno 10741 (MO). Chinandega: km 146.5 carretera a El Guasaule, Volcán Chonco, 12°42′N, 87°03′W, 500–600  m, 18 November 1982, Sandino 3831 (MO). Estelí: 4.9–7.6 km NE of Hwy. 1 at Estelí along road to Yalí, ca. 13°08′N, 86°19′W, 820–1100 m, 14 November 1979, Stevens & Grijalva 15518 (MO). Granada: W slope of Volcán Mombacho, road and trail above Finca Santa Ana, reservoir to somewhat above Plan del Flores, ca. 11°49′N, 85°58′W, 950–1150 m, 1 October 1977, Stevens 4346 (MO); environs de Granada, Levy 37 (G). Jinotega: El Cedro, 20 km al N del Valle del Cuá, 13°30′N, 85°37′W, 19 June 1980, Moreno 830 (MO); camino San Sebastián-Loc. Laurelels-Tuma, 25 November 1981, Téllez et al. 5214 (MEXU). León: along Hwy. 12, ca. 1 km SE of junction with Hwy. 28 (first quebrada Se of junction), ca. 12°15′N, 86°43′W, 30 m, 17 December 1978, Stevens 11138 (MO). Madriz: lower W slope of Cerro Volcán de Somoto, ca. 13°25′N, 86°35′W, 900–1300  m, 13 December 1979, Stevens & Grijalva 16351 (MO). Managua: Municipio San Francisco Libre, 1 km antes de llegar al poblado, 12°30′N, 86°17′W, 50 m, 21 October 1980, Moreno 3861 (MO). Masaya: Laguna de Apoyo, 11°56′N, 86°02′W, 100–140 m, 20 September 1981, Moreno 11157 (MO). Matagalpa: km 81 sobre la Carretera Panamericana, 1 km al N de Las Calabazas, 12°40′N, 86°05′W, 400 m, 1 November 1982, Moreno 18380 (MO). Nueva Segovia: 3 km west of Ocotal, 20 December 1968, Dudey 775 (MO). Rivas: camino entre La Virgen y San Juan del Sur, bosque tropical seco, 17 December 1980, Guzmán & Castro 1562 (MO); Sandy and gravelly shore of Lago de Nicaragua, S of dock at Moyogalpa, Isla de Ometepe, 11°32′N, 86°41′W, 50 m, 16 September 1983, Nee & Téllez 28161 (MO). San Carlos: en la Isla Donald Guevara, archipiélago de Solentiname, Lago Nicaragua, 18 September 1982, Martínez 2265 (MEXU, MO). Zelaya: Corn Island, N and W shore, Sand Fly Point to Southwest Bay, and NW part of Island, ca. 12°09′N, 83°02′W, 4 April 1981, Stevens 19948 (MO); Río Matis, 16 September 1982, Ortíz 188 (MO) Costa Rica. Puerto Viejo, Holdrigen Finca, Lathrop 5558 (COL). Cartago: Turrialba, Weston et al. 3309 (US). Guanacaste: Finca Taboga, 6 mi. S, 19 July 1968, Koch 4899 (MO); Río Potrero, c. 2 km west of the C.I.A., 200 m, 26 January

264

 

Taxonomic Treatment

1983, Garwood et al. 624 (MO); La Pacífica, 50 m, 4 km NW of Cañas, 21 November 1972, Opler 1578 (MO); Santa Rosa National Park, 30  km NW of Liberia, ca. 10°50′N, 85°35′W, 300  m, 27 January 1985, Janzen 12425 (MO); Cuajiniquil, Santa Elena Península, 30 m, 23 October 1968, Pohl & Davidse 11305 (MEXU). Heredia: Finca La Selva, the OTS Field Station on the Río Puerto Viejo just E of its junction with the Río Sarapiquí, 100 m, 14 May 1981, Folsom 10106 (MO); Finca La Selva, near Puerto Viejo, ca. 50  m, 25 August 1968, Pohl & Davidse 11013 (MEXU). Limón: quarry and edge of cacao grove, near Recope Refinery, 20 m, 15 June 1973, Pohl 12924 (MO); Hacienda Tapezco-Hacienda La Suerte, 29 air km W of Tortuguero, 10°30′N, 83°47′W, 18 August 1979, Davidson & Donahue 8481 (MO); Tortuguero, in primitive cornfield, 15 December 1974, Pohl & Lucas 13042 (MO). Puntarenas: hills near 27 Abril and along the road to Playa Tamarindo, Nicoya Peninsula, 10°16′N, 85°46′W, 9 November 1975, Burger & Baker 9920 (MO); Cantón de Puntarenas Monteverde, 10  km  S Monteverde on road to Inter American Highway, area of spring, 10°16′N, 84°50′W, 900 m, 10 December 1989, Haber & Zuchowski 9656 (MO). San José: vicinity of El General, 640 m, January 1939, Skutch 3977 (MO) Panama. Bocas del Toro: One cove N of Trouble Hole, 30–100 m, 16 February 1989, Peterson & Annable 6885 (MO, US); Isla Colón, N side of island, flat rock beach, 30 January 1989, Peterson & Annable 6453 (MO, US); Isla San Cristóbal, 1 km W of Bocatorito, 45 m, 3 February 1989, Peterson & Annable 6600 (MO, US). Chiriquí: vicinity of Puerto Armuelles, 28 July 1940, Woodson & Schery 865 (MO). Coclé: El Valle de Antón, edge of cloud forest and roadside, 2 December 1967, Lewis et al. 2601 (MO). Colón: vicinity of Sardinilla, 7-8 mi. E cement plant, 12 September 1965, Blum & Tyson 481 (MO); vicinity of Río Indio on road from Portobelo to Nombre de Dios, 50 m, 23 March 1976, Croat 33618 (MO). Darién: clearing at confluence of R.  Chucunaque and Río Canclones, 5 July 1962, Duke 5107 (MO); village of Mannene, 30 April 1968, Kirkbride Jr. & Bristan 1582 (MO); trail from El Real to crossing of Río Pirre, 10 July 1962, Duke 5181 (MO). Los Santos: Playa del Pueblo de Cambutal, 26 November 1983, Correa et  al. 4382 (MO). Panamá: Cuevas de Chilibre, Sucre 142 (SI); Pan-Am Hwy. at Río Mamoni ca. 4 mi. beyond Chepo, 11 September 1962, Duke 5585 (MO); Río Chagres, 1 mi. above Madden Lake, 7 October 1961, Duke 4470 (MO); tributary of Río Chagres, 5 miles SW of Cerro Brewster, sandy and rocky river banks, 14 December 1967, Lewis et al. 3416 (MO). Veraguas: Puerto Mutis, 12 miles south Santiago, common grass in shade, 27 December 1968, Tyson 6018 (MO). Zona del Canal: vicinity of Salamanca Hydrographic Station, Río Pequení, ca. 80 m, 28 July 1938, Woodson Jr. et al. 1621 (MO); in Government forest along Las Cruces Trail, 75 m, 25 February 1935, Hunter & Allen 727 (MO); Barro Colorado Island, Wheeler Trail at tower, July 1931, Starry 102 (MO); Barro Colorado Island, along shore west of dock, caespitose, 12 July 1960, Ebinger 386 (MO); border of thickets just inland from Playa Kobbe, 11 January 1971, Wilbur & Teeri 12997 (MO) Cuba. Havana: Habana, 10 April 1905, Curtiss 714 (MO, P). Oriente: lower northern slopes of main ridge above Río Yao, Sierra Maestra, 300–1000  m, 27 October 1941, Morton & Acuña 3507 (MO). Without locality, Wright 1538 (G, MO)

Ungrouped Species

265

Jamaica. Langley, 26 February 1961, Adams 9033 (MO); Castleton, Harris 11299 (MO, P), 11328 (MO, P); Cinchona, 17 October 1912, Harris 11309 (MO); Bog Walk, 17 December 1890, Hitchcock s.n. (MO); Ross Craig, 15 July 1960, Adams 7574 (MO) Haiti. Vicinity of Plaisance, Departmente du Nord, 400  m, 27 January 1926, Leonard 9207 (MO); vicinity of Jean Rabel, 27 January 1929, Leonard & Leonard 12556 (MO); vicinity of La Vallée, Tortue Island, 28 December 1928, Leonard & Leonard 11720 (MO) Dominican Republic. La Estrechura, Rancho Arriba, 700  m, 16 December 1973, Liogier 20856 (MO). La Vega: vicinity of Piedra Blanca, 200–500  m, 22 January 1946, Allard 14803 (MO). San Cristóbal: Cordillera Central, along road from Hato Damas to Mano Matuey, 15 June 1988, Croat 68576 (MO). Without locality, Eggers 1174 (P, W) Puerto Rico. Mayaguez, Sintenis 160 (G, P); on the Adjuntas road, seven miles from Ponce, 22 November 1902, Heller 6094 (MO, P); Caguas, Cerro Pelucho, 380–535 m, 14 April 1991, Axelrod & Axelrod 2334 (MO); El Yunque, March 1959, Martorell & Medina 429 (MO); Monte Montoso, 13 February 1915, Britton & Cowell 4142 (MO) Antigua. McCarthy valley, Box 90 (K) Guadalupe. Camp Jacob, P. Duss 674 (P) Martinica. Environs de St. Pierre, Belanger 389 (P), Hahn 481 (W). Without locality, Hahn 1047 (G, K, MO, P), Belanger 438 (P), Duss 1321 (MO), Sieber 28 (G, W), Stehle s.n. (P) St. Vincent. St. Patrick Parish, lower Bellewood, Cooley 8165 (K) Colombia. Amazonas: Trapecio amazónico, Loretoyacu river, October 1946, Schultes 8528 (COL). Antioquia: 1  km antes del Puente de Occidente, Scolnik et  al. 19jAn327 (F); Mun. San Luis, Cañón del Río Claro, sector norte, Parque Ecológico, 325  m, 13 August 1983, Cogollo 622 (COL, HUA, MO); vicinity of Planta Providencia, 26 km S & 23 km W (air) of Zaragoza, 7°13′N, 75°03′W, 400– 700 m, 31 May 1974, Denslow 2267 (MO); Mun. Dadeiba, 27 km después por la carretera al March, 250 m, 28 September 1986, Betancur et al. 261 (MO); Mun. Anorí, vía Dos Bocas-Providencia-Toma de Aljibes, a lo largo del rio Anorí, 310 m, 75°08′W, 7°18′N, 12 July 1987, Callejas et al. 4585 (MO); Brazuela de Perales, Río Magdalena, 10 January 1918, Pennell 3702 (MO); Quebrada “La Cristalina”, 6°N, 74°45′W, 770  m, 27 September 1987, Ramírez & Cárdenas López 1733 (MO). Atlántico: Barranquilla, Elias 560 (RB); entre Baranoa y Galapa, Dugand 5877 (COL). Bolívar: vicinity of Turbaco, 200–300  m, 22 November 1926, Killip & Smith 14179 (MO). Caldas: Santagueda, 7 April 1986, Gutiérrez 36 (COL). Cauca: Municipio de Guapi, Parque Natural Gorgona, Isla de Gorgonillo, 14 September 1987, Lozano et al. 5870 (COL, MO). Cesar: Mun. Valledupar, entre Atanquez y Guatapurí, 18 November 1985, Torres 2864 (COL). Chocó: Mun. Acandí, camino hacia el Darién, Quebrada del Limón, 200  m, 16 June 1976, Forero 407 (COL, MO); Río Jurado, 100  m, 5 September 1940, von Sneidern A162 (COL, MO). Córdoba: Mun. Tierralta, Cerro Paragüillo, Fonnegra 7937 (HUA). Guaviare: margen derecha del Río Guayabero, Pinto 345 (COL). Huila: Mun. El Agrado,

266

 

Taxonomic Treatment

quebrada La Yaguilda, September 1986, Fernández-Alonso 6844 (COL). La Guajira: Serranía La Macuira, entre el Arroyo y la duna Arehuara, Saravia 2360 (COL); intersection of Río Ranchería and Carbocol, 15 November 1980, Bunch et  al. 205 (HUA, MO). Magdalena: Foothill of the Sierra of Perijá, Manaure, Cuatrecasas & Romero Castañeda 24998 (COL); Santa Marta, Smith 167 (COL, MO, P). Meta: margen derecha del Río Gayabero, Pinto & Bischler 345 (COL). Nariño: Mun. Tumaco, vereda Tangarreal, corregimiento Jorge H.  Leal, 12 June 1998, Quevedo 93 COL). Quindío: Mun. Montenegro, Pueblo Tapado, Finca La Esmeralda, 25 June 1989, Zuloaga & Londoño 4185 (COL, MO, SI). San Andrés, Providencia y Santa Catalina: isla de Providencia, Freshwater Bay, 50  m, 25 January 1984, Wood 4241 (COL). Santander: between San Vicente et Barranca Bermeja, Fassett 25534 (USA). Sucre: Archipiélago de San Bernardo, Isla Mucura, Moreno & Lopez 10 (COL). Tolima: La Trinidad, Líbano, 21 December 1917, Pennell 3358 (MO). Valle del Cauca: Cisneros, open shale bank alon Río Dagua, 300–500 m, 5 May 1939, Killip 35596 (COL).Vaupés: desembocadura del Ariari en el Río Guayabero, Cabaña del Incora, Pinto & Sastre 933 (COL, P) Venezuela. Anzoátegui: alrededores de Pariaguán, Hernández 77 (VEN); Fundo El Saman, entre Quebrada El Onoto y el río Cabrutica, 11 September 1984, Montes 2584 (MO); carretera Puerto La Cruz a El Tigre, Muller 815 (VEN). Apure: 15 km NW of Achaguas along hwy. to San Fernando de Apure, 120 m, 9 November 1973, Davidse et al. 3930 (MO). Aragua: 16.3 km NW of Rancho Grande along road to Cata, 270 m, 15 November 1971, Davidse 3118 (MO, US, VEN); Mun. Autónomo Girardot, Parque Nac. H.  Pittier, entre La Trilla y la Estación Rancho Grande, 10°21′N, 67°43′W, 500  m, 23 August 1989, Zuloaga & Ortíz 4524 (MO, MY, PORT, SI, VEN); Colonia Tovar, Fendler 2499 (MO). Bolivar: on edge of plateau of Cerro Venamo, in vicinity of km 125 and campamento 125, between Luepa and El Dorado, 1100  m, Steyermark & Nilsson 810 (NY, VEN); Santa Marchía del Vapor, Distr. Roscio, Río Yuruan al E de El Dorado, Delascio et al. 8343 (VEN); Distr. Cedeño, vicinity of Panare village of Corozal, 6 km from Maniapure toward Caicara, 6°55′N, 66°30′W, 25 September 1985, Boom & Grillo 6106, 6115 (MO). Carabobo: Hacienda de Cura, cerca de San Joaquín, Pittier 8214 (VEN). Cojedes: carretera San Carlos-Tinaco, Delascio 3457 (VEN); 7  km  N of Los Colorados, which is just West of San Carlos, 220 m, 16 November 1971, Davidse 3136 (MO); Quebrada de Tierra Caliente, al N de Manrique, Trujillo 13904 (F); San Carlos, Box 3908 (VEN), 200 m. Distrito Federal: Caracas, Eggers 13006 (F); along Rio de las Caracas, above town of Caracas, 11 February 1973, Croat 21646 (MO, US); valley of río Chichiriviche, 0.5 km south of Chichiriviche, 10 October 1976, Steyermark & Carreño 112723 (MO, VEN). Falcón: Cerro Santa Ana, ascensión del lado sur desde el pueblo de Santa Ana, 750  m, 24 January 1966, SteyerMarchk & Braun 94643 (MO). Guárico: Río Orituco and road intersection, ca. 16 km S of Calabozo, 1 km downstream, 10 November 1971, Davidse 2972 (MO, VEN); sabanas al S de Calabozo, Bolaños s.n. (VEN); 15  km  SE of Tamaco along main hwy. between Altagracia de Orituco and Chaguaramas, 150 m, 19 November 1973, Davidse 4213 (MO, VEN). Miranda: Cerros del Bachiller, near east end, between Quebradas Corozal and Santa Cruz, south of Santa Cruz, 10°09′N, 65°48′W, 18 March 1978,

Ungrouped Species

267

Steyermark & Davidse 116427 (MO, VEN); Santa Lucía, Holt 561 (VEN); Distr. Paez, ca. 1–2 km upstream from the mouth of the rio Chiquito at its intersectecion with the rio Caura, S of El Guapo, 3 June1977, Davidse & González 13589 (MO, VEN); Distr. Brión, selva siempre verde a lo largo de la quebrada afluente del río Aricagua, 3.9 km oeste del Pueblo Seco, 1.6 km oeste de Aricagua, 75 m, 24 March 1973, Steyermark & Carreño 106943 (MO, VEN); alrededores de Capaya, Badillo 793a (VEN); Parque Nacional Guatopo, 22 km NNW Altagracia de Orituco, Nee 17751 (F, VEN); Parque Nacional Guatopo, near park headquarters at south entrance, 13 February 1973, Croat 21701 (MO). Monagas: La Pica near Maturín, Chase 12574 (VEN). Portuguesa: faldas pedregosas arriba del puente sobre el Río Maria, NE de Boca de Monte, 28–32  km al NNE de Guanare, 400–500  m, 2 November 1982, Steyermark et al. 127074 (MO, SI, VEN); Dtto. Guanare, terrenos de la UNELLEZ, Mesa de Cavacas, 200 m, 4 October 1988, Aymard et al. 7078 (MO, VEN); a 10 km al SO de Guanare, carretera Guanare-Papelón, 27 October 1982, Aymard & Ortega 1413 (MO, PORT); a lo largo de la carretera DesembocaderoSan Isidro, 32 km al NW de Guanare, 600–1000 m, 17 November 1986, Aymard & Ortega 4930 (MO, PORT). Sucre: Distr. Arismendi, Río Marchauco, sector Guayabero, 26 December 1983, Flores et al. 400 (MO, PORT); Península de Paria, dry hills between Cangua and San Juan de las Galdonas, 10°42′N, 62°51′W, 100 m, 2 December 1979, Steyermark & Liesner 120970 (MO, VEN). Táchira: Distr. Cordoba, Cerro La Camiri, just S of the town of Rio Negro, 7°36′N, 72°10′W, 430– 530 m, 6 November 1982, Davidse & González 21562 (MO, VEN). Yaracuy: San Felipe, Burkart 16389 (SI, VEN). Zulia: Distrito Colón, Hacienda La Rosita, km 20 de la carretera a Concha, Steyermark et al. 100143 (P, VEN); Sierra de Perijá, a lo largo del Río Yasa, vecindad de “Kasmera”, al SW de Machiquies, 300 m, 23 August 1967, Steyermark & Fernández 99665 (MO, VEN); Distr. Mara, cerca de Caño del Paso, Medina 847 (VEN); Dtto. Bolívar, entre El Pensado y Las Tres Marías, trayecto de 8 km al este de El Pensado, 22 July 1980, Bunting 9396 (MO, VEN); Dtto. Mara, cuenca del Río Guasare, alrededores del Destacamento Guasare 1 (La Yolanda), 10°52′N, 72°29′W, 200–250 m, 9 November 1982, Bunting et al. 12029 (MO, VEN); Los Ángeles de Tucuco, 23 June 1980, Davidse et al. 18474 (MO) Trinidad and Tobago. Maqueripe Bay, on sandy cliff base, 19 August 1976, Adams & Bhorai 14132 (MO); Arima, Eggers 960 (G, K, P, W); St. Ann’s Cascade, in banks, 12 November 1923, Broadway 5058 (MO); without locality, Fendler 929 (P) Guyana. Coast región, Jenman 7740 (NY, P, US), 7760 (NY, US); Akyma, Hitchcock 17431 (US); Issorora, Hitchcock 17547 (US); Aruka River, Bartlett 8577 (US); Demerara, Coast Región, Jenman 4403 (NY, US); Georgetown, Hitchcock 16598 (IAN, MO, NY, P, US); Tumamatari, Gleason 322 (NY, US); Wauna, Koriabo River, Archer 2360 (US); Waini River, Northwest District, 8°20′N, 59°40′W, 18 April 1923, de la Cruz 3772 (MO, NY, US); Rupununi, E of Meritiziero, Goodland 556 (NY, US) Suriname. Paramaribo, Samuels 455 (NY); without locality, Kappler 403 (NY) French Guiana. 46–48 km, route de Kourou, Hoock 134 (IAN, NY, P); Cayenne, Broadway 501 (NY, US); without locality, Leprieur s.n. (P)

268

 

Taxonomic Treatment

Ecuador. Guayas: El Progreso along road between Salinas and Guayaquil, 80°23′W, 02°13′S, 50 m, Barfod et al. 48489 (MO); Tenguel, 6 July 1946, Giler 1756 (MO). Los Ríos: SAn Antonio de Columa, km 39, on the road PuebloviejoGuaranda, 1°37′S, 79°18′W, 20 April 1980, Holm-Nielsen 22896 (MO); Hacienda Clementina on Río Pita, Asplund 5281 (P, US), 5575 (R); Represa Daule-Peripa, Río Conguillo, 2 June 1985, without collector  229 (MO); Jauneche Forest, Canton Vinces, between Mocachi and Palenque on Estero Peñafiel, 15 August 1978, Dodson et al. 7045 (MO); surrondings of Montalvo, 40 km E of Babahoyo, 1°47′S, 79°17′W, 100–200 m, 30 March 1973, Holm-Nielsen et al. 2630 (F, MO); 7 km E of Patricia Pilar, 28 July 1980, Hansen et al. 7789 (MO); Río Palenque Field Station halfway between Santo Domingo de Los Colorados & Quevedo, 200 m, 25 February 1974, Gentry 10185 (MO). Morona-Santiago: Centro Shuar Yukutais, 10 km S Sucua, Río Upano, 2°31′S, 78°09′W, 24 September 1988, Gómez 458 (MO). NapoPastaza: between Tiputini and Lagartococha, January 1953, Fagerlind & Wibom 2262 (MO). Pichincha: NE of Vicente Maldonado, reserva de ENDESA, 79°02′W, 00°06′N, 600 m, 2 June 1984, Laegaard 52215 (MO); Río Chiquilpe near junction with Río Baba, 7 km east of km 7 on highway Santo Domingo-Quevedo, 420 m, 12 July 1979, Dodson et al. 7966 (MO) Peru. Amazonas: Huánuco: across Río Huallaga from Tingo María, Leonicio Prado, 650 m, 17 January 1976, Gentry et al. 15966 (MO). Junín: San Francisco de Satipo, hillsides along Río Satipo, 700  m, 25 June 1977, Solomon 3305 (MO). Loreto: Maynas, Dpto. Iquitos, Padre Island, eastside, ca. 2  km above Entrada Amazonas, McDaniel & Rimachi 23032 (MO, US); Coronel Portillo, Ivita, km 59 from Pucallpa toward Tingo María, 270m, 21 January 1976, Gentry & Revilla 16127 (MO); Prov. Requena, trocha al Río Yavari, km 7, 27 August 1976, Revilla 1234 (MO); Prov. Maynas, Fatima, 3°40′S, 72°55′W, 118  m, 12 August 1980, Vásquez et  al. 446 (MO); Río Ampiyacu, Pebas and vicinity, Isla Gallinazos, Plowmann et al. 6550 (P); Prov. Maynas, carretera de Zungaro Cocha, cerca a la quebrada de Shushuna, en terrenos arenosos, 16 February 1983, Rimachi 6572 (MO); Prov. Maynas, Quebrada Yanomono, Explorama tourist camp, above mouth of Río Napo on Río Amazonas, 130  m, 5 November 1979, Gentry et  al. 27491 (MO). Madre de Dios: Prov. Tahuamanu, Río Tahuamanu, 180 m, 16 November 1973, Chávez Alfaro 1730 (MO); Cashucocha Camp, Río Manú, Parque Nacional de Manú, 380 m, 19 October 1979, Gentry et al. 26929 (MO); Prov. Manú, Parque Nacional del Manú, Río Manú, Cocha Cashu Station, 11°50′S, 71°25′W, 5 August 1984, Foster 9772 (MO, SI). Piura: hills of Chiarnique, ca. 20 km E.N.E. of Naupe, ca. 500 m, 5 June 1978, Barbour 2145 (MO). San Martín: on trail from Tarapoto to San Antonio, north-facing slope of Río Cumbasa, 1000  ft, 30 August 1937, Belshaw 3338 (MO); Puerto Pizana, margen derecha del río Huallaga, 350–400 m, 22 June 1974, Schunke Vigo 6991 (MO); Prov. of Tarapoto, along Río Cumbaza, 6°29′S, 76°24′W, 350 m, 7 November 1980, Croat 51131 (MO). Tumbes: Zarumilla, Infantes 5061 (P). Ucayali: Quebrada Shesha, tributary of Río Abujao, 1-2  days upriver by “Peki-peki”, ca. 60–70 km NE of Pucallpa, 8°02′S, 73°55′W, 250 m, 18 June 1987, Gentry & Díaz 58422 (MO)

Ungrouped Species

269

BRAZIL. Alagoas: Bebedoura, vicinity of Maceió, 5 December 1924, Chase 7835 (MO, US). Amazonas: Parana do Catua, Solimoes, Kuhlmann 1195 (SI); Foz do Rio Vaupes, Froes 28241 (IAN). Bahia: Aramarí, Alagoinhas, 15 July 1981, Bastos 152 (IBGE, MO); 90 km ao N de Bonfim, Mina Caraiba, Castellanos 71 (HB, R); 49 km N of S. do Bonfim, on the BA 130 highway to Juazeiro, 500 m, 40°13′W, 10°05′S, 26 February 1974, Harley et al. 16356 (CEPEC, MO); Espigąo Mestre, 5 km S of Cocos, sandy cerradao, 530 m, 16 March 1972, Anderson et al. 37074 (MO, P, US); Paratinga, Silva 173 (US). Ceará: Beira do Rio Imbuzeiro, Lofgren 492 (R); Mun. Quixada, estrada Quixada-Banabuiu, km 16, Coradin et al. 1987 (CEN); Barra do Coco, Black 55-18365 (IAN); Taperaoba, Black 55-18350 (IAN); without locality, Allemao & Cysneiros 1616 (R). Goiás: Corrego Estrema, 35 km NW of Formosa, 800 m, 18 April 1966, Irwin et al. 14984 (MO). Maranhão: Caxias to Barra do Corda, Swallen 3559 (SP); Carolina, Swallen 3895 (SP); Mun. de Loreto, city of Loreto, 7°06′S, 45°08′W, 150 m, 2 March 1970, Eiten & Eiten 10812 (MO); Mun. de Loreto, Ilha de Balsas región, between the Rios Balsas and Parnaíba, Facenda Morros, 7°20′S, 45°04′W, 300 m, 30 April 1962, Eiten & Eiten 4474 (MO, SP); S. Luis, Snethlage 150 (F). Mato Grosso: Aquidauana, 26 February 1930, Chase 11062 (MO, RB); entre Pimenta Bueno e Presidente Hermes, Exp. Rondon, Kuhlmann 1754 (SI); 85 km S of Xavantina, Irwin et al. 17157 (NY, SP, UB, US); Rio Areioes, 12  km  N of Xavantina on the São Felix road, Ratter & Fonseca 3295 (UB). Minas Gerais: S. Terezinha, 18 February 1950, Macedo 2169 (MO, US). Pará: Fazenda Camburupy, near Soure, Swallen 4955 (RB); vicinity of Pará, Baker 5 (F); without locality, Spruce Panicum 13 (P, US, W); Santarém, Spruce 466 (P); Marabá, Rio Tocantins, Ilha da Praia, Froes & Black 24665 (IAN); Pombal, Porto Agronómico, Parana do Tapora, Black 52-15490 ((IAN); Bolterra, beira do Capoeira, Black 47-994 (IAN); San Miguel do Guamá, June 1919, Goeldi 278 (MO); Mun. de Capanema, Rio Quatipuru, in vicinity of Miraselvas, ca. 30 km by road west of Braganca, 50  m, 1°04′S, 46°59′W, 9 April 1980, Davidse et  al. 18105 (MO); Mun. Tucuruí, Breu Branco, km 40 south of Represa Tucuruí along highway BR 422, 17 March 1980, Plowman et  al. 9658 (MO). Pernambuco: Campo de Criaçao de Rio Branco, Ramalho 37 (RB); Olinda, 11 November 1924, Chase 7649 (MO, US); Olinda, restinga do Rio Tapado, Leal & Da Silva 6 (SI); Rio Formoso Tamandaré, Falcao et al. 794 (RB, SI); Mun. Petrolina, BR 428, PetrolinaSalgueiro, km 32, Coradin et  al. 1263 (CEN); entre Serra Talhada e Petrolina, Heringer et al. 12 (R, RB); Escola de Agronomia do Nordeste, Coelho de Moraes 851 (IAN); Garanhuns, 850–900 m, 26 November 1924, Chase 7787 (MO, US). Piauí: Floriano to Oeiras, Swallen 4162 (R, RB, SP). Rio Grande do Norte: Angicos, Swallen 4709 (RB, SP). Rondonia: Mun. Costa Marques, ca. 5 km NW of Costa Marques, 12°25′S, 64°14′W, 150 m, 29 March 1987, Nee 34549 (MO) Bolivia. Beni: Ballivian, Estancia Espíritu, clearing in forest behind the residence, 224  m, 5 April 1987, Renvoize 4687 (MO). Chuquisaca: Ciudad Rincón Escala, 2200 m, 21 May 1987, Murguia 85 (MO, SI). La Paz: Prov. Sud Yungas, Alto Beni, subiendo a la concesión de Sapecho, al lado del arroyo Sapecho, 15°30′S, 67°21′W, 500  m, 21 April 1989, Seidel & Vargas 2695 (MO, SI). Santa Cruz: Cordillera, Abapo-Izozog, Agricultural Res. Str. Bush near Rio Grande, 400 m, 13

270

 

Taxonomic Treatment

March 1981, Renvoize & Cope 3932 (LPB, MO, US); Buena Vista, 4 March 1967, Steinbach 784 (F, GH, MO); Andres Ibañez, Río Piray, 20 km de Santa Cruz, 20 March 1981, Renvoize & Cope 4002 (K, LPB, MO, US); Ñuflo de Chavez, Estancia La Dolorida, 1/2 way betweeen San Javier and Concepción, 16°15′S, 62°15′W, 500 m, 16 April 1986, Killeen 1971 (MO); Ñuflo de Chavez, Concepción, 16°10′S, 62°00′W, 500 m, 20 November 1984, Killeen 604 (MO); Ñuflo de Chavez, logging road 17 km N of Concepción, 15°50′S, 62°00′W, 400 m, 29 March 1986, Killeen 1892 (MO); Ñuflo de Chavez, Estancia Alta Vista, 16°05′S, 61°55′W, 400 m, 15 November 1984, Killeen 552 (MO). Tarija: Arce, 29,2 km S of Emborozu- Sidras road on road to Bermejo, 21 April 1983, Solomon 9881A (MO, SI); Quebrada Tatarenda, 18 km N de Yacuiba, camino a Villa Montes, 63°37′W, 21°50′S, 600 m, 7 April 1977, Krapovickas & Schinini 30990 (CTES, MO) Argentina. Jujuy: Santa Bárbara, N de Palma Sola, Kiesling et al. 5355 (SI). Salta: Orán, Spegazzini s.n. (BAB 14120); Aguas Blancas, Villamil 2932 (SI); Río Bermejo y Orán, Ragonese & Covas s.n. (GH); without locality, Mársico 1217 (BAB); Dpto. Libertador San Martín, entre Coronel Cornejo y Arroyo Seco, 9 km de Coronel Cornejo, 11 March 1987, Zuloaga & Morrone 3039 (MO, SI) 63. Panicum venezuelae Hack., Oesterr. Bot. Z. 51: 368. 1901. Brachiaria venezuelae (Hack.) Henrard, Blumea 3(3): 435. 1940. Type. Venezuela. Without locality, 25 July 1891, H.  F. A. von Eggers 13471 (holotype, W 19160023492; isotypes, C 10017114, L 1223253, US 01231584, fragment ex W). Fig. 70 Panicum ineptum Hitchc. & Chase, Contr. U.S.  Natl. Herb. 17(6): 509, Fig.  98. 1915. Type. Dominican. Santo Domingo, C. Bertero 817 (holotype, B, destroyed; neotype: Venezuela, Loma de León, Dtto. Iribarren, 20 September 1950, Tamayo 3743, MO 1311991, here designated; isoneotypes, NY, VEN, n.v.) Panicum berteronianum Mez, Bot. Jahrb. Syst. 56, Beibl. 125: 5. 1921, nom. illeg. hom., non Panicum berteronianum (Schult.) Steud., 1854. Type. Venezuela. Distrito Federal. La Guayra, September 1852, J.  Gollmers s.n. (lectotype, B_10_0249079, here designated; isolectotype, US 00148177, fragment ex B). Other syntypes. Haiti. Without locality, C. Bertero s.n. Brazil. Without locality, F. Sellow s.n.(B_10_0249078, US 001148178, fragment ex B) Plants perennial, caespitose, with short and inconspicuous rhizomes; culms erect to scandent on the surrounding vegetation, 20–80 cm tall; long culms sometimes prostrate, rooting and tillering at the nodes, acting as stolons; internodes 3–8 cm long, cylindrical to compressed, hollow, sparsely hairy to glabrous; nodes densely hairy, with long, usually retrorse, whitish hairs. Sheaths 1–4 cm long, commonly shorter than internodes, pale, striate, densely to sparsely hispid, with long whitish hairs, margins ciliate. Ligule membranous-ciliate, reduced, 0.3–0.5 mm long, with long hairs beneath at the base of blade; collar pale, pilose. Blades ovate-lanceolate to lanceolate, acuminate, 3–9 × 0.5–1 cm, flat, shortly pseudopetiolate, pseudopetiole ca. 0.2 cm long; base cordate, densely to sparsely hirsute, the margins long ciliate toward the basal portion, otherwise ciliate or scabrous, cartilaginous, midrib not manifest. Inflorescence terminal, long exserted, lax, 2–11  ×  1–3  cm, formed by

Ungrouped Species

271

Fig. 70  Panicum venezuelae. (a) Plant. (b) Ligular region. (c) Branch of inflorescence. (d) Spikelet, ventral view. (e) Spikelet, dorsal view. (f) Lower lemma. (g) Spikelet, ventral view, lower lemma with “glands”. (h) Spikelet, lateral view, upper glume and lower lemma with “glands”. (i) Spikelet, dorsal view, upper glume with “glands”. (j) Anthers of a cleistogamous flower. (k) Anthers of a chasmogamous flower. (l) Upper anthecium, dorsal view. (m) Upper anthecium, ventral view. (n) Caryopsis, embryo view. (o) Caryopsis, hilum view. (From Pinto 307, US)

272

 

Taxonomic Treatment

4–10 alternate, distant, short, first-order branches with spikelets arranged unilaterally on short pedicels; main axis angulate, long hirsute, pulvini densely pilose, branch axes triquetrous, densely hirsute; axillary panicles shortly exserted, few flowered. Spikelets ellipsoid, 2.5–3 × 1.1–1.3 mm, in pairs, greenish, glumes and lower lemma sparsely or densely hirsute; cleistogamous flowers usually present, less commonly chasmogamous; lower glume ovate, 1.5–1.7 mm long, 1/2 or more the length of the spikelet, 3(–5) nerved, midrib scabrous; upper glume 2.6–2.9 mm long, gibbose, 5 nerved, with coarse tuberculate hairs toward the apex or over the entire surface, 2 “glands” occasionally present toward the middle portion of the outer surface, inner surface scabrous; lower lemma longer than upper glume, apex more or less incurved, sparsely pilose, midportion flattened and glabrous, 5 nerved, lateral nerves distant from midrib, with or without 2 “glands”; lower palea elliptic to obovate, 1.6–1.8  ×  0.7  mm, hyaline, shortly pilose toward the upper margins, otherwise glabrous; lower flower absent; upper anthecium ellipsoid to obovoid, obtuse, 1.6–1.8 × 0.8–1.1 mm, pale; indurate, smooth, glabrous, shortly stipitate; lemma 5 nerved, 3 anthers, those of cleistogamous flowers small, 0.2–0.3 mm long, those of cassmogamous flowers 0.9 mm long. Caryopsis 1.3 × 1 mm; hilum punctiform, embryo less than ½ the length of the caryopsis. Distribution and ecology (Fig. 71). It inhabits from the Caribbean and Mesoamerica to Venezuela and northern Brazil. It is found in humid and shady places, in forest margins and also along small watercourses, between 100 and 2200 m asl. Panicum venezuelae is a species of margins or interior of forests, stoloniferous and with flat an cordate blade. The species is also characterized by having hairy spikelets, arranged unilaterally in the inflorescences, with cleistogamous and chasmogamous flowers present, and with or without glands present in the lower lemma. The systematic position of the species has been controversial over time. Henrard (1940) transferred it to the genus Brachiaria without clarifying the reasons for this modification. Brown (1977) correctly indicates that it does not belong to this genus because it is a non-Kranz species; similarly, Sendulsky (1978) retains this species within Panicum. Zuloaga and Sendulsky (1988) include the species in section Stolonifera of Panicum. Subsequently, Sede et  al. (2009) place P. venezuelae in clade B of their phylogenetic analysis, separated from clade A, in which they classify species of the new genus Ocellochloa Zuloaga & Morrone, both within the clade currently considered as the tribe Paspaleae J. Presl, which includes taxa with a basic chromosomal number of x = 10; these authors point out that P. venezuelae should be excluded from Ocellochloa because it possesses axillary inflorescences, cleistogamous spikelets and depressed, non-crateriform lower lemma glands. Within clade B, P. venezuelae is related, with low support, to Echinolaena standleyi (Hitchc.) Stieber [= Oedochloa standleyi (Hitchc.) C. Silva & R.P. Oliveira] and to Echinolaena minarum (Nees) Pilg. [= Oedochloa minarum (Nees) C.  Silva & R.P. Oliveira]; it differs from species of the genus Oedochloa C. Silva & R.P. Oliveira (da Silva et al. 2015) by the characters mentioned above. Consequently, and taking into consideration the morphological characteristics and phylogenetic position, the

Ungrouped Species

273

Fig. 71  Distribution of Panicum venezuelae

placement of P. venezuelae within the Paspaleae is still uncertain (Silva et al., in prep.). Additional material examined. Guatemala. Guatemala: near Fiscal, Standley 59580, 80411, 80463 (F, US), 80630 (F) Honduras. El Paraíso: Quebrada de El Muro, between Las Mesas and Yuscarán, 5 November 1951, Standley 29263 (F, US); Quebrada del Muro, road to Yuscarán, 5 November 1951, Swallen 11329, 11333 (MO, US). Morazán: near El Jicarito, along road toward El Pedregal, Standley 20874, 21640 (F), 6 November 1951, Swallen 11377 (US); road toward San Antonio de Oriente, rocky thicket, 15 November 1950, Standley 27498 (MO, US); trail from La Quince, El Zamorano, to El Jicarito, Standley 14567 (US), 21272 (F, US); campus of Escuela Agrícola El Zamorano, Quebrada El Gallo, 800 m, 29 July 1970, Pohl & Davidse 12458 (MO); mountains above El Jicarito, 900–1300 m, 6 November 1951, Standley 29327 (F); 8 km S of La Venta by road, 850 m, 20 June 1970, Davidse & Pohl 2155a (MO); El Pedregal, 18 September 1943, Rodríguez 878 (F) Costa Rica. Guanacaste: La Cruz, 12 November 2004, Herrera Ch.. 10227 (INB, MO)

274

 

Taxonomic Treatment

Cuba. Oriente: Barbi, Loma del Gato, Sierra Maestra, 8 November 1922, Ekman 15661 (MO, NY, US); Loma del Gato, S of Loma San Juan, 900 m, July 1921, León et al. 10190 (NY, US) Dominican Republic. Monte Cristi: near Arroyo Seco, Ekman 12608, 13085 (US); Puerto Plata, Bajabónico, 25 May 1929, Ekman 14499 (US). Santiago: San José de las Matas, 11 April 1930, Ekman 14602 (US), Jiménez 950 (US) Venezuela. Distrito Federal: Colinas de Yaguara, Tamayo 1449 (F, VEN); lomas arriba de Antimano, 1200  m, 4 December 1927, Pittier 12581 (NY, US, VEN); entre Caracas y La Guayra, 23 June 1946, Burkart 17013 (SI, VEN), en bosquecillos y quebradas húmedas; Hacienda Sosa, Tamayo 110 (VEN), Badillo 60 (VEN). Falcón: Serranía de San Luis, Fila Las Playitas, 21 km SSO de Coro, 400 m, 11 November 1977, Ruiz 2543 (MO, VEN); Cerro Santa Ana, Paraguaná, Lasser & Aristeguieta, 3357, 3360 (VEN). Lara: en cerros arriba del caserío Simara, aproximadamente 8 km de Bobara, 20 May 1978, Burandt Jr. v0192 (MO); Dtto. Moran, carretera San Miguel-Cubiro, 1 km abajo de Agua Negra, Burandt Jr. V0688 (NY). Mérida: Loma El Cazadero, 1250 m, 20 October 1943, Luces 152 (VEN). Táchira: southern slope of Páramo Zumbador, 1750 m, Box & Alayón 3761 (VEN) Brazil. Alagoas: Tapera, Pickel s.n. (US-1645543), 2469 (USA). Bahia: Rodovia BA-052, km 148 cerca de Baixa Grande, entrada a Fazenda Olhos de Agua, 12°00′00″S, 40°18′00″W, 21 July 2006, Zuloaga et al. 9060, 9064 (SI); Alagoinhas, Chase 8135 (US); ca. 5 km SE of Feira da Santana along Highway BR-324, 250 m, 31 March 1976, Davidse et al. 11697 (MO); 17 km NW of Catu along the road to Alagoinhas, 150 m, 1 April 1976, Davidse et al. 11716 (MO); camino de Santa Ines a Rio Bahia, Pinheiro 7855 (MO); Espigao Mestre, 6 km S of Cocos, 520 m, 16 March 1972, Anderson et al. 37036 (F, MO, NY); Serra do Itiuba, 6 km E of Itiuba, 10°41′S, 39°48′W, 500 m, 19 February 1974, Harley et al. 16203 (K, MO, NY, US); Rodovia Santa Ines a Rio Bahia aos 10 km, capoeira, 7 July 1972, Pinheiro 1855 (CEPEC, MO, US); Santa Terezinha, Bondar 2609 (SP, US); Paraguaçu-valley, Muritiba, Pinto 307 (US); Feira de Santana, 100 m, 28 December 1924, Chase 8066 (F, MO, NY, RB, US), 8070 (US); Cachoeira, vale dos rios Paraguaçu e Jacuipe, Pedra do Cavalo 475 (CEPEC); basin of the upper São Francisco River, just beyond Calderão, ca. 32 km NE from Bom Jesus da Lapa, Harley 21497 (CEPEC, K, NY); ramal a 29 km na estrada Caetité/Brumado, ca. 3 ms ramal adentro, caatinga, 19 February 1992, de Carvalho et al. 3783 (MO). Ceará: Serra de Baturite, morro de Ceu, Eugenio 278 (RB). Paraiba: Moreno, Pickel 3848 (RB); Areia, Coelho de Moraes 850 (P). Pernambuco: Garanhuns, 850–900 m, 26 November 1924, Chase 7786 (F, MO, US), 7789 (F, MO, NY, US); Pesqueira, Pickel 1663 (US); Fazenda Recreio, Mun. de Rio Grande, Faria 2495 (RB). Rio Grande do Norte: Nova Cruz de Montanhas, 3 June 1934, Swallen 4813 (RB, US)

Species Excluded

The following list does not include valid names, or synonyms treated under those in the main treatment. Taxa recently excluded from Panicum are considered for the Flora Neotropica area, currently treated as valid in other genera of tribes Paniceae and Paspaleae. Panicum aciculare Desv. ex Poir.  =  Dichanthelium aciculare (Desv. ex Poir.) Gould & C.A. Clark P. acicularifolium Renvoize & Zuloaga = Renvoizea acicularifolia (Renvoize & Zuloaga) Zuloaga & Morrone P. acuminatum Sw. = Dichanthelium acuminatum (Sw.) Gould & C.A. Clark P. adenorhachis Zuloaga & Morrone = Dichanthelium adenorhachis (Zuloaga & Morrone) Zuloaga P. aequivaginatum Swallen = Dichanthelium aequivaginatum (Swallen) Zuloaga P. ammophilum Trin. ex Nees, nom. nud.  =  P. dichotomiflorum Michx. var. dichotomiflorum P. anceps Michx. = Coleataenia anceps (Michx.) Soreng P. andreanum Mez = Ocellochloa andreana (Mez) Zuloaga & Morrone P. animarum Renvoize = Apochloa animara (Renvoize) Zuloaga & Morrone P. antidotale Retz. = Janochloa antidotale (Retz.) Zuloaga & Delfini P. arctum Swallen = Trichanthecium arctum (Swallen) Zuloaga & Morrone P. arenicoloides Ashe = Dichanthelium arenicoloides (Ashe) LeBlond P. aristellum Döll = Canastra aristella (Döll) Zuloaga & Morrone P. arundinariae Trin. ex E. Fourn. = Morronea arundinariae (Trin. ex E. Fourn.) Zuloaga & Scataglini P. assurgens Renvoize = Dichanthelium assurgens (Renvoize) Zuloaga P. auricomum Nees ex Trin.  =  Trichanthecium auricomum (Nees ex Trin.) Zuloaga & Morrone P. auritum J. Presl ex Nees = Hymenachne aurita (J. Presl ex Nees) Balansa P. bahiense Renvoize = Apochloa bahiensis (Renvoize) Zuloaga & Morrone P. beyeri Hitchc. & Ekman = Coleataenia beyeri (Hitchc. & Ekman) J.R. Grande © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0_10

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276

Species Excluded

P. biglandulare Scribn. & J.G.  Sm.  =  Ocellochloa biglandularis (Scribn. & J.G. Sm.) Zuloaga & Morrone P. brachyanthum Steud. = Kellochloa brachyantha (Steud.) Lizarazu, M.V. Nicola & Scataglini P. brachystachyum Trin.  =  Ocellochloa brachystachya (Trin.) Zuloaga & Morrone P. bresolinii L.B. Sm. & Wassh. = Hymenachne bresolinii (L.B. Sm. & Wassh.) Zuloaga P. bulbosum Kunth = Zuloagaea bulbosa (Kunth) E. Bess P. caaguazuense Henrard = Trichanthecium caaguazuense (Henrard) Zuloaga & Morrone P. cabrerae Zuloaga & Morrone = Dichanthelium cabrerae (Zuloaga & Morrone) Zuloaga P. caparoense Zuloaga & Morrone  =  Dichanthelium caparoense (Zuloaga & Morrone) Zuloaga P. caricoides Nees ex Trin. = Coleataenia caricoides (Nees ex Trin.) Soreng P. cayoense Swallen = Morronea cayoensis (Swallen) Zuloaga & Scataglini P. chapadense Swallen = Ocellochloa chapadensis (Swallen) Zuloaga & Morrone P. chnoodes Trin. = Apochloa chnoodes (Trin.) Zuloaga & Morrone P. cipoense Renvoize & Send. = Apochloa cipoensis (Renvoize & Send.) Zuloaga & Morrone P. condensatum Bertol. = Hymenachne condensata (Bertol.) Chase P. congestum Renvoize = Dichanthelium congestum (Renvoize) Zuloaga P. cordovense E. Fourn. = Parodiophyllochloa cordovensis (E. Fourn.) Zuloaga & Morrone P. crateriferum Sonhs = Ocellochloa craterifera (Sohns) Zuloaga & Morrone P. cucaense Zuloaga & Morrone = Dichanthelium cucaense (Zuloaga & Morrone) Zuloaga P. cumbucana Renvoize = Dichanthelium cumbucanum (Renvoize) Zuloaga P. cupreum Hitchc. & Chase = Steinchisma cuprea (Hitchc. & Chase) W.V. Br. P. cyanescens Nees ex Trin.  =  Trichanthecium cyanescens (Nees ex Trin.) Zuloaga & Morrone P. davidsei Zuloaga & Morrone = Dichanthelium davidsei (Zuloaga & Morrone) Zuloaga P. decipiens Nees ex Trin. = Steinchisma decipiens (Nees ex Trin.) W.V. Br. P. deustum Thunb. = Batochloa deusta (Thunb.). Salariato & Zuloaga P. dichotomum L. = Dichanthelium dichotomum (L.) Gould P. dichotomum var. tenue (Muhl.) Zuloaga & Morrone = Dichanthelium dichotomum (L.) Gould var. tenue (Muhl.) Gould & C.A. Clark P. discrepans Döll = Cyphonanthus discrepans (Döll) Zuloaga & Morrone P. distichophyllum Spreng. = Trichanthecium distichophyllum (Spreng.) Zuloaga & Morrone P. durifolium Renvoize & Zuloaga = Renvoizea durifolia (Renvoize & Zuloaga) Zuloaga & Morrone P. elephantipes Nees ex Trin. = Louisiella elephantipes (Nees ex Trin.) Zuloaga

Species Excluded

277

P. elliottii Trin. ex Nees, nom. nud.  =  P. dichotomiflorum Michx. var. dichotomiflorum P. eligulatum N.E. Br. = Apochloa eligulata (N.E. Br.) Zuloaga & Morrone P. ensifolium Baldwin ex Elliott  =  Dichanthelium ensifolium (Baldwin ex Elliott) Gould P. euprepes Renvoize = Apochloa euprepes (Renvoize) Zuloaga & Morrone P. exiguiflorum Griseb. = Steinchisma exiguiflora (Griseb.) = W.V. Br. P. fonticolum Swallen = Trichanthecium fonticola (Swallen) Zuloaga & Morrone P. glaziovii Hack. = Renvoizea glaziovii (Hack.) Zuloaga & Morrone P. grande Hitchc. & Chase = Hymenachne grandis (Hitchc. & Chase) Zuloaga P. granuliferum Kunth  =  Trichanthecium granuliferum (Kunth) Zuloaga & Morrone P. grumosum Nees = Hymenachne grumosa (Nees) Zuloaga P. guatemalense Swallen  =  Morronea guatemalensis (Swallen) Zuloaga & Scataglini P. gymnocarpon Elliott = Phanopyrum gymnocarpon (Elliott) Nash P. harleyi Salariato, Morrone & Zuloaga  =  Hymenachne harleyi (Salariato, Morrone & Zuloaga) Zuloaga P. hebotes Trin. = Dichanthelium hebotes (Trin.) Zuloaga P. heliophilum Chase ex Zuloaga & Morrone  =  Dichanthelium heliophilum (Chase ex Zuloaga & Morrone) Zuloaga P. hemitomon Schult. = Hymenachne hemitomon (Schult.) C.C. Hsu P. hians Elliott = Steinchisma hians (Elliott) Nash P. hylaeicum Mez = Rugoloa hylaeica (Mez) Zuloaga P. ichunense Swallen = Trichanthecium ichunense (Swallen) Zuloaga & Morrone P. incumbens Swallen = Morronea incumbens (Swallen) Zuloaga & Scataglini P. irregulare Swallen = Ocellochloa irregularis (Swallen) Zuloaga & Morrone P. itatiaiae Swallen = Dichanthelium itatiaiae (Swallen) Zuloaga P. jaboncillo Hieron., nom. nud. = P. chloroleucum Griseb. P. jauanum Davidse = Apochloa jauana (Davidse) Zuloaga & Morrone P. lagostachyum Renvoize & Zuloaga  =  Renvoizea lagostachya (Renvoize & Zuloaga) Zuloaga & Morrone P. latissimum J.C. Mikan ex Trin. = Ocellochloa latissima (J.C. Mikan ex Trin.) Zuloaga & Morrone P. laxiflorum Lam. = Dichanthelium laxiflorum (Lam.) Gould P. laxum Sw. = Steinchisma laxum (Sw.) Zuloaga P. leptachne Döll = Hymenachne leptachne (Döll) Zuloaga P. longifolium Scribn. = Coleataenia longifolia (Scribn.) Soreng P. longiligulatum Nash = Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. longiligulatum (Nash) Gould & C.A. Clark P. longipedicellatum Swallen = Cnidochloa longipedicellata (Swallen) Zuloaga P. longum Hitchc. & Chase = Hymenachne longa (Hitchc. & Chase) Zuloaga P. loreum Trin. = Apochloa lorea (Trin.) Zuloaga & Morrone P. lutzii Swallen = Apochloa lutzii (Swallen) Zuloaga & Morrone

278

Species Excluded

P. machrisiana Swallen = Trichanthecium machrisianum (Swallen) Zuloaga & Morrone P. marauense Renvoize & Zuloaga = Renvoizea marauense (Renvoize & Zuloaga) Zuloaga & Morrone P. maximum Jacq. = Megathyrsus maximus (Jacq.) B.K. Simon & S.W.L. Jacobs P. mertensii Roth = Stephostachys mertensii (Roth) Zuloaga & Morrone P. micranthum Kunth = Trichanthecium micranthum (Kunth) Zuloaga & Morrone P. missionum Ekman  =  Parodiophyllochloa missiona (Ekman) Zuloaga & Morrone P. molinioides Trin. = Apochloa molinioides (Trin.) Zuloaga & Morrone P. nervosum Lam. = Trichanthecium nervosum (Lam.) Zuloaga & Morrone P. nitidum Lam. var. villosum A. Gray = Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. villosum (A. Gray) Gould & C.A. Clark P. noterophilum Renvoize = Trichanthecium noterophilum (Renvoize) Zuloaga & Morrone P. nutabundum Zuloaga & Morrone = Trichanthecium nutabundum (Zuloaga & Morrone) Zuloaga & Morrone P. obtusum Kunth = Hopia obtusa (Kunth) Zuloaga & Morrone P. orinocanum Luces = Trichanthecium orinocanum (Luces) Zuloaga & Morrone P. ovuliferum Trin. = Parodiophyllochloa ovulifera (Trin.) Zuloaga & Morrone P. pandum Swallen = Trichanthecium pandum (Swallen) Zuloaga & Morrone P. pantrichum Hack.  =  Parodiophyllochloa pantricha (Hack.) Zuloaga & Morrone P. parvifolium Lam. = Trichanthecium parvifolium (Lam.) Zuloaga & Morrone P. parviglume Hack. = Morronea parviglumis (Hack.) Zuloaga & Scataglini P. pedicellatum Vasey = Dichanthelium pedicellatum (Vasey) Gould P. penicillatum Nees ex Trin. = Parodiophyllochloa penicillata (Nees ex Trin.) Zuloaga & Morrone P. pernambucense (Spreng.) Mez ex Pilg.  =  Hymenachne pernambucensis (Spreng.) Zuloaga P. perystipum Zuloaga & Morrone  =  Dichanthelium perystipum (Zuloaga & Morrone) Zuloaga P. petersonii Hitchc. & Ekman = Coleataenia petersonii (Hitchc. & Ekman) Soreng P. petilum Swallen = Trichanthecium petilum (Swallen) Zuloaga & Morrone P. petrense Swallen = Trichanthecium petrense (Swallen) Zuloaga & Morrone P. petropolitanum Zuloaga & Morrone = Dichanthelium petropolitanum (Zuloaga & Morrone) Zuloaga P. piauiense Swallen = Ocellochloa piauiensis (Swallen) Zuloaga & Morrone P. pilosum Sw. = Rugoloa pilosa (Sw.) Zuloaga P. poliophyllum Renvoize & Zuloaga  =  Apochloa poliophylla (Renvoize & Zuloaga) Zuloaga & Morrone P. polycomum Trin. = Trichanthecium polycomum (Trin.) Zuloaga & Morrone P. polygonatum Schrad. = Rugoloa polygonata (Schrad.) Zuloaga P. portoricense Desv. ex Ham. = Dichanthelium portoricense (Desv. ex Ham.) B.F. Hansen & Wunderlin

Species Excluded

279

P. prionitis Nees = Coleataenia prionitis (Nees) Soreng P. pseudisachne Mez = Trichanthecium pseudisachne (Mez) Zuloaga & Morrone P. pulchellum Raddi = Ocellochloa pulchella (Raddi) Zuloaga & Morrone P. pycnoclados Tutin = Dichanthelium pycnoclados (Tutin) Davidse P. pyrularium Hitchc. & Chase = Trichanthecium pyrularium (Hitchc. & Chase) Zuloaga & Morrone P. restingae Renvoize & Zuloaga = Renvoizea restingae (Renvoize & Zuloaga) Zuloaga & Morrone P. rhizogonum Hack.  =  Parodiophyllochloa rhizogona (Hack.) Zuloaga & Morrone P. rigidulum Bosc ex Nees  =  Coleataenia longifolia (Scribn.) Soreng subsp. rigidula (Bosc ex Nees) Soreng P. rivale Swallen = Trichanthecium rivale (Swallen) Zuloaga & Morrone P. rude Nees = Ocellochloa rudis (Nees) Zuloaga & Morrone P. sabulorum Lam. = Dichanthelium sabulorum (Lam.) Gould & C.A. Clark P. sabulorum Lam. var. cordatum (Zuloaga & Morrone) Zuloaga = Dichanthelium sabulorum (Lam.) Gould & C.A.  Clark var. cordatum (Zuloaga & Morrone) Zuloaga. P. sabulorum Lam. var. polycladum (Ekman) Palacios = Dichanthelium sabulorum (Lam.) Gould & C.A. Clark var. polycladum (Ekman) Zuloaga P. sacciolepoides Renvoize & Zuloaga = Renvoizea sacciolepoides (Renvoize & Zuloaga) Zuloaga & Morrone P. scabridum Döll = Coleataenia scabrida (Döll) Zuloaga P. schwackeanum Mez  =  Trichanthecium schwackeanum (Mez) Zuloaga & Morrone P. sciurotis Trin. = Dichanthelium sciurotis (Trin.) Davidse P. sciurotoides Zuloaga & Morrone = Dichanthelium sciurotoides (Zuloaga & Morrone) Davidse P. scoparium Lam. = Dichanthelium scoparium (Lam.) Gould P. sendulskyae Zuloaga & Morrone  =  Dichanthelium sendulskyae (Zuloaga & Morrone) Zuloaga P. sipapoense Swallen = Apochloa sipapoensis (Swallen) Zuloaga & Morrone P. soderstromii Zuloaga & Send. = Ocellochloa soderstromii (Zuloaga & Send.) Zuloaga & Morrone ex Filg. & Sch. Rodr. P. spathellosum Döll = Steinchisma spathellosa (Döll) Renvoize P. sphaerocarpon Elliott = Dichanthelium sphaerocarpon (Elliott) Gould P. sphaerocarpon Elliott var. floridanum Vasey = Dichanthelium sphaerocarpon (Elliott) Gould var. floridanum (Vasey) Davidse P. stagnatile Hitch. & Chase  =  Hymenachne stagnatilis (Hitchc. & Chase) Zuloaga P. stenodes Griseb. = Coleataenia stenodes (Griseb.) Soreng P. stenophyllum Hack. = Steinchisma stenophyllum (Hack.) Zuloaga & Morrone P. stevensianum Hitchc. & Chase = Steinchisma stevensiana (Hitchc. & Chase) Zuloaga P. steyermarkii Swallen = Apochloa steyermarkii (Swallen) Zuloaga & Morrone

280

Species Excluded

P. stigmosum Trin. = Dichanthelium stigmosum (Trin.) Zuloaga P. stipiflorum Renvoize = Dichanthelium stipiflorum (Renvoize) Zuloaga P. stoloniferum Poir. = Ocellochloa stolonifera (Poir.) Zuloaga & Morrone P. strigosum Muhl. ex Elliott  =  Dichanthelium strigosum (Muhl. ex Elliott) Freckmann P. subtiramulosum Renvoize & Zuloaga = Apochloa subtiramulosa (Renvoize) Zuloaga & Morrone P. superatum Hack. = Dichanthelium superatum (Hack.) Zuloaga P. surrectum Chase ex Zuloaga & Morrone = Dichanthelium surrectum (Chase ex Zuloaga & Morrone) Zuloaga P. telmatum Swallen = Dichanthelium telmatum (Swallen) Zuloaga P. tenerum Beyr. ex Trin. = Coleataenia tenera (Beyr. ex Trin.) Soreng P. tepuianum Davidse & Zuloaga  =  Trichanthecium tepuianum (Davidse & Zuloaga) Zuloaga & Morrone P. teretifolium Hack. = Renvoizea teretifolia (Hack.) Zuloaga & Morrone P. tijucae Renvoize = Apochloa tijucae (Renvoize) Zuloaga & Morrone P. trichidiachne Döll = Morronea trichidiachnis (Döll) Zuloaga & Scataglini P. trinii Kunth = Renvoizea trinii (Kunth) Zuloaga & Morrone P. tuerckheimii Hack. = Aakia tuerckheimii (Hack.) J.R. Grande P. umbonulatum Swallen = Dichanthelium umbonulatum (Swallen) Davidse P. vaginiviscosum Renvoize & Zuloaga = Renvoizea vaginiviscosa (Renvoize & Zuloaga) Zuloaga & Morrone P. validum Mez = Osvaldoa valida (Mez) J.R. Grande P. verrucosum Muhl. = Kellochloa verrucosa (Muhl.) Lizarazu, M.V. Nicola & Scataglini P. viscidellum Scribn. = Dichanthelium viscidellum (Scribn.) Gould P. wettsteinii Hack. = Trichanthecium wettsteinii (Hack.) Zuloaga & Morrone P. proliferum Lam. var. Hackel ex Bertoni, nom. nud. = P. dichotomiflorum Michx. var. dichotomiflorum P. yavitaense Swallen = Trichanthecium yavitaense (Swallen) Zuloaga & Morrone

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© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0

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Scientific Index

C Chasea C. amara, 65 LT C. capillaris, 123 LT C. virgata, 101 LT E Eatonia, 37 Eriolytrum E. junceum, 90 LT L Leptoloma L. barbipulvinata, 123 L. capillaris, 123 LT L. miliacea, 175 LT M Milium M. barbipulvinatum, 123 M. capillare, 123 LT M. esculentum, 175 LT M. microspermum, 250 M. paniceum, 175 LT M. virgatum, 31*, 101 LT Monachne M. racemosa, 37, 88 LT M. urvilleana, 97 LT

P Panicum P. acutifolium, 259 P. alatum, 12, 15, 20, 31*, 106, 108 P. alatum var. alatum, 12, 13*, 14*, 15, 34, 108, 109*, 110* P. alatum var. minus, 12, 15, 26, 108, 111, 126* P. altum, 17, 19, 33, 62, 64* P. amarulum, 66 P. amarum Elliott var. amarum, 25, 64*, 65 P. amarum subsp. amarulum, 64*, 65, 66 P. amarum var. amarulum, 66 P. ammophilum, 275 P. amplectens, 46 P. aquarum, 9–12, 15, 20, 34, 104, 113, 114*, 115, 115* P. aquaticum, 17, 19, 20, 25, 31*, 33, 40, 42*, 48 P. aquaticum Poir. var. cartagoense, 41, 44 P. aquaticum var. aquaticum, 18*, 41, 41 P. aquaticum var. chloroticum, 46 P. axipilium, 233 P. aztecanum, 11, 12, 15, 20, 34, 105, 110*, 116, 117* P. bambusoides, 93 LT P. barbipulvinatum, 123 P. bartlettii, 10, 15, 19*, 20, 30*, 31, 39, 224, 225, 225*, 252

© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0

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290 Panicum (cont.) P. bartowense, 52 LT P. bergii, 10, 12, 15, 17, 19, 19*, 20, 31*, 82, 105, 118, 120, 149, 183 P. bergii fo. convoluta, 120 P. bergii fo. quadriglume, 189 P. bergii var. bergii, 26, 35, 115*, 118, 118, 119* P. bergii var. leiophyllum, 183 P. bergii var. pilosissimum, 35, 118, 122, 139*, 220 P. bergii var. quadriglume, 187 P. berteronianum, 270 LT P. beyrichii, 243 P. bobarti, 123 P. brachiatum, 46 P. brevifolium, 10, 19*, 27, 30*, 31, 39, 123, 225*, 227, 233, 259 P. buchingeri, 101 P. burkartii, 120 P. caespititium, 134 P. campestre, 14*, 15, 19, 27, 35, 183, 185, 204, 205, 205, 206*, 207*, 210 P. capillaceum, 257 P. capillaceum var. strictius, 259 P. capillare, 10, 12, 17, 20, 26, 34, 105, 110, 120, 123, 124*, 126* LT P. capillare subsp. barbipulvinatum, 26, 123 P. capillare var. agreste, 123 P. capillare var. barbipulvinatum, 123 P. capillare var. brevifolium, 123 P. capillare var. glabrum, 110 P. capillare var. hirticaule, 161 P. capillare var. miliaceum, 161 P. capillare var. stramineum, 190 P. capillarioides, 12, 15, 26, 105, 126*, 127, 128*, 129 P. cayennense, 9, 10, 15, 18*, 19, 20, 27, 35, 183, 204, 205, 207*, 208, 209*, 210 P. cayennense var. campestre, 183 P. cayennense var. curtatum, 210 P. cayennense var. divaricatum, 218 P. cayennense var. patulum, 208 P. cayennense var. quadriglume, 187 P. cervicatum, 12, 13*, 14*, 15, 18*, 20, 27, 31*, 35, 204, 205, 213, 213*, 215* P. chacoense, 154 LT P. chaseae, 10, 12, 35, 104, 130, 130, 131*, 132*, 145, 185 P. chloroleucum, 11, 15, 20, 24*, 31*, 33, 61, 64*, 67, 68*, 277 LT P. chloroticum, 25, 46, 47, 48

Scientific Index P. chloroticum var. agreste, 46 P. chloroticum var. luxurians, 46 P. chloroticum var. pingue, 46 P. chloroticum var. sylvestre, 41 LT P. coloratum, 19*, 20, 27, 31*, 196, 197, 200* P. complanatum, 15, 34, 61, 70, 70, 71*, 72* P. costaricense, 227 P. dasytrichum, 220 P. debile, 134 P. deciduum, 9, 11, 31*, 34, 61, 70, 72*, 73* P. decolorans, 10, 12, 15, 20, 26, 105, 115*, 133, 180, 181 P. dichotomiflorum, 9, 17, 20, 25, 31*, 33, 39–41, 43*, 44, 47*, 48 P. dichotomiflorum subsp. bartowense, 44, 52 LT P. dichotomiflorum subsp. dichotomiflorum, 44, 44 P. dichotomiflorum subsp. purinatorum, 47 P. dichotomiflorum var. bartowense, 52 LT P. dichotomiflorum var. geniculatum, 46 P. dichotomiflorum var. imperiorum, 46 P. dichotomiflorum var. purinatorum, 47 P. diffusum, 12, 13*, 15, 34, 106, 132, 132*, 134, 135 LT P. diffusum var. debile, 134 P. diffusum var. subcontractum, 135, 139 P. dispersum, 237 P. eccentricos, 189 P. elatum, 156 P. elliottii, 277 P. ephemeroides, 12, 15, 19, 20, 35, 107, 137, 138*, 139 P. ephemerum, 137 P. exiguum, 11, 12, 15, 19, 20, 35, 104, 132*, 135, 139, 140*, 141, 161, 185, 189 P. filamentosum, 257 P. filipes, 135, 149 P. flabellatum, 166, 168 P. francavillanum, 46 LT P. furvum, 9, 11, 12, 13*, 15, 34, 104, 132*, 143, 144 P. geniculatum, 46 P. ghiesbreghtii, 12, 15, 34, 115*, 130, 144, 161, 185 LT P. glandulopaniculatum, 30*, 227, 233 P. gouinii, 17, 19, 20, 23, 27, 35, 197, 197, 198, 200* P. gouinii var. pumilum, 197, 199 LT

Scientific Index P. granuliferum var. longifolium, 250, 252, 277 LT P. grupo Capillaria, 103 P. grupo Dichotomiflora, 39 P. grupo Diffusa, 103 P. grupo Urvilleana, 60 P. grupo Virgata, 60 P. guayaquilense, 250 P. haenkeanum, 10, 20, 23, 27, 31, 39, 227, 228*, 229, 230* P. hallii, 15, 105, 106, 148, 149 P. hallii subsp. filipes, 9, 10, 26, 34, 149, 150*, 195 P. hallii subsp. hallii, 9, 10, 12, 13*, 26, 34, 149, 150*, 151 P. hallii var. filipes, 149 P. halophilum, 199 P. hirsutum, 9, 12, 15, 19, 20, 26, 34, 105, 154, 155*, 156*, 235 P. hirticaule, 20, 26, 31*, 107, 108, 141, 145, 160, 161, 168, 192 P. hirticaule subsp. sonorum, 161 P. hirticaule subsp. stramineum, 190 P. hirticaule var. glabrescens, 161 LT P. hirticaule var. hirticaule, 12, 34, 160, 161, 162* P. hirticaule var. majus, 111 P. hirticaule var. miliaceum, 161 P. hirticaule var. minus, 111 P. hirticaule var. stramineum, 190 P. hirticaule var. verrucosum, 12, 13*, 14*, 15, 34, 110*, 160, 165 P. hirticaulon var. majus, 111 P. hirtivaginum, 144 P. hirtum, 10, 11, 15, 20, 30*, 31, 35, 39, 227, 230*, 231, 232*, 233 P. hispidifolium, 12, 14*, 15, 19*, 20, 26, 34, 107, 139*, 166, 167*, 168 P. hispidum, 166 P. hygrophilum, 41 P. ichnanthoides, 101 LT P. ineptum, 270 NT P. junceum, 93, 95 P. junceum var. strictius, 93, 95 P. junceum var. subnutans, 93, 95 P. lacustre, 33, 40, 53, 54*, 55* LT P. lasianthum, 237 P. lepidulum, 12, 15, 20, 34, 106, 116, 139*, 169, 170*, 171 P. ligulare, 12, 13*, 15, 31*, 35, 204, 205, 215*, 216, 217* LT P. litigosum, 227 P. littorale, 203 P. lundellii, 62

291 P. magnispicula, 10, 12, 15, 20, 35, 105, 139*, 173, 174* P. megastachyum, 97 P. microspermum, 250 P. miliaceum, 5, 12, 15, 19, 20, 26, 29, 31*, 37, 44, 103, 106, 175, 176*, 177* LT P. milium, 175 LT P. millegrana, 10, 15, 19*, 20, 30*, 31, 35, 39, 235, 236*, 238, 238*, 245, 260 P. mucronulatum, 12, 15, 19, 20, 35, 107, 156*, 178, 179*, 180 LT P. multiflorum, 46 LT P. multinodosum, 237 P. mystasipum, 10, 15, 34, 61, 79, 79, 80*, 81, 81* P. olyroides, 15, 31*, 34, 61, 70, 79, 81, 81*, 82, 82 P. olyroides var. denudatum, 213 P. olyroides var. hirsutum, 82, 87 P. olyroides var. olyroides, 24*, 25, 82, 82, 83* LT P. parcum, 11, 12, 15, 20, 26, 31*, 34, 106, 134, 150*, 171, 180, 181 P. patagonicum, 97 P. patentissimum, 235 P. pedersenii, 17, 35, 197, 200*, 201, 202* P. pedunculare, 210 P. peladoense, 11, 12, 15, 18, 19, 20, 35, 104, 130, 141, 144, 156*, 183, 184*, 185, 189 P. phragmitoides, 31*, 60 P. pilcomayense, 26, 120, 149 P. pilgeri, 31*, 93 P. pilosissimum, 220 P. pilosum, 239 P. polygamun var. hirticaule, 34, 160, 161, 162* P. preslii, 97 P. probandum, 243 P. proliferum var. chloroticum, 46 P. proliferum var. geniculatum, 46 P. proliferum var. pilosum, 53 LT P. proliferum var. richardi, 46 P. proliferum var. strictum, 41 P. puberulum, 243 P. quadriglume, 11, 12, 13*, 15, 20, 35, 37, 104, 139*, 145, 187, 188*, 189 P. racemosum, 10, 11, 15, 18*, 19, 20, 24*, 25, 31*, 33, 61, 69, 88, 89*, 91, 91* LT P. repens, 11, 17, 20, 23, 27, 31*, 35, 196, 197, 198*, 199, 200*, 203, 203 LT P. repens var. confertum, 199

292 Panicum (cont.) P. reptans, 37, 90 LT P. retrofractum, 46 LT P. rhigiophyllum, 220 P. rigiophyllum, 208 P. rotundum, 205 P. rudgei, 9, 10, 27, 31*, 35, 204, 205, 215*, 218, 219* P. rudgei var. brasiliense, 220 P. rugulosum, 235 P. rugulosum fo. effusum, 237, 238 P. rugulosum var. condensatum, 245 P. rugulosum var. glabrescens, 243 P. rugulosum var. hirtiglume, 243 P. rugulosum var. subvelutinum, 237 P. scoparium, 218, 279 P. sect. Capillare, 103 P. sect. Coloratae, 196 P. sect. Dichotomiflora, 25, 39 P. sect. Durae, 60 P. sect. Hiantes, 17, 25, 38, 60 P. sect. Miliaceae, 103 P. sect. Panicum, 15, 26, 38, 103, 115 P. sect. Repentia, 17, 27, 35, 38, 196 P. sect. Rudgeana, 9, 12, 15, 27, 35, 38, 185, 204 P. sect. Urvilleana, 60 P. sect. Virgata, 60 P. sellowii, 10, 15, 19*, 21, 27, 30*, 31, 35, 39, 237, 238, 243, 244*, 245, 246*, 260 P. sellowii var. longevaginatum, 239 P. sempervirens, 93 LT P. sessilicaule, 208 P. sonorum, 161 P. stramineum, 12, 15, 18*, 19, 20, 26, 31*, 34, 107, 132*, 137, 161, 190, 191*, 192 P. subgen. Eupanicum, 37 P. subglobosum, 237 P. tamaulipense, 9, 10, 34, 106, 132*, 195 P. trichanthum, 10, 15, 21, 27, 30*, 31, 35, 39, 225, 246*, 250, 251*, 259

Scientific Index P. trichanthum var. modestum, 250 P. trichoides, 10, 15, 21, 24*, 27, 30, 31, 35, 38, 257, 258*, 259, 260* LT P. tricholaenoides, 9, 17, 18*, 20, 25, 31*, 34, 61, 76, 91*, 92, 93, 95 P. tricholaenoides var. flavomarginatum, 93, 97 LT P. tricholaenoides var. tricholaenoides, 93, 93, 94* P. turgidum, 31*, 60 P. urvilleanum, 11, 14*, 15, 20, 24*, 25, 31*, 34, 60, 61, 69, 91, 97, 98*, 100* LT P. urvilleanum fo. subpilosum, 97 LT P. urvilleanum var. chloroleucum, 67 LT P. urvilleanum var. longiglume, 97 LT P. valenzuelanum, 243 P. venezuelae, 10, 15, 21, 31, 35, 39, 270, 271*, 272, 273, 273* P. venosum, 31*, 107, 177*, 196 P. vigoratum, 62 P. vinaceum, 214 P. virgatum, 23, 25, 31*, 33, 60, 62, 100*, 101 LT P. virgatum subsp. cubense, 101 LT P. virgatum var. cubense, 101 LT P. virletii, 151 LT S Saccharum reptans, 90 LT Sporobolus microspermus, 250 T Thalasium T. montevidense, 37, 90 U Urochloa venosa, 196

Subject Index

B Blue grass, 47 C Capim do resfriado, 141 Capim elefante, 218 Capim-lanudo, 192 Chiri-chiri, 161 Cocullito, 259 F Fall panicum, 47 Flying straw, 120 G Gamelote, 156 Guardasireno, 245 J Jaboncillo, 69 Jiwuasinaman, tosé, 84 K Kapi'í chu'i, 141 Kapi'í kokito, 245 Kapi'í tî, 185 Kituka watla, 252

M Marsh grass, 259 Millet, 175 P Penabo, 220 Pensrepens, 203 S Sacate Colorado, 151 Smooth witchgrass, 47 Switchgrass, 102 T Ta, 220 Tacuarita, 93 Torpedo grass, 203 Triguillo, 161 Tupe, 90 Y Yakani yĩñagopeha, 84 Z Zacahuastle, 166

© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 F. Zuloaga, S. Aliscioni, Panicum (Poaceae), Flora Neotropica 124, https://doi.org/10.1007/978-3-031-33768-0

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