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Phylogeny of sections Clavelligerae and Pectinatae of Panicum (Poaceae, Panicoideae, Paniceae): establishment of the new subtribe Dichantheliinae and the genus Adenochloa

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Abstract

Taxonomic features of African species of Panicum sections Clavelligerae and Pectinatae are reviewed and compared with Dichanthelium and other taxa in tribe Paniceae. The new subtribe Dichantheliinae is proposed on the basis of molecular and morphological characters: it includes non-Kranz species, growing at forests edges or in mountain grasslands with membranous-ciliate ligules, lax and open inflorescences, ellipsoid to oblongoid spikelets, and an indurate upper anthecium. Within the Dichantheliinae, the new genus Adenochloa is also established on the basis of chloroplast ndhF sequences and morphological characters, i.e., plants with clavellate multicellular, and glandular hairs present on blades, main axis, branches, and pedicels of inflorescences; ellipsoid to oblongoid, glabrous spikelets, with a lower glume 1/3 to more than 1/2 the length of the spikelet; a 7–13-nerved upper glume and a 5–9-nerved lower lemma; the upper anthecium is indurate, pilose or glabrous. Adenochloa includes 14 species from Africa and Madagascar. The new combinations: Adenochloa adenophylla, A. adenophora, A. bullockii, A. claytonii, A. ecklonii, A. flacciflora, A. habrothrix, A. hymeniochila, A. lukwangulense, A. nigromarginata, A. pectinella, A. pole-evansii, A. sadinii, and A. squarrosa are proposed and the new genus is compared with other genera of the tribe Paniceae. Also, lectotypes are designated for Brachiaria sadinii, Panicum adenophorum, P. adenophyllum, P. ecklonii, P. katentaniense, P. kisantuense, P. hymeniochilum, P. hymeniochilum var. glandulosum, P. snowdenii, and P. scandens; Panicum glanduliferum and P. omega are treated as new synonyms of Adenochloa hymeniochila and A. pectinella, respectively. Finally, new illustrations are provided for 6 species of the new genus.

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Acknowledgments

Funding of this research was provided by CONICET, grant 11220100100207. We like to express our deep gratitude to Francisco Rojas and Marcelo Moreno for the illustrations included in this contribution. We especially appreciate the help of the Editor-in-Chief Karol Marhold, Rob Soreng, and two additional anonymous reviewers who provided useful suggestions to improve an early version of this paper.

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  1. Instituto de Botánica Darwinion, Labardén 200, Casilla de Correo 22, San Isidro, Buenos Aires, B1642HYD, Argentina

    Fernando O. Zuloaga, L. Salomón & Maria A. Scataglini

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  1. Fernando O. Zuloaga
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  2. L. Salomón
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  3. Maria A. Scataglini
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Correspondence to Maria A. Scataglini.

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Handling editor: Karol Marhold.

Appendix: List of taxa used in the molecular analysis and GenBank accession numbers. For new sequences (*) voucher information is given

Appendix: List of taxa used in the molecular analysis and GenBank accession numbers. For new sequences (*) voucher information is given

Tribe Andropogoneae. Andropogon gerardii Vitman, AF117391; Bothriochloa bladhii (Retz.) S.T.Blake, AF117395; Chionachne koenigii (Spreng.) Thwaites, AF117397; Cymbopogon flexuosus (Nees ex Steud.) Will. Watson, AF117404; Phacelurus digitatus (Sibth. & Sm.) Griseb., AF117418. Sorghum bicolor (L.) Moench, U21981; Zea mays L., U21985. Tribe Arundinelleae. Arundinella hirta (Thunb.) Tanaka, AF117393. Tribe Chasmanthieae. Chasmanthium laxum (L.) H.O.Yates, U27296. Tribe Paniceae. Acroceras zizanioides (Kunth) Dandy, AY029618; Cenchrus ciliaris L., AY029625; C. compressus (R. Br.) Morrone, AY029672; C. setaceus (Forssk.) Morrone, AY029673; Chaetium bromoides (J. Presl) Benth. ex Hemsl., AY029626; Dichanthelium acuminatum (Sw.) Gould & C.A.Clark, AY188485; *D. boscii (Poir.) Gould & C.A.Clark, United States, Missouri, Cherokee Pass Quadrangle, Brant 5442 (MO) KP173703; D. clandestinum (L.) Gould, AY188461; D. cumbucana (Renvoize) Zuloaga, AY188464; *D. hebotes (Trin.) Zuloaga, Brazil, Ceará, Guaramiranga, Huber s.n. (IAN); D. koolauense (H.St.John & Hosaka) Gould & C.A.Clark, AY029627; *D. linaerifolium (Scribn.) Gould, United States, Missouri, Howell County, Summers 8082 (MO) KP173704; *D. malacophyllum (Nash) Gould, United States, Missouri, Vernon County, Camp Clark, Becker 1042 (MO) KP173705; *D. oligosanthes (Schult.) Gould, United States, Missouri, Vernon County, Camp Clark, Becker 900 (MO) KP173706; D. sabulorum (Lam.) Gould & C.A.Clark, AY029654; *D. scoparium (Lam.) Gould, United States, Missouri, Stoddard County, Gust 729 (MO) KP173707; *D. stigmosum (Trin.) Zuloaga, Brazil, Santa Catarina, Uribici, Silva and Silva 26 (SI) KP173708; *D. superatum (Hack.) Zuloaga, Brazil, Santa Catarina, Dalmoli et al. 62 (SI) KP173709; Brazil, Santa Catarina, Uribici, Dalmoli et al. 75 (SI) KP173710; *D. viscidellum (Scribn.) Gould, Colombia, Norte de Santander, vicinity of Toledo, Killip and Smith 20046 (US) KP173711; Digitaria setigera Roth ex Roem. & Schult., AY029629; Echinochloa colona (L.) Link, AY029631; Eriochloa punctata (L.) Desv., AY029634; Lasiacis sorghoidea (Desv.) Hitchc. & Chase, AY029639; Megathyrsus maximus (Jacq.) B. K. Simon and S. W. L. Jacobs, AY029649; Melinis repens (Willd.) Zizka, AY029675; Moorochloa eruciformis (Sm.) Veldkamp, AY188452; Oplismenus hirtellus (L.) P.Beauv., AY029644; Panicum. Section Dichotomiflora (Hitchc.) Honda: Panicum aquaticum Poir., AY029658; P. dichotomiflorum Michx., AY188466; P. elephantipes Nees ex Trin., AY029647; P. gouinii E.Fourn., AY188467; P. pedersenii Zuloaga, AY029646; P. repens L., AY029651. Section Panicum: P. bergii Arechav., AY188457; P. fauriei Hitchc., AY029650; P. miliaceum L., AY188472; P. nephelophilum Gaudich., AY029645; P. stramineum Hitchc. & Chase, AY188489. Section Rudgeana (Hitchc.) Zuloaga: P. cervicatum Chase, AY188459; P. rudgei Roem. & Schult., AY029661. Section Urvilleana (Hitchc.) Pilg: P. chloroleucum Griseb., AY188460; P. racemosum (P.Beauv.) Spreng., AY188481. Section Virgata Hitchc. & Chase ex Pilg.: P. tricholaenoides Steud., AY188493; P. virgatum L., U21986. Ungrouped. P. mystasipus Zuloaga & Morrone, AY188474; P. olyroides Kunth, AY188475. Panicum “incertae sedis”: Section Clavelligerae Stapf: P. adenophorum K.Schum., AY188454; P. claytonii Renvoize, AY188462; *P. bullockii, Zambia, Sunzu Mountain, 20 miles S of Abercorn, Phipps and Vesey-Fitzgerald 3314 (K) KP173693; *P. ecklonii, Tanzania, Iringa, Makete, Gereau 3123 (MO) KP173694; *P. flacciflorum, Tanzania, Mpanda, Uzondo Plateau, Bidgood et al. 5501 (K) KP173695; *P. habrothix, Burundi, Bujumbura, Muhira, Reekmans 8890 (K) KP173696; *P. hymeniochilum, Tanzania, Rukwa,, Sumbawanga, Harder et al. 1257 (MO) KP173697; *P. nigromarginatum, Tanzania, Bukoba, Minziro Forest Reserve, Bayona and Festo 9244 (K) KP173698; *P. pectinellum, Zambia, Mporokoso, 28 miles east of Mporokoso, Fitzgerald 3124 (MO) KP173699; *P. peteri, Mozambique, Sofala, southern tip of Chimanimani Mts., Muller and Kelly 1103 (K) KP173700; *P. pole-evansii, Zambia, Luapula, Lake Bangweulu, Renvoize 5513 (MO) KP173701; *P. sadinii; Zambia, Mwinilunga, river Lunga at Mwinilunga, Milne-Redhead 3339 (K) KP173702; Section Monticola Stapf: P. millegrana Poir., AY029660; P. sellowii Nees, AY188484; P. trichanthum Nees, AY188492. Section Verrucosa Hitchc. & Chase ex C. C. Hsu: P. verrucosum Muhl., AY188496. Ungrouped: P. antidotale Retz., AY188456; Parodiophyllochloa cordovensis (E.Fourn.) Zuloaga & Morrone, AY188463; P. missiona (Ekman) Zuloaga & Morrone, AY188473; P. ovulifera (Trin.) Zuloaga & Morrone, AY029653; P. penicillata (Nees ex Trin.) Zuloaga & Morrone, AY18847; Pseudechinolaena polystachya (Kunth) Stapf, AY029676; Sacciolepis indica (L.) Chase, AY029677; Setaria geminata (Forssk.) Veldkamp, AY029662; S. lachnea (Nees) Kunth, AY029683; S. macrostachya Kunth, AY029678; S. palmifolia (J.König) Stapf, AY029680; S. viridis (L.) Beauv., U21976; Stenotaphrum secundatum (Walter) Kuntze, AY029684; Trichanthecium cyanescens (Lam.) Zuloaga & Morrone, AY188465. T. parvifolium (Lam.) Zuloaga & Morrone, AY188476; T. schwackeanum (Mez) Zuloaga & Morrone, AY188483; T. wettsteinii (Hack.) Zuloaga & Morrone, AY188497; Urochloa acuminata (Renvoize) Morrone & Zuloaga AY029629; U. plantaginea (Link) R.D.Webster, AY029693; Zuloagaea bulbosa (Kunth) Bess, AY029648. Tribe Paspaleae. Aakia tuerckheimii (Hack.) J.R.Grande, AY188494; Altoparadisium chapadense Filg. et al., AY029619; Anthaenantia lanata (Kunth) Benth., AY029640; Anthaenantiopsis rojasiana Parodi, AY029620; Apochloa euprepes (Renvoize) Zuloaga & Morrone, AY029657; A. subtiramulosa (Renvoize & Zuloaga) Zuloaga & Morrone, AY188490; Arthropogon villosus Nees, AY029622; Axonopus anceps (Mez) Hitchc., AY029623; Axonopus hydrolithicus (Filg. et al.) A. López & Morrone, AY029642; Canastra lanceolata (Filg.) Morrone et al., AY029621; Coleataenia anceps (Michx.) Soreng, AY188455. C. caricoides (Nees ex Trin.) Soreng, GU253330; C. longifolia (Torr.) Soreng, AY188482. C. petersonii (Hitchc. & Ekman) Soreng, AY188479. C. prionitis (Nees) Soreng, AY029652; C. stenodes (Griseb.) Soreng, GU253333; C. tenera (Beyr. ex Trin.) Soreng, AY188491; Echinolaena inflexa (Poir.) Chase, AY029633; Homolepis glutinosa (Sw.) Zuloaga & Soderstr., AY029637; Hopia obtusa (Kunth) Zuloaga & Morrone, AY029659; Hymenachne donacifolia (Raddi) Chase, AY029635; H. grumosa (Nees) Zuloaga, AY188468; H. pernambucensis (Spreng.) Zuloaga, AY188478; Ichnanthus pallens (Sw.) Munro ex Benth., AY029638; Mesosetum chaseae Luces, AY029641; Ocellochloa chapadensis (Swallen) Zuloaga & Morrone, AY188486; O. piauiensis (Swallen) Zuloaga & Morrone, AY029656; O. stolonifera (Poir.) Zuloaga & Morrone, AY18848; Oncorachis ramosa (Zuloaga & Soderstr.) Morrone & Zuloaga, AY029686; Oplismenopsis najada (Hack. & Arechav.) Parodi, AY188453; Otachyrium versicolor (Döll) Henrard, AY029643; Osvaldoa valida (Mez) J.R.Grande, AY188495; Paspalum arundinellum Mez, AY029663; P. conjugatum Bergius, AY029669; P. glaziovii (A.G.Burm.) S.Denham, AY029689; P. remotum J.Remy, AY029668; P. vaginatum Sw., AY029665; Phanopyrum gymnocarpon (Elliott) Nash, AY188469; Plagiantha tenella Renvoize, AY029674; Rugoloa hylaeica (Mez) Zuloaga, AY188470; Rugoloa pilosa (Sw.) Zuloaga, AY188480; Steinchisma decipiens (Nees ex Trin.) W. V. Br., AY188499; S. hians (Elliot) Nash, AY029685; S. laxa (Sw.) Zuloaga, AY029655; S. spathellosa (Döll) Renvoize, AY188500; Stephostachys mertensii (Roth) Zuloaga & Morrone, AY188471; Streptostachys asperifolia Desv., AY029687; Tatianyx arnacites (Trin.) Zuloaga & Soderstr., AY029688. Tribe Thysanolaenae. Thysanolaena maxima (Roxb.) Kuntze, U21984. Tribe Tristachyideae. Danthoniopsis dinteri (Pilg.) C.E.Hubb., AY029695. Tribe Zeugiteae. Zeugites pittieri Hack., U21987.

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Zuloaga, F.O., Salomón, L. & Scataglini, M.A. Phylogeny of sections Clavelligerae and Pectinatae of Panicum (Poaceae, Panicoideae, Paniceae): establishment of the new subtribe Dichantheliinae and the genus Adenochloa . Plant Syst Evol 301, 1693–1711 (2015). https://doi.org/10.1007/s00606-014-1186-6

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