Abstract
The new genus Kellochloa is proposed, on the basis of morphological and molecular characters, to include two North American species of Panicum s.l., previously classified in sect. Verrucosa: P. brachyanthum and P. verrucosum. Both species are annual, with decumbent culms branching divaricately at the lower nodes, and erect culms with terminal and axillary inflorescences. The spikelets are pilose, with lower glume 1/6 to 1/4 the length of the spikelet, upper glume and lower lemma subequal, 3–5-nerved, the lower anthecium reduced to the lower lemma, and upper anthecium indurate, with bicellular microhairs and simple papillae evenly distributed all over the lemma and palea. Molecular phylogenetic studies, analyzing plastid ndhF and rpoA sequences, confirmed that these morphological characters are differential. The phylogenetic position and taxonomic features of the new genus are established and compared with Panicum s.str. and other taxa of the Sacciolepis–Trichanthecium clade. Full synonymies and descriptions, a distribution map, illustrations of both species, and a key are provided. Two new combinations are proposed. In addition, four African ungrouped species of Panicum s.l. were, for the first time, included in the phylogenetic analysis to analyze their relationship with the new genus and its positions in the Sacciolepis–Trichanthecium clade.
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References
Aliscioni SS, Giussani LM, Zuloaga FO, Kellogg EA (2003) A molecular phylogeny of Panicum (Poaceae: Paniceae). Tests of monophyly and phylogenetic placement within the Panicoideae. Amer J Bot 90:796–821. doi:10.3732/ajb.90.5.796
Barkworth ME (2004) Manual of grasses for North America and North of Mexico. University of Utah, Logan. http://herbarium.usu.edu/webmanual/
Brown WV (1977) The Kranz syndrome and its subtype in grass systematics. Mem Torrey Bot Club 23:1–97
Crins WJ (1991) The genera of Paniceae (Gramineae: Panicoideae) in the Southeastern United States. J Arnold Arb Suppl Ser 1:171–312
D’Ambrogio de Argüeso A (1986) Manual de técnicas en histología vegetal. Ed. Hemisferio Sur, Buenos Aires, p 83
Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nat Meth 9:772. doi:10.1038/nmeth.2109
Drummond AJ, Suchard MA, Xie D, Rambaut A (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molec Biol Evol 29:1969–1973. doi:10.1093/molbev/mss075
Duvall MR, Noll JD, Minn AH (2001) Phylogenetics of Paniceae (Poaceae). Am J Bot 88:1988–1992
Duvall MR, Saar DE, Grayburn WS, Holbrook GP (2003) Complex transitions between C3 and C4 photosynthesis during the evolution of Paniceae: a phylogenetic case study emphasizing the position of Steinchisma hians (Poaceae), a C3–C4 intermediate. Int J Pl Sci 164:949–958
Ellis RP (1976) A procedure for standardizing comparative leaf blade anatomy in the Poaceae. I. The leaf blade as viewed in transverse section. Bothalia 12:65–109
Ellis RP (1977) Distribution of the Kranz syndrome in the southern African Eragrostoideae and Panicoideae according to bundle sheath anatomy and cytology. Agroplantae 9:73–110
Ellis RP (1979) A procedure for standardizing comparative leaf blade anatomy in the Poaceae. II. The epidermis as seen in surface view. Bothalia 12:641–672
Ellis RP (1988) Leaf anatomy and systematics of Panicum (Poaceae: Panicoideae) in southern Africa. Monogr Syst Bot Missouri Bot Gard 25:129–156
Freckmann RW, Lelong MG (2003) Nomenclatural changes and innovations in Panicum and Dichanthelium (Poaceae: Paniceae). Sida 20:161–174
Giussani LM, Cota-Sánchez JH, Zuloaga FO, Kellogg EA (2001) A molecular phylogeny of the grass subfamily Panicoideae (Poaceae) shows multiple origins of C4 photosynthesis. Amer J Bot 88:1993–2012
Goloboff PA, Farris JS, Nixon KC (2008) TNT, a free program for phylogenetics analysis. Cladistics 24:774–786
Gómez-Martínez R, Culham A (2000) Phylogeny of the subfamily Panicoideae with emphasis on the tribe Paniceae: evidence from the trnL-F cpDNA region. In: Jacobs SWL, Everett JE (eds) Grasses: systematics and evolution. Commonwealth Scientific and Industrial Research Organization (CSIRO) Publishing, Collingwood, pp 136–140
Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symp Ser 41:95–98
Hattersley PW (1987) Variation in photosynthetic pathway. In: Soderstrom TR, Hilu KW, Campbell CS, Barkworth ME (eds) Grass systematics and evolution. Smithsonian Institution Press, Washington, DC, pp 49–64
Hattersley PW, Watson L (1976) C4 grasses: an anatomical criterion for distinguishing between NADP-Malic enzyme species and PCK or NAD-Malic enzyme species. Austral J Bot 24:297–308
Hitchcock AS, Chase A (1910) The North American species of Panicum. Contr US Natl Herb 15:1–396
Hsu CC (1965) The classification of Panicum (Gramineae) and its allies, with special reference to the characters of lodicule, style-base and lemma. J Fac Sci Tokyo Univ 3(9):43–150
Johansen DA (1940) Plant microtechnique. Mc Graw-Hill Bokk Company Inc, New York, p 523
Metcalfe CR (1960) Anatomy of the monocotyledons, I Gramineae. Clarendon Press, Oxford
Morrone O, Aagesen L, Scataglini MA, Salariato DL, Denham SS, Chemisquy MA, Zuloaga FO (2012) Phylogeny of the Paniceae (Poaceae: Panicoideae): integrating plastid DNA sequences and morphology into a new classification. Cladistics 28:333–356. doi:10.1111/j.1096-0031.2011.00384.x
Olmstead RG, Sweere JA (1994) Combining data in phylogenetic systematics: an empirical approach using three molecular data sets in the Solanaceae. Syst Biol 43:467–481
Petersen G, Seberg O (1997) Phylogenetic analysis of the Triticeae (Poaceae) based on rpoA sequence data. Molec Phylogen Evol 7:217–230
Rambaut A, Suchard MA, Xie D, Drummond AJ (2014) Tracer v1.6. Available from http://beast.bio.ed.ac.uk/Tracer
Ruzin SE (1999) Plant microtechnique and microscopy. Oxford University Press, New York
Simon BK, Jacobs SWL (2003) Megathyrsus, a new generic name for Panicum subgenus Megathyrsus. Austrobaileya 6:571–574
Thiers B (2014) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/. Accessed 2014
Webster RD (1988) Genera of the North American Paniceae (Poaceae: Panicoideae). Syst Bot 13:576–609
Zuloaga FO (1987) Systematics of New World Species of Panicum (Poaceae: Paniceae). In: Soderstrom TR, Hilu KW, Campbell CS, Barkworth ME (eds) Grass systematics and evolution. Smithsonian Institution Press, Washington, DC, pp 287–306
Zuloaga FO, Morrone O, Vega AS, Giussani LM (1998) Revisión y análisis cladístico de Steinchisma (Poaceae: Panicoideae: Paniceae). Ann Missouri Bot Gard 85:631–656
Zuloaga FO, Morrone O, Giussani LM (2000) A cladistic analysis of the Paniceae: a preliminary approach. In: Jacobs SWL, Everett JE (eds) Grasses: systematics and evolution. Commonwealth Scientific and Industrial Research Organization (CSIRO) Publishing, Collingwood, pp 123–135
Zuloaga FO, Morrone O, Davidse G, Filgueiras TS, Peterson PM, Soreng RJ, Judziewicz EJ (2003) Catalogue of New World Grasses (Poaceae): III. Subfamilies Panicoideae. Aristidoideae, Arundinoideae, and Danthonioideae. Contr US Natl Herb 46:1–662
Zuloaga FO, Scataglini MA, Morrone O (2011) A monograph of Trichanthecium, a new genus segregated from Panicum (Poaceae: Panicoideae: Paniceae). Syst Bot Monogr 94:1–98
Acknowledgments
This work was supported by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Grant #11220100100207. We would like to express our deep gratitude to Fernando Zuloaga for his help, guidance, and his invaluable feedback on the manuscript. We also acknowledge Lone Aagesen for her valuable and careful suggestions on the manuscript and assistance with English editing. We thank as well Francisco Rojas and Marcelo Moreno for preparing the illustrations, and two anonymous reviewers for helpful comments on the manuscript.
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Appendix
Appendix
List of species used in the phylogenetic analysis including the six new vouchers, in bold, and vouchers from Zuloaga et al. (2011). Voucher information and GenBank accession numbers (ndhF, rpoA) are shown.
Tribe Andropogoneae. Zea mays U21985, X07810. Tribe Paspaleae. Anthaenantiopsis rojasiana Zuloaga 6747 AY029620, JF804815; Echinochloa chacoensis Zuloaga 8573 EF218711, JF804820; E. cruspavonis Morrone 5369 EF218710, JF804819; Oplismenus burmanii Morrone 3625 EF218712, JF804821; Paspalum bertonii Zuloaga 7186 JF804854, JF804817; P. foliiforme Morrone 4669 JF804853, JF804816. Tribe Paniceae. Panicum aequinerve Kellogg 109 (MO) KR232095; P. bartlettii Alvarez 152 JF804882; P. brachyanthum Daniel et al . 25 (MO) KR232096, KR232090; P. brevifolium Gereau 5315 JF804860, JF804828; P. chionachne Wiehe 358 (US) KR232097, KR232091; P. eickii Robinson 5306 (US) KR232098, KR232092; P. heterostachyum Greenway 14150 JF804862, JF804830; P. hirtum Strudwick 4389 JF804861, JF804829; P. inaequilatum Phillips 61 (MO) KR232099, KR232093; P. millegrana Zuloaga 6766 AY029660, JF804852; P. sellowii Zuloaga 2123 AY188484, JF804822; P. trichanthum Morrone 5028 JF804858, JF804827; P. trichoides Zuloaga 7430 JF804859, JF804826; P. verrucosum Wheeler s.n. AY188496; P. verrucosum Kral 48168 (MO) KR232100, KR232094; Sacciolepis angustissima Morrone 4319 JF804856, JF804824; S. indica Bowen 3875 AY029677; S. myuros Longhi-Wagner 9390 JF804857, JF804825; S. vilvoides Zuloaga 6745 JF804854, JF804823; Trichanthecium arctum Liesner 15757 JF804881; T. auricomum Longhi-Wagner 9637 JF804870, JF804841; T. brazzavillense Fay 5891 JF804880, JF804851; T. caaguazuense Anderson 35724 JF804875, JF804846; T. cyanescens Morrone 4772 JF804867, JF804837; T. dinklagei Jenik & Hall 1129 JF804866, JF804834; T. distichophyllum Longhi-Wagner 9664 JF804869, JF804840; T. gracilicaule Renvoize 2229 JF804864, JF804832; T. granuliferum Jansen-Jacobs 4064 JF804871, JF804842; T. machrisianum Viana 1727 JF804874, JF804845; T. margaritiferum Milne-Redhead 4530 JF804863, JF804831; T. micranthum Morrone 4770 JF804877, JF804848; T. natalense Kellogg 117 JF804879, JF804850; T. nervosum Morrone 4789 JF804876, JF804847; T. parvifolium Zuloaga s.n. AY188476, JF804835; T. polycomum Morrone 4763 JF804873, JF804844; T. pseudisachne Longhi-Wagner 9606 JF804868, JF804839; T. pyrularium Swallen 4936 JF804878, JF804849; T. rivale Morrone 4765 JF804872, JF804843; T. schwackeanum Morrone 115 AY188483, JF804838; T. tenellum Richards 9306 JF804865, JF804833; T. wettsteinii Zuloaga 6972 AY188497, JF804836.
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Nicola, M.V., Lizarazu, M.A. & Scataglini, M.A. Phylogenetic analysis and taxonomic position of section Verrucosa of Panicum and its relationship with taxa of the Sacciolepis–Trichanthecium clade (Poaceae: Panicoideae: Paniceae). Plant Syst Evol 301, 2247–2260 (2015). https://doi.org/10.1007/s00606-015-1227-9
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DOI: https://doi.org/10.1007/s00606-015-1227-9