Published 2025-11-17
Keywords
- classification,
- Cottea,
- Enneapogon,
- Kaokochloa,
- molecular phylogenetics
- Schmidtia ...More
How to Cite
Copyright (c) 2025 Paul M. Peterson, Konstantin Romaschenko, Yolanda herrera Arrieta

This work is licensed under a Creative Commons Attribution 4.0 International License.
Abstract
To investigate the evolutionary relationships among 23 species in the Cotteinae we generated a DNA sequence-derived phylogeny utilizing three plastid (rps16-trnK spacer, rps16 intron, rpl32-trnL spacer) regions and the nuclear ribosomal internal transcribed spacer (ITS) region. The Bayesian tree provides strong support for the monophyly of the Cotteinae and its four genera, in order of divergence: Cottea, the Kaokochloa + Schmidtia clade, and the Enneapogon clade. Since the Eragrostideae clearly evolved from African ancestors and Kaokochloa, Schmidtia, and Enneapogon all share species from Africa, we hypothesize the ancestral area of Cotteinae to be Africa. Within Enneapogon we recovered four major clades, the first two splits with species primarily from Africa and the third split containing a basal species from Africa/Asia/Europe sister to all species from Australia. The ancestral area of Enneapogon is Africa and based on our current sample the 13 species from Australia appear to be derived from a single dispersal event from an ancestor shared with E. persicus.
References
- Aedo C. 2021. 121. Enneapogon Desv. ex P.Beauv. In: Romero Zarco C, Rico E, Crespo MB, Devesa JA, Buira A, Aedo C (Eds.), Flora Iberica Plantas vasculares de la Península Ibérica e Islas Baleares Vol. XIX(II) Gramineae (partim). Real Jardín Botánico, CSIC, Madrid, 1310‒1312.
- Bor NL. 1960. The grasses of Burma, Ceylon, India, and Pakistan (excluding Bambuseae). Pergamon Press, Oxford, 1‒767.
- Bor NL. 1970 Gramineae. In: Rechinger KH (Ed.), Flora Iranica 70/30. Akademische Druck und Verlagsanstalt, Graz, Austria, 1–573.
- Burbidge NT. 1941. A revision of the Australian species of Enneapogon Desv. Proceedings of the Linnean Society of London. 153(1): 52‒91. DOI: https://doi.org/10.1111/j.1095-8312.1941.tb01379.x
- Chen SL, Phillips SM. 2006. 21. Tribe Pappophoreae. In: Wu ZY, Raven PH, Hong DY (Eds.), Flora of China: Poaceae, volume 22. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis, 456−457.
- Clayton WD. 1970. Gramineae (Part 1). In: Milne-Redhead E, Polhill RM (Eds.), Flora of Tropical East Africa. Whitefriars Press, London, 1‒176.
- Clayton WD, Renvoize SA. 1986. Genera graminum: Grasses of the World. Kew Bulletin, Additional Series. 13: 1–389.
- Clayton WD, Vorontsova MS, Harman KT, Williamson H. 2016. GrassBase — The online World grass flora: The Board of Trustees, Royal Botanic Gardens. http://www.kew.org/data/grasses-db.html (accessed 29 July 2025).
- Cope, TA. 2007. Poaceae (Gramineae) In: Knees SG, Miller AG (Eds.), Flora of the Arabian Peninsula and Socotra, vol. 5 part 1. Edinburgh University Press and Royal Botanic Gardens, Kew. Edinburgh, 1‒387.
- Dávila P, Mejia-Saulés MT, Soriano-Martínez AM, Herrera-Arrieta Y. 2018. Conocimiento taxonómico de la Poaceae en México. Botanical Sciences. 96(3): 462–514. https://doi.org/10.17129/botsci.1894 DOI: https://doi.org/10.17129/botsci.1894
- Edgar R. 2004. MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research. 32: 1792–1797. DOI: https://doi.org/10.1093/nar/gkh340
- Felsenstein J. 1985. Confidence limits on phylogenies: An approach using the bootstrap. Evolution. 39: 783–791. DOI: https://doi.org/10.1111/j.1558-5646.1985.tb00420.x
- Fish L, Mashau AC, Moeaha MJ, Nembudani MT. 2015. Identification guide to Southern African grasses, an identification manual with keys, descriptions and distributions. Strelitzia. 36(ix): 1–798.
- Gallaher TJ, Peterson PM, Soreng RJ, Zuloaga FO, Li DZ, Clark LG, Tyrrell CD, Welker CAD, Kellogg EA, Teisher JK. 2022. Grasses through space and time: An overview of the biogeographical and macroevolutionary history of the Poaceae. Journal of Systematics and Evolution. 60(3): 522‒569. https://doi.org/10.1111/jse.12857 DOI: https://doi.org/10.1111/jse.12857
- Grass Phylogeny Working Group (GPWG) III Arthan W +72 additional authors, 2024. A nuclear phylogenomic tree of grasses (Poaceae) recovers current classification despite gene tree incongruence. New Phytologist. 245(2) 818‒834. https://doi.org/10.1111/nph.20263 DOI: https://doi.org/10.1111/nph.20263
- Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS. 2018. UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution. 35(2): 518‒522. https://doi.org/10.1093/molbev/msx281 DOI: https://doi.org/10.1093/molbev/msx281
- Huelsenbeck JP, Ronquist FR. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics (Oxford, England). 17(8): 754–755. DOI: https://doi.org/10.1093/bioinformatics/17.8.754
- Kunth KS. 1929. Révision des Graminées, Vol. 1. Dessins de Madame Eulalia Delile, Paris.
- Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T, Ashton B, Meintjes P, Drummond A. 2012. Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics. 28: 1647–1649. https://pmc.ncbi.nlm.nih.gov/articles/PMC3371832/ DOI: https://doi.org/10.1093/bioinformatics/bts199
- Minh BQ, Nguyen MAT, von Haeseler A. 2013. Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution. 305: 1188‒1195. https://doi.org/10.1093/molbev/mst024 DOI: https://doi.org/10.1093/molbev/mst024
- Palisot de Beauvois AMFJ. 1812. Essai d’une nouvelle agrostographie; ou nouveaux genres des Graminées; avec figures représentant les caractères de tous les genres. Fain, Paris, France, 1‒82. DOI: https://doi.org/10.5962/bhl.title.474
- Peterson PM, Romaschenko K, Herrera Arrieta Y. 2015a). A molecular phylogeny and classification of the Eleusininae with a new genus, Micrachne (Poaceae: Chloridoideae: Cynodonteae). Taxon. 64(3): 445–467. https://doi.org/10.12705/643.5 DOI: https://doi.org/10.12705/643.5
- Peterson PM, Romaschenko K, Herrera Arrieta Y. 2015b. Phylogeny and subgeneric classification of Bouteloua with a new species, B. herrera-arrietae (Poaceae: Chloridoideae: Cynodonteae: Boutelouinae). Journal of Systematics and Evolution. 53(4): 351–366. https://doi.org/10.1111/jse.12159 DOI: https://doi.org/10.1111/jse.12159
- Peterson PM, Romaschenko K, Herrera Arrieta Y. 2016. A molecular phylogeny and classification of the Cynodonteae (Poaceae: Chloridoideae) with four new genera: Orthacanthus, Triplasiella, Tripogonella, and Zaqiqah; three new subtribes: Dactylocteniinae, Orininae, and Zaqiqahinae; and a subgeneric classification of Distichlis. Taxon. 65(6): 1263–1287. https://doi.org/10.12705/656.4 DOI: https://doi.org/10.12705/656.4
- Peterson PM, Romaschenko K, Johnson G. 2010a. A classification of the Chloridoideae (Poaceae) based on multi-gene phylogenetic trees. Molecular Phylogenetics and Evolution. 55: 580–598. https://doi.org/10.1016/j.ympev.2010.01.018 DOI: https://doi.org/10.1016/j.ympev.2010.01.018
- Peterson PM, Romaschenko K, Johnson G. 2010b. A phylogeny and classification of the Muhlenbergiinae (Poaceae: Chloridoideae: Cynodonteae) based on plastid and nuclear DNA sequences. American Journal of Botany. 97: 1532–1554. https://doi.org/10.3732/ajb.0900359 DOI: https://doi.org/10.3732/ajb.0900359
- Peterson PM, Romaschenko K, Soreng RJ. 2014. A laboratory guide for generating DNA barcodes in grasses: A case study of Leptochloa s.l. (Poaceae: Chloridoideae). Webbia. 69(1): 1−12. https://doi.org/10.1080/00837792.2014.927555 DOI: https://doi.org/10.1080/00837792.2014.927555
- Peterson PM, Soreng RJ, Davidse G, Filgueiras TS, Zuloaga FO, Judziewicz EJ. 2001. Catalogue of New World grasses (Poaceae): II. subfamily Chloridoideae. Contributions from the United States National Herbarium 41: 1−255. https://www.biodiversitylibrary.org/item/13801#page/1/mode/1up
- Phillips SM. 1995. Poaceae (Gramineae). In: Hedberg I, Edwards S (Eds.), Flora of Ethiopia and Eritrea. The National Herbarium, Addis Abada University, Addis Abada, Ethiopia and Department of Systematic Botany, Uppsala Sweden, 1–420.
- Plants of the World Online (POWO). 2025. Facilitated by the Royal Botanic Gardens, Kew http://www.plantsoftheworldonline.org (accessed 29 July 2025).
- Reeder JR. 1965. The tribe Orcuttieae and the subtribes of the Pappophoreae (Gramineae). Madroño. 18(1): 18‒28. https://www.jstor.org/stable/41423175
- Reeder JR. 2003. 18. Pappophoreae Kunth In: Barkworth ME, Capels KM, Long S, Piep MB (Eds.), Flora of North America North of Mexico, Vol. 25 Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press, New York, 285‒289.
- Rambaut A, Drummond A, Xie D, Baele G, Suchard M. 2018. Posterior summarization in Bayesian phylogenetics using tracer 1.7. Systematic Biology. 67(5): 901‒904. https://doi.org/10.1093/sysbio/syy032 DOI: https://doi.org/10.1093/sysbio/syy032
- Renvoize SA. 1968. The Afro-Asian species of Enneapogon P. Beauv. (Gramineae). Kew Bulletin. 22: 393–401. DOI: https://doi.org/10.2307/4108332
- Reutemann AG, Tivano JC, Lucero LE, Vegetti AC, Giussani LM. 2011. Phylogenetic relationships within Pappophoreae s.l. (Poaceae: Chloridoideae): Additional evidence based on ITS and trnL-F sequence data. South African Journal of Botany. 77: 693‒702. https://doi.org/10.1016/j.sajb.2011.03.010 DOI: https://doi.org/10.1016/j.sajb.2011.03.010
- Romaschenko K, Peterson PM, Soreng RJ, Garcia-Jacas N, Futorna O, Susanna A. 2012. Systematics and evolution of the needle grasses (Poaceae: Pooideae: Stipeae) based on analysis of multiple chloroplast loci, ITS, and lemma micromorphology. Taxon. 61(1): 18–44. https://doi.org/10.1002/tax.611002 DOI: https://doi.org/10.1002/tax.611002
- Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP. 2012. MrBayes 3.2: Efficient bayesian phylogenetic inference and model choice across a large model space. Systematic Biology. 61: 539–542. https://academic.oup.com/sysbio/article/61/3/539/1674894 DOI: https://doi.org/10.1093/sysbio/sys029
- Roodt-Wilding R, Spies JJ. 2006. Phylogenetic relationships in southern African chloridoid grasses (Poaceae) based on nuclear and chloroplast sequence data. Systematic and Biodiversity. 4(4): 401–415. https://doi.org/10.1017/S1477200006002131 DOI: https://doi.org/10.1017/S1477200006002131
- Sanchez-Ken JG. 2018. Riqueza de especies, clasificación y listado de las gramíneas (Poaceae) de México. Acta Botánica Mexicana. 126(126): 1–115. https://doi.org/10.21829/abm126.2019.1379 DOI: https://doi.org/10.21829/abm126.2019.1379
- Soreng RJ, Peterson PM, Romaschenko K, Davidse G, Zuloaga FO, Judziewicz EJ, Filgueiras TS, Davis JI, Morrone O. 2015. A worldwide phylogenetic classification of the Poaceae (Gramineae). Journal of Systematics and Evolution. 53(2): 117–137. https://onlinelibrary.wiley.com/doi/10.1111/jse.12150/epdf DOI: https://doi.org/10.1111/jse.12150
- Soreng RJ, Peterson PM, Romaschenko K, Davidse G, Teisher JK, Clark LG, Barberá B, Gillespie LJ, Zuloaga FO. 2017. A worldwide phylogenetic classification of the Poaceae (Gramineae) II: An update and a comparison of two 2015 classifications. Journal of Systematics and Evolution. 55(4): 259–290. https://doi.org/10.1111/jse.12262 DOI: https://doi.org/10.1111/jse.12262
- Soreng RJ, Peterson PM, Zuloaga FO, Romaschenko K, Clark LG, Teisher JK, Gillespie LJ, Barberá P, Welker CAD, Kellogg EA, De-Zhu L, Davidse G. 2022. A worldwide phylogenetic classification of the Poaceae (Gramineae) III: An update. Journal of Systematics and Evolution. 60(3): 476–521. https://doi.org/10.1111/jse.12847 DOI: https://doi.org/10.1111/jse.12847
- Stebbins GL, Crampton B. 1961. A suggested revision of the grass genera of temperate North America. Recent Advances in Botany: 133‒145. University of Toronto Press.
- Trinius CB. 1830.Graminum Genera quadam speciesque complures definitionibus novis ilustravit. Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1(1): 54‒93.
- Weiller CM, Lazarides M. 2005. Enneapogon. In: Mallett K (Ed.), Flora of Australia Volume 44B Poaceae 3. CSIRO Publishing, Melbourne, 1‒486.
- Zwickl DJ. 2006. Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. University of Texas, Austin.
