Taxonomic revision of Rinorea ilicifolia (Violaceae) from Africa and Madagascar


The uncertain taxonomic circumscription between Rinorea ilicifolia (Welw. ex Oliv.) Kuntze and R. spinosa (Boivin ex Tul.) Baill. has led to considerable confusion when attempting to identify the two species. The plants from mainland Africa have typically been determined as R. ilicifolia while the plants in Madagascar and the Comoro Islands have been variously assigned to R. spinosa, R. ilicifolia, or R. ilicifolia subsp. spinosa (Boivin ex Tul.) Grey-Wilson. A third taxon from east Africa, R. ilicifolia var. amplexicaulis Grey-Wilson, has auriculate leaf bases and a mostly discrete distribution from R. ilicifolia, suggesting a greater degree of divergence than is indicated by its taxonomic rank of variety. To better understand the circumscription among these three taxa, we performed multivariate and discriminant analyses of six quantitative variables measured from herbarium specimens. Based on results from these analyses, we deemed there was insufficient morphological divergence to maintain R. ilicifolia and R. spinosa as separate species. Instead, we recognise a single species, R. ilicifolia, composed of three subspecies: R. ilicifolia subsp. ilicifolia in mainland Africa, R. ilicifolia subsp. spinosa in Madagascar and the Comoro Islands, and R. ilicifolia subsp. amplexicaulis (Grey-Wilson) Wahlert, comb. et stat. nov. from Tanzania and Burundi. Here we provide a taxonomic revision of R. ilicifolia, a distribution map and preliminary risk of extinction assessment using IUCN Red List categories and criteria for the three subspecies. We also propose a change of status for Rinorea [unranked] Ilicifoliae Engl. to the rank of section, an action supported by phylogenetic and morphological evidence. The new section includes R. ilicifolia and 13 other taxa from mainland Africa. Lectotypes are designated for the section and for Alsodeia spinosa Boivin ex Tul., and a second-step lectotype is selected for Alsodeia ilicifolia Welw. ex Oliv.

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  1. Achoundong, G. & Cheek, M. (2005). Two further new species of Rinorea (Violaceae) from Cameroon. Kew Bull. 60: 581 – 586.

    Google Scholar 

  2. Albuquerque, S., Brummitt, R. K. & Figueiredo, E. (2009). Typification of names based on the Angolan collections of Friedrich Welwitsch. Taxon 2: 641 – 646.

    Article  Google Scholar 

  3. Ballard Jr., H. E., Paula-Souza, J. & Wahlert, G. A. (2014). Violaceae. In: K. Kubitzki (ed.), The Families and Genera of Vascular Plants, pp. 303 – 322. Springer-Verlag, Berlin.

    Google Scholar 

  4. Bachman, S., Moat, J., Hill, A. W., de la Torre, J. & Scott, B. (2011). Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. In: V. Smith & L. Penev (eds), e-Infrastructures for Data Publishing in Biodiversity Science. ZooKeys 150: 117 – 126 (version beta).

  5. Baillon, M. (1886). Violaceae. Bull. Mens. Soc. Linn. Paris 73: 582 – 584.

  6. Bâthie, H. Perrier de la. (1954). Violacées. In: H. Humbert (ed.), Flore de Madagascar et des Comores, pp. 1 – 55. Muséum National d’Histoire Naturelle, Paris.

  7. Carrisso, L. W. (1937). Violaceae. In: A. W. Exell & F. A. Mendonça, Conspectus Florae Angolensis, pp. 68 – 76. Agência Geral do Ultramar, Lisbon.

  8. Engler, A. (1900). Beiträge zur Flora von Afrika. Bot. Jahrb. Syst. 28: 332 – 510.

    Google Scholar 

  9. Engler, A. (1904). Violaceae africanae. Bot. Jahrb. Syst. 33: 132 – 147.

    Google Scholar 

  10. Exell, A. W. (1935). Mr John Gossweiler's Portuguese West African plants. Dicotyledones: Polypetalae — Addendum. J. Bot. 73 (suppl.): 11 – 12.

  11. Grey-Wilson, C. (1981). Notes on African Violaceae. Kew Bull 36: 103 – 126.

    Article  Google Scholar 

  12. Grey-Wilson, C. (1986). Violaceae. Flora of Tropical East Africa. Balkema, Rotterdam.

    Google Scholar 

  13. Henderson, A. (2006). Traditional morphometrics in plant systematics and its role in palm systematics. Bot. J. Linn. Soc. 151: 103 – 111.

    Article  Google Scholar 

  14. IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1. Second edition. Gland and Cambridge.

  15. James, F. C. & McCulloch, C. E. (1990). Multivariate analysis in ecology and systematics: panacea or Pandora’s box? Annual Rev. Ecol. Evol. Syst. 21: 129 – 166.

    Google Scholar 

  16. Kuntze, C. E. O. (1891). Revisio generum plantarum 1: 40 – 42. Arthur Felix, Leipzig.

  17. Madagascar Catalogue (2018). Rinorea. Catalogue of the plants of Madagascar. Missouri Botanical Garden, St. Louis, Missouri and Antananarivo. Available from:

  18. Oliver, D. (1868). Flora of Tropical Africa 1: 1 – 149. L. Reeve & Co., London.

  19. R Development Core Team (2013). R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. Available at:

  20. Rietz, G. E. (1930). The fundamental units of biological taxonomy. Svensk Bot. Tidskr. 24: 332 – 428.

  21. Robson, N. K. B. (1960). Violaceae. In: A. W. Exell & H. Wild (eds), Flora Zambesiaca, pp. 246 – 260. University Press, Glasgow.

    Google Scholar 

  22. Tennant, J. R. (1963). Notes on tropical African Violaceae. Kew Bull. 16: 409 – 435.

    Article  Google Scholar 

  23. Tulasne, L. R. (1868, publ. 1869). Violarieae. Ann. Sci. Nat., Bot. sér. 5, 9: 299 – 321.

  24. Turland, N. J., Wiersema, J. H., Barrie, F. R., Greuter, W., Hawksworth, D. L., Herendeen, P. S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, T. W., McNeill, J., Monro, A. M., Prado, J., Price, M. J. & Smith, G. F. (2018). International Code of Nomenclature for Algae, Fungi, and Plants (Shenzhen Code), Adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Veg. 159. Koeltz Botanical Books, Glashütten. Available at:

  25. van Velzen, R., Wahlert, G. A., Sosef, M. S. M., Onstein, R. E. & Bakker, F. T. (2015). Phylogenetics of Rinorea (Violaceae): elucidating infrageneric relationships using plastid and nuclear DNA sequences. Syst. Bot. 40: 174 – 184.

    Article  Google Scholar 

  26. Venables, W. N. & Ripley, B. D. (2002). Modern Applied Statistics with S. Fourth Edition. Springer, New York.

  27. Wahlert, G. A. (2010). Phylogeny, Biogeography, and a Taxonomic Revision of Rinorea (Violaceae) from Madagascar and the Comoro Islands. Ph.D. thesis, Ohio University, Athens, Ohio.

  28. Wahlert, G. A. & Ballard Jr., H. E. (2012). A phylogeny of Rinorea (Violaceae) inferred from plastid DNA sequences with an emphasis on the African and Malagasy species. Syst. Bot. 37: 964 – 973.

    Article  Google Scholar 

  29. Wahlert, G. A., Marcussen, T., Ballard, Jr., H. E., de Paula-Souza, J. & Feng, M. (2014). A phylogeny of the Violaceae (Malpighiales) inferred from plastid DNA sequences: implications for generic diversity and infrafamilial groupings. Syst. Bot. 39: 239 – 252.

  30. Wuertz, D. (2010). fBasics: Rmetrics – Markets and Basic Statistics. R package version 2110.79. Available at:

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We appreciate the many herbarium curators at K, MO, P, PRE, TAN, TEF, and WAG who made specimens available for this study. Two anonymous reviewers generously provided detailed feedback which helped to improve the accuracy of this paper. We also thank Jacek Wajer (BM) for providing high resolution images of herbarium material. Peter Phillipson (MO, P) provided assistance with mapping and protected areas in Madagascar. Financial support was provided by an Ohio University Graduate Student Senate Grant for Original Research and Student Enhancement Award (GAW) and an Ohio University Baker Grant (HEB).

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Correspondence to Gregory A. Wahlert.

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Appendix. List of specimens used in morphometric analysis

Appendix. List of specimens used in morphometric analysis

Rinorea ilicifolia subsp. ilicifolia: Y. S. Abeid & O. A. Kibure 706 (MO); Y. S. Abeid & O. A. Kibure 735 (MO); J.-G. Adam 3181 (MO, P); J.-G. Adam 20793 (P); J.-G. Adam 23589 (MO); J.-G. Adam 29747 (MO); J.-G. Adam 30109 (MO); M. Aubréville 1801 (P); L. Bernardi 8304 (P); L. Bernardi 8414 (P); J. P. M. Brenan 8491 (P); F. J. Breteler 6105 (MO, P); F. B. Dhego et al. 592 (MO); A. Chevalier 14820 (P); A. Chevalier 16641 (P); A. Chevalier 16741 (P); A. Chevalier 16854 (P); A. Chevalier 16921 (P); A. Chevalier 22601 (P); K. O. Darko 461 (MO, P); J. de Koning et al. 9747 (MO); J. J. F. E. de Wilde 6361 (MO); J. J. F. E. de Wilde & A. G. Voorhoeve 3736 (P); W. J. J. O. de Wilde & B. E. E. Duyfjes-de Wilde 1625 (MO); A. A. Enti 435 (MO); A. A. Enti 644 (MO); A. A. Enti 1683 (MO); P. Ezrom et al. 189 (MO); R. B. Faden et al. 77/415 (MO); L. Festo 817 (MO); I. Friis et al. 3919 (MO); Frontier-Tanzania 1420B (MO); Frontier-Tanzania 2700 (MO); L. Gautier et al. LG2265 (MO); L. Gautier et al. LG2863 (MO); D. J. Harris & J. M. Fay 76 (MO); D. J. Harris & J. M. Fay 228 (MO); D. J. Harris & J. M. Fay 534 (MO); C. J. Kayombo et al. 4678 (MO); O. A. Kibure 84 (MO); O. A. Kibure 491 (MO); O. A. Kibure 514 (MO); J. Koechlin 3914 (P); M. G. Latilo 22 (P); J. Lewalle 4900 (MO); Maclaud 182 (P); M. Merello et al. 1337 (MO); M. A. Mwangoka 1257 (MO); M. A. Mwangoka & S. Ali 2188 (MO); L. B. Mwasumbi & P. Mponda 12643 (MO); L. B. Mwasumbi & P. Munyenyembe 13848 (MO); P. B. Phillipson et al. 4947 (MO); G. E. Pilz 2667 (MO); M. Reekmans 3892 (MO [2 sheets]); S. A. Robertson & W. R. Q. Luke 4544 (MO); P. Schnell 5328 (P); V. Simon et al. 9 (MO); A. R. da Torre & J. Paiva 9888 (MO); L. White 881 (MO).

Rinorea ilicifolia subsp. spinosa: L. H. Boivin 2122bis (P); L. H. Boivin 3296 (P); J. M. Bosser 5899 (P); J. M. Bosser 18012 (P); J. M. Bosser 18111 (P [2 sheets]); G. Cremers 2443 (P); P. Coulon 88 (MO); R. Decary 1356 (BM, P); R. Decary 2361 (P); R. Decary 15808 (P); L. Gautier et al. LG3134 (P); J.-L. Guillaumet 2256 (P); P. Hoffmann et al. 435 (MO); H. Humbert 18845 (P); L. Humblot 151 (P); M. Keraudren-Aymonin & G. G. Aymonin 25629 (P); J.-N. Labat et al 2230 (P); J. D. Leandri 263 (P); J. D. Leandri 666 (P); J. D. Leandri 953 (P); J. D. Leandri 1067 (P); J. D. Leandri & P. Saboureau 2907 (P); C. Mas 294 (P); H. Perrier 355 (P); H. Perrier 1409 (P); H. Perrier 12117 (P); H. Perrier 12118 (P); M. Pignal et al. 1926 (MO); C. H. O. Pobéguin 68 (P); R. Randrianaivo et al. 228 (P); J. M. C. Richard 185 (P); J. M. C. Richard 210 (P); J. M. C. Richard 328 (P); J. M. C. Richard 555 (P); RN 2008 (P); RN 2548 (P); RN 2714 (P); RN 3011 (P); RN 4917 (P); RN 5664 (P); RN 8153 (P); RN 10331 (P); RN 11418 (P); G. Rouhan & F. Bernier 38 (MO); SF 3169 (P); SF 14197 (P); SF 15513 (P); SF 16343 (P); SF 16571 (P); SF 18034 (P); SF 26211 (P); SF 26398 (P); L. Tarnaud 150 (P); H. Tinguy 804 (P); S. Wohlhauser & H. Andriamalaza 60267 (P).

Rinorea ilicifolia subsp. amplexicaulis: Y. S. Abeid et al. 1274 (MO); Y. S. Abeid et al. 2282 (MO); G. Gobbo 344 (MO); W. R. Q. Luke et al. 8185 (MO); B. E. Mhoro 33 (MO); D. W. Thomas 3691 (MO).

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Wahlert, G.A., Gilland, K.E. & Ballard, H.E. Taxonomic revision of Rinorea ilicifolia (Violaceae) from Africa and Madagascar. Kew Bull 75, 12 (2020).

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Key Words

  • Comoro Islands
  • conservation
  • morphometrics
  • Rinorea spinosa
  • taxonomy