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Article
Published: 2022-07-26

The nomenclature and taxonomy of Crassulaceae subfam. Kalanchooideae resolved

Ria Olivier Herbarium, Department of Botany, Nelson Mandela University, P.O. Box 77000, Gqeberha, 6031 South Africa
Australian National Herbarium, Centre for Australian National Biodiversity Research (joint venture between Parks Australia and CSIRO), GPO Box 1700, Canberra ACT 2601, Australia
classification Crassulaceae subfam. Cotyledonoideae Crassulaceae subfam. Kalanchooideae circumscription amended subfamily Eudicots

Abstract

The two subfamily names Crassulaceae subfam. Kalanchooideae and Crassulaceae subfam. Cotyledonoideae were simultaneously validly published by Alwin Berger in 1930. In a long-overlooked contribution, René C.J.E. Maire was the first to combine these two taxa and when he did so, in 1977, adopted the name Crassulaceae subfam. Cotyledonoideae for the united subfamily and unambiguously included Crassulaceae subfam. Kalanchooideae in synonymy. However, crassuloid systematists have continued to use the name Crassulaceae subfam. Kalanchooideae at this rank virtually exclusively for the past ca. 30 years. This subfamily, as presently circumscribed, consists of the genera Adromischus, Cotyledon (nom. cons.), Kalanchoe, and Tylecodon, and in phylogenetic studies is also widely referred to as the Kalanchoe clade. We do not follow Maire and rather propose maintenance of the name Crassulaceae subfam. Kalanchooideae, with a proposal to reject the name Crassulaceae subfam. Cotyledonoideae being under consideration. The circumscription of Crassulaceae subfam. Kalanchooideae is here formally amended to reflect its current circumscription as a monophyletic unit.

References

  1. Adanson, M. (1763) Familles des Plantes. Seconde Partie. Etat actuel de la Botanike. Chez Vincent, Imprimeur-Libraire de Mgt le Comte de Provence, Paris, 640 pp. [https://www.biodiversitylibrary.org/page/3611216]

  2. Baker, J.G. (1881 [21 February]) Notes on a collection of flowering plants made by L. Kitching, Esq., in Madagascar in 1879 (Plates VII. & VIII.). Journal of the Linnean Society, Botany 18: 264–281. [https://www.biodiversitylibrary.org/item/8373#page/271/mode/1up]

  3. Baker, J.G. (1893 [27 May]) New or noteworthy plants. Cotyledon barbeyi, Schweinf. ined.* The Gardeners’ Chronicle. [A weekly illustrated journal of Horticulture and Allied Subjects], 3rd series, XIIII(No. 335): 624–625. [https://www.biodiversitylibrary.org/item/83804#page/646/mode/1up]

  4. Barrie, F.R. (2015 [mayo]) 98. Crassulaceae. In: G.M. Davidse, M.S. Sousa Sánchez, S. Knapp & F. Chiang Cabrera (Editores Generales), Flora mesoamericana. Suaruraceae a Zygophyllaceae. Volumen 2, Parte 3. Instituto de Biología, Universidad Nacional Autónoma de México, Missouri Botanical Garden Press, St Louis, and The Natural History Museum (London), London, pp. 11–19; whole work 347 [348] pp.

  5. Battandier, J.A. & Trabut, L.C. (1905 [April 1905; 1902 on title page; 1904 on wrapper]) Flore analytique et synoptique de l’Algérie et de la Tunisie. [“Famille 41. Crassulacées” on pp. 131–133, Mucizonia additionally on p. 441.] Ive Giralt, Alger, 460 pp. [https://bibdigital.rjb.csic.es/viewer/13171/?offset=#page=131&viewer=picture&o=bookmark&n=0&q=]

  6. Berger, A. (1930 [Mai]) Crassulaceae. Unterfam. II. Kalanchoideae [sic, Kalanchooideae]. 6. Kalanchoe Adanson. 7. Kitchingia Baker. 8. Bryophyllum Salisb. In: A. Engler & K. Prantl (Eds.) Die natürlichen Pflanzenfamilien. edn 2, 18a. Verlag von Wilhelm Engelmann, Leipzig, pp. 378–386, 402–412.

  7. Bruyns, P.V., Hanáček, P. & Klak, C. (2019 [February]) Crassula, insights into an old, arid-adapted group of southern African leaf-succulents. Molecular Phylogenetics and Evolution 131: 35–47. https://doi.org/10.1016/j.ympev.2018.10.045

  8. Candolle, A.P. de. (1801) Histoire naturelle. Botanique. Extrait d’un mémoire sur la famille des Joubarbes. Bulletin des Sciences, par la Société Philomathique (de Paris) 3 (49): 1–2. [https://www.biodiversitylibrary.org/page/5495103#page/9/mode/1up]

  9. Candolle, A.P. de (1828 [mid-March]) Prodromus systematis naturalis regni vegetabilis, sive enumeratio contracta ordinum, generum, specierumque plantarum huc usque cognitarum, juxta methodi naturalis normas digesta. Pars Tertia. Sistens Calyciflorarum Ordines XXVI. Sumptibus Sociorum Treuttel et Wurtz, rue de Bourbon, no 17, Paris, & Strasbourg & London, 494 pp. [https://www.biodiversitylibrary.org/page/152793#page/410/mode/1up]

  10. Carrillo-Reyes, P., Sosa, V. & Mort, M.E. (2009) Molecular phylogeny of the Acre clade (Crassulaceae): dealing with the lack of definitions for Echeveria and Sedum. Molecular Phylogenetics and Evolution 53: 267–276.  https://doi.org/10.1016/j.ympev.2009.05.022

  11. Eggli, U. (2003) Crassulaceae. In: U. Eggli (Ed.) Illustrated handbook of succulent plants: Crassulaceae. Springer Verlag, Berlin, pp. 5–8; whole work 458 pp. and XLVIII plates.  https://doi.org/10.1007/978-3-642-55874-0_2

  12. Hamet, R. (1907) Monographie du genre Kalanchoe. Bulletin de l’Herbier Boissier, sér. 2, 7: 869–900. [https://www.biodiversitylibrary.org/item/105267#page/939/mode/1up]

  13. Hamet, R. (1908) Monographie du genre Kalanchoe (Suite et fin.). Bulletin de l’Herbier Boissier, sér. 2, 8: 17–48. [https://www.biodiversitylibrary.org/item/104945#page/31/mode/1up]

  14. Haworth, A.H. (1812) Synopsis plantarum succulentarum, cum descriptionibus, synonymis, locis; observationibus anglicanis, culturaque. Typis Richardi Taylor et Socii, Shoe-Lane, Londoni [London], 334 pp., including 14 numbered “APPENDIX” pages.  https://doi.org/10.5962/bhl.title.9462

  15. IPNI. (2022+) [continuously updated] The International Plant Names Index. Available from: https://www.ipni.org/ (accessed April 2022).

  16. Koorders, S.H. (1919 [Mai] [24 October 1918]) Beitrag zur Kenntniss der Flora von Java No. 18. Beschreibung von Kalanchoë Schumacheri vom Idjen-Plateau und Revision der Crassulaceae von Java. (Mit 2 lithographischen Tafeln.). Bulletin du Jardin Botanique de Buitenzorg sér. 3, 1 (3): 169–180, Figures 14 and 15. [https://www.biodiversitylibrary.org/item/27173#page/205/mode/1up (text); https://www.biodiversitylibrary.org/item/27173#page/243/mode/1up (figures)]

  17. Ledebour, C.F. von. (1843) Flora rossica sive enumeratio plantarum in totius imperii rossici provinciis europaeis, asiaticis, et americanis hucusque observatarum. Voluminis II. Pars I. [Umbilicus sect. Chiastophyllum on p. 176]. Sumtibus Librariae E. Schweizerbart, Stuttgartiae [Stuttgart], 937 pp.  https://doi.org/10.5962/bhl.title.6606

  18. Lemaire, [A.]C. (1852) Genre nouveau de la familie des Crassulacées. Adromischus. Jardin Fleuriste; Journal General des Progres et des Interets Horticoles et Botaniques. Ghent. [Miscellanées]. 2: 58–61. [https://www.biodiversitylibrary.org/item/274797#page/294/mode/1up]

  19. Linnaeus, C. (1753) Species plantarum, exhibentes plantas rite cognitas, ad generarelatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Vol. 1. Salvius, Stockholm, 560 pp.  https://doi.org/10.5962/bhl.title.669

  20. Linnaeus, C. [filius]. (1782 [“1781”, published in April 1782]) Supplementum plantarum systematis vegetabilium editionis decimae tertiae, generum plantarum editiones sextae, et specierum plantarum editionis secundae. Editum a Carolo a Linné. Impensis Orphanotrophei, Brunsvigæ [Braunschweig], 467 pp. + one unnumbered page headed “EMENDANDA”. [https://www.biodiversitylibrary.org/page/411019#page/256/mode/1up]

  21. Lu, M., Fradera-Soler, M., Forest, F., Barraclough, T.G. & Grace, O. (2021) Evidence linking life-form to a major shift in diversification rate in Crassula. American Journal of Botany 109: 272–290.  https://doi.org/10.1002/ajb2.1797

  22. Maire, R. (1977) Kalanchoe Adanson (1763) Rosales: Crassulaceae. In: Quézel,P.[A.] (Ed.) Flore de l’Afrique de Nord (Maroc, Algérie, Tunisie, Tripolitaine, Cyrénaïque et Sahara). Vol. XIV. Rhoeadales: Cruciferae fin Resedaceae. Sarraceniales: Droseraceae. Rosales: Crassulaceae. Éditions Paul Lechevalier éditeur S.A.R.L., 19, rue Augereau, 19, Paris, pp. 239–241, 251–252, 263. [https://archive.org/details/flore-afrique-du-nord/Vol%2014%20Flore%20de%20l%27Afrique%20Du%20Nord%20Maroc%20Algerie%20Tunisie%20Tripolitaine%20Cyrenaique%20Et%20Sahara%20R-Maire-1977/page/n119/mode/2up]

  23. Messerschmid, T.F.E., Klein, J.T., Kadereit, G. & Kadereit, J.W. (2020) Linnaeus’s folly—phylogeny, evolution and classification of Sedum (Crassulaceae) and Crassulaceae subfamily Sempervivoideae. Taxon 69 (5): 892–926.  https://doi.org/10.1002/tax.12316

  24. Metzing, D. (2017) The scientific heritage of the gardener, botanist, and succulent researcher Alwin Berger. [In: M. Mariotti & L. Minuto (Eds.) Alwin Berger and others. The signs of German culture in the gardens and in the Riviera landscape before and after the Great War. Proceedings of a conference held on 28 October 2016 organised by Università degli Studi di Genova, Centro Servizi di Ateneo per i Giardini Botanici Hanbury, with the support of the Alwin Berger Archiv, Möschlitz, Germany. Giardini Botanici Hanbury, La Mortola, Ventimiglia, Italy.] Bollettino dei Musei e degli Istituti biologici dell’Università di Genova 79: 17–44.

  25. Miller, A.G. (1996) Family 60. Crassulaceae. 5. Cotyledon L. In: Miller, A.G., Cope, T.A. & Nyberg, J.A. (Eds.) Flora of the Arabian Peninsula and Socotra. Volume 1. Edinburgh University Press in association with the Royal Botanic Garden Edinburgh and the Royal Botanic Gardens, Kew, Edinburgh, p. 477; whole work 586 pp.

  26. Moran, R.V. (2009) Crassulaceae J. Saint-Hilaire. Stonecrop or Orpine Family. In: Flora of North America Editorial Committee (Eds.) Flora of North America north of Mexico. Magnoliophyta: Paeoniaceae to Ericaceae. Volume 8. Oxford University Press, New York and Oxford, pp. 147–150 [whole family to p. 229]; whole work 585 pp.

  27. Mort, M.E., Levsen, N., Randle, C.P., Van Jaarsveld, E.[J.] & Palmer, A. (2005 [July]) Phylogenetics and diversification of Cotyledon (Crassulaceae) inferred from nuclear and chloroplast DNA sequence data. American Journal of Botany 92 (7): 1170–1176.

  28. Mort, M.E., O’Leary, T.R., Carillo-Reyes, P., Nowell, T., Archibald, J.K. & Randle, C.P. (2010) Phylogeny and evolution of Crassulaceae: past, present and future. Schumannia 6 / Biodiversity and Ecology 3: 69–86.

  29. Mort, M.E., Soltis, D.E., Soltis, P.S., Francisco-Ortega, J. & Santos-Guerra, A. (2001) Phylogenetic relationships and evolution of Crassulaceae inferred from matK sequence data. American Journal of Botany 88: 76–91.

  30. Raymond-Hamet. (1933) Observations sur le Kalanchoe uniflora (Stapf) Raymond-Hamet. Bulletin de la Société Botanique de France 80 (4): 542–548.  https://doi.org/10.1080/00378941.1933.10833878

  31. Salisbury, R.A. (1805 [1 June 1805]) The Paradisus Londinensis: or coloured figures of plants cultivated in the vicinity of the metropolis 1. Published by William Hooker, No. 6, Frith Street, city not stated, presumably London, 157 pp. & 117 plates.  https://doi.org/10.5962/bhl.title.53520

  32. Schmalfuß, L. (2017) Alwin Berger: his life, his family and friends. [In: M. Mariotti & L. Minuto (Eds.) Alwin Berger and others. The signs of German culture in the gardens and in the Riviera landscape before and after the Great War. Proceedings of a conference held on 28 October 2016 organised by Università degli Studi di Genova, Centro Servizi di Ateneo per i Giardini Botanici Hanbury, with the support of the Alwin Berger Archiv, Möschlitz, Germany. Giardini Botanici Hanbury, La Mortola, Ventimiglia, Italy.] Bollettino dei Musei e degli Istituti biologici dell’Università di Genova 79: 9–16.

  33. Silva, V., Figueiredo, E. & Smith, G.F. (2015) Alien succulents naturalised and cultivated on the central west coast of Portugal. Bradleya 33: 58–81.  https://doi.org/10.25223/brad.n33.2015.a10

  34. Smith, G.F. (2020a) Typification of the infrageneric names published in Kalanchoe (Crassulaceae subfam. Kalanchooideae) by Alwin Berger. Phytotaxa 453 (1): 52–54.  https://doi.org/10.11646/phytotaxa.453.1.6

  35. Smith, G.F. (2020b) Nomenclature of the partial infrageneric classification proposed for Kalanchoe (Crassulaceae subfam. Kalanchooideae) by René C.J.E. Maire in 1976 [sic; 1977]. Phytotaxa 468 (2): 231–235.  https://doi.org/10.11646/phytotaxa.468.2.8

  36. Smith, G.F. (2020c) The kalanchoes (Crassulaceae subfam. Kalanchooideae) of Raymond-Hamet (1890–1972). Bradleya 38: 120–140. https://doi.org/10.25223/brad.n38.2020.a15

  37. Smith, G.F. (2021) Reinstatement of Kalanchoe [subg. Bryophyllum] sect. Alatae (Crassulaceae subfam. Kalanchooideae) for K. porphyrocalyx and K. uniflora, two Madagascan endemics. Bradleya 39: 177–181.  https://doi.org/10.25223/brad.n39.2021.a17

  38. Smith, G.F. (2022) Notes on Kalanchoe subg. Bryophyllum (Crassulaceae subfam. Kalanchooideae) and the kalanchoes of Sijfert H. Koorders (1919). Phytotaxa 549 (1): 87–96.  https://doi.org/10.11646/phytotaxa.549.1.7

  39. Smith, G.F. & Figueiredo, E. (2018a) The infrageneric classification and nomenclature of Kalanchoe Adans. (Crassulaceae), with special reference to the southern African species. Bradleya 36: 162–172.  https://doi.org/10.25223/brad.n36.2018.a10

  40. Smith, G.F. & Figueiredo, E. (2018b) Kalanchoe crouchii Gideon F.Sm. & Figueiredo (Crassulaceae), a new species from the dolomites of the Wolkberg Centre of Endemism, Mpumalanga Province, South Africa. Haseltonia 25: 84–90.  https://doi.org/10.2985/026.025.0107

  41. Smith, G.F., Figueiredo, E. & Van Wyk, A.E. (2019) Kalanchoe (Crassulaceae) in southern Africa. Classification, biology, and cultivation. Academic Press, an imprint of Elsevier, London, San Diego, Cambridge (U.S.A.), and Oxford, 328 pp.  https://doi.org/10.1016/C2017-0-00602-X

  42. Smith, G.F., Figueiredo, E. & Walker, C.C. (2021c) Kalanchoe beharensis. Crassulaceae subfam. Kalanchooideae. Madagascar. Flowering Plants of Africa 67: 50–59, plate 2365.

  43. Smith, G.F., Klein, D.-P. & Shtein, R. (2021a) A revision of the Malagasy Kalanchoe subg. Kitchingia (Crassulaceae subfam. Kalanchooideae): history, taxonomy, and nomenclature. Phytotaxa 507 (1): 67–80.  https://doi.org/10.11646/phytotaxa.507.1.3

  44. Smith, G.F. & Monro, A.M. (2022) (2908) Proposal to reject the name Crassulaceae subfam. Cotyledonoideae (Crassulaceae). Taxon 71. In press.

  45. Smith, G.F., Shtein, R. & Klein, D.-P. (2021b) Kalanchoe subg. Alatae (Crassulaceae subfam. Kalanchooideae), a new rank for K. sect. Alatae. Phytotaxa 509 (2): 255–258.  https://doi.org/10.11646/phytotaxa.509.2.9

  46. Smith, G.F., Shtein, R., Klein, D.-P., Parihar, B., Almeida, A., Rodewald, S. & Kadereit, G. (2021d) Special Review. Sexual and asexual reproduction in Kalanchoe (Crassulaceae subfam. Kalanchooideae): a review of known and newly recorded strategies. Haseltonia 28: 2–22.  https://doi.org/10.2985/026.028.0101

  47. Smith, G.F., Van Jaarsveld, E.J., Arnold, T.H., Steffens, F.E., Dixon, R.D. & Retief, J.A. (Eds.) (1997) List of southern African succulent plants. Contributed and supported by the National Botanical Institute of South Africa and the Succulent Society of South Africa. Umdaus Press, Pretoria, 175 pp.

  48. Stapf, O. (1930) Index londinensis to illustrations of flowering plants, ferns and fern allies []. Volume II. [Chiastophyllum on pp. 176, 316, and 317.] Clarendon Press, Oxford. 548 pp.

  49. Stekl, R. (2022) Ein wunderschönes Geburtstagsfest für Alwin Berger. / A wonderful birthday party for Alwin Berger. Avonia 40 (1): 74–77.

  50. Stevens, P.F. (2001 onwards) Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Available from: http://www.mobot.org/MOBOT/research/APweb/ (Accessed April 2022)

  51. ’t Hart, H. (1995) Chapter 10. Infrafamilial and generic classification of the Crassulaceae. In: ’t Hart, H. & Eggli, U. (Eds.) Evolution and systematics of the Crassulaceae. Backhuys Publishers, Leiden, pp. 159–172; whole work 192 pp.

  52. Thiede, J. & Eggli, U. (2007) Crassulaceae. In: Kubitzki, K. (Ed.) The families and genera of vascular plants. IX. Flowering Plants. Eudicots. Springer-Verlag, Berlin, pp. 83–118.

  53. Thorne, R.F. (1983) Proposed new realignments in the angiosperms. Nordic Journal of Botany 3 (1): 85–117.

  54. Thorne, R.F. (1992) An updated phylogenetic classification of the flowering plants. Aliso 13 (2): 365–389.

  55. Tölken, H.R. (1978) New taxa and new combinations in Cotyledon and allied genera. Bothalia 12: 377–393. [https://journals.abcjournal.aosis.co.za/index.php/abc/article/view/1794/1762]

  56. Tölken, H.R. (1985) Crassulaceae. 3166, 4. Kalanchoe. 3166a, 5. Bryophyllum. In: Leistner, O.A. (Ed.) Flora of Southern Africa 14. Botanical Research Institute, Department of Agriculture and Water Supply, place of publication not stated, likely Pretoria, pp. 61–74; whole work 244 pp.

  57. 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. (Eds.) (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Koeltz Botanical Books, Glashütten. [Regnum Vegetabile 159], 254 pp.  https://doi.org/10.12705/Code.2018

  58. Uhl, Ch. H. (1948) Cytotaxonomic studies in the subfamilies Crassuloideae, Kalanchoideae, and Cotyledonoideae of the Crassulaceae. American Journal of Botany 35: 695–697.

  59. Uhl, Ch. H. (1956) The Crassulaceae and cytotaxonomy. Cactus and Succulent Journal (U.S.) 28 (1): 28–30. [https://archive.org/details/sim_cactus-and-succulent-journal_jan-feb-1956_28_1/page/28/mode/1up]

  60. Uhl, Ch. H. (1963) Chromosomes and phylogeny of the Crassulaceae. Cactus and Succulent Journal (U.S.) 35 (3): 80–84. [https://archive.org/details/sim_cactus-and-succulent-journal_may-june-1963_35_3/mode/2up]

  61. Van Ham, R.C.H.J. (1995) Chapter 2. Phylogenetic relationships in the Crassulaceae inferred from chloroplast DNA variation. In: ’t Hart, H. & Eggli, U. (Eds.) Evolution and systematics of the Crassulaceae. Backhuys Publishers, Leiden, pp. 17–29; whole work 192 pp.

  62. Van Ham, R.C.H.J. & ’t Hart, H. (1998) Phylogenetic relationships in the Crassulaceae inferred from chloroplast DNA restriction-site variation. American Journal of Botany 85: 123–134.

  63. Wickens, G.E. (1987) Crassulaceae. In: Polhill, R.M. (Ed.) Flora of Tropical East Africa: Crassulaceae. Published on behalf of the East African Governments by A.A. Balkema, Rotterdam, 66 pp.