Abstract
The nomenclature and taxonomy of the Malagasy Kalanchoe macrochlamys (Crassulaceae [subfam. Cotyledonoideae] tr. Kalanchoeae), one of the kalanchoes described by the French botanist Henri Perrier de la Bâthie in 1928, are discussed and clarified. The species has remained little-known for nearly 100 years, likely because it had not been illustrated until 2006, and then as K. maromokotrensis. The typification of the name K. macrochlamys is corrected and it is shown that K. maromokotrensis belongs in the synonymy of K. macrochlamys. A map and discussion of the natural geographical distribution range of K. macrochlamys are provided, along with an amplified description of the species, and illustrations of it from several localities as well as its multiple reproductive strategies. Kalanchoe macrochlamys is also compared to K. cymbifolia.
References
- Adanson, M. (1763 [July–August]) Familles des plantes. Contenant une Préface Historike sur l’état ancien & actuel de la Botanike, & une Téorie de cette Science. II. Partie. Chez Vincent, Imprimeur-Libraire de Mgr le Comte de Provence, Paris, 640 pp. [https://www.biodiversitylibrary.org/item/6958#page/794/mode/1up]
- Baker, J.G. (1890) Further contributions to the Flora of Madagascar. (Plates L.–LIII.). Journal of the Linnean Society, Botany (London) 25: 294–350. [https://www.biodiversitylibrary.org/item/8381#page/350/mode/1up]
- Berger, A. (1930 [Mai]) Crassulaceae. Unterfam. II. Kalanchoideae [sic, Kalanchooideae]. 6. Kalanchoe Adanson. 7. Kitchingia Baker. 8. Bryophyllum Salisb. In: Engler, A. & Prantl, K. (Eds.) Die natürlichen Pflanzenfamilien, edn 2, 18a. Verlag von Wilhelm Engelmann, Leipzig, pp. 378–386, 402–412.
- Boiteau, P. (1947 [novembre-décembre]) Les plantes grasses de Madagascar. [Overview of sections in Kalanchoe.] Cactus (Paris) 1947 (12): 5–10, figures 1–4.
- Boiteau, P. & Allorge-Boiteau, L. (1995) Kalanchoe (Crassulacées) de Madagascar. Systématique, Ecophysiologie et Phytochimie. ICSN, CNRS, 91198 Gif-sur-Yvette, Éditions Karthala, Paris, 252 pp.
- Castillon, J.-B. & Castillon, J.-P. (2010 [juillet]) Les Aloe de Madagascar / The Aloe of Madagascar. Published by the authors, Etang-Salé, Île de la Réunion, France, 399 [400] pp.
- Chauvet, B. (1972) The forests of Madagascar. In: Battistini, R. & Richard-Vindard, G. (Eds.) Biogeography and ecology of Madagascar. [Monographiae biologicae vol. 21. Series Editor: J. Illies, Schlitz.] Dr. W. Junk B.V., Publishers, The Hague, pp. 191–199. https://doi.org/10.1007/978-94-015-7159-3_5
- Clements, J. (2009) Darwin’s notebook. The life, times and discoveries of Charles Robert Darwin. The History Press, Cheltenham, Gloucestershire, 160 pp.
- Cremers, G. (1976) Aloe nouveaux du nord de Madagascar. Adansonia sér. 2, 15 (4): 497–503. https://doi.org/10.5962/p.297047
- Delbourgo, J. (2018 [‘2017’]) Collecting the world. The life and curiosity of Hans Sloane. Penguin Books, Penguin Random House, United Kingdom, place of publication not stated, 504 pp.
- Descoings, B. (1997) Note sur quelques espèces nouvelles de Kalanchoe (Crassulaceae) de Madagascar. Le Journal de Botanique de la Société Botanique de France 4: 79–90. https://doi.org/10.3406/jobot.1997.1692
- Descoings, B. (2005) Crassulaceae madecassae novae. Société Botanique de l’Ardèche (éd.) Aubenas, 18 pp. + four colour plates.
- Descoings, B. (2003) Kalanchoe. In: Eggli, U. (Ed.) Illustrated handbook of succulent plants: Crassulaceae. Springer Verlag, Berlin, Heidelberg, New York, pp. 143–181; whole work 458 pp. + XLVIII plates.
- Descoings, B. & Rebmann, N. (2006 [août]) Une très belle espèce nouvelle de Kalanchoe (Crassulaceae): K. maromokotrensis. Succulentes 2006 (3): 10–17. https://doi.org/10.3406/jobot.2006.2006
- Edwards, A. (2021) The plant hunter’s atlas. A tour of botanical adventures, chance discoveries and strange specimens. Greenfinch, an imprint of Quercus Editions Ltd, an Hatchette U.K. company, London, 304 pp.
- Eggli, U. (Ed.) (2003) Illustrated handbook of succulent plants: Crassulaceae. Springer-Verlag, Berlin & Heidelberg, 458 pp + plates I–XLVIII. https://doi.org/10.1007/978-3-642-55874-0
- Farr, J. [with L. Carter] (2004) The gardens of Emily Dickinson. Harvard University Press, Cambridge, Massachusetts, U.S.A. and London, England, 350 [351] pp. https://doi.org/10.4159/9780674036727
- Gallo, L. & Smith, G.F. (2024) A review of the genus Kalanchoe (Crassulaceae subfam. Cotyledonoideae) in the Euro-Mediterranean area, with an annotated catalogue and identification key. Phytotaxa 674 (2): 171–198. https://doi.org/10.11646/phytotaxa.674.2.2
- Ganzhorn, J.U., Wilmé, L. & Mercier, J.-L. (2014) 2. Explaining Madagascar’s biodiversity. In: Scales, I.R. (Ed.) Conservation and environmental management in Madagascar. [Earthscan conservation and development series. Series Editor: W.M. Adams.] Routledge, Taylor & Francis Group, London and New York, pp. 17–43.
- Goodman, S.M., Raherilalao, M.J. & Wohlhauser, S. (2018) Les aires protégées terrestres de Madagascar: leur histoire, description et biote. Tome II. Le nord et l’est de Madagascar / The terrestrial protected areas of Madagascar: their history, description, and biota. Volume II. Northern and eastern Madagascar. Association Vahatra, Antananarivo, in three volumes, vol. I [Introduction.] pp. 1–424, vol. II. [Northern and eastern Madagascar] pp. 425–1231, vol. III [Western and southwestern Madagascar—Synthesis] pp. 1233–1716.
- Grinand, C. & Nourtier, M. (2022) Forest dynamics: carbon, drivers, and trends. In: Goodman, S.M. (Ed.) The new natural history of Madagascar. Volume 1. Princeton University Press, Princeton, New Jersey, U.S.A. and Oxford, Oxfordshire, United Kingdom, pp. 105–113, [whole work 2246 pp., in two volumes]. https://doi.org/10.2307/j.ctv2ks6tbb.21
- Hamet, R. (1907 [novembre]) Monographie du genre Kalanchoe. (A suivre. [To be continued]) Bulletin de l’Herbier Boissier, 2me sér., 7 (11): 869–900 [also independently numbered (1)–32]. [https://www.biodiversitylibrary.org/item/105267#page/939/mode/1up]
- Hamet, R. (1908 [janvier]) Monographie du genre Kalanchoe (Suite et fin. [Continuation and end], with Errata on p. 17). Bulletin de l’Herbier Boissier, 2me sér., 8 (1): 17–48 [also independently numbered (33)–64]. [https://www.biodiversitylibrary.org/item/104945#page/31/mode/1up]
- Hamet, R. & Perrier de la Bâthie, [J.M.]H.[A.] (1912) Contribution à l’étude des Crassulacées malgaches. [Kalanchoe bouvieri Raym.-Hamet & H.Perrier on p. 366.] Annales des Sciences Naturelles. Botanique, sér. 9, 16: 361–377. [https://www.biodiversitylibrary.org/item/24564#page/771/mode/1up]
- Hooker, W.J. (1859) Bryophyllum proliferum. Proliferous Bryophyllum. Nat. Ord. Crassulaceae—Octandria Monogynia. Curtis’s Botanical Magazine 85: Tab. 5147. [https://www.biodiversitylibrary.org/item/14225#page/175/mode/1up]
- Humbert, J.H. (1972) Espèces nouvelles de Begonia de Madagascar. Bulletin du Muséum National d’Histoire Naturelle sér. 3, Botanique 47 [Botanique 3]: 75–83.
- IPNI. (2025 onwards) The International Plant Names Index. Available from: https://www.ipni.org/ (accessed January 2025)
- Klein, D.-P., Shtein, R., Nusbaumer, L. & Callmander, M.W. (2021) Kalanchoe darainensis (Crassulaceae), a new species from northeastern Madagascar. Candollea 76 (1): 117–123. https://doi.org/10.15553/c2021v761a12
- Koechlin, J. (1972) Flora and vegetation of Madagascar. In: Battistini, R. & Richard-Vindard, G. (Eds.) Biogeography and ecology in Madagascar. [Monographiae biologicae vol. 21. Series Editor: J. Illies, Schlitz.] Dr. W. Junk B.V., Publishers, The Hague, pp. 145–190. https://doi.org/10.1007/978-94-015-7159-3_4
- 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; https://www.biodiversitylibrary.org/item/27173#page/243/mode/1up]
- Lamarck, J-B.P.A. de M. de. (1786 [16 October]) Encyclopédie méthodique. Botanique. Tome second, 1. [Cotyledon L. sensu lato, i.e., including Kalanchoe, treated on pp. 139–142.] Chez Panckoucke, Libraire, Hôtel de Thou, rue des Poitevins, Paris, and Chez Plomteux, Imprimeur des Etats, Liège, 774 pp. [https://www.biodiversitylibrary.org/item/15260#page/141/mode/1up]
- Linnaeus, C. (1753) Species plantarum, exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Volumes 1 and 2. Salvius, Stockholm, 560 pp. [volume 1] and 1200 pp. [total]; indices not paginated. https://doi.org/10.5962/bhl.title.669
- Miyata, K., Ikeda, H., Fujiki, T. & Ohba, H. (2016) A supplementary description of Kalanchoe humifica (Crassulaceae), with special reference to the floral characters and its chromosome number. Journal of Japanese Botany 91: 13–18.
- Miyata, K., Ikeda, H. & Ohba, H. (2018 [10 August]) Lectotypification of Kalanchoe humifica (Crassulaceae). Novon 26 (2): 214–216. https://doi.org/10.3417/2018128
- Moat, J. & Smith, P. (Eds.) (2007) Atlas of the vegetation of Madagascar / Atlas de la vegetation de Madagascar. Kew Publishing, Royal Botanic Gardens, Kew, Richmond, Surrey, 124 pp.
- Perrier de la Bâthie, [J.M.]H.[A.] (1928 [Février]) Observations nouvelles sur le genre Kalanchoe. Archives de Botanique II, Bulletin Mensuel. Caen 2 (2): 17–31.
- Persoon, C.H. (1805 [1 April–15 June 1805]) Synopsis plantarum, seu enchiridium botanicum, complectens enumerationem systematicam specierum hucusque cognitorum. Pars prima. Apud Carol. Frid. Cramerum, Parisiis lutetiorum [Paris] et apud J.G. Cottam, Tubingæ [London], pp. 445–446; whole work 546 pp. https://doi.org/10.5962/bhl.title.638
- Ranirison, P., Andriamiarantsoa, Y., Gautier, L. & Wohlhauser, S. (2018) Site 6. Loky Manambato. Vegetation. In: Goodman, S.M., Raherilalao, M.J. & Wohlhauser, S. (Eds.) Les aires protégées terrestres de Madagascar: leur histoire, description et biote / The terrestrial protected areas of Madagascar: their history, description, and biota. Volumes I–III. Association Vahatra, Antananarivo, Madagascar, [Volume II], pp. 532–534.
- Raymond-Hamet & Marnier-Lapostolle, J. (1964) Le genre Kalanchoe au Jardin Botanique “Les Cèdres”. Archives du Muséum National d’Histoire Naturelle, Septième Série, Tome VIII: 1–110; Plates I–XXXVII. [https://www.biodiversitylibrary.org/item/280857#page/21/mode/1up]
- Raymond-Hamet & Perrier de la Bâthie, [J.M.]H.[A.] (1915) Troisième contribution a l’étude des Crassulacées Malgaches. Annales du Musée Colonial de Marseille, sér. 3, 3: 63–117. [https://www.biodiversitylibrary.org/item/22992#page/85/mode/1up]
- Rodewald, S.E., Klein, D.-P., Shtein, R., Smith, G.F., Letsara, R. & Kadereit, G. (2024) S.267.5 A phylogenetic framework for the model genus Kalanchoe (Crassulaceae). [S.267 Phylogenetics and phylogenomics. Session 2. pp. 816–819.] In: XXth International Botanical Congress (IBC). July, 21st–27th, 2024. Book of Abstracts. Oral presentations. Fase 20 Ediciones, C/ Narváez, 15- 1o Izqda.- 28009, Madrid, p. 818 of 883 pp.
- Rodewald, S.E., Klein, D.-P., Shtein, R., Smith, G.F., Joyce, E., Morales Briones, D.F., Bernhard, S., Letsara, R., Mertes, H., Hühn, P. & Kadereit, G. (2025) A new phylogenetic framework for the genus Kalanchoe (Crassulaceae) and implications for infrageneric classification. Annals of Botany 135 (7): 1311–1328. https://doi.org/10.1093/aob/mcaf004
- Salisbury, R.A. (1805 [1 June]) 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
- Shtein, R. & Smith, G.F. (2021a) A revision of the climbing kalanchoes (Crassulaceae subfam. Kalanchooideae) of Madagascar including the description of Kalanchoe sect. Invasores and K. ser. Vilana. Phytotaxa 482 (2): 93–120. https://doi.org/10.11646/phytotaxa.482.2.1
- Shtein, R. & Smith, G.F. (2021b) Taxonomy and nomenclature of the southern Madagascan Kalanchoe peltigera, with reference to K. rosei var. seyrigii (Crassulaceae subfam. Kalanchooideae). Phytotaxa 490 (1): 47–59. https://doi.org/10.11646/phytotaxa.490.1.4
- Shtein, R. & Smith, G.F. (2022) Lectotypification of the name Kalanchoe peltigera (Crassulaceae subfam. Kalanchooideae). Phytotaxa 539 (1): 74. https://doi.org/10.11646/phytotaxa.539.1.9
- Small, J.K. (1933) Manual of the southeastern flora, being descriptions of the seed plants growing naturally in Florida, Alabama, Mississippi, Eastern Louisiana, Tennessee, North Carolina, South Carolina and Georgia. [Illustrated.] The University of North Carolina Press, Chapel Hill, 1554 pp. https://doi.org/10.5962/bhl.title.696
- Smith, G.F. (2020) The kalanchoes (Crassulaceae subfam. Kalanchooideae) of Raymond-Hamet (1890–1972). Bradleya 38: 120–140. https://doi.org/10.25223/brad.n38.2020.a15
- 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
- Smith, G.F. (2023) A review of exotic Kalanchoe taxa (Crassulaceae subfam. Kalanchooideae) naturalised and cultivated in Africa and the Gulf of Guinea islands, with their nomenclature updated. Phytotaxa 613 (1): 1–28. https://doi.org/10.11646/phytotaxa.613.1.1
- Smith, G.F. (2025) A nomenclator botanicus for the bryophyllous and related species published in, or transferred to, Bryophyllum, now included in Kalanchoe (Crassulaceae subfam. Cotyledonoideae). Phytotaxa 706 (1): 1–33. https://doi.org/10.11646/phytotaxa.706.1.1
- Smith, G.F. & Monro, A.M. (2024a) “Parting is such sweet sorrow [...]”, or the perils of publishing papers in parts: the case of the nomenclatural novelties published by Raymond Hamet (1890–1972) in his two-part monograph (1907, 1908) of Kalanchoe (Crassulaceae subfam. Kalanchooideae), with special reference to the new names and new combinations treated (but not first published) in part 2. Phytotaxa 645 (2): 149–162. https://doi.org/10.11646/phytotaxa.645.2.4
- Smith, G.F. & Monro, A.M. (2024b) The correct name for the subfamily containing Kalanchoe is Crassulaceae subfam. Cotyledonoideae, and not Crassulaceae subfam. Kalanchooideae, with amendment of the circumscription of Crassulaceae subfam. Cotyledonoideae. Phytotaxa 672 (3): 293–298. https://doi.org/10.11646/phytotaxa.672.3.7
- Smith, G.F. & Monro, A.M. (2025) A revised tribal classification of Crassulaceae subfam. Cotyledonoideae: separating Kalanchoe from Adromischus, Cotyledon, and Tylecodon. Phytotaxa 719 (1): 55–64. https://doi.org/10.11646/phytotaxa.716.1.6
- Smith, G.F. & Shtein, R. (2021a) Nomenclature and taxonomic placement of the infraspecific taxa recognised in the Madagascan Kalanchoe miniata (Crassulaceae subfam. Kalanchooideae) by Perrier de la Bâthie and Boiteau and Mannoni in the 20th century. Phytotaxa 514 (2): 118–128. https://doi.org/10.11646/phytotaxa.514.2.3
- Smith, G.F. & Shtein, R. (2021b) Typification and homotypic synonymy of the names published in Kalanchoe (Crassulaceae subfam. Kalanchooideae) by Raymond-Hamet and Henri Perrier de la Bâthie in the 1910s and in 1948. Phytotaxa 520 (1): 40–56. https://doi.org/10.11646/phytotaxa.520.1.3
- Smith, G.F., Shtein, R., Klein, D.-P., Parihar, B., Almeida, A., Rodewald, S. & Kadereit, G. (2021) 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
- Thiers, B. (2025 [continuously updated]) Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/ih/ (accessed January 2025)
- Turland, N.J., Wiersema, J.H., Barrie, F.R., Gandhi, K.N., Gravendyck, J., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Klopper, R.R., Knapp, S., Kusber, W.-H., Li, D.-Z., May, T.W., Monro, A.M., Prado, J., Price, M.J., Smith, G.F. & Zamora Señoret, J.C. (2025) International Code of Nomenclature for algae, fungi, and plants (Madrid Code). Regnum Vegetabile 162. The University of Chicago Press, Chicago and London, 303 pp. https://doi.org/10.7208/chicago/9780226839479.001.0001
