A review of growth form and plant duration (life cycle) in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in an evolutionary and classificatory framework

GIDEON F. SMITH; BHARTI PARIHAR; ANA M.R. ALMEIDA

doi:10.11646/phytotaxa.592.3.3

Issue: Vol. 592 No. 3: 20 April 2023

Type: Article

Published: 2023-04-20

DOI: 10.11646/phytotaxa.592.3.3

Page range: 217-254

Abstract views: 384

PDF downloaded: 7

**A review of growth form and plant duration (life cycle) in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in an evolutionary and classificatory framework**

GIDEON F. SMITH⁺⁻
BHARTI PARIHAR⁺⁻
ANA M.R. ALMEIDA⁺⁻

Ria Olivier Herbarium, Department of Botany, Nelson Mandela University, P.O. Box 77000, Gqeberha, 6031 South Africa.

Department of Biological Sciences, California State University East Bay (Hayward), 25800 Carlos Bee Blvd. Hayward, California, 94110, U.S.A.

annual arborescent biennial herbaceous multi-annual perennial shrubby woodiness Eudicots

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Abstract

Representatives of Kalanchoe (Crassulaceae subfam. Kalanchooideae) are well-known for having evolved a range of reproductive mechanisms, at least some of which have contributed to certain species and nothospecies having become weeds in places remote from their natural habitats. In addition, kalanchoes have evolved a range of different growth forms in response to a diversity of historical environmental pressures prevalent in the locations where they grow naturally. In terms of growth form, many species are herbaceous, soft- to firm-stemmed perennials that remain small to medium-sized. In some of these same species, variants that are annuals, biennials, or multi-annuals are found. A further group of species are perennial, multi-stemmed shrubs with slightly woody but rather brittle stems and branches. A small group of Malagasy species have woody stems and branches, to the extent that the plants are truly arborescent or nearly so. The distribution of growth form, with special reference to stem and branch morphology, across the four subgenera at present formally recognised in Kalanchoe is reviewed in an evolutionary and classificatory framework, and illustrated.

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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) Kalanchoe torrejacqii (Crassulaceae subfam. Kalanchooideae), a new species in K. subg. Bryophyllum from the Namorona River valley, Madagascar. Phytotaxa 498 (3): 205–212. https://doi.org/10.11646/phytotaxa.498.3.6

Shtein, R. & Smith, G.F. (2021c) 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. & Ikeda, J. (2021) Aspects of the taxonomy of the Kalanchoe daigremontiana species complex (Crassulaceae subfam. Kalanchooideae) and associated interspecific hybrids in southern Madagascar, with the description of a new nothospecies, K. ×descoingsii (=K. laetivirens × K. tubiflora). Phytotaxa 524 (4): 235–260. https://doi.org/10.11646/phytotaxa.524.4.2

Smith, G.F. (2019) Notes on Kalanchoe ×houghtonii (Crassulaceae subfam. Kalanchooideae), an early hybrid between two species of K. subg. Bryophyllum. Haseltonia 26: 78–85. https://doi.org/10.2985/026.026.0111

Smith, G.F. (2020a) Kalanchoe Adans. subg. Fernandesiae Gideon F.Sm. (Crassulaceae subfam. Kalanchooideae), a new subgenus from southern and south-tropical Africa. Haseltonia 27: 2–7. https://doi.org/10.2985/026.027.0101

Smith, G.F. (2020b) Kalanchoe ×estrelae (Crassulaceae subfam. Kalanchooideae): a new nothospecies for the hybrid between K. luciae and K. sexangularis. Phytotaxa 441 (2): 225–228. https://doi.org/10.11646/phytotaxa.441.2.11

Smith, G.F. (2020c) Zwei Neuzugänge im Kalanchoe-Gartenbau: K. ×estrelae and K. ×gunniae. / Two recent additions to Kalanchoe horticulture: K. ×estrelae and K. ×gunniae. Avonia 38 (4): 322–327.

Smith, G.F. (2020d) Plant smart. The big easy. Gardening with Kalanchoe. Veld & Flora 106 (3): 40–43.

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

Smith, G.F. (2020f) 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

Smith, G.F. (2020g) Taxonomy and nomenclature of Kalanchoe ×laetivirens (Crassulaceae subfam. Kalanchooideae), a further invasive nothospecies from Madagascar. Phytotaxa 460 (1): 97–109. https://doi.org/10.11646/phytotaxa.460.1.6

Smith, G.F. (2020h) Kalanchoe ×hankeyi [= K. longiflora × K. sexangularis] (Crassulaceae subfam. Kalanchooideae), a new nothospecies from South Africa. Phytotaxa 451 (1): 93–96. https://doi.org/10.11646/phytotaxa.451.1.9

Smith, G.F. (2020i) Character inheritance in Kalanchoe ‘Vivien’ Bischofberger (Crassulaceae subfam. Kalanchooideae), a cultivar derived from an interspecific hybrid between two southern African species (K. luciae Raym.-Hamet × K. sexangularis N.E.Br.). Bradleya 38: 3–13. https://doi.org/10.25223/brad.n38.2020.a1

Smith, G.F. (2020j) The taxonomy and nomenclature of Kalanchoe humilis, K. prasina, and K. figueiredoi (Crassulaceae subfam. Kalanchooideae). Phytotaxa 454 (4): 267–276. https://doi.org/10.11646/phytotaxa.454.4.4

Smith, G.F. (2020k) Kalanchoe ×hummeliae Gideon F.Sm.: the hybrid between K. beharensis Drake and K. millotii Raym.-Hamet & H.Perrier (Crassulaceae subfam. Kalanchooideae). Bradleya 38: 89–93. https://doi.org/10.25223/brad.n38.2020.a12

Smith, G.F. (2021a) 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

Smith, G.F. (2021b) Kalanchoe ×leistneri (K. sexangularis × K. winteri) (Crassulaceae subfam. Kalanchooideae), a new southern African nothospecies. Phytotaxa 509 (2): 248–254. https://doi.org/10.11646/phytotaxa.509.2.8

Smith, G.F. (2022a) 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. (2022b) Kalanchoe sect. Raveta (Crassulaceae subfam. Kalanchooideae) of Raymond-Hamet (1916): a long-overlooked name in K. subg. Kalanchoe. Phytotaxa 550 (2): 208–212. https://doi.org/10.11646/phytotaxa.550.2.10

Smith, G.F. (2022c) Date and place of publication and author attribution of the combination Kalanchoe sect. Raveta (Crassulaceae subfam. Kalanchooideae). Phytotaxa 560 (1): 131–134. https://doi.org/10.11646/phytotaxa.560.1.12

Smith, G.F. (2022d) Kalanchoe ×trageri, at last a name for K. blossfeldiana × K. pumila (Crassulaceae subfam. Kalanchooideae), a horticulturally successful nothospecies. Phytotaxa 550 (1): 45–58. https://doi.org/10.11646/phytotaxa.550.1.3

Smith, G.F. (2022e) Identity of Kalanchoe ‘Fang’ and K. ‘Rose Leaf’, two cultivars derived from K. ×edwardii (Crassulaceae subfam. Kalanchooideae), with notes on aspects of interspecific hybridisation in the genus. Bradleya 40: 151–160. https://doi.org/10.25223/brad.n40.2022.a14

Smith, G.F. (2022f) A review of the ecology and natural history of Kalanchoe (Crassulaceae subfam. Kalanchooideae) in southern Africa. Bradleya 40: 161–184. https://doi.org/10.25223/brad.n40.2022.a15

Smith, G.F. (2022g) Of paddles, soup plates, and clubs: the taxonomy of the southern and south-tropical African Kalanchoe sect. Raveta (Crassulaceae subfam. Kalanchooideae; Kalanchoe subg. Kalanchoe) and its constituent species. Phytotaxa 568 (1): 8–26. https://doi.org/10.11646/phytotaxa.568.1.2

Smith, G.F. (2022h) Kalanchoe ×verdoorniae [K. luciae × K. winteri] (Crassulaceae subfam. Kalanchooideae), a new, large-growing nothospecies from South Africa. Phytotaxa 568 (1): 72–82. https://doi.org/10.11646/phytotaxa.568.1.5

Smith, G.F. (2022i) A review of the cytotaxonomy of Kalanchoe (Crassulaceae subfam. Kalanchooideae) with reference to evolutionary trends in the genus. Phytotaxa 560 (2): 153–185. https://doi.org/10.11646/phytotaxa.560.2.2

Smith, G.F. (2023) A reassessment of combinations previously proposed or published for Kalanchoe subg. Alatae (Crassulaceae subfam. Kalanchooideae) and its valid publication at the rank of subgenus. Phytotaxa 585 (1): 61–66. https://doi.org/10.11646/phytotaxa.585.1.7

Smith, G.F., Bernard, S., Thiede, J. & Nickol, M. (2020b) Notes on the natural geographical distribution range of the Madagascan Kalanchoe dinklagei Rauh (Crassulaceae). Haseltonia 27: 21–28. https://doi.org/10.2985/026.027.0104

Smith, G.F., Chesselet, P., Van Jaarsveld, E.J., Hartmann, H., Hammer, S., Van Wyk, B-E., Burgoyne, P., Klak, C. & Kurzweil, H. (1998) Mesembs of the world. Illustrated guide to a remarkable succulent plant group. Briza Publications, Pretoria, 405 pp.

Smith, G.F. & Figueiredo, E. (2017a) The taxonomy of Kalanchoe longiflora Schltr. ex J.M.Wood (Crassulaceae), an endemic of South Africa’s Maputaland-Pondoland Region of Endemism. Bradleya 35: 122–128. https://doi.org/10.25223/brad.n35.2017.a11

Smith, G.F. & Figueiredo, E. (2017b) Kalanchoe fedtschenkoi Raym.-Hamet & H.Perrier (Crassulaceae) is spreading in South Africa’s Klein Karoo. Bradleya 35: 80–86. https://doi.org/10.25223/brad.n35.2017.a7

Smith, G.F. & Figueiredo, E. (2018) 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

Smith, G.F. & Figueiredo, E. (2019) Taxonomic and nomenclatural notes on Kalanchoe millotii Raym.-Hamet & H.Perrier and the cultivar Kalanchoe ‘Fern Leaf’ (Crassulaceae). Bradleya 37: 79–86. https://doi.org/10.25223/brad.n37.2019.a30

Smith, G.F. & Figueiredo, E. (2020a). The nomenclature and taxonomy of Kalanchoe nyikae (Crassulaceae subfam. Kalanchooideae), a little-known species from East Africa. Phytotaxa 454 (2): 153–158. https://doi.org/10.11646/phytotaxa.454.2.7

Smith, G.F. & Figueiredo, E. (2020b) Kalanchoe ×gildenhuysii (Crassulaceae subfam. Kalanchooideae), a new nothospecies derived from K. millotii and K. tomentosa. Phytotaxa 442 (1): 43–46. https://doi.org/10.11646/phytotaxa.442.1.8

Smith, G.F. & Figueiredo, E. (2022a) Nomenclatural and taxonomic notes on the names published in Kalanchoe (Crassulaceae subfam. Kalanchooideae) by Drake del Castillo in 1903. Phytotaxa 547 (2): 158–166. https://doi.org/10.11646/phytotaxa.547.2.3

Smith, G.F. & Figueiredo, E. (2022b) Nomenclatural notes on the economically important Flaming Katy, Kalanchoe blossfeldiana (Crassulaceae subfam. Kalanchooideae). Phytotaxa 549 (1): 104–110. https://doi.org/10.11646/phytotaxa.549.1.9

Smith, G.F. & Figueiredo, E. (2022c) Clarifying the typification of the name Kalanchoe orgyalis (Crassulaceae subfam. Kalanchooideae). Phytotaxa 543 (1): 99–102. https://doi.org/10.11646/phytotaxa.543.1.11

Smith, G.F. & Figueiredo, E. (2022d) Refining the circumscription of a variable species: epitypification of the name Kalanchoe rotundifolia (Crassulaceae subfam. Kalanchooideae). Phytotaxa 555 (1): 109–112. https://doi.org/10.11646/phytotaxa.555.1.9

Smith, G.F. & Figueiredo, E. (2022e) Kalanchoe klopperae (Crassulaceae subfam. Kalanchooideae), a southern African species newly distinguished from K. rotundifolia. Novon 30: 104–109. https://doi.org/10.3417/2022780

Smith, G.F. & Figueiredo, E. (2023) Kalanchoe orgyalis. Madagascar. Flowering Plants of Africa 68. [in press]

Smith, G.F., Figueiredo, E. & Bernhard, S. (2019c) Notes on the taxonomy and nomenclature of Kalanchoe brevisepala (Humbert) L.Allorge and its basionym, K. millotii Raym.-Hamet & H.Perrier subsp. brevisepala Humbert, and Kalanchoe dinklagei Rauh (Crassulaceae). Bradleya 37: 87–96. https://doi.org/10.25223/brad.n37.2019.a31

Smith, G.F., Figueiredo, E. & Crouch, N.R. (2017) Kalanchoe leblanciae. Crassulaceae. South Africa. Mozambique. Flowering Plants of Africa 65: 56–66, plate 2328.

Smith, G.F., Figueiredo, E. & Crouch, N.R. (2021e) Kalanchoe millotii. Crassulaceae: Kalanchooideae. Madagascar. Flowering Plants of Africa 67: 80–89, plate 2368.

Smith, G.F., Figueiredo, E., Loureiro, J. & Crouch, N.R. (2019b) Kalanchoe ×gunniae Gideon F.Sm & Figueiredo (Crassulaceae), a new South African nothospecies derived from Kalanchoe paniculata Harv. × Kalanchoe sexangularis N.E.Br. Bradleya 37: 141–150.

Smith, G.F., Figueiredo, E. & Silva, V. (2015) Kalanchoe ×houghtonii (Crassulaceae) recorded near Lisbon, Portugal. Bouteloua 20: 97–99.

Smith, G.F., Figueiredo, E. & Van Wyk, A.E. (2019a) Kalanchoe (Crassulaceae) in southern Africa. Classification, biology, and cultivation. Academic Press, an imprint of Elsevier, London, San Diego, Cambridge (USA), and Oxford, 328 pp. https://doi.org/10.1016/B978-0-12-814007-9.00003-7

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

Smith, G.F. & Hahn, N. (2019) Kalanchoe crundallii I.Verd. (Crassulaceae), the chameleon kalanchoe, a little-known South African endemic from the Soutpansberg, Limpopo Province, with notes on its conservation status. Bradleya 37: 112–117. https://doi.org/10.25223/brad.n37.2019.a5

Smith, G.F. & Hankey, A. (2021) Kalanchoe ×forbesiae (Crassulaceae subfam. Kalanchooideae) derived from K. lubangensis × K. sexangularis: towards improved horticultural material. Phytotaxa 528 (5): 290–300. https://doi.org/10.11646/phytotaxa.528.5.3

Smith, G.F., Klein, D.-P. & Shtein, R. (2021b) 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

Smith, G.F. & Monro, A.M. (2022a) The nomenclature and taxonomy of Crassulaceae subfam. Kalanchooideae resolved. Phytotaxa 555 (3): 241–251. https://doi.org/10.11646/phytotaxa.555.3.3

Smith, G.F. & Monro, A.M. (2022b) (2908) Proposal to reject the name Crassulaceae subfam. Cotyledonoideae (Crassulaceae). Taxon 71 (4): 910–911. https://doi.org/10.1002/tax.12778

Smith, G.F. & Shtein, R. (2020) Kalanchoe ×edwardii (Crassulaceae subfam. Kalanchooideae), a nothospecies derived from K. beharensis and K. tomentosa. Phytotaxa 475 (2): 117–124. https://doi.org/10.11646/phytotaxa.475.2.6

Smith, G.F. & Shtein, R. (2021a) A review of horticulturally desirable characters in Kalanchoe (Crassulaceae subfam. Kalanchooideae): variable and deviating vegetative and reproductive morphologies useful in breeding programmes. Haseltonia 28: 106–119. https://doi.org/10.2985/026.028.0110

Smith, G.F. & Shtein, R. (2021b) Typification of the name Kalanchoe daigremontiana (Crassulaceae subfam. Kalanchooideae). Phytotaxa 494 (2): 244–249. https://doi.org/10.11646/phytotaxa.494.2.8

Smith, G.F. & Shtein, R. (2022) Nomenclature and taxonomy of the partial infrageneric classification systems proposed for Kalanchoe (Crassulaceae subfam. Kalanchooideae) by Boiteau and Boiteau & Mannoni in the mid-20th century. Phytotaxa 536 (2): 141–153. https://doi.org/10.11646/phytotaxa.536.2.3

Smith, G.F., Shtein, R. & Klein, D.-P. (2021a) 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

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

Smith, G.F., Wolff, E. & Thoumin, L. (2020a) The taxonomy and nomenclature of Kalanchoe gastonis-bonnieri Raym.-Hamet & H.Perrier (Crassulaceae subfam. Kalanchooideae), with biographical notes on Gaston Eugène Marie Bonnier (1853–1922). Bradleya 38: 94–103. https://doi.org/10.25223/brad.n38.2020.a13

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Smith, G.F., Parihar, B. & Almeida, A.M. (2023)

A review of growth form and plant duration (life cycle) in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in an evolutionary and classificatory framework

. Phytotaxa 592 (3): 217–254. https://doi.org/10.11646/phytotaxa.592.3.3

A review of growth form and plant duration (life cycle) in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in an evolutionary and classificatory framework

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**A review of growth form and plant duration (life cycle) in Kalanchoe (Crassulaceae subfam. Kalanchooideae) in an evolutionary and classificatory framework**