Kalanchoe, Adanson, 1763

Smith, Gideon F. & Figueiredo, Estrela, 2024, A review and analysis of the early nomenclature and taxonomy associated with the genus name Kalanchoe (Crassulaceae subfam. Kalanchooideae), published by Michel Adanson (1727 - 1806) in 1763, with notes on the late- 18 century taxonomy of the genus, Phytotaxa 633 (2), pp. 108-124 : 109-118

Valid publication of the genus name Kalanchoe :

—In the nomenclatural system that Adanson (1763) used, species were named in the same way as genera, i.e., by means of a single name—also referred to as a unitary name (see Bai et al. 2023: 152). Parkinson (1987a) argued that the genus names of Adanson were used simultaneously at two different ranks (those of genus and species) and that they were therefore not validly published, i.e., that they were contrary to the 1983 International Code of Botanical Nomenclature (IBCN) (see Voss et al. 1983: Art. 20.4(b )), currently the International Code of Nomenclature for algae, fungi, and plants (ICN) (see Turland et al. 2018: Art. 20.4(b )). This resulted in the validity of Adanson’s generic names being widely questioned more than 30 years ago [see for example Parkinson (1987a, b, c) in general, and Dorr & Parkinson (1990: 131) on Ravenala Adanson (1763: 67) in particular].

Parkinson (1987a: 85–86) submitted a proposal for consideration by the then Committee for Spermatophyta for Kalanchoe and other genus names of Adanson to be regarded as not having been validly published. These names, depending on the circumstances, would consequently require a positive conservation decision to allow their usage to continue. In the case of Kalanchoe , its conservation against Vereia Andrews (1798 : t. 21) was proposed formally ( Parkinson 1987a: 85–86).

However, in the interests of nomenclatural stability, the views of Parkinson (1987a, b, c) were not approved, with Brummitt (1990: 295–296) recording that: “The committee [i.e., the then Committee for Spermatophyta] unanimously rejects Parkinson’s proposition on Adanson’s names, and the proposals […] [including on conserving Kalanchoe against Vereia ] are therefore to be rejected as unnecessary.” As a result Adanson’s genus names, including Kalanchoe , were ruled as in fact validly published (see Brummitt 1993: 874 and Nicolson 1993: 443).

Adanson’s description of Kalanchoe :—When Adanson (1763: 248) validly published the genus name Kalanchoe , in “XXXIII. Famille Les Jourbarbes. Seda.”, i.e., what is known today as the Crassulaceae , he provided the following description for the genus in an analytical Table: “ Feuilles. Oposées, simples & ailées. Fleurs. Corymb. Calice. 4 f. [feuilles]. Corolle. Tube long. 4 divis. Etamines. 8, inégales sur 2 rangs. Pistil. 4[,] ovaires. Graines. Nombreuses, menues, cylindriques”. This description, in translation, is here interpreted as: “ Leaves opposite, simple, winged [further discussed below under ‘ Identity of kalanchooid material at Adanson’s disposal:—’]. Flowers [carried in a] corymb. Calyx 4 leaved [i.e., consisting of 4 sepals]. Corolla long tube. 4-divided [i.e., elongated and consisting of 4 petals]. Stamens 8, unequal, in two ranks. Pistil [consisting of] 4 ovaries. Seed numerous, small, cylindrical.”

Although this description of Kalanchoe was subsequently inevitably expanded, refined, and clarified, it still well defines the genus in the context of the Crassulaceae , and confirms, at least to some degree, Adanson’s approach of using multi-character descriptions when classifying biological organisms.

The tetramerous nature of the flowers of at least one species, Cotyledon laciniata Linnaeus (1753: 430) , which, pre-Adanson, was included in Cotyledon Linnaeus (1753: 429) , nom. cons., was well-known. Linnaeus himself had noted this deviation from the pentamerous flowers of Cotyledon , as understood at present, in the first ( Linnaeus 1753: 430) and second ( Linnaeus 1762: 615) editions of his Species plantarum , and after 1753 in the 5 th edition of his Genera plantarum ( Linnaeus 1754a: 196), the first edition of which had appeared in 1737. However, more vaguely, the corolla and calyx of a broadly defined Cotyledon were noted as “semiquinquefidum” in Linnaeus (1737: 136).

Additional information provided by Adanson when he described Kalanchoe :—For the genus name Kalanchoe [“ Kalanchoè .”], Adanson (1763: 248) provided the following three statements in the first column of the analytical Table: [1.] “ Chin.”; [2.] “ Tjsacarbebe. Rumph. 5. t. 95.”; and [3.] “ Cotyledon . Boerh. I. B. t. 288.” ( Table 1 ). These statements were added to on p. 530 and p. 614 of Adanson (1763).

For ease of reference all the statements, sequentially numbered, that were associated with the name Kalanchoe by Adanson (1763: 248, 530, 614) are included in Table 1 and discussed in the text that follows.

“ Kalanchoè .” ( Adanson 1763: 248, 614):—[ Table 1 rows A.1, B.5 and C.8] Since it was the genus name published by Adanson, on p. 248 “ Kalanchoè ” was printed in bold and in a slightly larger font size than the rest of the text associated with it. On p. 530 the genus name was printed in small capitals and the ‘E’ lacked a grave accent, and on p. 614 it was again given with a grave accent on the ‘ è ’.

“ Chin. / “ Sin.” ( Adanson 1763: 248, 530):—[ Table 1 rows A.2 and B.5] Of the 11 genera that Adanson (1763: 248–249) recognised in the “Famille Les Jourbarbes. Seda.”, it was only for Kalanchoe that Adanson added a statement, (“ Chin.”), about the origin of the name [ Kalanchoe , not of “ Tjsacarbebe ”, see ‘“ Tjsacarbebe. Rumph. 5. t. 95.” ( Adanson 1763: 248, 530, 614):—’, below]. The work ( Adanson 1763) was written mostly in French, in which “ China ” translates as “ Chine ”, hence the abbreviation “ Chin.”. The statement “ Chin.” is not regarded as an indication of the origin of the material that Adanson included in Kalanchoe , but rather as the linguistic or geographical origin of the name Kalanchoe . In contrast to Adanson (1763: 248), where “ Chin.” is given, the abbreviation “ Sin.” is used on p. 530, i.e., the Latin abbreviation for “ Sina ” [ China] (further discussed below under ‘“ Tjsacarbebe. Rumph. 5. t. 95.” ( Adanson 1763: 248, 530, 614):—’).

“ Tjsacarbebe. Rumph. 5. t. 95.” ( Adanson 1763: 248, 530, 614):—[ Table 1 rows A.3, B.6, and C.8, in Adanson (1763: 530, 614) as “Tsjakarbebe”, i.e., spelled with a ‘k’, not a ‘c’, to so provide the phonetical transcription of the name; and only “ Rumph.” given in Adanson (1763: 530)] Adanson was not enamoured with the binomial system of nomenclature introduced by Linnaeus 10 years before his own Familles des plantes was published (also see for example McOuat 2001). This contributed to contemporary fellow botanists and natural historians failing to appreciate the significance of Adanson’s work, which, in the case of the publication of the name Kalanchoe , was not adopted for several decades. For example, K. crenata ( Andrews 1798: t. 21) Haworth (1812: 109), which was first described as Vereia crenata Andrews (1798 : t. 21) 25 years after Adanson’s Familles des plantes was published, is one indication that Adanson’s publication of the genus name Kalanchoe in 1763 was either overlooked (unlikely) or ignored (probably). In addition, Adanson’s preference to not follow Linnaeus’s binomial system has unfortunately resulted in considerable nomenclatural confusion concerning Adanson’s genus names and some of these are still being typified 260 years later (see for example Bai et al. 2023: 156).

For species names, Adanson had a preference for using one-word vernacular names instead of the Latin binomials introduced by Linnaeus and “ Tjsacarbebe ” is the common name of the (apparently but, as argued further on, very likely not single) species he included in Kalanchoe .

“ Rumph.” is a reference to the German-born, Dutch naturalist Georg Eberhard Rumphius [Rumpf] (1628–1702) who is best known for having authored the posthumously published, “ Herb. amboin.” ( Rumphius 1747 [1741–1750]). The ‘ Index universalis in sex tomos et Auctuarium Herbarii amboinensis Cl. Georgii Everhardi Rumphii ’ ( Rumphius 1755), edited by Dutch botanist Johannes Burman (1706–1779) is credited as the first place in the literature in which an author other than Linnaeus or his students used Linnaean binomials, following their introduction by Linnaeus (1753) two years earlier ( Stafleu & Cowan 1983: 990, Jarvis 2019).

Rumphius (1747: vol. 5: 276 and “ Tab. XCV ” on the facing p.), which was also posthumously “ nagezien en uitgegeven ” [edited and published] by “Joannes Burmannus”—Rumphius had been deceased for 45 years—included the following text: “Tabula Nonagesima Quinta ( Fig. 1A ). Plantam exhibet Anatis Rumphio dictam, cum ejus caule florifero A separato, qui quam rarissime prodit. Observatio. Haec est Cotyledon foliis laciniatis H. Cliffort. p. 175 & Cotyledon Afra, folio crasso, lato, laciniato, flosculo aureo. Boerh. Ind. H. L. B. part. I. p. 288. ubi vide iconem; & Telephium Africanum , angustiore folio, flore aurantiaco. Plukn. Almag. p. 362. Tab. 228. Fig. 3. [English: “Ninetyfifth Plate. Shows Rumphius’s Planta A[Á]natis, with its flower-carrying stem [peduncle] which is rarely borne. Observation. This is the Cotyledon foliis laciniatis of Hort. Cliffort. p. 175. and Cotyledon Afra, folio crasso, lato, laciniato, flosculo aureo of Boerh. Ind. H. L. B. part. I. p. 288. where the Figure can be seen, and the Telephium Africanum , angustiore folio, flore aurantiaco of Plukn. Almag. p. 362. Tab. 228. Fig. 3.”] As noted further on, the reference to “ Telephium […] flore aurantiaco” was an error.

The description included in Rumphius (1747: 276), as well as the material illustrated, coincide with what is today known as K. laciniata , which is not indigenous to Ambon ( Indonesia), where Rumphius worked, but according to him it was found in cultivation. The plant was named “ Planta Anatis ” by Rumphius (1747: 276). Linnaeus (1754b: 21), who attempted to provide binomial names for some of the species described by Rumphius (see Jarvis 2007: 93), equated “ Planta Anatis ” with Cotyledon laciniata . Note that the designations listed in the left-hand column of Linnaeus (1754b) were not names accepted by Linnaeus upon publication and are not validly published ( Turland et al. 2018. Ex. 2 under Art. 36). Merrill (1917: 243) noted that the description of Rumphius might also apply to K. pinnata , but “primarily, however, Planta anatis is certainly Kalanchoe laciniata ”.

Rumphius (1747: 276) also noted the Malay vernacular name of the species as “Tsjaccarbebe” [spelled with two ‘c’s].

The citation of “ Plukn. ” in Rumphius (1747: 276) is a reference to Leonard Plukenet (1642–1706) [Plukenetius], a British botanist and physician (see Stafleu & Cowan 1983: 300). In the context of Kalanchoe , two of Plukenet’s publications are of relevance:

• Plukenet’s Phytographia of 1692; and

• Plukenet’s Almagestum of 1696.

“ Plukn. Almag. p. 362. Tab. 228. Fig. 3.” is a reference to the “ Almagestum ” ( Plukenet 1696). On p. 362 of Plukenet (1696) there is mention of: “ Telephium maximum Africanum flore aurantiaco [orange], ex Cod. Benting Phytogr. Tab. 228 fig. 3. Itèm ex Hollandiâ habuimus.” In turn, “ex Cod. Benting Phytogr. Tab. 228. fig. 3” is a reference to Plukenet (1692), i.e., to his “ Phytographia ”, which was published four years earlier. Figure 3, in the centre of the plate “Tab. 228”, i.e., “TAB. CCXXVIII”, is distinctly a rendition of a species of Kalanchoe , but certainly not of K. laciniata , which has characteristically incised leaves ( Fig. 1B ). In the caption to “TAB. CCXXVIII”, Figure 3 is described as “ Telephium max, Africanum flore aurantiaco. ex Cod. Benting.” This text clearly coincides closely with the phrase name and text included in Plukenet (1696: 362).

Apart from where Kalanchoe had been validly published on p. 248 of Adanson (1763), this genus name was also referenced on pp. 530 and 614. On p. 530 the following information was provided: “ KALANCHOE . Sin. [here given as the Latin abbreviation for “ Sina ” [ China], and not as “ Chin.”, as on p. 248]. Camell. Tsjakarbebe [note the spelling with a ‘k’, not a ‘c’, as on p. 248]. Rumph. Cotyledon, Boerh. ”

“ Camell.” ( Adanson 1763: 530) [ Table 1 row B.5] is a reference to Georg Joseph Camel (1661–1706), a Jesuit missionary and pharmacist who worked in the Philippines ( Gicklhorn & Gicklhorn 1954: 20–25, Cullum 1956, Stafleu & Cowan 1979: 491, Kroupa et al. 2020: 174–179). Camel’s name is variously spelled, as Kamel, Cameli, Camelli, Camello, or Camellus. He himself signed his name as ‘Camel’ ( Reyes 2009: 273, Note 1) and this spelling is adopted in this paper, but “Camello” is used in the reference to “Camello, G.J. (1704)”.

Camel corresponded with several notable botanists and plant collectors in Great Britain and continental Europe, including John Ray (1627–1705) and James Petiver (ca. 1665–1718), himself a pharmacist ( Gicklhorn & Gicklhorn 1954: 76–78, Kroupa 2016), and through Petiver, some of Camel’s preserved specimens, drawings, and texts were taken up in the herbarium of Hans Sloane (1660–1753). Sloane bequeathed his collections to the British nation, so providing an early nucleus for the British Museum, later the Natural History Museum, London, where some of Camel’s Philippine materials still survive ( Cristini 2022: 25). Camel died of dysentery in the Philippines at the age of 45 ( Cullum 1956: 336, Kroupa et al. 2020: 177). The horticulturally popular genus Camellia Linnaeus (1753: 698) commemorates Camel. Tea, the globally popular beverage consumed hot or cold, also is at present included in Camellia , as C. sinensis ( Linnaeus 1753: 515) Kuntze (1887: 195) . Camel is botanically best known for his work Herbarium aliarumque stirpium in insula Luzone Philippinarum: […] ( Camello 1704). “ Herb. philipp.”, as it is known when abbreviated, was published as an appendix to volume three of John Ray’s Historiae plantarum tomus tertius ( Stafleu & Cowan 1979: 492, 1983: 609).

As recorded by Kroupa et al. (2020: 178–179), citing Gossypium arboreum Linnaeus (1753: 693) , tree cotton, as an example, which is indigenous to the Indian subcontinent, there existed “[…] early modern networks of commerce and migration that brought the two regions [the Indian subcontinent and the Philippines] together” and that “Through the lens of Kamel’s plants, we can recreate the bustling, cosmopolitan world of early modern Manila ”. It is therefore clear that plant material from mainland Asia, including from the Indian subcontinent and China, could easily have reached the Philippines. Kalanchoe laciniata ( Linnaeus 1753: 430) Candolle (1802 : t. 100) was indeed recorded from southern India by Wight & Walker-Arnot (1834: 360) (see also Philcox 1999: 63, Descoings 2003: 160).

A further significant aspect of Camel’s work in the Philippines is that he recorded the indigenous, vernacular names of the plants he collected and investigated ( Turner & Veldkamp 2009: 190, Kroupa et al. 2020: 179). In Camello (1704: 6, no. 18) he describes a plant as “ Telephium Sempervivum , seu Sinarum Kalanchauhuy ” where “ Sinarum ” indicates that the genus name Kalanchoe has a Chinese origin ( Don 1834: 108, Wittstein 1856: 137, Harvey 1862: 378 and, more recently, Bayton 2020: 173). Although Cristini (2022: 25) correctly noted that Adanson (1763: 248) derived “ Kalanchoè ” from “ Kalanchauhuy ”, Cristini (2022) further noted that the latter name was a vernacular name obtained from “[…] the Chinese people (“ Sinarum ”)”, a statement for which we found no evidence. “ Sinarum ” means “of China ” and here it refers to the common name “ Kalanchauhuy ” having a geographical origin in China. We concur with Pattock (2022: 148) that the name had an origin in one of the Chinese languages. Further to this, Eggli & Newton (2004: 125) speculated that Kalanchoe might have been derived from “ancient Indian”, as in “‘kalanka-’, spot, rust; and ‘chaya’, gloss; perhaps for the glossy and perhaps sometimes reddish leaves of the Indian K. laciniata .” (see also Smith & Figueiredo 2023: 231).

“ Cotyledon . Boerh. I. B. t. 288.” ( Adanson 1763: 248, 530):—[ Table 1 rows A.4 and B.7, with the book and plate references not given on p. 530 of Adanson (1763) and Boerhaave’s work not having been mentioned on p. 614 of the same)] “ Boerh.” is a reference to the Dutch physician, chemist, and botanist Herman Boerhaave (1668–1739), who is best known botanically for his two works that appeared in 1710 (Index plantarum, quae in horto academico Lugdano Batavo reperiuntur. [Ind. pl. hort. Lugd. Bat.]) and in 1720 (Index alter plantarum quae in horto academico Lugduno-Batavo aluntur. [Ind. alter hort. Lugd.-Bat.]), respectively ( Stafleu & Cowan 1976: 250). Post-1720, these two works were sometimes bound and distributed or sold together.

Specifically, “ Cotyledon . Boerh. I. B. t. 288.” and “ Cotyledon . Boerh.”, as included in Adanson (1763) on p. 248 and p. 530, respectively, are references to the entry: “10. Cotyledon ; Afra; folio crasso, lato, laciniato, flosculo aureo [golden]. Telephium , maximum , Africanum , flore aurantio. Ex Cod. Bent. Plukn. Phyt. 228 3? H. R. D.” ( Boerhaave 1720: 288). The black-and-white line drawing linked with entry no. 10 and similarly captioned appears on the page facing p. 288 in Boerhaave (1720). The material illustrated in Boerhaave (1720: [unnumbered] page facing p. 288) clearly corresponds with material of what today is known as K. laciniata ( Fig. 1C ). The “?” that appears in entry no. 10 on p. 288 of Boerhaave (1720) is an indication that, as discussed above under ‘“ Tjsacarbebe. Rumph. 5. t. 95.” ( Adanson 1763: 248, 530, 614):—’, the plate in Plukenet (1692: “TAB. CCXXVIII, Fig. 3”), while of a species of Kalanchoe , cannot be reconciled with material today known as K. laciniata .

Notably, mention of “ Cotyledon ” in “ Cotyledon . Boerh. I. B. t. 288.” is not a reference to the broadly conceived Cotyledon of Linnaeus (1753: 429–430). In fact, nowhere where dealing with Kalanchoe did Adanson (1763: 248, 530, 614) include a direct reference to Linnaeus (1753). However, the protologue of the name Cotyledon laciniata Linnaeus (1753: 430) included the following information: “ laciniata , 6. COTYLEDON foliis laciniatis, floribus quadrifidis. Hort. cliff 175. Roy. lugdb. 454. Cotyledon afra , folio crasso lato laciniato, flosculo aureo. Boerh. lugdb. I. p. 288. t. 288. Telephium africanum , angustiori folio, flore aurantiaco. Pluk. alm. 362. t. 228. f. 3. Habitat in AEgypto.”

Identity of kalanchooid material at Adanson’s disposal:—Material with deeply invaginated, i.e., ‘laciniated’, leaf blades that is referenced or mentioned in Adanson (1763), Rumphius (1747), and Linnaeus (1738), in the latter two instances typically with phrase names ( Table 2 ), very likely refers to the species at present known as K. laciniata ( Figs 1D and 1E ). For example, reference to the “winged” leaves [as “ Feuilles. Oposées, simples & ailées.”] of [one of] the kalanchoes that Adanson (1763: 248) had at his disposal when he described the genus is an indication that he was very likely referring to K. laciniata . Especially the lower leaves of the species create the impression that they consist of a central blade with two or more lateral “wings” ( Figs 1B and 1D ).

It may be tempting to argue that Adanson (1763: 248, 530, 614) had material of K. ceratophylla Haworth (1821: 24) at his disposal, rather than of K. laciniata , when publishing Kalanchoe and the vernacular name Tjsacarbebe. [Note that the protologue associated with the name K. ceratophylla was published in Haworth (1821: 23), and not in Haworth (1819), as stated by Descoings (2003: 144).] However, with the exception of the material from Ambon ( Indonesia) cited by Rumphius (1747), in the references cited by Adanson (1763), the material was clearly stated as of African origin. Some 40 years after Kalanchoe was described, Haworth (1821: 24) noted, in contrast, that K. ceratophylla originated from China, when he stated for this species “ Habitat in Sina [ China]”. However, Descoings (2003: 144) treated the name K. ceratophylla as being of uncertain application, which probably gave rise to Eggli & Newton (2004: 125) rather proposing K. laciniata as (one of) the species referenced by Adanson (1763: 248), when suggesting one of two possible derivations for the genus name. The other species mentioned by Eggli & Newton (2004: 155) was K. spathulata Candolle (1801 : t. 65), which was introduced earlier from China. However, the latter does not have incised leaves.

When early in his career Adanson spent six years conducting natural history explorations in Senegal he might have gained first-hand experience of African kalanchoes. However, no kalanchoes have thus far been recorded as indigenous to that country. Kalanchoe crenata [ Sierra Leone and Guinea (see Smith et al. 2019: 148–154)] and K. laciniata [ Ivory Coast, Benin, and Morocco (see Maire 1977: 258–260 and Smith et al. 2019: 165–169)] are the two species of Kalanchoe whose known natural geographical distribution ranges come closest to Senegal. Kalanchoe laciniata , the type species of Kalanchoe , was cited by Haworth (1829: 302) as doubtfully also occurring in Egypt [“AEgypto”], indicated with a “?”, apart from it having been found “in India ”. In its very broad definition currently widely upheld, the species possibly occurs in Egypt (but see Dobignard & Chatelain 2011: 369) and has been recorded from neighbouring Sudan ( Pickering & Darbyshire 2015: 155–156). Kalanchoe crenata is a widely used medicinal species and has been recorded in infant healthcare in western, central, and eastern Africa, for example ( Sibeko et al. 2023: 125).

Of the African kalanchoes, K. faustii Font Quer (1935: 149) from Morocco, a K. laciniata look-alike, is the species with the most northwesterly natural distribution range. Some sources, for example Dobignard & Chatelain (2011: 369), uphold K. faustii as a species distinct from K. laciniata , while other sources, for example Descoings (2003: 160) place it, as well as K. laciniata subsp. faustii (Font Quer) Maire (1977: 260) , in the synonymy of K. laciniata .

Shortly after K. faustii was described, Maire (1938: 418), while still regarding it as an accepted species, and not as warranting recognition at the rank of subspecies, noted a relationship between K. laciniata and K. crenata , the two species here suggested as having been known to Adanson (1763: 248, 530, 614): “Les fleurs de cette be’le plante, qui atteint 1 m 50 de hauteur, sont jaune d’or et diurnes; elles se ferment la nuit. Les espèces les plus affines sont les K. crenata Haw. et K. laciniata (L.) D.C. de l’Afrique tropicale. Nous cultivons cette plante à Alger où elle se développe vigoureusement en pleine terre; le Jardin Botanique de l’Université d’Alger en distribue les graines et des boutures.” [English: “The flowers of this beautiful plant, which reaches a height of 1.5 m, are golden-yellow and diurnal; they close at night. The most closely related species are K. crenata Haw. [sic, (Andrews) Haw.] and K. laciniata (L.) DC. from tropical Africa. We cultivate this plant in Algiers where it grows vigorously in the ground; the Botanical Garden of the University of Algiers distributes its seeds and cuttings.”]

Both K. crenata and K. laciniata are widely cultivated as ornamentals and at least K. crenata ( Figs 2A and B ) and possibly also K. laciniata , are known to escape and become naturalised with ease in places well away from where they occur naturally ( Maire 1977: 260, Vargas et al. 2022: 3–4, 6, 11–12 of 22), especially through the germination of copiously produced seed, and it is possible that Adanson had access to both species while based in Senegal.

Kalanchoe crenata , of which the basionym is Vereia crenata , was described from a plant cultivated in the garden of James Vere, a wealthy merchant based at Kensington Gore, London, England, and a patron of botany; Vere was commemorated in the genus name Vereia Andrews (1798 : t. 21). The plant that was cultivated in Vere’s garden originated from Sierra Leone, where it had been collected in 1793 by Adam Afzelius (1750–1837), a Swedish botanist. Incidentally, Afzelius is commemorated in K. afzeliana Britten (1871: 393) , which also is today included in the synonymy of K. crenata . However, as noted by Fernandes (1980: 338) it is possible—in fact, very likely—that plants of K. crenata were known in Europe long before it was described in 1798, given that the figure published by Plukenet (1692: t. 228, Fig. 3) that he later described as ‘ Telephium maximum Africanum flore aurantiaco, ex Cod. Benting. Phytogr. Tab. 228. fig. 3. Item ex Hollandiâ habuimus’ ( Plukenet 1696: 362) very likely refers to K. crenata and was published more than 100 years before the species was finally described post-1753.

According to Fu et al. (2001a: 204, b), K. ceratophylla is the only species of Kalanchoe indigenous to China that has “pinnately lobed” leaves. In this regard, Ohba (2003: 248) drew attention to similarities between it and K. laciniata in terms of the laciniate leaves of both species. However, K. ceratophylla was only “introduced [to the Royal Botanic Gardens, Kew,] from China in 1820 by Captain Nisbett” ( Wickens 1982: 674), i.e., almost 60 years after Adanson (1763) published the genus name Kalanchoe . This is the material that Haworth (1821: 23) described, from cultivation at Kew, as K. ceratophylla . Material of K. spathulata was earlier introduced from China, but this species does not have laciniate leaves. Two years after Fu et al. (2001a: 204–205) was published, K. ceratophylla var. indochinensis Ohba (2003: 254) was described for incised-leaved material “[…] with many elongate lateral branchlets […]”from Laos and Vietnam.

Fernandes (1980: 377) had doubts about the identification of Rumphius’s plant from Ambon as K. laciniata because of a reported lack of pubescence of specimens from Malaysia, as opposed to the pubescent type material of the species name, and suggested that only the examination of further material from the region might confirm it. In this instance it is possible that Fernandes (1980: 377) might have referred to material of K. ceratophylla . However, as we argue here, there is sufficient evidence that probably placed K. laciniata in cultivation in the Philippines.

The first species of “ Cotyledon ” that Linnaeus (1738: 175), i.e., pre-1753, included under “ COTYLEDON g. pl. [Genera plantarum ] 382 [the genus number, not the page number].” was “1. COTYLEDON foliis laciniatis.”, which he noted as “ Crescit in [grows in]Africa.” He also provided the following two references: “ Cotyledon afra , folio crasso lato laciniato, flosculo aureo. Boerh. lugdb. 1. p. 288 t. 288.” and “ Telephium africanum , angustiori folio, flore aurantiaco Pluk. alm. 362. t. 228. f. 3.” However, “ Telephium africanum , angustiori folio, flore aurantiaco”, as cited by Linnaeus (1738: 175) with reference to “ t. 228. f. 3” is not convincingly of a species today referable to Kalanchoe . Fernandes (1980: 375) identified it as a species of Sedum Linnaeus (1753: 430) . In fact, “ Telephium africanum , angustiori folio, flore aurantiaco”, was illustrated in Plate 216, Figure 2 , and not in Plate 228, Figure 3 ( Plukenet 1692: t. CCXVI)”. Four years after “Phytogr.” ( Plukenet 1692) was published, Plukenet (1696: 362) stated for the latter species that it had been previously considered to be a Portulaca Linnaeus (1753: 445) : “ Telephium Africanum , angustiori folio, flore aurantiaco è Cod. Bent. sub Portulacae titulo in Phytogr. Tab. 216 fig. 2. hujus Icon exhibetur.” Like Linnaeus (1738), Rumphius (1747: 26), earlier, also cited Plukenet’s Plate 228, Figure 3, as being associated with “ Telephium africanum , angustiori folio, flore aurantiaco”.

The legitimacy of Kalanchoe :—To determine the legitimacy of the genus names that Adanson (1763) published, including of Kalanchoe , it is necessary to check whether he cited any previously published, legitimate genus names under his own genus names, in his analytical Tables or in the associated indices. This is additionally because in several instances Adanson (1763) did cite such earlier, legitimate genus names under his genera, which would make his names superfluous under Turland et al. (2018: Art. 52.1 and Art. 52.2). For example (see Turland et al. 2018: Ex. 1 under Art. 52.2), Adanson (1763: 166) cited the genus name Chrysophyllum Linnaeus (1753: 192) as a synonym under Cainito Adanson (1763: 166) so rendering Cainito Adans. illegitimate because it was superfluous when published—Adanson should have adopted Chrysophyllum L. A similar situation applies in the “Famille Les Jourbarbes. Seda.” of Adanson (1763), i.e., the Crassulaceae , when “ Rhodiola . Lin.” was included in the synonymy of Rhodia Adanson (1763: 248) — Adanson should have adopted Rhodiola Linnaeus (1753: 1035) .

Turland et al. (2018: Art. 52.1 and 52.2) state, respectively, that: “A name, unless conserved […], is illegitimate and is to be rejected if it was nomenclaturally superfluous when published, i.e. if the taxon to which it was applied, as circumscribed by its author, definitely included the type […] of a name that ought to have been adopted […]” and that “For the purpose of Art. 52.1, definite inclusion of the type of a name is effected […]” (italicised emphasis on ‘ name ’ introduced here).

In the case of Kalanchoe, Adanson (1763: 248 , 530, 614) included references to “ Cotyledon . Boerh. I. B. t. 288” [on p. 248] and to “ Cotyledon . Boerh. ” [on p. 530], with both of these being references to the same pre-Linnaean phrase name (of Boerhaave). The other references cited by Adanson, i.e., “ Rumph. 5. t. 95” and “ Camell.”, which did not include mention of “ Cotyledon ”, but rather referenced “ Telephium ” ( Plukenet 1696: 362) and “Sempervivum” ( Camello 1704: 6, no. 18), were also to publications that predated 1 May 1753, the starting point for the valid publication of names for non-fossil Spermatophyta (and some other groups, for that matter) ( Turland et al. 2018: Art. 13.1). The pre-Linnaean ‘names’ published in the works cited by Adanson (1763: 248, 530, 614) were not validly published and are therefore not “names” in the sense of Turland et al. (2018: Art. 6.3).

None of the elements included in Kalanchoe by Adanson (1763: 248, 530, 614) is the type of a genus name that ought to have been adopted instead of Kalanchoe . Fernandes (1980: 376) designated the specimen Herb. Clifford 175 [“l’Hortus Siccus Cliffortianus”] held at Herb. BM under barcode BM000628567 as the type of the name K. laciniata . It was corrected to lectotype by Jarvis (2007: 451). Kalanchoe laciniata is the type of the genus Kalanchoe . This element represents the type of a legitimate species name, which is in fact a representative of Kalanchoe as presently circumscribed.

The two elements, “ Cotyledon afra , folio crasso lato laciniato, flosculo aureo. Boerh. lugdb. i. p. 288. t. 288” and “ Telephium africanum , angustiori folio, flore aurantiaco. Pluk. alm. 362. t. 228. f. 3.”, that were cited in the protologue of the name Cotyledon laciniata by Linnaeus (1753: 430), were applied to the same species, i.e., to Cotyledon laciniata . No other elements were cited for C. laciniata by Linnaeus (1753: 430), and Cotyledon , i.e., the genus name itself, was eventually conserved and typified on C. orbiculata Linnaeus (1753: 429) , a species with pentamerous flowers (see [Tölken in Jarvis in] Nicolson 1992: 561, and Barrie 2006: 795–796, Jarvis 2007: 451, and Wiersema et al. 2018 + [continuously updated] for summaries). Note, however, that, as discussed above, the wrong phrase name, which does not apply to “ Tab. 228 fig. 3” was still used by Linnaeus (1753: 430), after this error was introduced in Linnaeus (1738: 175). It should have been “ Telephium maximum Africanum flore aurantiaco, ex Cod. Benting Phytogr. Tab. 228 fig. 3. Itèm ex Hollandiâ habuimus”. However, Linnaeus (1753: 430) cited the correct Figure number.

In summary, in the protologue of Kalanchoe ( Adanson 1763) , reference to Cotyledon was not to the genus Cotyledon sensu Linnaeus (1753) , and no types of species names currently included in Cotyledon , as conserved and typified with a pentamerous-flowered species, were mentioned. The genus name Kalanchoe Adans. , which dates from 1763, is not homotypic with Cotyledon L., nom. cons., which dates from 1753, and Kalanchoe is a legitimate genus name.

The name “ Tjsacarbebe ” cited by Adanson (1763) was merely a Malay vernacular name and not intended as a genus name, nor as a species name (a binary combination, therefore) in the sense of Turland et al. (2018: Art. 23.1).

The 18 th century taxonomy of species today included in Kalanchoe :—The taxonomy of Kalanchoe started off conservatively in the 18 th century with only Cotyledon laciniata , i.e., K. laciniata , having been included in Linnaeus (1753: 430). Apart from K. laciniata having been known at the time, K. crenata [first published as Vereia crenata in 1798] was also, as discussed above, very likely available in Europe since pre-Linnaean times, but it was not included in Linneaus (1753) as a taxon distinct from his C. laciniata .

As far as could be determined, only two further species that are today included in Kalanchoe were described in the 1700s, by Lamarck (1786), also as species of Cotyledon following the broadly conceived Linnaean concept of Cotyledon , rather than Adanson’s narrower recognition of the tetramerous-flowered ‘cotyledons’ as kalanchoes. However, Lamarck (1786: 139, 141), like Linnaeus, realised the disparate nature of Cotyledon in terms of the number of flower parts of the species that he included in the genus and, in a rudimentary identification key, he separated the species with pentamerous flowers [“*Fleurs quinquefides”] from the three that he recognised with tetramerous flowers [“**Fleurs quadrifides”].

Species no. 12 included in Cotyledon by Lamarck (1786), C. pinnata Lamarck (1786: 141) , is the present-day K. pinnata ( Figs 2C and D ). This species is often in the vernacular known as Goethe’s kalanchoe ( Goethe 1857: 161–162) and is well-known in mild-climate parts of the world for easily becoming established—even naturalised and invasive—in places well-beyond its natural geographical distribution range in Madagascar ( Smith & Figueiredo 2018a: 220, Smith et al. 2019: 263–268, Smith et al. 2021: 9–12, 18, Smith 2023b: 12). The species was eventually transferred to the genus Bryophyllum Salisbury (1805 : t. 3), as B. pinnatum (Lam.) Oken (1841: 1966) , but Bryophyllum is at present included in Kalanchoe , as K. subg. Bryophyllum (see Smith & Figueiredo 2018b: 169).

Species no. 14 included in Cotyledon by Lamarck (1786), as C. aegyptiaca Lamarck (1786: 142) , was the second new species of Kalanchoe described in that work. A combination for this species in Kalanchoe , as was the case for K. laciniata and K. crenata , was only published in the early 1800s, as K. aegyptiaca (Lam.) Candolle (1801 : t. 64). For the “Kalanchée d’Égypte”, referenced by Candolle (1801: t. 64) as “Cotylet d’Egypte. LAM. Dict. n. 14 [1786, species number 14 on p. 142]”, it was stated that: “Elle est originaire d’Egypte. On l’y cultive dans les jardins, où elle fleurit à la fin de l’hiver; on la nomme Vudni, Forskahl dit qu’elle est indigène du mont Melhân, où on lui donne le nom de Odejn.” [English: “It originates from Egypt. It is grown there in gardens, where it flowers at the end of winter; it is called Vudni. Forskahl [see under Cotyledon deficiens Forsskål (1775: 89) ] says it is indigenous to the Melhân Mountains, where it is called Odejn.” The description and plate of K. aegyptiaca immediately preceded the text and plate of K. spathulata Candolle (1801 : t. 65) that originated from China.

Figures

FIGURE 1. A. The plate of the “Eende-Plant, zoo als die van Rumphius genaamt werd” included in Rumphius (1747: “Tab. XCV” on the page facing p. 276). The material figured coincides well with what is today known as Kalanchoe laciniata with its laciniate leaf blades, shown in B. C. The material illustrated in Boerhaave (1720: [unnumbered] page facing p. 288) also corresponds with what today is known as K. laciniata. D. The leaves of K. laciniata are variously and virtually throughout deeply incised. E. The flowers of K. laciniata are yellow and quite small, which coincide with what Rumphius (1747: 276 and “Tab. XCV” on the facing page) described as “[…] gout-geel bloempie[s] […]”. All photographs, except 1A and 1C, by Gideon F. Smith.

FIGURE 2. A. The non-laciniate leaves of Kalanchoe crenata are crenate along the margin, as in the material illustrated in Plukenet (1692: t. 228, Figure 3). B. Several forms of K. crenata have orangey-yellow flowers, or, as described in “Plukn. Almag. p. 362. Tab. 228. Fig. 3.”, i.e., (Plukenet 1696), “golden-yellow flowers”. C. Kalanchoe pinnata, in the vernacular often known as Goethe’s kalanchoe, was, at first, in 1786, described as Cotyledon pinnata. D. Close-up of flowers of K. pinnata. All photographs by Gideon F. Smith.