identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
092F87C1064FDB0FFF28CFDA2E8AF84A.text	092F87C1064FDB0FFF28CFDA2E8AF84A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kalanchoe houghtonii Ward 2006	<div><p>1. Kalanchoe × houghtonii Ward (2006: 94)</p> <p>Homotypic synonym:— Bryophyllum × houghtonii (D. B. Ward) Forster (2006: 383). Type:— U.S.A., Florida, central Merritt Island, Brevard County, 10 February 2000, D. B. Ward 10700 [collected with R. B. Huck] (holotype, consisting of three parts, FLAS!; isotypes FTG, NY!, SD, US!, USF).</p> <p>Designations not validly published:—‘ Bryophyllum tubimontanum ’ Houghton (1935: 44); “ Kalanchoe × hybrida HORT. ”</p> <p>Taxonomic notes:—The first documented synthesis of material today known as K. × houghtonii (= K.daigremontiana × K. tubiflora) was when Arthur Duvernoix Houghton (1870–1938) recorded the hybrid as having been produced in the USA (Houghton 1935: 44). By 1954, i.e., within 20 years, material with the parentage K. daigremontiana and K. tubiflora was featured and illustrated in the first German edition of Jacobsen (1986: 615, Fig. 860) as “ Kalanchoe × hybrida HORT. ”, and also treated (but not illustrated) as “ K. cv. Hybrida” by Jacobsen (1977: 285). Whether the material that Jacobsen had access to originated from the laboratory of Resende, who created the hybrid in Lisbon, Portugal (Resende 1956: 241–242), or from Houghton in the USA, or from elsewhere, is not known.</p> <p>Material that was eventually described as K. × houghtonii has become successfully naturalised in Florida, USA, although it is not known whether the material so recorded originated from beyond Florida and was carried there, or whether the hybrid material arose in the state where the putative parents were growing socially (Ward 2006: 94). Kalanchoe × houghtonii has also become established and sometimes weedy in mild-climate regions on six continents (Smith 2019, Herrando-Moraira et al. 2020). We therefore deliberately clarify and elaborate on previous taxonomic treatments of this nothospecies (see for example Shaw 2008). In addition, some confusion has existed regarding the correct application of the name K. × houghtonii, with, for example, Maire (1976: 266–267) and Sajeva &amp; Costanzo (2000: 172, top left image) illustrating and describing material of this nothospecies under the name K. daigremontiana.</p> <p>Overall, we recognise four morphotypes in what are likely K. × houghtonii, two of which are of known synthetic origin, and another two seem to occur naturally and may represent evidence of gene flow and introgression between K. daigremontiana and K. tubiflora where they grow sympatrically in their natural habitat in Madagascar.</p> <p>A. Kalanchoe × houghtonii Morphotype A (Fig. 1A–C)</p> <p>Plants of Morphotype A have triangular-lanceolate leaf blades that are attenuate (when young) to subpeltate (when mature) basally, and large, inflated flowers reminiscent of those of K. tubiflora. The corolla is usually magenta, red in strong sun, rather than orange, and the leaves are much larger than those of Morphotype B (see [2.] below), so resulting in confusion between Morphotype A and K. daigremontiana (one of the parents of the nothospecies). A further similarity with K. daigremontiana is evidenced in the slower transition in leaf arrangement from decussate to tricussate, with plants remaining decussate for a longer period, before an alternate or tricussate arrangement can be observed (whereas K. tubiflora is tricussate). Leaves of plants of this group are generally only purplish brownmaculate abaxially and along the petiole.</p> <p>Plants of Morphotype A have been produced artificially more than once on different continents (continental Europe and the USA; see Resende 1956: 241–242 and Ward 2006, 2008), and at least in the USA they seem to be derived from one of two clones originally produced by Houghton, according to Shaw (2008: 22) (cited by Herrando-Moraira et al. 2020). However, Houghton (1935: 44) only presented a single clone of this formula, which, based on the single figure he provided, matches rather K. × houghtonii Morphotype B (see [2.] below). As noted by Shaw (2008: 22), K. × houghtonii was illustrated in Baldwin (1949: 344, Fig. 1); however, Baldwin (1949: 343) only reported the chromosome number of a single clone of the nothospecies (presumably the one illustrated), as being 2 n = 51. i.e., a triploid, given that the chromosome numbers earlier reported for K. daigremontiana and K. tubiflora [as K. verticillata Scott Elliot (1891: 14)] were 2 n = 34 (diploid) and 2 n = 68 (tetraploid), respectively (Baldwin 1938: 576). Baldwin (1938) did not report a chromosome count for the hybrid for which the name K. × houghtonii was eventually published. Note that some cytological variation was later found in K. daigremontiana, with chromosome numbers of 2 n = 34, 60, 68, 176 reported by Sharma &amp; Gosh (1967: 318–319). However, we were unable to verify the identity of the material investigated; the counts may have been based on artificially raised selections or on forms of the nothospecies, such as K. × houghtonii Morphotype A, which is often confused with K. daigremontiana as stated above.</p> <p>The plant investigated by Baldwin (1949) likely represents the material recorded by Houghton (1935: 44), i.e., K. × houghtonii Morphotype B (see [2.] below). Although Shaw (2008: 22) notes that the illustration provided by Baldwin (1949: 344) does not allow placement of the plant in a given cultivar of the nothospecies, we speculate that the triploid material investigated by Baldwin (1949) was derived from plants as figured in Houghton (1935: 44), i.e., K. × houghtonii Morphotype B. Separately, Resende (1956: 241–242) reported producing two hybrid races in K. × houghtonii; one sterile, an F 1 triploid, and the other fertile, F 1 and F 2 tetraploids. It is therefore likely that K. × houghtonii Morphotype A rather represents the tetraploid race of the nothospecies.</p> <p>Plants of K. × houghtonii Morphotype A are very invasive, so far naturalised in numerous mild-climate regions around the world, and due to their larger, K. daigremontiana -like leaves, produce significant quantities of bulbils along the margins. The cultivars K. × houghtonii ‘J.T. Baldwin’ (of which we consider K. × houghtonii ‘Jaws of Life’ to likely be a synonym) and K. × houghtonii ‘Garbí’, naturalised in Florida, USA, and Portugal and Spain, respectively (Shaw 2008, Guillot Ortiz et al. 2014, Smith et al. 2015), belong to this group. Photos of plants of K. × houghtonii Morphotype A can be found at http://sven-bernhard.de/anzeige_bilder_en.htm, under “ K. × houghtonii B_255-10-93-30”.</p> <p>Despite the fact that the one clone originally produced by Houghton very likely does not belong to Morphotype A, the type of K. × houghtonii clearly belongs to this Morphotype based on its size and leaf morphology. As stated above, it is this Morphotype that is typically found naturalised in various mild-climate regions, including in Florida, USA, from where the type was collected.</p> <p>B. Kalanchoe × houghtonii Morphotype B (Fig. 1E–F)</p> <p>Plants we regard as representative of Morphotype B have triangular-ovate leaf blades that are basally cuneate to subpeltate, and their large, inflated flowers are reminiscent of those of K. tubiflora. The corolla is usually orange rather than magenta, and the triangular-ovate leaves are much shorter than in Morphotypes A, C, and D, so giving the plants an overall smaller appearance when compared to the other Morphotypes, as well as both parents. Leaf arrangement transitions from decussate (as in K. daigremontiana) in younger plants, to tricussate (as in K. tubiflora) in mature plants. Leaves of plants of this group are adorned with darker purplish brown maculations both ab- and adaxially, and also along the petiole.</p> <p>Plants of K. × houghtonii Morphotype B so far have been only documented with certainty in cultivation, and based on the ovate leaf blades, which are barely longer than the petiole, as figured in Houghton (1935: 44), we suggest that it is likely that Morphotype B, specifically, the cultivar K. × houghtonii ‘Hybrida’ (Fig. 1E), was the one originally produced by Houghton. It is therefore likely that Morphotype B is the sterile triploid reported by Baldwin (1949: 343), who only reported the cytology of a single hybrid (but see Shaw 2008: 22).</p> <p>Other than K. × houghtonii ‘Hybrida’, the first cultivar in K. × houghtonii to be artificially produced and described, the variegated K. × houghtonii ‘Pink Butterflies’ (Fig. 1F), also belongs to K. × houghtonii Morphotype B and was likely vegetatively selected from K. × houghtonii ‘Hybrida’. Probable renamings of K. × houghtonii ‘Pink Butterflies’ include: K. × houghtonii ‘Pink Teeth’, K. × houghtonii ‘Pink Sparkler’, and K. × houghtonii ‘Fujicho’.</p> <p>Perhaps as a result of their sterility, but more likely because of their smaller leaves that inevitably produce fewer bulbils per leaf, plants of Morphotype B appear to be less invasive than those of Morphotype A, though material representative of Morphotype B has become naturalised in some places, for example in Spain.</p> <p>C. Kalanchoe × houghtonii Morphotype C (Fig. 1G)</p> <p>Plants representative of Morphotype C have linear leaf blades that are attenuate apically and basally, and the large, inflated flowers are reminiscent of those of K. tubiflora. The corolla is usually orange, and the plants transition relatively quickly from a decussate to a tricussate leaf arrangement. Plants of this Morphotype are therefore hardly distinguishable from K. tubiflora, especially when grown under strong solar irradiation, which results in the linear leaves becoming more terete rather than more lanceolate. Leaves of plants of this group are generally only purplish brown-maculate abaxially and along the petiole.</p> <p>To our knowledge, this Morphotype has yet to become naturalised successfully in foreign habitats, perhaps as a result of its leaf blade being shorter compared to its long petiole, bearing fewer bulbils per leaf. The cytology and fertility of material of Morphotype C are not known. Plants of this Morphotype exist in several Botanical Gardens that we visited, and also seem to occur naturally in Madagascar, for example along the Onilahy River, where the ranges of K. daigremontiana and K. tubiflora intersect [see for example https://www.inaturalist.org/observations/70546264]. It is not known if the naturally occurring plants of this Morphotype are transitional between the two parent species or whether they represent a natural hybrid between them. Photographic images of plants of this Morphotype in culture can be found at http://sven-bernhard.de/anzeige_bilder_en.htm, under “ K. × houghtonii HD_141070”. Due to their rarity in cultivation, no cultivar names exist for selections of K. × houghtonii Morphotype C.</p> <p>...Continued on the next page</p> <p>*When grown under the same conditions, no difference can be observed between the two cultivars.</p> <p>D. Introgressed K. daigremontiana / K. × houghtonii Morphotype D (Fig. 2; see also Smith &amp; Shtein 2021: 246, Fig. 2C)</p> <p>Plants representative of Morphotype D have apically long-attenuate, lanceolate leaf blades that are auriculate to peltate basally, and the small, funnel-shaped flowers are reminiscent of K. daigremontiana. The corolla is usually pink-purple, rather than magenta-red (as in Morphotype A) or orange (as in Morphotypes B and C), and the leaves are much larger in mature plants, resulting in confusion between material of Morphotype D and K. daigremontiana. A further similarity to K. daigremontiana is the leaf arrangement that does not transition to tricussate—in plants of Morphotype D leaf arrangement generally remains decussate (Smith &amp; Shtein 2021: 246, Fig. 2C) or becomes alternate as flowering approaches [https://www.inaturalist.org/observations/12919993]. Young plants of Morphotype D are generally very similar in appearance to representatives of Morphotype C in that they have linear, though almost entire, leaves, while mature plants are hardly distinguishable from K. daigremontiana, as a result of their large, often peltate, leaves.</p> <p>Considerable morphological variation exists in K. × houghtonii Morphotype D, with material having been observed that ranges from almost indistinguishable from K. daigremontiana (e.g., Fig. 2A–D), to plants more clearly transitional between K. daigremontiana and K. tubiflora, so highlighting the possibility that the material is indeed rather of hybrid origin [https://www.inaturalist.org/observations/12919993]; either K. × houghtonii or K. daigremontiana is variously applied to such transitional forms. To our knowledge, Morphotype D has yet to become naturalised successfully in foreign habitats, and the cytology and pollen and seed fertility of these plants are not known. Plants of K. × houghtonii Morphotype D exist in several Botanical Gardens that we visited, and also seem to occur naturally in Madagascar, for example in Toliara and along the Onilahy River, where the ranges of K. daigremontiana and K. tubiflora intersect [e.g., https://www.inaturalist.org/observations/70546286]. Again, it is not known if the plants of Morphotype D that occur naturally in Madagascar are transitional between the species (K. daigremontiana and K. tubiflora) or represent natural hybrids between them. Photographs of plants of Morphotype D in cultivation can be found at http://svenbernhard.de/anzeige_bilder_en.htm, under “ K. daigremontiana SSZ _96-2255-0”, a clone collected in 1996 from Toliara, Madagascar, by Dieter Supthut†, at the time of the Städtische Sukkulenten Sammlung, Zürich.</p> <p>Plants referable to K. × houghtonii Morphotype D seem to arise occasionally in cultivation, for example, ISI 2007- 25 K. ‘Parsel Tongue’ (Fig. 2E–H), a cultivar distributed by the International Succulent Institute through the Huntington Botanical Gardens, Pasadena, California, USA (Trager 2007). Kalanchoe ‘Parsel Tongue’ is morphologically very similar to K. daigremontiana, except for its leaves that are apically long-attenuate; the funneliform-peltate (rather than saddle-shaped-peltate) leaf blade base; the finely-maculate abaxial leaf blade colouration; and the alternate leaf arrangement when mature. It is possible that this cultivar represents a back-cross of K. daigremontiana with a fertile cultivar of K. × houghtonii, perhaps K. × houghtonii ‘J.T. Baldwin’.</p> <p>Summary:—Our studies and observations confirm that K. daigremontiana and K. tubiflora hybridise with ease, and even in their natural habitat some transitional or hybrid forms are present where the species grow sympatrically. Such forms that grow naturally in Madagascar are morphologically more diverse than the forms of K. × houghtonii that arose in cultivation. However, thus far only artificially created forms show strong invasive tendencies. Furthermore, many of the naturally occurring transitional or hybrid forms appear very similar to either K. daigremontiana or K. tubiflora, suggesting rather limited introgression. Nevertheless, where the natural geographical distribution ranges of K. daigremontiana and K. tubiflora overlap, several vegetative and reproductive morphological characters show considerable variation (Table 1).</p> </div>	http://treatment.plazi.org/id/092F87C1064FDB0FFF28CFDA2E8AF84A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shtein, Ronen;Smith, Gideon F.;Ikeda, Jun	Shtein, Ronen, Smith, Gideon F., Ikeda, Jun (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, DOI: 10.11646/phytotaxa.524.4.2
092F87C10647DB0BFF28CFB02C44F99F.text	092F87C10647DB0BFF28CFB02C44F99F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kalanchoe laetivirens , Descoings 1997	<div><p>2. Kalanchoe laetivirens Descoings (1997: 85).</p> <p>Treated as K. × laetivirens by Smith (2020g: 105) (Fig. 7).</p> <p>Homotypic synonym:— Bryophyllum laetivirens (Desc.) Byalt (2008: 463). Type:— MADAGASCAR. Environs de Toliara (Tulear), à quelques km au sud sur la route d’Anantsono (Saint Augustin), bord de chemin, plante échappé du jardin de l’auberge “La Mangrove”, 25 novembre 1994, [English: “ MADAGASCAR. Surroundings of Toliara (Tulear), a few km to the south on the road to d’Anantsono (Saint Augustine), roadside, plant escaped from the garden of the inn “La Mangrove”, 25 November 1994 ”] [B.M.] Descoings 28234 (holotype P †). Neotype:—Ex hort. South Africa. Gauteng Province, Pretoria, 17 May 2020, G.F. Smith 1107 (PRU), designated here.</p> <p>Notes on the type:—The holotype of the name K. laetivirens is a specimen, B.M. Descoings 28234, dated 25 November 1994, which Descoings (1997) stated to have deposited at Herb. P (Descoings 1997: 85). However, this specimen is not extant at that repository (Florian Jabbour, pers. comm.). Apart from the specimen B.M Descoings 28234, no other material that would qualify as original in the sense of Turland et al. (2018: Art. 9.4) could be traced. We accordingly here neotypify (see above) the name K. laetivirens on G.F. Smith 1107, a specimen held at Herb. PRU.</p> <p>The holotype of K. laetivirens was prepared from plants located a few km to the south of Toliara, Madagascar, on the road to Saint Augustine, which likely escaped from the garden of the inn “La Mangrove” (Descoings 1997: 85). The inn “La Mangrove” probably refers to the Sarodrano Hotel, located in a mangrove forest on a small peninsula near Saint Augustine. Indeed, as of 2018, K. laetivirens is still growing along the road adjacent to the peninsula just north of Saint Augustine, and is still being cultivated at the nearby lodge (pers. obs. by one of us, JI). The neotype material designated here was possibly derived from clonal material of the garden-based Arboretum d’Antsokay, Toliara. This garden-based population of K. laetivirens was reportedly originally derived from the Isalo Massif (see “Natural distribution”, below), and later sampled by O. Olsen and introduced into cultivation around the world as ISI 95-36, HBG 70004 (Shaw 2008: 25, and see also http://media.huntington.org/ISI/corrections.html). It is likely that the populations that escaped from the garden of the inn “La Mangrove” from which the holotype of the name K. laetivirens was prepared, were also propagated from populations cultivated in Toliara, and therefore originated from the same population that was later distributed globally.</p> <p>Vegetative reproduction through the copious number of leaf marginal bulbils is the most common reproductive strategy in this species and seed fertility is low. It is likely therefore that the material preserved as both the holotype † and as the neotype originated from the same natural population in the Isalo Massif, Madagascar (see below).</p> <p>Natural distribution:—Southern Madagascar, Isalo Massif (Fig. 9). At the time that Descoings (1997) described K. laetivirens he had not seen material of this species in the wild. Regarding the natural distribution of K. laetivirens, Descoings (1997: 85) stated the following: “ RÉPaRTITION: la localité d’origine se situe au col des Tapia, à environ 170 km au nord-est de Toliara sur la route d’Ihosy et Fianarantsoa, aux portes du massif gréseux de l’Isalo. Cette information nous a été donnée par Monsieur Hermann PETIGNAT qui cultive cette espèce à Toliara, dans son jardin botanique de l’Arboretum d’Antsokai. Les plantes vues à Saint Augustin proviennent de ce jardin. [English: “DISTRIBUTION: the place of origin is located at the Tapia Pass, about 170 km northeast of Toliara on the road to Ihosy and Fianarantsoa, at the entrance to the sandstone massif of Isalo. This information was given to us [B.M. Descoings] by Mr Hermann PETIGNAT who cultivates this species in Toliara, in his botanical garden in the Arboretum d’Antsokay. The plants seen in Saint Augustine [from where material deposited as the holotype was collected] come from this garden.”] One of us (JI) observed the species at the locality from where Mr Hermann Petignat reported it (Fig. 7A–B), as well as at the Arboretum d’Antsokay (Fig. 5A–C).</p> <p>Numerous recent observations additionally place K. laetivirens in populous or urban areas across Madagascar, specifically, in Ifaty [https://www.inaturalist.org/observations/56394567], and Manasoa [https:// www.inaturalist.org/observations/12919994] in southwestern Madagascar, as well as in Anjajavy [https://www. inaturalist.org/observations/2633188] in northwestern Madagascar, and Antsiranana [https://www.inaturalist.org/ observations/55629658] in northeastern Madagascar. These observations suggest that K. laetivirens may now be established, possibly as escapes from cultivation across Madagascar, well beyond the range communicated to Descoings by Hermann Petignat (Descoings 1997).</p> <p>Taxonomic notes:—The proposed connection between K. laetivirens and the hybrid with the formula K. daigremontiana × K. laxiflora, which was created and illustrated by Resende &amp; Viana (1965: 180, 195 [Fig. 35], 196 [Fig. 36]) and referred to as ‘ Bryophyllum × crenodaigremontianum ’, was mentioned in Shaw (2008: 25) and later investigated by Smith (2020g: 105). As noted in Smith (2020g), material of K. laetivirens and K. daigremontiana × K. laxiflora are indeed remarkably similar, especially in their broad, large, bright green leaves that, at least adaxially, are generally unmarked by any maculation or colour patterning other than towards the margins, as well as in the general size of mature leaves, the thick, herbaceous stems of the material, and the extensive inflorescences of the two entities.</p> <p>Nevertheless, several diagnostically significant differences can be observed between these plants (Smith 2020g: 99). To allow a better comparison of K. laetivirens with the hybrid K. daigremontiana × K. laxiflora created by Resende &amp; Viana (1965), one of us (RS) recreated this combination in cultivation in Israel (Fig. 10). In addition, we were able to access a similar hybrid, K. daigremontiana × K. fedtschenkoi Raymond-Hamet &amp; Perrier de la Bâthie (1915: 75), for examination. The latter material was created by Sheng Jian Lu and Hung I Lu, separately, in Taiwan (Fig. 11). Significant differences observed between K. laetivirens and the two hybrids mentioned above are:</p> <p>1. Resende and Viana’s hybrid (K. daigremontiana × K. laxiflora) is not constitutively bulbiliferous, whereas K. laetivirens is (see Garcês et al. 2007). Kalanchoe laetivirens is one of the most strongly phyllo-bulbiliferous species, producing many dozens of large-sized, many-foliated bulbils on each mature leaf. Bulbil production in K. laetivirens occurs on dedicated, well-developed pedestals that are spathulate in shape and down-curved towards the abaxial leaf blade surface to facilitate the placement, especially once severed from the pedestals, of the bulbils on, or at least closer to, the ground and therefore below the substantial dentations along the leaf blade margins, which are subducted by the bulbil pedestals. However, the hybrid of Resende and Viana is at most induced or semi-constitutively bulbiliferous, and pedestals and/or bulbil formation cannot be readily observed (Resende &amp; Viana 1965: Figs 35 &amp; 36). Observations of live F 1 material of K. daigremontiana × K. laxiflora and K. daigremontiana × K. fedtschenkoi shows that these combinations produce induced phyllo-bulbiliferous hybrids, that form very reduced, sessile, and indistinct pedestals after bulbil formation, and never ahead of it.</p> <p>2. The F 1 hybrids of K. daigremontiana × K. laxiflora and K. daigremontiana × K. fedtschenkoi that we observed, as well as K. laxiflora and K. fedtschenkoi, are perennials. In contrast, both K. daigremontiana and K. laetivirens are multi-annuals that only flower once in their life cycles, following the culmination of one to three years of vegetative growth. However, it is not clear whether Resende and Viana’s hybrid was perennial or (multi-)annual.</p> <p>3. Though impossible to tell from the illustrations provided by Resende and Viana, the live F 1 material we observed do not share the unique growth habit and phenological development of K. laetivirens (and of K. ×descoingsii, for that matter; see below). Kalanchoe laetivirens and hybrids derived from it are low-growing for most of their vegetative life span, forming wide, densely foliated, solitary pseudo-rosettes, with the leaves presenting especially thick, short petioles, and large, broad leaf blades. Then, during the year in which they will flower, the plants can increase in height for up to more than a factor of ×4, developing leaves bearing longer petioles and shorter, narrower leaf blades. The hybrids we observed, as well as K. laxiflora, and K. fedtschenkoi, gradually elongate in a manner more typical of Kalanchoe in general during their development, do not remain as low-growing rosettes for most of their vegetative stage, and often branch just prior to, or after, flowering.</p> <p>4. Kalanchoe laetivirens is unique among representatives of K. sect. Invasores by the basal portion of the stem being extremely wide. Furthermore, the lower leaves are distinctly thick-petiolate and amplexicaul, resulting in nodal scars that give the stems a quadrangular appearance. These traits are absent in Resende and Viana’s hybrid and the hybrids we observed, as well as in K. daigremontiana, K. laxiflora, and K. fedtschenkoi.</p> <p>5. In Resende and Viana’s hybrid and the ones with the same or a similar parentage that we observed, the leaves transition more gradually towards the peduncle when compared to those of K.laetivirens.Though also usually auriculate, in the hybrids the auricules are often shorter, especially in lower leaves. The basal leaves of K. laetivirens tend to also be extremely broad, more so than in the hybrid of Resende and Viana, and the hybrids we observed. Notably, K. laetivirens does not produce lobed leaves, whereas some of Resende’s F 2 hybrids are conspicuously trilobate (Resende &amp; Viana 1965: 196, Fig. 36). Moreover, we observed basally trilobate leaves, though only barely so, in F 1 hybrids of both K. daigremontiana × K. laxiflora and K. daigremontiana × K. fedtschenkoi, as well as in K. ×descoingsii (see below). This suggests that while the leaves of K. daigremontiana and K. laetivirens are not trilobate, their hybrids most often tend to be so, at least obscurely.</p> <p>6. Kalanchoe laetivirens further differs from the hybrid of Resende and Viana and the similar hybrids we observed in some flower traits, including colour. The calyx of K. laetivirens is particularly short and small, similar to that of K. daigremontiana and K. sanctula, whereas the calyx and calyx tube of the hybrids are intermediate between those of the larger ones of K. laxiflora and K. fedtschenkoi, and is therefore longer, with the calyx tube of the hybrids being especially elongated in comparison. The succulent calyx texture of K. laetivirens further differs from the flimsier calyx texture in the hybrids. Finally, K. laetivirens, K. daigremontiana, and K. sanctula flower in shades of pink-purple, often with whitish green infusion, but lacking orange pigmentation (Figs 7D and 8C–D, and as Smith 2020g: 102, 104 recorded and illustrated for K. laetivirens), whereas the flower colour of the hybrids we observed is often salmon- to deep red (Figs 10D and 11C–D), a combination of pink-purple with the orange flower colour of K. laxiflora and K. fedtschenkoi. Note though that some forms of K. laetivirens produce flowers with substantial white sections (Smith 2020g: Figs 5, 6, 8, and 9).</p> <p>We conclude that K. laetivirens not only differs from the hybrid of Resende and Viana and combinations similar to it in several diagnostic traits, but also possesses some unique traits among representatives of K. sect. Invasores, for example its unique phenology, thickened lower stem and leaves, as well as its calyx morphology (discussed above). Many of the traits present in K. laetivirens are not intermediate between any two related species.</p> <p>Given their multiple, specialised reproductive strategies, including the production of large numbers of bulbils on leaf margins, multi-annual representatives of K. sect. Invasores, including K. daigremontiana and K. laetivirens, do not rely exclusively on sexual reproduction to ensure the establishment of the next generation. Aspects of the reproductive biology of these species, such as the reduced viability of seed and the occurrence of a high frequency of deformed flowers (Smith 2020g, reported for K. laetivirens), also commonly occur in K. daigremontiana (Garcês et al. 2007, Herrera &amp; Nassar 2009, pers. obs. by the authors), and in K. sanctula. A decreased reliance on sexual reproduction, as opposed to vegetative reproduction, likely resulted in a reduction in the maintenance of uniform and viable fertile structures. However, the evolution of asexual reproduction in the genus, particularly in constitutively phyllo-bulbiliferous species, such as K. daigremontiana and K. laetivirens, is not yet fully understood. Regardless, as noted by Smith (2020g: 97) and Smith &amp; Shtein (2021: 247, Fig. 3D) for K. daigremontiana and as shown in this paper for K. laetivirens, both K. daigremontiana and K. laetivirens do produce at least some viable seed, and ample interspecific hybrids.</p> <p>The type specimen of K. laetivirens was collected as a garden escape and not in the wild (Smith 2020g: 100). However, Descoings (1997: 85) was informed that the species originated from the Isalo Massif (see ‘Natural distribution’ of K. laetivirens, above), where one of us (JI) photographed it (Fig. 7A–B). The natural distribution range of K. laetivirens is therefore not known to be sympatric with those of either K. daigremontiana (see below regarding the occurrence of K. daigremontiana at “Isalo”.) or K. laxiflora, two species offered as possible parents of K. × laetivirens (Smith 2020g: 105). Note though that hybridisation between species grown in a garden is a common occurrence (see ‘Introduction’ for references). While both K. daigremontiana and K. laetivirens occur naturally in southwestern Madagascar, the original habitat of K. laetivirens in the Isalo Massif, as reported to Descoings (1997: 85), is over 100 km away from the closest recorded K. daigremontiana occurrence to the south, the type locality of the latter species, the Androhiboalava Mountain, near Benenitra, as well as from the Marosavoha Mountain in the same region (Cristini 2020), and over 150 km away from the closest recorded occurrence of K. daigremontiana to the north, in the Makay Massif (Smith &amp; Shtein 2021: 246, Fig. 2A)—three out of four occurrences mentioned by Raymond-Hamet &amp; Perrier de la Bâthie (1914: 131) in the protologue of K. daigremontiana. A fourth occurrence observed by Perrier de la Bâthie, in the Isalo region, was also mentioned. However, no exsiccata or recent photographic evidence or other observations exist to corroborate this (see Supplementary file). It is possible that Perrier de la Bâthie (in Raymond-Hamet &amp; Perrier de la Bâthie 1914) might in fact have referred to K. laetivirens, which was then still undescribed, when he documented this locality for K. daigremontiana, or that the latter species is indeed also present there (Fig. 9). In contrast, K. laxiflora is only known to naturally occur in east-central and southeastern Madagascar, though some forms of the related species, K. fedtschenkoi, do occur in the Isalo Massif.</p> <p>These morphological and geographical considerations suggest that K. laetivirens should be treated as an accepted species, closely related to K. daigremontiana and K. sanctula.</p> </div>	http://treatment.plazi.org/id/092F87C10647DB0BFF28CFB02C44F99F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shtein, Ronen;Smith, Gideon F.;Ikeda, Jun	Shtein, Ronen, Smith, Gideon F., Ikeda, Jun (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, DOI: 10.11646/phytotaxa.524.4.2
092F87C10640DB08FF28C94C2B58FE1E.text	092F87C10640DB08FF28C94C2B58FE1E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kalanchoe sanctula Descoings 1997	<div><p>3. Kalanchoe sanctula Descoings (1997: 87) (Fig. 8).</p> <p>Homotypic synonym:— Bryophyllum sanctulum (Desc.) Byalt (2008: 464). Type:— MADAGASCAR. Sud-est, région de Tôlanara (Fort Dauphin), environs d’Amboasary, route d’Ambovombé, collines boisées, sol léger, en exposition éclairée, 13 novembre 1994, [English: “ MADAGASCAR. South-east, Taolagnaro region, surroundings of Amboasary, Ambovombé road, wooded hills, light soil, illuminated exposure, 13 November 1994 ”] [B.M.] Descoings 28180 (holotype P †). Neotype:— Clonally derived material from Descoings 28180, first cultivated in France, by Bernard Descoings, then by Philippe Richaud, then cultivated and collected for preservation in Israel, R. Shtein 832 (neotype TELA [TELA937]; isoneotype TELA [TELA938]), designated here.</p> <p>Notes on the type:—The holotype of the name K. sanctula is a specimen, B.M. Descoings 28180, dated 13 November 1994, which Descoings stated to have deposited at Herb. P (Descoings 1997: 87). However, as is the case with the type specimen of the name K. laetivirens, this specimen is not extant at that repository (Florian Jabbour, pers. comm.). Apart from the specimen B.M. Descoings 28180, no other material that would qualify as original in the sense of Turland et al. (2018: Art. 9.4) could be traced. We accordingly here neotypify (see above) the name K. sanctula on R. Shtein 832, a specimen held at Herb. TELA. This specimen was clonally derived from the original gathering from which the holotype was prepared.</p> <p>Unlike K. laetivirens, K. sanctula so far remains rare in cultivation, possibly because it has not been offered for sale or exchange through large-scale international horticultural introduction programmes. As a result, K. sanctula has yet to be reported as having successfully colonised locations outside Madagascar.</p> <p>Natural distribution:—Southern Madagascar, region of Amboasary (Fig. 9).</p></div> 	http://treatment.plazi.org/id/092F87C10640DB08FF28C94C2B58FE1E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shtein, Ronen;Smith, Gideon F.;Ikeda, Jun	Shtein, Ronen, Smith, Gideon F., Ikeda, Jun (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, DOI: 10.11646/phytotaxa.524.4.2
092F87C10643DB17FF28CD032A2FFE3A.text	092F87C10643DB17FF28CD032A2FFE3A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kalanchoe Adanson 1763	<div><p>Kalanchoe ×descoingsii Shtein, Gideon F.Sm. &amp; J.Ikeda sp. nov. (Figs 3 and 4).</p> <p>Type:— MADAGASCAR. Arboretum d’Antsokay, southeastern Madagascar, in a sparse forest floor, collected on 04 October 2018 by Jun Ikeda s.n., R. Shtein 501 (holotype TELA [TELA935]; isotype TELA [TELA936]). Material harvested as the holo- and isotypes was cultivated in Israel (Figs 3 and 4).</p> <p>Suggested parentage:— Kalanchoe laetivirens Descoings (1997: 85) × K. tubiflora (Harvey 1862: 380) Hamet (1912: 44).</p> <p>Diagnosis:— Kalanchoe × descoingsii differs from all other representatives of K. subg. Bryophyllum by being lowgrowing for most of its vegetative life span; by its densely foliated, solitary pseudo-rosettes that drastically elongate as flowering approaches; by being constitutively phyllo-bulbiliferous; by its oblong leaves that are apically obtuse to rounded; by having deeply dentate leaf blade margins subducted by recurved, spathulate bulbil-carrying pedestals; by its distal leaves and peduncular leaves gradually transitioning from conspicuously petiolate, deeply auriculate, basally broadened, oblong basal leaves, into being subsessile, attenuate, entire bar the apex, and generally not bulbil-forming; by its peduncle indistinctly transitioning from the distal vegetative part of the stem into the flower-bearing portion of inflorescence; by the corolla that is somewhat pinkish to whitish green when emerging, to being predominantly vibrant pink to orange-pink at anthesis; by the free sepal segments being longer than wide, ovate-deltoid, and about as long as to somewhat longer than the fused portion; by the calyx tube being wider than long; and by having nectar scales that are about as wide as long, spreading, and rounded-notched at the apex.</p> <p>Description:— Plants multi-annual, entirely glabrous, non-glaucescent throughout, unbranched, erect, often basally decumbent, about 0.8–1.2 m tall when in flower. Stem mostly herbaceous, lenticular, cylindrical; lower portion 25–35 cm long, 10–13 mm in diam., dense, green to pinkish brown; upper portion 50–80 cm long, 5–8 mm in diam., extremely sparse, purple; leaf scars narrowly crescentiform, conspicuous, slightly protruding, not connate; internodes 0.7–1.5 cm long basally, 4.5–6.0 cm long in distal half, distinctly lighter green to pink. Leaves opposite, decussate, succulent; basal leaves distinctly petiolate, densely arranged, wilting by anthesis; petiole 3.0– 5.5 cm long, 4–6 mm in diam. medially, up to 6–8 mm in diam. at base, subcylindrical, often slightly concave above and convex below, slightly widened around stem, green to purple, distinctly maculate with large dark brown or dark purple blotches; blade 8–12 cm long, 3.5–5.8 cm wide at base, 2.0– 3.5 cm wide elsewhere, adaxially green to green-brown, sometimes ornate with purple veining patterns centrally and basally, minutely dark-maculate, abaxially yellowish grey, strongly striped in dark brown or dark purple, narrowly elliptic to oblong, sometimes basally obtusely trilobate, widened distinctly in basal &lt;⅓, widest at basal ¼, strongly guttered lengthwise, concave above and convex below; margins green-grey to reddish pink, regularly dentate, darkly blotched between dentations, below blotches marked by pale organogenic and/or embryogenic notches, later developing pedestals; dentations 2.0–5.5 × 3–6 mm, acute, broad, increasing in acuity basally and apically; bulbil production fully constitutive, occurs on dedicated pedestals, at least on apical ½ of each leaf blade margin or on its entirety; bulbils large, bearing 3–5 leaf pairs and sometimes their own bulbils while still attached to mother plant, almost round leafed, minutely maculate; pedestals 2.0–5.5 × ± 1.5 mm, spathulate, bent to abaxial blade surface, incurved apically; base strongly auriculate, extending up to 17 mm behind petiole-blade connection, often ornate with two erect, dentate, bulbiliferous auricles; apex obtuse to rounded; apical leaves petiolate to subsessile; petiole 1.5–2.5 cm long; blade 5.0–7.5 × 0.8–1.2 cm, equally wide throughout to widest apically, narrowly elliptic-oblanceolate, mostly entire, few-dentate apically, not pedestalate, apically bluntly obtuse to rounded, basally minutely auriculate-cuneate to attenuate. Inflorescence a compact, many-flowered, capitate thyrse, lacking leaves or barely leafed at anthesis, non-bulbiliferous, i.e., dense bulbil clusters do not develop, but a few, large, discrete plantlets may form post-flowering, dense, terminal, 7.5–15.0 cm wide, consisting of 5–7 short dichasial cymes, each 5–9 cm long; peduncle 14–21 cm long, 4–6 mm in diam., single, cylindrical, indistinctly transitioning from distal vegetative part of stem into flower-bearing portion of inflorescence; bracts 3–5 × ± 1 cm, sessile, attenuate, narrowly oblong, mostly entire, few-dentate and bluntly obtuse apically, wilting soon; floral bracts ± 2 × ± 1 cm, bract-like, wilting soon; pedicels 12–18(–25) mm long, curved, cylindrical, tapering towards flowers from 1.2–1.4 mm to ± 0.60–0.75 mm in diam., purple. Flowers numerous, pendent, campanulate. Calyx 12–15 × 7–8 mm, light or grey-green to purple, dark purple-spotted where free sepal segments fuse, succulent, thick, campanulate-urceolate, rounded, basally ± flat, widest in middle of tube, then gradually constricted upwards; tube 5.5–6.5 mm long; free sepal segments 6.5–8.0 mm long, widest at base at ± 5 mm, ovate-deltoid, very acute, apiculate, adpressed to corolla to slightly spreading. Corolla 30–33 mm long, vibrant pink to orange-pink, whitish green when emerging and where obscured by calyx, non-stipitate; tube 22–24 mm long, especially basally ± 4-angled, basally adpressed around and above carpels, up to 5.5 mm wide, then diminishing to ± 4 mm wide at its thinnest, then gradually inflating to base of petals where it is generally widest at a width of 7–8 mm; lobes 8.0–9.5 × 6–7 mm [at base], widest at 8–9 mm at ⅓ from tip, broadly obovate, apically rounded, very slightly apiculate, somewhat spreading. Stamens 8, inserted low down in corolla tube, at ± level of carpels, slightly exserted, visible between, and reaching apex of petals; filaments of two similar lengths, fused basally, then adpressed, protruding inside base of corolla, free higher up for ± 20–22 mm, thin, yellow-green basally and apically, strongly infused with purple-pink elsewhere; anthers ± 1.0–1.2 × 0.8 mm, dark grey, producing yellow pollen, ovate-cordate, somewhat longer than wide. Pistil consisting of 4 carpels; carpels ± 7–8 × 1.7–2.0 mm, convergent, fused basally, free above, uniformly dark green, conical lanceolate, widest in lower ⅓, cuneate towards styles; styles ± 15–17 mm long, uniformly yellowish green; stigmas very minutely capitate, dark green; scales ± 1.5–2.0 × 1.5–2.0 mm, yellow-green, free, spreading, rounded-square, about as wide as long, thickened especially basally, incurved, upper corners rounded, emarginate. Follicles not seen. Seeds not seen. Chromosome number: 2 n = unknown.</p> <p>Eponymy:— Kalanchoe ×descoingsii commemorates Bernard Marie Descoings (born Paris, France, 7 September 1931 –died Largentière, Ardèche, France, 23 October 2018) (Madin 2018). He studied agronomy at Montpellier and in 1953 graduated as an agricultural engineer. From 1953 to 1954 he completed a training course at the Phanerogam Laboratory of the Muséum national d’Histoire naturelle, in Paris, where, under the guidance of Jean Henri Humbert, he developed an interest in tropical botany. Descoings (Fig. 12) subsequently spent time in Madagascar (1954–1958) and after having been recruited by the ‘Office de la Recherche Scientifique et Technique Outre-Mer’ (ORSTOM), spent four years in the Congo, where he directed ORSTOM’s Botanical Laboratory from March 1960 to 1965. Between 1965 and 1985 Descoings was variously based in Montpellier, French Guiana, and Burkina Faso, and from 1970 until his retirement in 1995 he additionally held various positions in Montpellier, France. In 1976 he defended his doctoral dissertation, ‘Approche des formations herbeuses tropicales par la structure de la végétation’, in Natural Sciences in the Faculty of Sciences of the University of Montpellier. While Descoings was based at the Paris Herbarium as a young man, the exposure he had to Jean Henri Humbert likely sparked his interest in Kalanchoe, as Humbert himself had more than a passing interest in the genus (Humbert 1933a, b). From the late 1990s to mid-2000s, Descoings described 16 new species and nothospecies in the genus from Madagascar, including K. laetivirens and K. sanctula, both reviewed in this paper. In 2003 Descoings treated Kalanchoe in the Crassulaceae volume of the Illustrated Handbook of Succulent Plants project (Eggli 2003), which in several respects does not coincide taxonomically with the earlier treatment of Boiteau &amp; Allorge-Boiteau (1995) of the genus as found in Madagascar. In 2005, Descoings paid special attention to the nothospecies of southern Madagascar, being the first to recognise their relatively common occurrence (Descoings 2005a). A year later he published an infrageneric classification that proposed the recognition of a morphologically diverse K. subg. Calophygia Descoings (2006: 24), but not K. subg. Kitchingia, a view with which we do not agree. Smith (2019) was dedicated to Descoings.</p> <p>Distribution:—Southern Madagascar, Arboretum d’Antsokay (Fig. 9). Mr Andry Petignat, Hermann Petignat’s son, an author and the current owner of the Arboretum d’Antsokay informed us that K. × descoingsii was not imported to his garden and therefore most likely spontaneously arose there. He further noted the similarities this plant bears to K. laetivirens, especially in habit and colouration, and that it rarely flowers, remaining rather short, though it has successfully spread in the garden, presumably vegetatively.</p> <p>Kalanchoe tubiflora and K. laetivirens, the parents of K. × descoingsii, are not known to grow sympatrically in the natural distribution range of K. laetivirens (discussed above in detail). In cultivation and in the Arboretum d’Antsokay, K. × descoingsii is able to rapidly reach its bulbil producing phase and spreads easily at a rate on par with that of K. × houghtonii. Detached bulbils may start bulbil production on their fourth leaf pair, or even carry bulbils of their own while still attached to the mother plant. Kalanchoe × descoingsii therefore poses a high invasiveness potential.</p> </div>	http://treatment.plazi.org/id/092F87C10643DB17FF28CD032A2FFE3A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shtein, Ronen;Smith, Gideon F.;Ikeda, Jun	Shtein, Ronen, Smith, Gideon F., Ikeda, Jun (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, DOI: 10.11646/phytotaxa.524.4.2
