Kalanchoe darainensis (Crassulaceae), a new species from northeastern Madagascar

Abstract

A new species of small, rosulate and upright-flowered Kalanchoe Adans. (Crassulaceae) from northeastern Madagascar
is described and illustrated: Kalanchoe darainensis D.-P. Klein & Callm. Morphologically, Kalanchoe darainensis is most
similar to Kalanchoe blossfeldiana Poelln. and Kalanchoe globulifera H. Perrier which are known from the northwestern
Tsaratanana Massif, c. 100 km southwest of Daraina. Detailed notes on the habitat and ecology of Kalanchoe darainensis
are provided, as well as a short taxonomical overview of allied species of the Kalanchoe subg. Kalanchoe from Madagascar,
including an identification key to those species. Despite its restricted distribution in the protected Antsahabe massif, the
new species is preliminary assessed as “Least concern” [LC] using the IUCN Red List Criteria.

Full text

Kalanchoe darainensis (Crassulaceae),
a new species from northeastern
Madagascar
David-Paul Klein, Ronen Shtein, Louis Nusbaumer & Martin W. Callmander
Abstract
KLEIN, D.P., R. SHTEIN, L. NUSBAUMER & M.W. CALLMANDER (). Kalanchoe darainensis (Crassulaceae), a new species from
northeastern Madagascar. Candollea :  – . In English, English and French abstracts. DOI: http://dx.doi.org/./cva
A new species of small, rosulate and upright-owered Kalanchoe Adans. (Crassulaceae) from northeastern Madagascar
is described and illustrated: Kalanchoe darainensis D.-P. Klein & Callm. Morphologically, Kalanchoe darainensis is most
similar to Kalanchoe blossfeldiana Poelln. and Kalanchoe globulifera H. Perrier which are known from the northwestern
Tsaratanana Massif, c.km southwest of Daraina. Detailed notes on the habitat and ecology of Kalanchoe darainensis
are provided, as well as a short taxonomical overview of allied species of the Kalanchoe subg. Kalanchoe from Madagascar,
including an identication key to those species. Despite its restricted distribution in the protected Antsahabe massif, the
new species is preliminary assessed as “Least concern” [LC] using the IUCN Red List Criteria.
Résumé
KLEIN, D.P., R. SHTEIN, L. NUSBAUMER & M.W. CALLMANDER (). Kalanchoe darainensis (Crassulaceae), une nouvelle espèce du
nord-est de Madagascar. Candollea :  – . En anglais, résumés anglais et français. DOI: http://dx.doi.org/./cva
Une nouvelle espèce de Kalanchoe Adans. (Crassulaceae), petite, rosulée et à eurs dressées du nord-est de Madagascar
est décrite et illustrée: Kalanchoe darainensis D.-P. Klein et Callm. Morphologiquement, Kalanchoe darainensis est plus
proche de Kalanchoe blossfeldiana Poelln. et Kalanchoe globulifera H. Perrier qui sont connues du nord-ouest du massif de
Tsaratanana, c.km au sud-ouest de Daraina. Des notes détaillées sur l’habitat et l’écologie de Kalanchoe darainensis
sont fournies, ainsi qu’un bref aperçu taxonomique des espèces alliées de Kalanchoe subg. Kalanchoe de Madagascar, y
compris une clé d’identication de ces espèces. Malgré sa distribution restreinte dans le massif protégé d’Antsahabe,
la nouvelle espèce est préalablement évaluée comme «Préoccupation mineure» [LC] en utilisant les critères de la Liste
Rouge de l’IUCN.
Keywords
CRASSULACEAE – Kalanchoe – Madagascar – Daraina – Loky-Manambato – New species
Addresses of the authors:
DPK: Humboldt-Universität zu Berlin, Späthstrasse /, Berlin D-, Germany. E-mail: david-paul.klein@agrar.hu-berlin.de
RS: e Steinhardt Museum of Natural History, Tel Aviv University, Klausner St , Tel Aviv-Yafo, Israel.
LN, MWC: Conservatoire et Jardin botaniques de Genève, ch. de l’Impératrice , C.P. ,  Chambésy, Switzerland.
Submitted on December , . Accepted on February , . First published online on March , .
© CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE ISSN : - – Online ISSN : - – Candollea () :  –  ()

Introduction
The genus Kalanchoe Adans. (Crassulaceae, Kalanchooideae)
comprises more than  species known to date and occurs
in Madagascar, Africa, Arabia and tropical Asia. Currently
species and nothospecies are recognized in Madagas-
car, all but one of which are endemic to the Great Island
(M C, ). e genus of Kalanchoe
consists of three subgenera: the autonymic Kalanchoe subg.
Kalanchoe which is known from Madagascar, Africa, Arabia
and Asia, as well as Kalanchoe subg. Bryophyllum (Salisb.)
Koorders and Kalanchoe subg. Kitchingia (Baker) Gideon F.
Sm. & Figueiredo, both endemic to Madagascar (S &
F, ).
Kalanchoe subg. Kalanchoe includes species that share the
following characters: plants annual or perennial, herbaceous or
woody-arborescent, calyx unfused for most of its length with
a short to indistinct tube, sepal segments often free, usually
adpressed to the corolla tube, laments inserted ±above the
middle of the corolla tube, rarely below, owers typically erect,
rarely omni-directional or pendent, leaves and inorescences
never bulbiliferous, carpels convergent, usually much longer
than the styles, scales elongated to linear, with a length/width
ratio usually >, anthers included in corolla-tube or very
slightly exserted, often dressed with a ±spherical connective
gland on the anthers (B & A-B, ;
D, ; S & F, ).
Among the Malagasy representatives of Kalanchoe subg.
Kalanchoe, several morphologically similar species were his-
torically placed in “Group ” of R-H () to
so emphasize the morphological similarities they share with
species indigenous to mainland Africa, in contrast to similari-
ties shared with other species from Madagascar. ese species
are K.boisii Raym.-Hamet & H. Perrier, K.briquetii Raym.-
Hamet, K.chapototii Raym.-Hamet, K.globulifera H. Perrier
and K.globulifera var. coccinea H. Perrier (K.blossfeldiana
Poelln.). Later, B & A-B () and
A-B () proposed to further subdivide
these species into the two informal groups “Occidentales” and
“Globuliferae”, along with K. aromatica H. Perrier, K. bouvetii
Raym.-Hamet & H. Perrier and K. lanceolata (Forssk.) Pers.
e group “Globuliferae” was dened based on the spherical
connective glands (“globules”) on the anthers, whereas species
placed in “Occidentales” have terminal inflorescences, are
glandular pilose (except K. blossfeldiana) and do not have the
aforementioned connective glands. However, they added that
the latter group “ne présente pas beaucoup d’homogénéité”
[do not present much homogeneity] (B & A-
B, : ). ey also underlined the morphological
anity of both groups to some continental African species, as
opposed to other Malagasy representatives of Kalanchoe subg.
Kalanchoe, such as the species they placed in the informal
“Lanigerae” group.
Excluding Kalanchoe aromatica and K. bouvetii, that have
omni directional, zygomorphic flowers and distinct corolla
tubes, we consider the species of both informal groups, “Occi-
dentales” and “Globuliferae”, to form a common species group.
ese are small statured shrubs to medium sized herbs, usually
at least partially covered with a glandular indumentum, that
have erect, actinomorphic owers, highly reduced to indistinct
calyx tubes, long, deltoid to narrowly lanceolate sepals, apically
acute to acuminate and linear to liform nectary scales. Two
further recently described species, K. antennifera Desc. and K.
pareikiana Desc. & Lavranos can also be placed in this species
group, based on their morphology. Kalanchoe antennifera origi-
nates from the vicinity of Antsalova in the Tsingy de Bema-
hara in western Madagascar (Mangelsdorff, pers. comm.).
e precise type locality was unknown to D ()
who stated “Africa” in the protologue. Moreover, despite its
occurrence in Arabia, virtually across continental Africa, and in
Madagascar, the exact natural geographical distribution range
of K. lanceolata remains open to conjecture. Kalanchoe lanceolata
rather shares character expressions with species of Kalanchoe
subg. Kalanchoe from beyond Madagascar and accordingly we
refrain from placing it in this otherwise Malagasy group.
During inventories of the Daraina region (now called
Loky-Manambato, see P et al., ), four speci-
mens of small, rosulate, glandular pilose, succulent plants
were collected in the massif of Antsahabe (Fig.). These
specimens have the erect owers, linear to liform nectary
scales, apically inserted laments and reduced calyx tube that
are found in representatives of Kalanchoe subg. Kalanchoe, and
were all originally determined as representing K. pumila Baker.
However, K.pumila diers from the collected specimens by
being taller (
– cm) and oury-pruinose, by lacking an
indumentum, and by having large, pink-coloured, campanulate
corolla tubes (B, ; B & A-B,
118 – A new species of Kalanchoe (Crassulaceae) from Madagascar Candollea 76, 2021
Antananarivo
13.3°S 13.3°S
13.0°S
49.5°E 49.8°E
49.8°E 50.0°E
50.0°E
Kalanchoe darainensis
Natural Peak
Village
Capital
Loky-Manambato
Road RN5a
N
Maromokotra
Daraina
1171 m
1138 m
1099 m
TYPE
0 6 12 km
Fig. 1. – Distribution map of Kalanchoe darainensis D.-P. Klein & Callm.
[map derived from open data sources: https://earthdata.nasa.gov,
OpenStreetMap® and M & S 

). It is here demonstrated that these collections from the
Daraina region are unique among all other known Kalanchoe
and are described below as a new species: K. darainensis D.-P.
Klein & Callm. Kalanchoe darainensis is most similar to K.
blossfeldiana and K.globulifera, and represents yet another
Malagasy member of Kalanchoe subg. Kalanchoe with strong
similarities to continental African species.
Key to Kalanchoe darainensis
and allied species
. Plants perennial, soboliferous; indumentum absent to
partial; leaves ovate-oblong, obovate or orbicular, never
lobed; owers golden yellow or scarlet red .................. 
a. Plants predominantly annual, non-soboliferous; indumen-
tum partial to complete; leaves ovate-lanceolate to linear,
often lobed, rarely bi-lobed; owers whitish to golden
yellow .......................................................................... 
. Flowers yellow, few ( to ); leaves subsessile; indumen-
tum sparsely glandular, restricted to stem and peduncle .
................................................................. K. globulifera
a. Flowers scarlet, numerous; leaves petiolate to subses-
sile; indumentum absent or partially densely glandular,
restricted to leaves, stem and peduncle ........................ 
. Indumentum absent; leaf blades ovate-oblong, apically
obtuse to acute; inorescences numerous, terminal and
axillary, many-owered thyrses ............. K. blossfeldiana
a. Indumentum densely glandular, restricted to leaves, stem
and peduncle; leaf blades ovate, elliptic to orbicular, api-
cally rounded-obtuse; inflorescences single, terminal,
many-owered dichasial cymes .............. K. darainensis
. Sepals and corolla especially thick and eshy; glandular
hair thick-stalked, extends to the upper part of the stem,
young leaves, inorescence, and sepals; owers whitish
cream-yellow ........................................... K. pareikiana
a. Sepals and corolla not thick or fleshy; glandular hair
slender-stalked, complete, i.e. extends from the base of
the stem to the petals; owers golden yellow ............... 
. Leaves alternate basally, decussate higher up; petals sub-
oblong, retuse, i.e. rounded-obtuse and notched; laments
.
– .mm in length, free for >⅓ of their length, papil-
lose; anthers fully exserted, wider than long, lacking con-
nective glands ............................................. K. briquetii
a. Leaves decussate; petals ovate, apically acute to acumi-
nate; laments 
– mm in length, free for < of their
length, non-papillose; anthers partially to fully included,
longer than wide, dressed with ±spherical connective
glands .......................................................................... 
. Plants perennial; leaves - to -partite, occasionally bi-
lobed and petiolulate; leaf or leaflet blades oblong to
linear; margins obtusely crenate-sinuate; petals strongly
acuminate with an acumen .
– .mm long; styles much
longer than carpels .................................... K. chapototii
a. Plants annual; leaves - to -partite, if lobed not bi-lobed
nor petiolulate; leaf or leaet blades ovate to lanceolate;
margins entire to minutely crenate-sinuate; petals acumi-
nate with an acumen .
– mm long; styles much shorter
than carpels ................................................................. 
. Leaves unlobed; leaf or leaet blades ovate, apically sub-
obtuse; styles > times shorter than carpel ...... K. boisii
a. Leaves -partite; leaf or leaet blades ovate-lanceolate,
apically acuminate-acute; styles < times shorter than
carpel ...................................................... K. antennifera
Taxonomy
Kalanchoe darainensis D.-P. Klein & Callm., sp. nov. (Fig.,
).
Holotypus: Madagascar. Reg. SAVA [Prov.
Antsiranana]: Daraina, forêt d’Antsahabe, [°'S
°'E], m, .IV., fl., Ranirison 749
(G[G]!; iso-: TEF image!, P).
Kalanchoe darainensis D.-P. Klein & Callm. differs from
other representatives of Kalanchoe subg. Kalanchoe by its dense
indumentum of medium to long glandular hairs, covering all
parts of the plant except owers and bracteoles, its basally near-
ligneous stems, soboliferous above, developing a single, erect,
terminal inflorescence per rosette, rarely two, with a single
peduncle each, composed in the apical half of a single node, its
indistinct calyx tube, narrowly lanceolate and acute sepals,
a corolla tube gradually constricted above to a subcylindrical
throat in the apical half, with scarlet red, erect owers and
anthers dressed with attened spheroid connective glands.
Plants small, perennial, rosulate and succulent, .
– .cm
high, mostly erect, creeping to decumbent at base, sobolifer-
ous, bearing one to few lateral basal shoots, almost entirely
densely covered with medium to long glandular hairs. Stems
.
– .mm in diameter, terete, basally near-ligneous and
without indumentum when mature, bearing roots and lateral
shoots, densely covered with medium (.
– mm) white-
brownish glandular hairs. Leaves opposite, decussate, succu-
lent, light bright green to olive with slightly reddish margins,
arising from a rosette bearing up to  pairs of leaves, entirely
covered with whitish to brownish, medium (.mm) to long
(.mm) glandular hairs, long-petiolate to subsessile; petioles
.
– .×.
– mm, subcylindrical, somewhat flattened
above, convex below, broadening towards the leaf base, densely
covered with whitish to brownish, short (.mm) to medium
(mm) glandular hairs. Leaf blades 
– ×
– mm, ovate,
elliptic to orbicular, sometimes wider than long, adaxially
canaliculate along the midrib, densely covered with long (up
Candollea 76, 2021 A new species of Kalanchoe (Crassulaceae) from Madagascar – 119

120 – A new species of Kalanchoe (Crassulaceae) from Madagascar Candollea 76, 2021
Fig. 2. – Kalanchoe darainensis D.-P. Klein & Callm. A. Habit; B. Basal leaf; C. Dissection of corolla, showing androecium and gynoecium;
D. Flower; E. Inflorescence; F. Anther with flattened spheroid connective gland.
Ranirison 749, TEF Drawings: R.L. Andriamiarisoa]
1cm
1cm
1mm
5mm
0.6mm
1mm
A
B
C
D
E
F

to .mm) whitish glandular hairs; tip obtuse-rounded; base
obtuse to ±rounded or near-truncate, often oblique; margins
irregularly wide-sinuate to wide-crenate. Inorescences a few- to
many-owered dichasial cyme with up to  secondary branches
bearing up to  owers, erect, typically one terminal inores-
cence per rosette, rarely two, sparsely covered with medium (up
to .mm) whitish-brownish glandular hairs up to the second
node of the inorescence; peduncle 
– ×.
– .mm,
terete, thinning apically, bearing 
– few pairs of peduncular
bracts, apical half composed of a single node, densely covered
with medium (up to .mm) whitish-brownish glandular hairs
peduncular bracts .
– ×.
– mm, borne in the lower third
or middle of the peduncle, subsessile, densely covered with
medium (up to .mm) whitish-brownish glandular hairs,
tip obtuse, base cuneate, margins irregularly sinuate to crenate.
Flowers 
– ×.
– mm, erect. Bracts .
– .×.
– .mm,
sessile, lanceolate, slightly incurved, apically acute, sparsely
glandular hairy. Pedicels 
– .×.
– .mm, erect, widen-
ing towards the flower. Bracteoles .
– .×.
– .mm,
sessile, narrowly lanceolate, incurved, apically acute, gla-
brous. Calyx tubes .
– .mm long, indistinct, green. Sepals

– .×.
– .mm, free, sometimes slightly adpressed to
corolla tube in lower , narrowly lanceolate, apically acute.
Corolla tubes 
– mm long, at its widest part .
– mm, sc arlet
red, ±-angled where it is widest and adpressed against the
carpels, gradually constricted above to a subcylindrical throat,
very slightly widening towards the petals. Petals 
– .×mm,
ovate-obovate, apex obtuse, short-mucronate (.mm long).
Stamens , arranged in two rows,  stamens each, fused to the
corolla tube internally for the most part, free segments short,
anthers of upper row at most reaching to slightly exserting the
corolla mouth; upper laments (oppositipetalous) inserted at
c..mm above corolla base, free for .
– .mm; lower la-
ments (alternipetalous) inserted at c.mm above corolla base,
free for 
– .mm. Anthers dull brownish-yellow, each locule
(theca) .×.×.mm, elliptic, base emarginate, apex
rounded, dressed with a attened spheroid connective gland;
connective gland erythrocyte-shaped, distinctly present only
on the upper stamens, .×.×.mm. Pistils consisting
of  carpels; carpels .
– ×
– .mm, greatest width towards
the middle, convergent; styles .
– .mm long; stigmas very
slightly capitate. Scales .
– .×.mm, long and thin,
Candollea 76, 2021 A new species of Kalanchoe (Crassulaceae) from Madagascar – 121
Fig. 3. – Kalanchoe darainensis D.-P. Klein & Callm. A. Rosulate growth with succulent, bright green to olive leaves;
B. Terminal, scarlet red and many-flowered dichasial cymes; C. Leaves entirely covered with short to long glandular hairs.
A  B: Ranirison 749; C: Callmander et al. 241 Photos: A  B: P. Ranirison; C: M. Callmander]
B
A
C

attened, adpressed to carpel, whitish, ligulate / strap-shaped,
apex truncate to rounded. Seeds .×.mm, brown, longi-
tudinally furrowed (striate).
Distribution, ecology and phenology. – Kalanchoe darainensis
is known only from the Antsahabe forest in the Loky-Manam-
bato region (Daraina) in northeastern Madagascar (Fig.).
Being situated between the Eastern humid and the Western
dry phytogeographical domains, with elevation levels partially
resembling those of the Central domain, this area comprises
particularly diverse vegetation types (R et al., ).
Only four individuals of the new species were found among all
the localities visited during a vegetation study, which included
more than , records of plant occurrences in the  main
forest areas of that region (N, ). ese individu-
als were observed at elevations between  and m where
the annual mean precipitation reaches 
– mm and
annual mean temperatures vary from .
– °C, depending on
the elevation level (N, ). Kalanchoe darainensis
grows on the lower slopes of rocky outcrops on raw mineral
soils (lithic erosion soils on leucocratic granite or skeletal soils
on cristalin domes) in rupicolous vegetation mainly surrounded
by primary mesophilous and ombrophilous forest (R
et al., ), but also by matrix habitats on a ridge covered with
sparse vegetation with a low canopy. e rupicolous vegetation
reaches heights of m with occasionally emerging treelets up
to 
– (– ) m tall. e most frequently recorded species occur-
ring with K. darainensis include: Artabotrys darainensis Deroin
& L.Gaut. (Annonaceae), Aulotandra trigonocarpa H.Perrier
(Zingiberaceae), Chasechloa humbertiana A.Camus (Poaceae),
Doryopteris cordifolia (Baker) Diels (Pteridaceae), Dracaena
reexa var. linearifolia Baker (Asparagaceae), Emilia humifusa
DC. (Asteraceae), Eugenia viguieriana H.Perrier (Myrtaceae),
Pandanus cf. grallatus B.C. Stone (Pandanaceae) and Xerophyta
sp. (Velloziaceae).
Other species of Kalanchoe collected in the Loky Manam-
bato region include K. a. miniata Hilsenb. & Bojer ex Tul.,
K. maromokotrensis Desc. & Rebmann, K. pinnata (Lam.) Pers.
and K. porphyrocalyx (Baker) Baill.
Kalanchoe darainensis was collected in flowers between
April and October.
Conservation status. – While Kalanchoe darainensis is known
only from four collections representing four locations (sensu
IUCN, ), it is situated within a large block of forest in the
Loky-Manambato protected area. No future decline is expected
in terms of Area of Occupancy (AOO), Extent of Occurrence
(EOO), or quality of habitat. We therefore assign a prelimi-
nary risk of extinction status of “Least concern” [LC] using the
IUCN Red List Categories and Criteria (IUCN, ).
Notes. – Kalanchoe darainensis is morphologically most
similar to K. blossfeldiana and K. globulifera (Table). e two
latter species are restricted to the high-altitude humid forests
of the Tsaratanana mountain range, c.km south-west
from Daraina. Like K. darainensis, these species are perennial,
small-sized, soboliferous, rosulate plants with simple leaves
and a subcylindrical corolla-throat. Kalanchoe blossfeldiana and
K. darainensis share the same ower characters, i.e. the scarlet
red ower color and the petals being ovate-obovate, with an
obtuse, mucronate apex. Kalanchoe globulifera and K. darainensis
both produce few- to many-owered dichasial cymes, are at
least partially glandular pilose and are dressed with connective
glands on the anthers. Among the northern representatives of
Kalanchoe subg. Kalanchoe in Madagascar, a scarlet red ower
color is unique to K. blossfeldiana and K.darainensis.
Additional specimens examined. – Madagascar. Reg. SAVA [Prov.
Antsiranana]: Daraina, massif d’Antsahabe, °'S °'E, m,
.X., Callmander et al. 241 (G, K, MO, P, TEF); ibid. loco, °'S °'E,
m, .XII., Gautier & Nusbaumer LG 4843 (G, K, MO, P, TEF); ibid.
loco, °'S °'E, m, .XI. Razatsalama et al. 771 (MO, P, TAN).
Table 1. – Diagnostic characters distinguishing Kalanchoe darainensis D.-P. Klein & Callm. from K. blossfeldiana Poelln. and K. globulifera H. Perrier.
K. darainensis K. blossfeldiana K. globulifera
Habit height [cm] 8  13.5 9  30 6.5  9( 22.5)
Indumentum dense, partial, glandular absent sparse, partial, glandular
Leaf blade shape ovate, elliptic to orbicular ovate-oblong obovate-orbicular
Leaf blade tip rounded to obtuse obtuse to acute rounded
Type of inflorescence terminal, few- to many-flowered
dichasial cymes
terminal and axillary,
many-flowered thyrses
terminal, few- to many-flowered
dichasial cymes
Number of inflorescences typically 1 numerous 1 to few
Flower colour scarlet red scarlet red golden yellow
Calyx tube [mm] indistinct, 0.1  0.2 short, 1 short, 0.5
122 – A new species of Kalanchoe (Crassulaceae) from Madagascar Candollea 76, 2021

Acknowledgements
We are grateful to RogerLalaAndriamiarisoa for the ne
drawings, and to PatrickRanirison and LaurentGautier for
granting the permission of publishing their eld photos. DPK
gladly thanks omasJanßen for providing multiple and ver-
satile support in the examination of herbarium specimens.
He especially thanks Lucile Allorge-Boiteau for encourag-
ing him to carry out the description of this new species. LN
thanks SébastienWohlhauser, the NGO Fanamby, Con-
servation International, Fondation Jean-MarcelAubert,
the Direction générale des Eaux et Forêts de Madagascar
and the Département de Biologie et Ecologie Végétales de
l’Université d’Antananarivo for their technical and nancial
support for the inventory of the Loky-Manambato region.
MWC thanks SébastienWohlhauser for the unforgettable
eld trip to Darainain . Finally, PatrickRanirison and
GideonF.Smith are thanked for their reviews and comments
which greatly helped improving an earlier version of this
manuscript.
References
A-B, L. (). Madagascar centre de spéciation et
d’origine du genre Kalanchoe (Crassulaceae). In: L,
W.R. (ed.), Biogéographie de Madagascar: 
– . ORSTOM,
Paris.
B, J.G. (). Contributions to the ora of Madagascar. Part I.
Polypetalae. J. Linn. Soc., Bot. : 
– .
B, P. & L. A-B (). Kalanchoe (Crassulacées)
de Madagascar. Systématique, écophysiologie et phytochimie. Karthala,
Paris.
D, B. (). Kalanchoe. In: E, U. (ed.), Illustrated
handbook of succulent plants: Crassulaceae: 
– . Springer-Ver lag,
Berlin.
D, B. (). Kalanchoe antennifera, Crassulacée africaine
nouvelle. Acta Bot. Gallica : 
– .
IUCN (). IUCN Red List Categories and Criteria. Version 3.1.
Ed.. IUCN Species Survival Commision, Gland and Cam-
bridge.
M C (). Catalogue of the plants of
Madagascar. Missouri Botanical Garden, St. Louis & Antanana-
rivo. [http://www.tropicos.org/project/mada]
M, J. & P. S (). Atlas of the vegetation of Madagascar.
Atlas de la végétation de Madagascar. Royal Botanic Gardens, Kew.
N, L. (). Species distribution patterns in steep
environmental gradients: downscaling of a biogeographical framework
(Loky-Manambato Region, NE Madagascar). Ph.D. thesis, Uni-
versity of Geneva.
P, P.B., L.D. A, L. G &
P.P.L II (). Flora. Site . Loky Manambato. In:
G, S.M. et al. (ed.), The terrestrial protected areas of
Madagascar: eir history, description, and biota: 
– . Asso-
ciation Vahatra, Antananarivo.
R, P., Y. A, L. G &
S.W (). Vegetation. Site . Loky Manambato.
In: G, S.M. et al. (ed.), e terrestrial protected areas of
Madagascar: eir history, description, and biota: 
– . Associa-
tion Vahatra, Antananarivo.
R-H (). Monographie du genre Kalanchoe. Bull.
Herb. Boissier ser. , : 
– .
S, G.F. & E. F (). e infrageneric classica-
tion and nomenclature of Kalanchoe Adans. (Crassulaceae), with
special reference to the southern African species. Bradleya :

– .
Candollea 76, 2021 A new species of Kalanchoe (Crassulaceae) from Madagascar – 123