ISSN: 0075-5974 (print)
ISSN: 1874-933X (electronic)
KEW BULLETIN (2020) 75: 50
DOI 10.1007/S12225-020-09904-5
New species and nomenclatural changes in Cynorkis (Orchidaceae)
from Madagascar, the Comoros and the Mascarenes
Johan Hermans1,2 , Landy Rajaovelona3, Phillip Cribb1, Jean-Michel Hervouet4, Anton Sieder2 &
Jacky Andriantiana5
Summary. Four new species, Cynorkis kiehnii, C. marianneae, C. marmorata and C. murex, are described for the first
time. Cynorkis frappieri and C. raymondiana are compared, with the latter reduced to a synonym of the former.
Cynorkis gaesiformis, C. galeata, C. nutans and C. stenoglossa are compared and considered one variable species, with
the name C. nutans having priority. Additional information is provided about C. mammuthus following the discovery
of living material in N Madagascar.
Key Words. Descriptions, distribution, IUCN Red List assessments, taxonomy, types.
Introduction
The genus Cynorkis Thouars (Orchidaceae:
Orchidoideae) currently comprises around 200 species, subspecies and varieties in total, 183 of which are
largely endemic to Madagascar and the surrounding
islands (Mascarenes, Comoros), and fewer than 20
species are restricted to mainland Africa (Govaerts
et al. 2019). Since the major work on the orchid flora
of Madagascar by Schlechter (1924) and Perrier de la
Bâthie (henceforth Perrier) (1939), when 78 species
were recognised, a number of novelties in the genus
have been described from the region by ToilliezGenoud & Bosser (1961, 1962), Bosser & Veyret
(1970), Bosser (1969, 1980, 1998, 2002, 2015),
Hermans & Cribb (2007, 2014) and Hermans et al.
(2007, 2017).
assessments which will be completed and submitted
for review and publication by IUCN once the species
names are validly published and therefore available
for assessment. All the assessments have been compiled based on current knowledge of these taxa, by
one of the authors (Landy Rajaovelona), who is an
IUCN Red List assessor, using the IUCN Red List
Categories and Criteria (2012).
New Species
Cynorkis kiehnii Hermans, Sieder & Prehsler sp. nov.
Type: Madagascar, Mahajanga prov., Bealanana area,
1227 m, 31 Jan. 2018, Gamisch, Sieder, Prehsler &
Andriantiana 7586 (holotype WU!).
http://www.ipni.org/urn:lsid:ipni.org:names:77211716-1
Materials & Methods
As part of ongoing research and fieldwork on the
orchid flora of Madagascar at the Royal Botanic
Gardens Kew, the University of Vienna and PBZT
Antananarivo, specimens of putative new taxa were
compared with descriptions, herbarium material and
drawings of all the type specimens of the genus to
ascertain their novelty. This has resulted in the
recognition of a number of new species and a better
understanding of the nomenclature of others.
The conservation status of the new species given in
this paper are summaries of the full IUCN Red List
Erect lithophytic herb up to 45 cm high including the
inflorescence. Tubers 1 – 2, elongate, ovoid (15 – 25 × 6
– 9 mm), clustered, surface velvety; roots slightly hairy,
fleshy, wiry, emerging from the base of the new
growth, c. 1.5 mm in diam. Stem up to 3.5 cm, c.
4 mm in diam., enveloped by an acuminate sheath,
white spotted with lavender-pink, 30 × 4.3 mm. Leaves
1 – 2, erect to arching, longly lanceolate, conduplicate,
acuminate, somewhat fleshy, a little shiny on top, up to
16 × 1.5 cm, pale green with the veins darker, spotted
with brownish-pink, base white spotted lavender-pink.
Inflorescence erect, emerging from the leaf axil, up to 34
Accepted for publication 21 November 2019. Published online 9 December 2020
1
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK. e-mail: j.hermans@kew.org
2
Core Facility, Botanical Garden, University of Vienna, Rennweg 14, 1030 Vienna, Austria.
3
Kew Madagascar Conservation Centre, Lot II J 131 Ambodivoanjo, Ivandry, Madagascar.
4
61 rue du Lieutenant Ricard, Chatou, France.
5
Parc Botanique et Zoologique de Tsimbazaza, Rue Fernand Kasanga, Tsimbazaza, Antananarivo, Madagascar.
© The Author(s), 2020
50 Page 2 of 25
cm, 1.8 – 2.1 mm in diam., pale green spotted
brownish-pink, glabrous at the base, becoming finely
pubescent towards the apex, sometimes with a long
leaf-like lanceolate-subulate sheath in the lower half,
45 – 52 mm long, 5 – 8 mm wide, with a second and
third peduncle sheath further up, of the same shape
but shorter, 21 – 24 mm long, 2 – 3 mm wide. Rachis
loosely racemose, with 1 – 3 flowers. Floral bracts
reddish-brown spotted purple, ovate-lanceolate, acuminate, 7 – 8.5 mm long, 1.7 – 1.9 mm wide. Pedicel
with ovary densely hirsute, pale brownish-red, flecked
longitudinally with purple-red, almost straight to
arching, 22 – 27 × 1.8 – 2.3 mm in diam. Flowers c.
3 cm long, 3.3 cm wide not including the spur, overall
white more or less tinted purple-pink, lateral sepals
and petals pale, dorsal sepal darker with the veins
more intense, lip becoming darker towards the tip,
column white, spur pale brownish-red. Dorsal sepal
ovate, 10.1 – 10.4 × 4.1 – 4.4 mm, forming a hood with
the petals over the column, strongly gibbose, the apex
recurved, the base finely pubescent. Lateral sepals
spreading, ovate-spathulate, 10.2 – 10.5 × 6.6 – 6.8
mm, slightly concave, the margins involute, the apex
auriculate and reflexed. Petals broadly falcate, 8.8 – 9.1
× 2.4 – 2.6 mm, appressed to the inside of the dorsal
sepal margins. Lip entire, narrowly lanceolate, 14.7 –
14.9 × 2.7 – 2.9 mm, with the base shortly auriculate,
attenuate, very strongly recurved in the apical half,
margins strongly incurved, the base with a short
rounded ridge; spur lengthening as the flowers
develop, slightly infundibuliform at the base and
parallel with the ovary, then pendent, subulate, the
apical third a little thickened and flattened, in mature
flowers 35 – 39 × 1.8 – 2.9 mm. Column in a horizontal
to angular plane 11 – 14 mm long, 5 – 6.5 mm wide,
with two long rostellum arms for over half its length,
each with a rounded side-lobe for half the length,
staminodes underlying the side lobes forming ovate
auricles, a large nose-like mid-lobe, anther loculi
elongate, longer than wide, narrowed at the base.
Pollinia including the long caudicle c. 11 mm long.
Fruit fusiform with raised ridges, verrucose 35 × 3 mm.
Figs 1, 2.
RECOGNITION. With its entire mid-lobe of the rostellum
and two long arms, each with a rounded side-lobe
Cynorkis kiehnii belongs in section Gibbosorchis
H.Perrier ex Hermans (Hermans et al. 2007: 290). It
is recognised by its long lanceolate leaves, a large leaf
sheath, a tall 1 – 3-flowered inflorescence, and
glabrous flowers with a densely hirsute ovary, an entire
strongly incurved lip, a spur subulate in the middle
and the apical half thickened and flattened, and long
divided rostellum arms. It is undoubtedly very close to
but also very different from two Cynorkis described
recently from the same area: C. mammuthus Hermans
& P.J.Cribb (Hermans et al. 2017: 13) and
© The Author(s), 2020
KEW BULLETIN (2020) 75: 50
C. elephantina Hermans, Andriant. & Sieder (Hermans
et al. 2017: 7) (compared in Table 1). They share the
entire lip, hirsute ovary and column shape but
C. kiehnii differs from the former by the fewer flowers
almost a third the size, the broader petals, the
narrowly lanceolate lip (vs broadly oblanceolate) and
the differently shaped spur which is subulate in the
middle and the apical half thickened and flattened (vs
subulate) along its length. It differs from C. elephantina
in particular by its fewer elongate lanceolate leaves (vs
obliquely lanceolate), 1 – 3 flowers (vs 15 – 22), the
flowers that are about 1 3 smaller with a spur that is
about half the length and subulate in the middle with
the apical half thickened and flattened (vs subulate
along its entire length), and its purple-pink tinted
whitish flowers (vs ivory-white spotted with burgundy).
DISTRIBUTION. Known only from the Bealanana area in
northern Madagascar.
SPECIMENS EXAMINED. MADAGASCAR. Mahajanga prov.,
Bealanana area, 1227 m, Jan. 2018, Gamisch, Sieder,
Prehsler & Andriantiana 7586 (holotype WU!).
HABITAT. On top of inselberg, on basalt rock in a thin
layer of moss, also in cracks in the rock, growing with
Nematostylis anthophylla (A.Rich. ex DC.) Baill. (Baillon
1879: 198) (Rubiaceae) and Cynorkis siederi Hermans &
Andriant. (Hermans et al. 2017: 18). 1200 – 1250 m.
CONSERVATION STATUS. Cynorkis kiehnii is only known
from a single unprotected locality on top of inselbergs in Sofia region, Mahajanga province. It was
found alongside a road and such fragments are
known to be particularly at risk from human
development. The species is assessed as Critically
Endangered CR based on Criterion D, with the
number of mature individuals likely to be fewer than
50 (around 20 plants were found on a single
inselberg), and under criterion B2ab(i,ii,iii) with a
continuing decline in the value of the AOO and the
EOO estimated to be less than 10 km2 and the
habitat quality due to human activities such as land
clearing and fire for agricultural uses.
FLOWERING TIME. January – February.
ETYMOLOGY. The name honours Dr Michael Kiehn;
the inspiring Professor of Botany and Director of the
Department of Biogeography and the Botanical Garden, University of Vienna.
Cynorkis marianneae Hermans sp. nov. Type: Madagascar, Antananarivo prov., Ankazobe area, along
RN4, in grass verge, in shade beneath a pine copse,
Jan. 1996, Hermans 8135. (holotype K!).
http://www.ipni.org/urn:lsid:ipni.org:names:77211717-1
Erect, single-leafed herb c. 20 cm tall. Tubers 2 – 3,
elongate, clustered, surface velvety; roots villous,
fleshy, wiry c. 1 mm in diam. Stem short, 1.5 cm long,
KEW BULLETIN (2020) 75: 50
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Fig. 1. Cynorkis kiehnii. A plants in habitat; B flower, lateral view; C detail of flower.
enveloped by a membranous sheath, 8 mm in diam.
Leaf elongate, ligulate-linear, 12.5 × 1.3 cm, somewhat
fleshy, solitary, pale green. Inflorescence glabrous, up to
13 × 0.1 cm, arching, with a short membranous sheath
at the base and then towards the middle a 32 × 3 mm
50
PHOTOS: ANTON SIEDER.
leaf-like sheath, caudate at the apex. Rachis racemose,
lax, with 3 – 4 flowers. Floral bracts narrowly lanceolate,
av. 15 × 3 mm, becoming shorter towards the apex.
Flowers 2.5 × 1.5 cm, with pale olive green petals and
sepals, a white lip with a crimson purple base,
© The Author(s), 2020
50 Page 4 of 25
KEW BULLETIN (2020) 75: 50
Fig. 2. Cynorkis kiehnii. A habit; B inflorescence; C flower; D floral bract; E dorsal sepal, front view; F dorsal sepal, oblique view; G
lateral sepal; H petal; J lip; K column and lip, side view; L column, side view. From Gamisch, Sieder, Prehsler & Andriantiana 7586.
DRAWN BY JUDI STONE.
© The Author(s), 2020
KEW BULLETIN (2020) 75: 50
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50
Table 1. Comparison of Cynorkis elephantina, C. mammuthus and C. kiehnii.
Cynorkis elephantina
Plant height
Leaves
Inflorescence
Dorsal sepal (mm)
Lateral sepals (mm)
Petals (mm)
Lip
spur
Altitude (m)
Cynorkis mammuthus
Cynorkis kiehnii
up to 35 cm
obliquely lanceolate,
up to 23 × 9 cm
up to 29 cm, 15 – 22 flowers
15 – 16.2 × 7 – 7.8
16 – 17 × 6.0 – 6.2
14 – 14.5 × 2.1 – 2.2
entire, obovate-lanceolate,
20.1 – 21.2 × 5 – 5.2 mm
subulate along its entire length, 7 –
7.8 cm long
up to 75 cm
lanceolate, funnel-shaped
up to 12 × 6.5 cm
up to 40 cm, 11 – 20 flowers
32 – 37 × 14 – 18
29 – 35 × 12 – 15
31 – 35 × 2.5 – 2.9
entire, oblanceolate,
36 – 41 × 12 – 16 mm
subulate along its entire length, 10.5
– 27 cm long
319
c. 250
becoming darker within the throat and a brownish red
spur, a white column with the margins purple, and a
green ovary with purple-red grooves. Pedicel with ovary
fusiform, slightly arched, glabrous except for the join
with the base of the column which has a few scattered
glandular hairs, 21 – 23 × 1 2.1 mm. Dorsal sepal ovate,
8.2 – 8.5 × 3.5 – 3.6 mm, forming a hood over the
column with the petals obtuse, with glandular hairs on
the exterior. Lateral sepals spreading, oval-lanceolate,
9.9 – 10.2 × 4.6 – 5.1 mm, obtuse, with a few hairs at
the base. Petals narrowly lanceolate, falcate, 8.2 – 8.4 ×
1.9 – 2 mm. Lip unequally 4-lobed, 21 – 23 × 11 – 13
mm; lateral lobes elliptic, rounded; mid-lobe dividing
around the middle into two oblanceolate, obtuse
lobes, the margins a little reflexed; the disc raised a
little in the centre forming a rounded ridge; spur
slightly sigmoid, parallel with the ovary, 22 – 23 × 0.9 –
1.1 mm, gradually narrowing towards the apex, the
base with a few glandular hairs. Column horizontal,
overall c. 5 × 3 mm, with two 2 mm attenuate rostellum
arms for 1 3 its length, with a narrow nose-like mid-lobe
recurved at the apex, anther loculi ovoid. Pollinia with
1.5 mm long caudicles. Figs 3, 4.
RECOGNITION. Cynorkis marianneae is recognised by its
long ligulate leaf, few-flowered inflorescence, and
flowers with a four-lobed lip with the elliptic lateral
lobes with rounded margins, a mid-lobe divided
into two oblanceolate, obtuse lobes with the
margins a little reflexed, a spur about the same
length as the ovary and the column with a narrow,
recurved mid-lobe.
There are strong similarities with both Cynorkis
flexuosa Lindl. (Lindley 1835: 331) (Fig. 5A) and
C. speciosa Ridl. (Ridley 1886: 122) (Fig. 5B): it is likely
that it is a natural hybrid of these two common and
variable species. All three were found flowering in the
same locality, a few paces apart, with C. flexuosa the
most common. Table 2 shows a comparison of the
up to 35 cm
Long-lanceolate,
up to 16 × 1.5 cm
up to 32 cm, 1 – 3 flowers
10.1 – 10.4 × 4.1 – 4.4
10.2 – 10.5 × 6.6 – 6.8
8.8 – 9.1 × 2.4 – 2.6
entire, narrowly lanceolate,
14.7 – 14.9 × 2.7 – 2.9 mm
subulate in the middle, the
apical half thickened
and flattened 3.5 – 3.9 cm long
1227
three species, measurements for C. flexuosa and
C. speciosa are based on averages of 42 specimens from
various herbaria and descriptions. Cynorkis marianneae,
is similar in general plant habit and size to both
species, it shares the overall lip and spur shape with
both but has an intermediate petal, lip and spur size
between the two. Both putative parents are very
variable in colour but they share the green dorsal
sepal–petal hood; the lip of C. speciosa is normally palepink to white with an undefined darker pink area at its
base, C. flexuosa invariably has a yellow lip with
generally a defined dark red-maroon spot at its base,
it is therefore possible that the colour of C. marianneae
is a combination of these.
Cynorkis marianneae differs from C. speciosa by the
fewer and smaller flowers, the smaller petals, lip and
slightly shorter spur. The lobes of the lip do not
overlap and the lateral lobes are rounded vs angular
sinuate, the anterior lobes are obtuse (vs angular) and
slightly repand (vs undulate). The column structure is
also different, with a narrow recurved mid-lobe of the
rostellum (vs a broader truncate one in C. speciosa). It
differs from C. flexuosa by its slightly larger flowers with
especially the lip and petals over the maximum
recorded size of C. flexuosa, it also lacks the yellow
colour of the lip which is consistent in C. flexuosa. The
spur is sigmoid (vs straight) and over a third longer.
The rostellum arms are less deeply divided and the
narrow recurved mid-lobe is intermediate between
C. flexuosa and C. speciosa.
All the described varieties of Cynorkis flexuosa and
C. speciosa were compared but none matched the new
entity.
DISTRIBUTION. Endemic to the Highlands of Madagascar, in Antananarivo province.
SPECIMENS EXAMINED. MADAGASCAR. Antananarivo
prov. Ankazobe area, along RN4, Jan. 1996, in grass
verge, in shade beneath scrub in a pine copse,
Hermans 8135 (holotype K!).
© The Author(s), 2020
50 Page 6 of 25
KEW BULLETIN (2020) 75: 50
Fig. 3. Cynorkis marianneae. A flower, face view; B flower, lateral view.
HABITAT. Seasonally dry grassland, in shade of pine
trees.
CONSERVATION STATUS. Cynorkis marianneae is only
known from a single unprotected locality in
Analamanga region, Antananarivo province. The species is assessed as Critically Endangered CR based on
Criterion D, with the number of mature individuals is
likely to be fewer than 50 (only a single plant was
observed in the type locality), and under criterion
B2ab(i,ii,iii) with a continuing decline in the value of
the AOO and the EOO estimated to be less than
10 km2 and the habitat quality due to human activities
such as land clearing and fire for agricultural use.
FLOWERING TIME. January.
ETYMOLOGY. Named for Marianne Cribb who discovered
a plant of this species in a secluded position.
NOTES. Natural hybrids in the genus Cynorkis have
been recorded a number of times and two include
C. speciosa as a parent: C. ×syringescens Hermans &
P.J.Cribb (Hermans et al. 2017: 21) (C. speciosa × C.
fastigiata Thouars (1822: t. 3)) and C. ×ranaivosonii
Hermans & Rajaovelona (2019: 547) (C. speciosa × C.
uniflora Lindl. (Lindley 1835: 331)). Other putative
© The Author(s), 2020
PHOTOS: JOHAN HERMANS.
crossings in the genus include C. ×mirabile Hermans &
P.J.Cribb (2007 : 252) ((C. lowiana Rchb.f.
(Reichenbach f. 1888: 150) × C. gibbosa Ridl. (Ridley
1883: 331)) and C. paradoxa (Frapp. ex Cordem.)
Schltr. (Schlechter 1915: 401) ((C. lilacina Ridl.
(Ridley 1885: 515) × C. ridleyi T.Durand & Schinz,
(1895: 92)). There is no absolute certainty that
C. marianneae is a natural hybrid. Further study of
populations and other occurrences could confirm its
status but meanwhile it is prudent to treat it as a
species with similarities to C. flexuosa and C. speciosa.
Cynorkis marmorata Hermans sp. nov. Type: Madagascar, Antananarivo prov., Anjozorobe area, degraded humid
evergreen forest, in undergrowth, in shade of small trees,
1305 m, May 1998, Hermans 8257 (holotype K!).
http://www.ipni.org/urn:lsid:ipni.org:names:77211718-1
Small single-leafed terrestrial herb up to 6 cm tall
without the inflorescence. Tubers 1 – 2, clustered,
ovoid, woolly on the exterior; roots villous, wiry, c.
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50
Fig. 4. Cynorkis marianneae. A leaf; B inflorescence; C flower; D dorsal sepal; E lateral sepal; F petal; G lip; H column, ovary and lip
spur (lip removed); J column, side view; K column from above. From Hermans 8135. DRAWN BY JUDI STONE.
© The Author(s), 2020
50 Page 8 of 25
KEW BULLETIN (2020) 75: 50
Fig. 5. A Cynorkis flexuosa, plant in habitat, and B C. speciosa, part of inflorescence, for comparison with C. marianneae.
PHOTOS:
JOHAN HERMANS.
1 mm in diam. Leaf on a short, 4 – 7 mm, stem-like
petiole, blade lanceolate, attenuate, 28 – 53 × 10 – 12
mm, serrate at the margins, pale green with characteristic interrupted, silvery-white horizontal to transversal bands in between the veins, giving it a marbled
appearance, brownish-green underneath. Inflorescence
erect, slender, up to 25 cm long. Peduncle angular,
hirsute, brownish-green, 1 – 1.2 mm in diam., with 1
– 2 lanceolate sheaths, the tip acuminate and the
mid-vein thickened, 3.5 – 3.9 × 1 mm. Rachis loosely
racemose, around half the length of the inflorescence, hirsute, reddish-brown, developing fruits
Table 2. Comparison of Cynorkis flexuosa, C. speciosa and C. marianneae.
Cynorkis flexuosa
Leaves, average size (cm)
Inflorescence average length (cm)
Dorsal sepal (mm)
Lateral sepals (mm)
Petals (mm)
Lip
Spur
Column
© The Author(s), 2020
15 × 1
25
12 – 20 × 2 – 3
8 – 11 × 4 – 4.5
6.5 – 8 × 1.5 – 2
16 – 21 × 8 – 15 mm,
lobes not overlapping.
Yellow
8 – 14 × 1.5 – 2 mm,
straight
c. 4 × 3.5 mm, mid-lobe
nose-like, slightly recurved
Cynorkis speciosa
Cynorkis marianneae
17 × 1.5
30
9 – 12 × 4 – 5.3
9 – 18 × 4 – 6
10 – 12 × 2.5 × 3.5
23 – 28 × 14 – 19 mm,
lobes partly overlapping.
Pinkish-white
20 – 30 × 2 mm, sigmoid
14 × 1.2
14
8.2 – 8.5 × 3.5 – 3.6
9.9 – 10.2 × 4.6 – 5.1
8.2 – 8.4 × 1.9 – 2
21 – 23 × 11 – 13 mm,
lobes not overlapping. White
tinted pink
22 – 23 × .09 – 1.1 mm, slightly
sigmoid
c. 5 × 3 mm, mid-lobe narrow,
recurved
c. 6 × 4 mm, mid-lobe broad,
arching
KEW BULLETIN (2020) 75: 50
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50
whilst the apical flowers are still developing. Floral
bracts lanceolate, acuminate, 2.4 – 2.6 × c. 1 mm,
brownish-green. Flowers c. 10 × 8 mm, non-resupinate, divaricate, white to pale pink with pale
brownish-green sepals and spur, and a lip with the
disc with 3 – 4 longitudinal reddish-brown bands, a
small pink patch on the mentum underneath,
continuing along the mid-vein. Pedicel with ovary
fusiform, straight, 8.1 – 8.7 × 1.2 – 1.4 mm, very
hirsute, some of the hairs glandular. Dorsal sepal
forming a loose hood with the petals, broadly ovate,
2.3 – 2.4 × 2.2 – 2.4 mm, obtuse, glandular on the
exterior. Lateral sepals spreading, lanceolate-falcate,
3.5 – 3.7 × 2.2 – 2.3 mm, attenuate, glandular on the
exterior. Petals lanceolate, acute, 2.5 – 2.7 × 0.8 – 0.9
mm. Lip 3-lobed with a short concave hypochile
towards the base, 3.9 – 4.2 × 4 – 4.2 mm, the lateral
lobes triangular, attenuate, the midlobe broadly
ovate, with undulate margins, glabrous except for
the exterior of the hypochile which has a few hairs;
spur curved, 3.3 – 3.5 × 0.7 – 0.8 mm, broadened at
the base, then tubular, then clavate at the tip. Column
in a flat blade, c. 1.3 × 1.2 mm, the rostellum angular
and entire for almost its whole length; anther loculi
ovoid. Pollinia c. 1.2 mm, with a long caudicle. Fruit
fusiform, purple, c. 9 × 3 mm. Figs 6, 7.
SPECIMENS EXAMINED. MADAGASCAR. Antananarivo
prov., Anjozorobe area, May 1998, 1305 m, Hermans
8257 (holotype K!).
HABITAT. In humid evergreen forest, in deep shade, in
small scattered clumps, amongst leaf humus, mosses
and lichens. c. 1300 m.
CONSERVATION STATUS. Cynorkis marmorata is only
known from a single unprotected locality in the
Analamanga region, Antananarivo province. The species is assessed as Critically Endangered CR based on
Criterion D, with the number of mature individuals
likely to be fewer than 50 (around 25 plants were seen
in one locality), and under criterion B2ab(i,ii,iii) with
a continuing decline in the value of the AOO and the
EOO estimated to be less than 10 km 2 and the habitat
quality due to human activities such as shifting
agriculture and forest logging.
FLOWERING TIME. April to May.
ETYMOLOGY. Referring to its marbled leaf.
RECOGNITION. With its small flowers with the lip
Erect terrestrial herb up to 15 cm high excluding the
inflorescence. Tubers 2 – 3, elongate, ovoid, 2 – 3 × 0.5
– 1 cm, clustered, surface velvety; roots villous, fleshy,
wiry c. 1.5 mm in diam. Stem short, 1 – 2 cm long. Leaf
solitary, broadly lanceolate, caudate at the apex, 52 –
75 × 23 – 31 mm, attenuate at the apex, with five
prominent veins, margins undulate, pale green with
the veins a little darker, the mid-vein bordered by a
broader dark area, on a long 30 – 52 × 3 – 7 mm
petiole enveloped for a third of its length by a sheath.
Inflorescence up to 30 cm long, 1.5 – 2 mm in diam.,
finely pubescent. Peduncle with 4 – 6 equally spaced
sheaths 6 – 8 × 1.4 – 1.6 mm. Rachis racemose, c. 5 cm
long, loosely 12 – 17-flowered. Floral bracts lanceolate,
acuminate, irregularly spinose at the margins and
along the mid-vein, bright purple when young,
becoming reddish-brown with age. Flowers porrect,
spreading, c. 10 × 15 mm, pale pinkish-white with
much darker purple-magenta markings, a pale
pinkish-white dorsal sepal, lateral sepals with parallel
dark purple-magenta bands, sometimes fused into
one, petals with purple margins, a lip with small
purple spots at the base and several triangular spots
alongside the paler central vein, a spur with a darker
purple tip, rostellum arms edged purple, and purplishred anther loculi. Pedicel with ovary fusiform, 13 – 14.5
× 1.8 – 1.9 mm, hirsute, white. Dorsal sepal ovate, 4.1 –
4.3 × 1.9 – 2 mm forming an arching hood over the
column with the petals, the apical part narrowed, the
uppermost and column characteristics Cynorkis
marmorata belongs in sect. Hemiperis (Frapp. in
Cordem.) Perrier in Humbert (1939: 74). It is
recognised by its small lanceolate leaf with serrate
margins and the silvery reticulate markings on the
blade, its long hirsute inflorescence, and small hairy
flowers with an unequally 3-lobed lip with a rounded
hypochile at the base, and an upturned spur clearly
clavate at the tip.
Cynorkis marmorata is somewhat similar to
C. henrici Schltr. (Schlechter 1924: 52), C. ridleyi
T.Durand & Schinz (1895: 92) and C. sororia Schltr.
(Schlechter 1913: 154) with which it shares the
general lip shape, short thin spur and patterned
leaves but they all have much larger flowers and
bigger ovate leaves. It is closest to C. flabellifera
H.Perrier (1951: 144) with which it shares a similar
general habit, lip and spur shape but it consistently
has a narrower leaf with an up to 7 mm petiole (vs
45 mm), and flowers that are about 1 3 its size, a
dorsal sepal that is broadly ovate (vs lanceolate), a
lip with an ovate mid-lobe (vs flabellate),
C. flabellifera also lacks the serrate leaf margins
and has purple-white flowers and has only been
found in Antsiranana province in the far north of
Madagascar.
DISTRIBUTION. Endemic to Antananarivo province of
Madagascar.
Cynorkis murex Hermans sp. nov. Type: Madagascar,
Toamasina prov., near Andasibe, c. 1150 m, May 1996,
Hermans 8240 (holotype K!).
http://www.ipni.org/urn:lsid:ipni.org:names:77211719-1
© The Author(s), 2020
50 Page 10 of 25
Fig. 6. Cynorkis marmorata. A plant in habitat; B leaf; C rachis.
exterior longly and densely hirsute. Lateral sepals
spreading, parallel with the lip, ovate-acute, 5.8 – 6 ×
3.1 – 3.2 mm, margins incurved, hirsute on the
exterior. Petals partly covered by the dorsal hood,
oblong-lanceolate, 3.2 – 3.3 × 1.1 – 1.2 mm, the apex
with a small auricle, glabrous. Lip 5-lobed, 6.3 – 6.5 ×
4.9 – 5 mm, the basal lobes small, rounded deltate, the
lateral lobes strongly attenuate-angular, incurved, the
mid-lobe strongly attenuate-angular recurved at the
tip, the disk with a longitudinal raised ridge along
its
length; spur 8.9 – 9.1 × 2.1 – 2.3 mm, in the basal 2 3
parallel to the ovary, narrowing, hirsute, then abruptly
narrowed, descending and glabrous. Column angular
3.2 – 3.3 × 2.4 – 2.5 mm, with two 3 mm rostellum arms
for over half its length, with a small nose-like midlobe, unevenly obovoid-auricular staminodes towards the base, anther loculi elongate. Pollinia
finely granular, c. 3.2 mm long including the long
caudicle. Figs 8, 9.
RECOGNITION. Cynorkis murex is recognised by its
single broadly lanceolate leaf on a petiole almost as
long as the blade, its tall loosely racemose inflorescence, and hirsute, brightly coloured flowers with a
5-lobed lip and a descending, angular spur.
Its closest relative is undoubtedly Cynorkis densserpens Hermans & P.J.Cribb (2014: 1) (Fig. 10),
described recently from the same area: it shares the
plant habit, some of the floral characteristics, and it
also flowers at the same time. The new species is very
different in that the leaf margins are undulate and
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KEW BULLETIN (2020) 75: 50
PHOTOS: JOHAN HERMANS.
entire (vs serrate). The lip is consistently smaller and
has rounded-deltate (vs auriculate) basal lobes, strongly attenuate-angular lateral lobes and mid-lobe (vs
rounded-obtuse), an angular, descending spur (vs
ascending, subulate), it is also a little shorter, 3 mm
long rostellum arms (vs 2 mm) and staminodes that
are a different shape (they are compared in Table 3).
The colour of the flowers is also consistently and
considerably different.
DISTRIBUTION. Endemic to the Andasibe area in
Toamasina province in Eastern Madagascar.
SPECIMENS EXAMINED. MADAGASCAR. Toamasina prov.,
near Andasibe, c. 1150 m, May 1996, Hermans 8240
(holotype K!).
HABITAT. In humid evergreen forest, on wooded banks
near small streams, in moist humus, in dappled shade.
c. 1150 m.
CONSERVATION STATUS. Cynorkis murex is endemic to
Madagascar and distributed in Alaotra-Mangoro region in Toamasina province. The value of the area of
occupancy (AOO) is estimated to be less than 10 km2.
The species is located outside protected areas. In
addition, the EOO, the AOO and the habitat of the
species are suffering a continuing decline due to
shifting agriculture. It is therefore assessed as
Endangered (EN). Further research is
recommended, particularly on population size of this
species.
FLOWERING TIME. May.
ETYMOLOGY. Referring to Murex, a genus of molluscs,
often with a prickly exterior, once used to produce the
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50
Fig. 7. Cynorkis marmorata. A base of habit; B part of stem, enlarged; C inflorescence; D flower side view; E flower, front view; F
floral bract; G dorsal sepal; H lateral sepal; J petal; K lip; L column and lip, side view; M column. From Hermans 8257. DRAWN BY JUDI
STONE.
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KEW BULLETIN (2020) 75: 50
Fig. 8. Cynorkis murex. A plant in habitat; B rachis; C flower, face view.
Tyrian purple dye, similar in colour to the markings
on the flower of the orchid.
NOTES. Cynorkis dens-serpens and C. murex are known
from several substantial colonies, about 10 km apart in
separate valleys: all plants within each of these
colonies were consistent in shape, size and colour
and no intermediate forms have been found.
The identity of Cynorkis frappieri and C.
raymondiana
Cynorkis frappieri Schltr. (Schlechter 1915: 400) as
Cynosorchis frappieri; Bernet (2010: 150); Szelengowicz
& Tamon (2013: 269). Type: Réunion, 'Tampon, rive
gauche du bras de Jean Payet' (Frappier 1895: 237),
probably not preserved. Neotype designated here: île
de Bourbon [Réunion], Herbier A. Delteil s.n. (Herb.
Drake) (neotype P00693021!).
Hemiperis tenella Frapp. ex Cordem. (Frappier 1895:
237); Frappier nom. nud. (1880: 11). Type as above.
Cynorkis raymondiana H.Perrier ex Hermans (Hermans
et al. 2007: 292); Cribb & Hermans (2009: 80)
synon. nov. Lectotype: Madagascar, nr FortDauphin [Tôlanaro], Decary 10142 (lectotype
P00102022!), designated here. Paratype: Decary
10019 (P00102023!).
Cynosorchis raymondiana H.Perrier in Arch. Bot. Bull.
Mens. 5 (1931: 48) (unpublished, see Hermans
2007: 290): Perrier (1939: 97).
Bicornella raymondiana (H.Perrier.) Szlach. & Kras
(Szlachetko & Kras 2006: 145), nom. invalid.
© The Author(s), 2020
PHOTOS: JOHAN HERMANS.
Small squat herb up to 7 cm high excluding the
inflorescence. Tubers 1 – 2, clustered, ovoid, 1 – 2 ×
1.3 – 1.8 mm, woolly on the exterior; roots villous,
wiry, c. 1 mm in diam. Leaves 2 – 3, flat, radical, ovate
to lanceolate, 3.7 – 8.2 × 1 – 2.8 cm, with a very short
conduplicate petiole attenuate at the tip, margins
entire, green. Inflorescence erect, racemose, up to
22 cm long, glabrous, green with up to 13 flowers
but generally fewer. Peduncle slender, 1.5 – 2 mm in
diam., over 2 3 the length of the inflorescence, with 3 –
5 distant lanceolate peduncle sheaths, 5 – 12 × 1.1 –
2.2 mm, the basal one longer and more acuminate
than the others. Rachis laxly flowered, 2 – 6 cm. Floral
bracts lanceolate, acuminate, 3.8 – 4.9 × 1.3 – 1.5 mm.
Flowers very small for the genus, erect to arching, up to
7 × 6 mm, glabrous, crystalline white with the exterior
of the sepals more or less tinted pinkish green, the lip
with up to 6, more or less merging, magenta spots,
with magenta-marked basal lobes, and yellow pollinia,
(Szelengowicz & Tamon (2013: 270) report forms
without distinct spots on the flowers and other forms
with an overall pink to mauve pigmentation). Said to
be scented of goat (d’odeur hircine) by Frappier
(1895: 238) but this has not been verified in the
field. Pedicel with ovary fusiform, 6 – 10 × 1 – 1.2
mm, glabrous, green. Dorsal sepal forming a tight
hood with the petals, broadly ovate, obtuse, concave,
2.1 – 3 × 1.5 – 1.8 mm. Lateral sepals spreading,
ovate, concave, 2.8 – 3.0 × 1.5 – 1.8 mm. Petals
lanceolate to elliptic, acute, 2 – 2.5 × 0.9 – 1.2 mm.
Lip obcordate, unequally 5-lobed, 3.2 – 4 × 2.5 – 2.9
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50
Fig. 9. Cynorkis murex. A habit; B stem, enlarged view; C floral bract; D flower front view; E flower, side view; F dorsal sepal, front
view; G dorsal sepal, oblique view; H lateral sepal; J petal; K lip; L column from above. From Hermans 8240. DRAWN BY JUDI STONE.
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Fig. 10. Cynorkis dens-serpens. A rachis, B close up of flower, for comparison with C. murex.
mm, the anterior lobe triangular acute, the lateral
lobes a little larger, obovate obtuse, the basal lobes
small, triangular acute; spur thickened in the lower
half to club-shaped, 1.5 – 1.8 × 0.4 – 0.6 mm. Column
in a horizontal blade to somewhat ascending, 1 – 1.7 × 1 –
1.1 mm, anther bilobed, midlobe of the rostellum and
staminodes small, rounded, lateral lobes of the rostellum
bidentate at the front, with the midlobe of the rostellum
appearing tridentate, the pollinia ending in a distinct
viscidium. Figs 11, 12.
PHOTOS: JOHAN HERMANS.
RECOGNITION. With its slender plant habit, laxly fewflowered inflorescence, small flowers and rostellum
with a short toothed horizontal blade Cynorkis frappieri
belongs in sect. Hemiperis (Frapp. ex Cordem.) Perrier
in Arch. Bot. Bull. Mens. 5 (1931: 36 unpublished);
Perrier (1939: 74). It is recognised by its glabrous habit
and flowers, two or rarely three ovate to lanceolate
leaves flattened on the ground, a long peduncle at
2
least 3 the length of the inflorescence, and small
white with distinct magenta spotted flowers with a
Table 3. Comparison of Cynorkis dens-serpens and C. murex.
Cynorkis dens-serpens
Cynorkis murex
Inflorescence
Dorsal sepal (mm)
Lateral sepals (mm)
Petals (mm)
Lip
up to 25 cm tall, 8 – 15 flowers
3.8 – 5 × 2 – 3
5.7 – 6 × 3.9 – 4.5
3.6 – 4.3 × 1.5 × 2
5-lobed, basal lobes auriculate, lateral &
mid-lobes rounded-obtuse, 7 – 10 × 7 – 9 mm
Spur
Column
ascending, subulate, straight, up to 12 mm long
rostellum arms c. 2 mm, staminodes globose
up to 30 cm tall, 12 – 17 flowers
4.1 – 4.3 × 1.9 – 2
5.8 – 6 × 3.1 – 3.2
3.2 – 3.3 × 1.1 – 1.2
5-lobed, basal lobes rounded deltate,
lateral & mid-lobes strongly
attenuate-angular, 6.3 – 6.5 × 4.9 – 5 mm
angular descending, narrowing, up to 9 mm
rostellum arms c. 3 mm, staminodes obovoid, auricular
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Page 15 of 25
Fig. 11. Cynorkis frappieri. A plant in habitat; B part of inflorescence.
short spur swollen in the lower half. In habit and
flower shape it is similar to C. sigmoidea (K.Schum.)
Kraenzl. (Kraenzlin 1898: 490) from the Comoros but
the flowers of that species are almost double the size,
the lip segments narrower and the raceme and ovary
pubescent (vs glabrous). It is also similar to C. tenella
Ridl. (Ridley 1886: 124) from Madagascar and the
Comoros but the flowers of that species are a third
larger, the spur double the length and the lip 3-lobed (vs
5-lobed), the raceme, ovary and back of the dorsal sepals
are pubescent (vs glabrous). Examination of all the
herbarium specimens of C. frappieri and C. raymondiana
combined with field observations confirm that these taxa
are identical in all respects with the former name having
priority. A drawing and manuscript notes, accompanying
the Delteil s.n. (P00693021) herbarium sheet, contain an
annotation by Bosser that it is ‘similar and may be the
same species as Cynorkis raimondiana’.
DISTRIBUTION. Widely distributed on Madagascar from
the central area in Antananarivo province, through
the NE mountains in Fianarantsoa province to the far
south in Toliara province. Also widespread on Réunion ranging from the east, the south-west and
Cirque de Cilaos in the centre. Known from one
locality in western Mauritius.
50
PHOTOS: JOHAN HERMANS.
SPECIMENS EXAMINED. MADAGASCAR. Fort-Dauphin,
littoral forest, 18 July 1932, R. Decary 10142
(P00102022!); nr Fort-Dauphin, littoral forest, 8 July 1932, R. Decary 10019 (P00102023!); Ambondrombe
area, moss forest remnants, c. 1200 m, 10 April 1941,
Herb. Jard. Bot. Tan. 4572 (P00102024!, P00102025!);
Antananarivo prov. nr Anjozorobe, Andrianarivo, 1305
m, May 2018, Hermans 8226 (K!). MAURITIUS. Corps de
Garde, June 1976, J. Bosser 22267 (P00693026!). RÉUNION. île Bourbon, Herbier A. Delteil s.n. (Herb.
Drake) P (P00693021!); Mare Longue, Saint-Philippe,
Dec. 1971, Bosser 20848 (P-spirit!); Basse Vallée,
Dec. 1972, J. Bosser 21602 (P0069992!).
HABITAT. On Réunion in humid forest, montane forest
and drier areas, terrestrial or very rarely epiphytic in
semi-shade at an altitude from 400 – 1300 m (Bernet
(2010: 150). On Madagascar in humid forest, ‘Tapia’
forest and littoral forest in semi-shade at an altitude
from 0 – 1305 m.
CONSERVATION STATUS. Cynorkis frappieri has a large
distribution in Madagascar in Analamanga, HauteMatsiatra and Anosy regions from Antananarivo,
Fianarantsoa and Toliara regions. It is also widespread
in Réunion, ranging from East, the SW and Cirque de
Cilaos in the centre and known from one locality in
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KEW BULLETIN (2020) 75: 50
Fig. 12. Cynorkis frappieri. A habit; B tip of inflorescence; C flower, side; D dorsal sepal; E lateral sepals; F petals; G lip; H column
and spur, side; J column, top. From Decary 10141. DRAWN BY LINDA GURR after sketches by J. Hermans.
© The Author(s), 2020
KEW BULLETIN (2020) 75: 50
Mauritius. The extent of occurrence (EOO) is estimated to be more than 20,000 km2. The species is not
recorded in any Protected Areas in Madagascar it is
therefore inferred that most of its subpopulations are
threatened by habitat destruction due to slash and
burn agriculture (Tavy) and fires set for land-clearing
and pastureland. The species is therefore assessed as
Near Threatened (NT).
FLOWERING TIME. Madagascar: May to June. Mauritius:
June. Réunion: March to July.
ETYMOLOGY. Tenella: from the Latin tenellus, small and
delicate, refers to the small and delicate nature of the
plant; raymondiana refers to Raymond Decary (1891 –
1973), French colonialist and naturalist who collected
a large number of plants, described by Perrier; frappieri
refers to the French botanist Charles Frappier de
Montbenoist (1813 – 1885).
NOTES. Frappier first listed Hemiperis tenella (1880: 11)
from Réunion but this was without description or
indication of a type. The first description of Hemiperis
tenella was by Frappier in Cordemoy’s Flore de l’île de la
Réunion (1895: 237) and had an extensive protologue
but did not refer to type material, it only mentioned
the specific locality of Saint-Pierre (Tampon, rive gauche
du bras de Jean Payet) where a plant was seen (vu viv.)
by the author. No herbarium material has been found
so far of the species directly associated with Frappier
or Cordemoy but there is a good specimen at P
originating from A. Delteil s.n. ex herbier Drake
(P00693021), which is contemporary with the description and is selected here as the neotype for the
species. Schlechter transferred H. tenella, together with
many other Hemiperis, to Cynorkis in 1915 but the name
C. tenella could not be transferred as it had been used
by Ridley in 1886 (p. 124) for a different species from
Madagascar and the Comoros. Cynosorchis raymondiana
H.Perrier (1931: 48), mentioned by Perrier in the Flore
de Madagascar (1939: 97), was never effectively published but later validated, as explained in Hermans
et al. (2007: 290). Perrier (1939: 97) cited two Decary
specimens (10019 and 10142), collected a few days
apart near Fort-Dauphin [Tôlanaro] in southern
Madagascar, no holotype was indicated in Perrier’s
protologue, in the validation of the species in
Hermans et al. (2007: 292) Decary 10141 was listed as
the Holotype in error. The herbarium sheet Decary
10142 in P was annotated in Perrier’s hand as ‘sp. nov.
type’, Decary 10019 was annotated ‘sp. nov. identical to
Decary 10141!’. Under the International Code of
Nomenclature for algae, fungi and plants (Turland
et al.: 2018: article 9.3 & 9.11) a lectotype had to be
designated: Decary 10142 was chosen here because it
was indicated as ‘type’ by Perrier and the specimen,
containing five plants is more complete and representative. Decary 10019 is a paratype. It is interesting to
note that Perrier used the spelling ‘raimondiana‘ on
the herbarium sheets.
Page 17 of 25
50
Szlachetko & Kras (2006: 145) in their unsubstantiated fragmentation of the genus invalidly transferred
Cynorkis raymondiana to the genus Bicornella (Lindley
1835: 335) and incorrectly referred to Perrier’s
basionym as published in Archivio Botanico per la
Sistematica, Fitogeografia e Genetica 5: 48 (1931).
The identity of Cynorkis gaesiformis, C. galeata,
C. nutans C. stenoglossa
Cynorkis nutans (Ridl.) H.Perrier (1936: 582); Perrier
(1939: 156); Hermans et al. (2007 158); Cribb &
Hermans (2009: 111); Bosser & Lecoufle (2011: 293);
Szelengowicz & Tamon (2013: 277); Hervouet (2018:
299). Types (former syntypes): Madagascar, Central,
Baron 1703 (lectotype BM000032250!; isolectotype
K000415611! designated in Hermans et al. (2007:
158); P00102284!); Madagascar, Imerina, Deans Cowan
s.n. (paratype BM000911499!); Madagascar,
Andrangoloaka, Parker s.n. (paratype K!).
Habenaria nutans Ridl. (Ridley April 1885: 507);
Kraenzlin (1901: 923).
Imerinorchis nutans (Ridl.) Szlach. (Szlachetko 2005:
69) synon. nov.
Cynorkis galeata Rchb.f. (Reichenbach f. Oct. 1885: 536)
(as Cynosorchis galeata). synon. nov.; Kraenzlin (1901:
923); Perrier (1939: 157); Hermans et al. (2007: 149).
Type: Mayotte, Combani forest, Humblot 209 (1209)
(holotype W! R5342; isotype P00102270!, P00102271!,
P00102272!, BM000034836!).
Imerinorchis galeata (Rchb.f.) Szlach, (Szlachetko 2005:
69) synon. nov.
Cynosorchis nutans var. campenoni H.Perrier (1939: 156),
nom. nud.; Hermans et al. (2007: 158). Based upon:
Madagascar, Imerina, Nov. 1881, R. P. Campenon
s.n. (P00102291!).
Cynorkis stenoglossa Kraenzl. (Kraenzlin 1893: 63);
Kraenzlin (1901: 494); Schlechter (1913: 154);
Schlechter (1915: 402); Schlechter (1924: 72);
Perrier (1939: 158); Hermans et al. (2007: 166);
Cribb & Hermans (2009: 112); Bosser & Lecoufle
(2011: 300). synon. nov. Type: Madagascar, S
Betsileo, Ankafina forest, March 1881, J. M.
Hildebrandt 4204 (syntype W! R14896).
Imerinorchis stenoglossa (Kraenzl.) Szlach. (Szlachetko
2005: 69). synon. nov.
Cynorkis stenoglossa H.Perrier var. pallens H.Perrier ex
Hermans (Hermans et al. 2007: 293); Perrier (1939:
159); Cribb & Hermans (2009: 112); Hermans et al.
(2007: 166); synon. nov. Type: Madagascar, Mt
Ambre, H. Perrier 17733 (holotype P00102296!,
isotype P102297!).
Cynorkis stenoglossa var. pallens H.Perrier (1931: 72):
Perrier (1930: 18). nom. nud.
Cynorkis gaesiformis H.Perrier (1951: 146) synon. nov.
Type: Madagascar, S of Moramanga, Decary 18419
(holotype P00102285!).
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50 Page 18 of 25
Terrestrial, epiphytic or rarely lithophytic herb up to
30 cm tall including the inflorescence, with 3 – 6
elongate pubescent tubers 1 – 4.5 × 0.5 – 1.5 cm; roots
dense, woolly. Leaves 1 or rarely 2, radical, broadly
elliptic to oblong, 4.5 – 19.5 × 2.5 – 6.2 cm, tip
acuminate, narrowed into a more or less attenuate
petiole at the base, blade flat, arching towards the tip,
green with the veins often a little paler, with a thin
sheath enveloping the base, acuminate to caudate, 1 –
3 cm long. Inflorescence erect to divergent, 6 – 26 cm
tall, with up to 30 flowers. Peduncle about 2 3 of the
inflorescence, 3 – 4 mm in diam., densely hirsute
along its length, generally with a prominent lower
peduncle sheath 12 – 42 × 3 – 19 mm and 2 – 3
additional sheaths 6 – 12 × 2.5 – 3.2 mm. Rachis subdensely to densely racemose, often opening in succession with the lower ones fading before the upper ones
open. Floral bracts narrowly lanceolate, 4.8 – 8.9 × 1.5 –
2.1 mm, attenuate, hirsute on the exterior. Flowers
variable in size, up to 16 × 29 mm, very variable in
colour, ranging from pale pink to deep mauve-purple,
some forms having pink, lilac to purple sepals, lip and
petals, the anther loculi brownish-purple. Pedicel with
ovary fusiform, 12 – 24 × 1.7 – 3.5 mm, densely hirsute,
white to pink. Dorsal sepal lanceolate, 4.5 – 8.1 × 2 – 3.3
mm, helmet-shaped, the tip recurved, forming a tight
hood with the petals, hirsute on the exterior. Lateral
sepals broadly lanceolate to ovate, 7.7 – 17.3 × 4 – 7.9
mm, flat, asymmetric, spreading, attenuate, often
hirsute on the exterior, especially towards the attenuate base. Petals adnate to the dorsal sepal, lanceolate,
4.5 – 8.4 × 2.2 – 3.9 mm, expanded at the outer margin
towards the base becoming broadly falcate and from
that point narrowing towards the obtuse apex. Lip
ligulate to narrowly lanceolate, 5.3 – 10.1 × 1.1 – 2.9
mm, a little narrowed toward the middle, with more or
less pronounced lateral lobes near the base ranging
from minutely expanded through lobular to dentate,
tip sub-acute to attenuate; spur curved to sigmoid, 8.5 –
19.3 × 1.9 – 3.1 mm, parallel with the ovary first
becoming more pendent, tubular at the base then
expanded, then narrowing into a more or less curved
tip. Column 2 – 3.9 × 1.7 – 2.8 mm, with distinct rostellum
arms, the anther loculi subglobose, staminodes small,
slightly protruding below the lateral lobes of the
rostellum, rostellum deeply notched in the middle,
mid-lobe reduced to a small swollen blade, lateral arms
c. 1 mm long, thick, incurved, the caudicles of the
pollinia ending in a distinct viscidium. Fig. 13.
RECOGNITION. With its small mid-lobe of the rostellum
and distinct elongate anther canals, Cynorkis nutans
belongs in sect. Imerinorchis H.Perrier ex Hermans
(Hermans et al. 2007: 290). It is recognised by its large
leaf, tall hirsute inflorescence with a prominent lower
peduncle sheath, the ovary and flowers hirsute on the
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KEW BULLETIN (2020) 75: 50
exterior, the dorsal sepal and the petals forming a
hood over the column, the flat, spreading, asymmetric, broadly lanceolate to ovate lateral sepals, and
ligulate lip with small lobes at the base and a curved
spur which is thickened around the middle.
It is similar in habit and lip shape to Cynorkis
zaratananae Schltr. (Schlechter 1924: 75) but that
species has a longer leaf petiole, slightly smaller
flowers by c. 1 3 and shorter spur, inflated at the tip
(vs narrowing). It has a similar hirsute inflorescence
and flowers and a similar lip shape to C. rosellata
(Thouars) Bosser (1997: 188) and C. saxicola Schltr.
(Schlechter 1924: 70) but those species have several
leaves arranged in a rosette (vs single), slightly smaller
flowers with a shorter spur half the length of the ovary
(vs about the same length) which is thickened towards
the tip (vs narrowing).
DISTRIBUTION. Widespread in Madagascar, mainly in
the Eastern forest but also in the north and south, the
Comoros (on Ngazidja [Grande Comore], Nzwani
[Anjouan] and Mayotte). Records from Mauritius
and Réunion are referable to Cynorkis inermis
(Thouars) Hermans & P.J.Cribb (Hermans et al.
2017: 28).
SPECIMENS EXAMINED. MADAGASCAR. Andrangoloaka,
comm. 1880, Parker s.n. (K!); S Betsileo, Ankafina
forest, March 1881, Hildebrandt 4204 (W! R14896);
Imerina, rec. Nov. 1881, R. P. Campenon s.n.
(P00102291!); Imerina, 1881, Deans Cowan s.n.
BM000911499!); Central, comm. Oct. 1882, Baron
1703 (lectotype BM000032250!; isolectotype
K000415611!; P00102284!); April 1890, Scott Elliot s.n.
K000415612!; Mandraka valley, Oct. 1905, C. d’Alleizette
478 (P00102299!); Manongarivo massif, c. 1500 m,
May 1909, Perrier 1941 (P000102309!); Andringitra,
1600 m, Sept. 1911, Perrier 8005 (P00102310!); E,
Maningory basin, 700 m, Sept. 1912, Perrier 11377
(P00102311!); Analamazaotra, 1000 m, Aug. 1924,
Humbert 2266 (P00102307!); Mt Ambre, Sept. 1926,
Perrier 17733 (P00102296!, P102297!); S of
Fianarantsoa, 1000 m, Oct. 1926, Decary 5834
(P00102303!); East, between Moramanga & Anosibe,
eastern forest, on trunks amongst moss, c. 600 m,
Oct. 1927, Perrier 18028 (P00102312!, P00102313!);
Fort-Dauphin distr., Col de Manangotry, 600 – 700 m,
Aug. 1932, R. Decary 10355 (P00102286!); Matitantana,
Tsianovoha forest, Sept. 1934, Heim s.n. (P00102304!);
Andrangovalo massif, 1200 – 1400 m, Oct. 1937,
Humbert & Cours 17784 (P00102288-90!); S of
Moramanga, Sept. 1942, Decary 18419 (holo.
P00102285!); Montagne d’Ambre forest, 1000 – 1200
m, Oct. 1944, Homolle 44 (P00102305-6!); Marojejy
massif, 1000 m, Nov. – Dec. 1959, Humbert & Saboureau
31479 (P00102308!); Ankeramadinika, Aug. 1962,
Bosser 16214 (P00102300!); road to Lakato,
KEW BULLETIN (2020) 75: 50
Fig. 13. Cynorkis nutans. A & B plants in habitat; C & D rachises.
Page 19 of 25
50
PHOTOS: JOHAN HERMANS.
© The Author(s), 2020
50 Page 20 of 25
Aug. 1963, Bosser 17233 – 4 (P00102301 – 2!); Mt
d’Ambre, 1200 – 1300 m, Nov. – Dec. 1964, Morat 1211
(P!); W of Ranomafana, 900 – 1100 m, Sept. 1992,
Malcomber & Rakoto 1554 (G!, K!, MO); Montagne
d’Ambre, 1400 – 1475 m, April 1993, Malcomber et al.
22331 (K!, MO!); Fianarantsoa, Ranomafana S of
Ambohimiera, 880 – 1100 m, Sept. 1993, Kotozafy &
Randriananantena 240 (K!, MO); Antananarivo prov,
Ankazobe, Hort., 1993, Hermans 2550 (K!); Toamasina
prov., nr Maromiza, 1180 m, Jan. 2000, Hermans 5102
(K!); Toamasina prov., road to Lakato, 1043 m,
Feb. 2000, Hermans 5195 & 6615 (K!); Toamasina
prov., Andasibe area, 1229 m, Sept. 2000, Hermans
5022 (K!); Antsiranana, Andapa, Doany, Betsomanga,
Marojejy, 1250 m, Oct. 2001, Gautier et al. LG3942
(G!); Fianarantsoa, Atsimo-Atsinanana, SW Befotaka,
1228 m, Sept. 2005, Toamasina, Lakato, Manasamena,
1064 m, Sept. 2007, Razanatsima et al. 417 (MO, P!,
TAN!); Rakotovao et al. 2015 (MO, P!, TAN!); Hermans
5022 (K!); Toamasina prov., Andasibe area, Sept. 2007,
Hermans 6768 (K!); Antsiranana, Montagne d'Ambre,
1460 m, Nov. 2007, Gautier et al. 5113 (G!, K!, MO, P!,
TEF, WAG); Antsiranana, Daraina, forêt de Binara, 1015
m, Nov. 2005, Nusbaumer & Ranirison 1577 (G!); Alaotra
Mangoro, Ambatovy. Ampirarazona forest, 1072 m,
Sept. 2008, Antilahimena et al. 6523 (MO, P!, TAN!);
Montagne d'Ambre area, Hermans 2776 (K!); AlaotraMangoro, Ambatovy, 1141 – 1222 m, Sept. 2017,
Ambatovy Spirit Collection, 3264A7115 – 3268A7115 &
3711A6515 (TAN!); Maralambo, Herb. Inst. Sc. Mad. 822
(TAN!); Andapa, Duran 1039 (P00102287!); Montagne
d’Ambre, Saboureau s.n. (P00102314!); Mora, Anosibe,
Millot. 334 (TAN!). COMOROS. Mayotte, Humblot 209
(1209) (W! R5342); (P00102270!, P00102271!,
P00102272!, BM000034836!); Mayotte, Combani forest,
Sept. 1884, Humblot 1320 (BM!, W!); Grande Comore,
May 1886, Humblot 1520 (P00102273 – 5!) Grande
Comore, May 1886, Humblot 1520 (P00102273 – 5!);
Grande Comore, La Convalescence to Boboni,
Sept. 1958, Benson s.n. (BM000034808!, P00102276!);
Anjouan, Mt Tingui, Oct. 1959, Benson 145 (BM!, TAN!);
Grande Comore, Karthala Forest, 120 m, Sept. 1996,
Moulaert 019 (P!); Grand Comore, Karthala, 1095 m,
Aug. 2000, Brionnaud et al. 57 (CNDRS, K!, MO, P!);
Mayotte, Chirongui, Mont Chongui, Feb. 2008, Barthelat
et al. 1300 (P!); Karthala, piste de Boboni à Convalescence, Oct. 2008, 1200 – 1700 m, Pignal et al. 3350 (P!);
Hort., Hermans 4392 (K!).
HABITAT. Terrestrial, epiphyte or lithophyte in evergreen forest, montane forest, lichen forest, in humus
and moss, generally close to the ground on mossy tree
trunks when epiphytic. 600 – 1700 m.
CONSERVATION STATUS. Cynorkis nutans is widespread
from Northern to Southern Madagascar in Diana,
Sava, Analamanga, Alaotra-Mangoro, VatovavyFitovinany, Haute-Matsiatra and Anosy regions in
Antsiranana, Antananarivo, Toamasina and Toliara
© The Author(s), 2020
KEW BULLETIN (2020) 75: 50
provinces. It also occurs in the Comoros. The extent of
occurrence (EOO) is estimated to be largely more than
20,000 km2. In addition, the species is located in many
protected areas such as Analamazaotra-Mantadia,
Marojejy, Montagne d’Ambre, Ranomafana. Part of the
population may be threatened by habitat destruction
due to charcoal production, firewood collection,
agricultural clearing but these may not affect the
survival of this species. Cynorkis nutans is therefore
assessed as Least Concern (LC).
FLOWERING TIME. August to April but mainly September to December.
ETYMOLOGY. Campenoni refers to Pierre Campenon
(1844 – 1909), a Catholic missionary who found the
type of the variety; gaesiformis refers to the halberd —
gaesum spear-shape of the lip; galeata refers to the
helmet-shape of the joined dorsal sepal and petals;
nutans refers to the loosely nodding head of flowers as
perceived by Ridley; pallens is derived from pallidus
referring to the pale colour of the flowers; stenoglossa
refers to the narrow lip.
NOTES. The species was first described as Habenaria
nutans by Ridley in the Journal of the Linnean Society,
Botany, based on a paper read to the Society in
January 1885. The volume (21) itself is dated 1886
on the title-page but the following page lists the
actual dates of publication with Ridley’s paper
being effectively published on April 29th 1885. In
the protologue Ridley mentioned three different
collections from Madagascar: ‘Central, Baron no
1703; Imerina, Deans Cowan Herb. Brit. Mus. and
Andrangoloaka, Parker! Herb. Kew’. Later the same
year H. G. Reichenbach f. described Cynorkis galeata
in Flora, with the particular issue (31) dated 21
October 1885. Reichenbach cited specimen Humblot
209, collected on Mayotte. The holotype in W
consists of two plants accompanied by a sketch of
the flower and a detailed manuscript description by
Reichenbach; there are four other sheets at BM
and P. The P specimen is also labelled ‘1209’;
Humblot’s collection numbers were preceded by ‘1’
in label transcriptions, following a re-numbering by
Humbert in P. In 1893 Kraenzlin (: 63) described
C. stenoglossa, based on a Hildebrandt specimen
from Central Madagascar: the W specimen is
labelled in error as Arnottia inermis but seen by
Kraenzlin in 1938; it is likely that other material in
B was destroyed; there are several specimens in P
(Perrier 1941, 11377, 18028) annotated as syntypes,
in error. Ridley’s Habenaria nutans was transferred
to the genus Cynorkis by Perrier in 1936 (: 582). In
1939 Perrier referred to unpublished 1931 names
of C. nutans var. campenoni and C. stenoglossa var.
pallens, both being validated by Hermans in 2007.
The type material in P of C. stenoglossa var. pallens
consists of two sheets of Perrier 17733, clearly
labelled as ‘type’ (P00102296) and ‘isotype’
KEW BULLETIN (2020) 75: 50
Page 21 of 25
50
Table 4. Summary of measurements and characteristics of specimens of Cynorkis gaesiformis, C. galeata, C. nutans, C. nutans var.
campenoni, C. stenoglossa and C. stenoglossa var. pallens.
C. nutans & vars
C. stenoglossa & vars
15 – 25
15 – 25
oblong, 10 – 16 × 3 – 5
cm
17 – 20 cm,
hirsute
5–8×2
elliptic-elliptic, 10 – 14 × 2.5 –
3.8 cm
26 cm,
glandular-hirsute
5 – 9 × 1.5 – 2
oblong, 8 – 12 × 2.5 – 3 cm
5.2 – 3 mm,
hooded
9.2 – 11 × 4 – 5.2
mm, asymmetric
5 × 2.5
6 – 10 × 2 mm,
ligulate halberd
shape at base
c 15 × 1 mm,
thickened in the
middle
2 × 2.1
Madagascar:
Toamasina prov.
6 – 8.2 × 2.1 × 3.5 mm,
hooded
7.5 -12 × 4.2 – 6 mm,
semi-oval
6 – 8.6 × 2 – 3
6 – 10 × 1 – 2.1 mm,
ligulate, dentate at base
15 – 19 × 0.7 – 1 mm,
sigmoid, expanded in
the middle
3 – 3.5 × 2 – 3.2
Comoros
4.5 – 7.5 × 2 -3 mm,
hooded
7 – 14 × 4 – 7 mm,
semi-orbicular
4.5 – 8 × 2 – 4
5.6 – 9 × 1.5 – 2 mm,
ligulate, lobular at base
4.5 – 8 × 2 – 3 mm,
hooded
7.5 – 16 × 4 – 8 mm,
semi-orbicular
5–8×2–4
5 – 10 × 1 – 3.8 mm,
narrow ligulate with basal
teeth
8.5 – 14 × 1 mm,
expanded in the middle
600 – 800
Sept.
1095 – 1700
Aug. – Oct.
C. gaesiformis
Plant height
(cm)
Leaf
25 – 30
Inflorescence
32 cm,
hirsute
5×2
Floral bracts
(mm)
Dorsal sepal
Lateral
sepals
Petals (mm)
Lip
Spur
Column (mm)
Distribution
Altitude (m)
Flowering
time
oblong, 16 × 4 cm
C. galeata
25
(P00102297). In 1951 Perrier (: 146) described
C. gaesiformis from a collection by Decary in the
Madagascan Highlands, the type sheet was annotated in 1950 as ‘Cynosorchis caesumiformis sp. nov’. In
2005 Szlachetko transferred Cynorkis galeata,
C. nutans and C. stenoglossa to his segregate genus
Imerinorchis but did not include C. gaesiformis in the
transfer.
Several authors and botanists have already indicated some similarity between Cynorkis gaesiformis, C.
galeata, C. nutans and C. stenoglossa: the BM type
specimen of C. galeata (Humblot 209) was annotated
by Ridley as ‘Habenaria nutans Ridley!’; Kraenzlin
(1901: 920 & 923) listed ‘Cynorkis galeata Rchb.f. huc
Habenaria nutans Ridley’ under his Species subdubiae v.
dubiae, Bosser determined the Decary 18419 type
specimen of C. gaesiformis as C. nutans in 1968 and
Hervouet (2018: 299) remarked that C. galeata, C.
nutans and C. stenoglossa are to be compared.
Following field observations and comparing all the
available herbarium material identified under these
four names and their varieties it has become clear
that they are all part of one variable species with
Ridley’s C. nutans having priority over Reichenbach’s
C. galeata by just a few months, C. gaesiformis and
C. stenoglossa are later additions of 1893 and 1951
respectively. Table 4 shows the great range in size of
plant and flower segments of the different species
and varieties but also indicates their overlapping
11 – 20 × 1 – 1.2 mm, curved,
expanded in the middle
2 – 4 × 1.5 – 2
Madagascar: Antananarivo,
Antsiranana, Fianarantsoa,
Toamasina & Toliara
provinces. Comoros
600 – 1460
Aug. – April
6 – 12 cm,
glandular-hirsute
5 – 9 × 1.5 × 2
2–4×2
Madagascar: Antananarivo,
Antsiranana, Fianarantsoa,
Toamasina provinces
600 – 1300
Aug. – March
characteristics; field observations have also shown
great variations within colonies. Plants consistently
have a single (rarely two), comparatively large leaf, a
long hirsute inflorescence, an hirsute ovary and
exterior of the flower, broad flat lateral sepals which
are spreading and asymmetric, a ligulate lip which
has variable but always small lobes at the base, a
more or less sigmoid spur which is thickened in
the middle and a rostellum with a reduced midlobe and separate incurved arms. Colour of the
flowers is also very variable and may have led to
some of the proliferation of names; field observations and photographic records have shown that
there are a number of consistently different colour
forms but it has also shown that the diverse colour
variations are mixed within colonies and localities
(see Fig. 13).
Following its general plant and flower morphology, Cynorkis inermis (Thouars) Hermans & P.J.Cribb
(Hermans et al. 2017: 28) from Mauritius and
Réunion could also be considered part of this species
but the lack of a spur makes it distinct. A few plants
of C. inermis have been found on Réunion that have a
short spur or remnant thereof (Bernet 2010: 122 &
pers. comm. and Szelengowicz & Tamon 2013: 277)
but the lack or reduction of a spur indicates a
change in pollinator interaction and its endemism
to the islands of Mauritius and Réunion warrant its
recognition as a distinct species. The genetics and
© The Author(s), 2020
50 Page 22 of 25
Fig. 14. Cynorkis mammuthus. A & B plants in habitat; C rachis.
© The Author(s), 2020
KEW BULLETIN (2020) 75: 50
PHOTOS: ANTON SIEDER.
KEW BULLETIN (2020) 75: 50
nomenclature of this species are discussed in
Hermans et al. (2017: 27).
Notes on Cynorkis mammuthus
Cynorkis mammuthus Hermans & P.J.Cribb (Hermans
et al. 2017: 13), described recently from northern
Madagascar, was based on a herbarium specimen,
without much information, collected 25 years ago
and consisted of part of an inflorescence only. In
March 2019, a team from the University of Vienna
and the Parc Botanique et Zoologique de
Tsimbazaza Antananarivo, visited localities in N
Madagascar and discovered it flowering in the
Antsohihy area, not far from its type locality. This
gives the opportunity to add more information to the
description of the species, especially details about the
plant, colour of the flowers and the lengthening of
the spur.
Cynorkis mammuthus Hermans & P.J.Cribb (Hermans
et al. 2017: 13). Type: Madagascar, Mahajanga prov.,
road to Antsohihy, near Anjalazala, fl. Feb. 1994,
Hermans 2699 (holotype K!).
Erect terrestrial or lithophytic plant, up to 75 cm high,
including the inflorescence. Tubers 2 – 3, elongate,
clustered, surface velvety; roots a little hairy, fleshy,
wiry c. 2 mm in diam. Stem elongate, dark green, c. 10
– 20 cm long, 8 – 18 mm in diam., enclosed by 1 – 2
leaf-like sheaths and the basal leaf petiole; 3 – 4 large
distant opposite leaves, followed by 2 – 4 leaf-like
sheaths above, decreasing in size. Leaves obliquely
lanceolate, narrowed at the base into a long funnelshaped petiole annular at the base, with a strong midvein and 2 – 3 parallel veins, 5 – 12 cm long, 2.5 –
6.5 cm wide, leaves and leaf sheaths pale green
spotted with brownish-maroon. Inflorescence emerging
from the centre of the developed growth, dark green
becoming paler towards the rachis, hirsute towards
the apex, up to 40 cm, 1.9 – 3.5 mm in diam., with 2 –
3 lanceolate leaf-like sheaths strongly amplectant
towards the base, caudate at the tip, c. 2 – 4.5 cm
long, 1 cm wide; rachis up to 20 cm, densely hirsute,
somewhat ridged, lengthening as the flowers open,
laxly 11 – 20-flowered; floral bracts funnel-shaped,
serrate-hirsute at the margin, dorsally keeled, acuminate, then apiculate at the apex, about ¼ the length of
the ovary 21 – 28 mm long, 8 – 14 mm wide; pedicel
with ovary densely hirsute, almost straight, 60 – 72 mm
long, 1.8 – 2.2 mm wide. Flowers overall c. 4.5 – 6 cm
long, 2.9 – 4 cm wide, not including the spur, at first
yellowish-green then becoming pure white on opening, tips of the rostellum arms pale pink, pollinia
Page 23 of 25
50
caudicles brown. Dorsal sepal 32 – 37 × 14 – 18 mm
when flattened, forming a hood with the petals
arching over the column, gibbose, expanded and
hirsute at the base. Lateral sepals spreading, oblongelliptic with a distinctive notched wing on the upper
margin towards the tip, 29 – 35 × 12 – 15 mm. Petals
appressed to the inside of the dorsal sepal margins,
linear-lanceolate, 31 – 35 × 2.5 – 2.9 mm. Lip entire,
oblanceolate, naviculate, acuminate at the tip, 36 – 41
× 12 – 16 mm; spur down-curved then pendent,
subulate, lengthening with age 10.5 – 27 cm long, c.
2 mm in diam. Column in an angular plane 33 –
36 mm long, 9 – 12 mm wide, with two very long
rostellum arms for more than half its length, a small
nose-like mid-lobe, staminodes towards the base,
forming triangular auricles, anther loculi ovate,
pollinia including the long caudicle c. 26 mm long.
Fruit fusiform, densely hirsute, c. 6 × 0.5 cm. Fig. 14.
SPECIMENS EXAMINED . MADAGASCAR . Mahajanga
prov., road to Antsohihy, fl. Feb. 1994, Hermans
2699 (holotype K!); Befandriana, Feb. 1942, Jard.
Bot. Tana 6242 (P!); near Antsohihy, 257 m, 22
March 2019, T. & C. Sieder, D. Prehslet, W. Knirsch &
J. Andriantiana 7849 (W!).
HABITAT. In full sun on rock, in a thin layer of humus
and moss. Short evergreen scrubland with Kalanchoe
spp. and other geophytes. It is interesting to note
that the spotted foliage of Cynorkis mammuthus is
somewhat similar to that of Kalanchoe gastonis-bonnieri
R.Hamet & H.Perrier (1912: 364) (Crassulaceae),
with which it grows together.
CONSERVATION STATUS. Cynorkis mammuthus is endemic to Madagascar, restricted to the Sofia region in
Mahajanga province at low elevation. It is only
known from a few individuals, the Area of Occupancy is estimated to be less than 500 km2, in three
unprotected areas. The species is threatened by
habitat degradation due to frequent fires. Cynorkis
mammuthus is therefore assessed as Endangered EN,
according to the IUCN Red List Categories and
Criteria. Further research is needed to check the
population size and status.
Acknowledgements
We are most grateful to the directors and staff at BM,
BR, BRLU, DBN, G, HEID, K, M, MO, P, SZU, TAN,
TEF, W, WU, M, SZL, ZSS and their relevant
libraries. We would like to thank the anonymous
reviewers for their constructive suggestions and
corrections, Judi Stone for the detailed drawings,
Julian Shaw for taxonomic advice and Michael Kiehn
for his support. We are also very grateful to Solo
© The Author(s), 2020
50 Page 24 of 25
Rapanarivo, head of the flora department at PBZT
Antananarivo, for his permission to use the collection at the PBZT herbarium and for providing the
necessary collecting authorisation.
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this licence, visit http://creativecommons.org/
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References
Baillon, M. H. (1879). Sur Quelques genres de
Rubiacées dont la place est douteuse. Bull. Mens.
Soc. Linn. Paris 1: 198 – 200.
Bernet, P. (2010). Orchidées de la Réunion. Naturae
Amici éditions.
Bosser, J. (1969). Contribution à l'Étude des Orchidaceae
de Madagascar X. Adansonia 9: 343 – 359.
____ (1980). Contribution à l'Étude des Orchidaceae
de Madagascar. Adansoni, 20: 257 – 261.
____ (1997). Contribution à l’étude des Orchidaceae
de Madagascar et des Mascareignes. XXVII.
Adansonia 19: 181 – 188.
____ (1998). Contribution à l’étude des Orchidaceae
de Madagascar et des Mascareignes. XXVIII.
Adansonia 20: 281 – 283.
____ (2002). Contribution à l’étude des Orchidaceae
de Madagascar et des Mascareignes. XXXII.
Adansonia 24: 21 – 25.
____ (2015). Description de trois Cynorkis Thouars
nouveaux (Orchidaceae) du Nord de Madagascar.
Adansonia 37: 19 – 23.
____ & Lecoufle, M. (2011). Les Orchidées de Madagascar. Biotope, Mèze.
____ & Veyret, Y. (1970). Contribution a l'Étude des
Orchidaceae de Madagascar XIII. Adansonia 10:
213 – 217.
Cribb, P. & Hermans, J. (2009). Field Guide to the
Orchids of Madagascar. Kew Publishing, Royal
Botanic Gardens, Kew.
© The Author(s), 2020
KEW BULLETIN (2020) 75: 50
Durand, T. A. & Schinz, H. (1895). Conspectus Florae
Africae 5. Charles van der Weghe, Bruxelles.
Frappier C. de Monbenoist (1880). Orchidées de l'Île de
la Réunion, Catalogue des Espèces Indigènes Découvertes
Jusqu'à ce Jour. Réunion.
____ in J. E. de Cordemoy (1895). Flore de l'Ile de la
Réunion. Orchidées: 165 – 262. Klincksieck, Paris.
Govaerts, R., Bernet, P., Kratochvil, K., Gerlach, G.,
Carr, G., Alrich, P., Pridgeon, A. M., Pfahl, J.,
Campacci, M. A., Holland Baptista, D., Tigges, H.,
Shaw, J., Cribb, P., Kreuz, K. & Wood, J. (2019).
World Checklist of Selected Plant Families. Facilitated by
the Royal Botanic Gardens, Kew. Published on the
Internet: http://apps.kew.org/wcsp/.
Hamet, R. & Perrier, H. (1912). Contribution a l’étude
des Crassulacées Malgaches. Ann. Sci. Nat. Bot. 10:
361 – 377.
Hermans, J., Andriantiana, J. L., Sieder A., Kiehn,
M., Cribb, P., Rajaovelona, L. & Gardiner, L.
(2017). New species and nomenclatural changes
in Cynorkis (Orchidaceae: Orchidoideae) from
Madagascar and the Mascarenes. Kew Bull. 72,
38: 1 – 31.
____ & Cribb, P. J. (2007). Cynorkis in Madagascar.
Orchid Rev. 115: 248 – 255.
____ & ____ (2014). New species and new names in
Malagasy orchids (Orchidaceae). Kew Bull. 69, 9517:
1 – 16.
____, Hermans, C., Du Puy, D., Cribb, P. & Bosser, J.
(2007). Orchids of Madagascar 2nd edition. Kew
Publishing. Royal Botanic Gardens, Kew.
____ & Rajaovelona, L. (2019). A new Natural Hybrid
in Cynorkis from Madagascar. Amer. Orchid Soc. Bull.
88: 547 – 550.
Hervouet, J-M. (2018). A la recherche des Orchidées de
Madagascar. Biotope, Mèze.
Humbert, H. (1939). Flore de Madagascar. 49e. Famille. –
Orchidées. Tananarive Imprimerie Officielle, Madagascar.
IUCN (2012). IUCN Red List Categories and Criteria:
Version 3.1. 2nd edition. Gland and Cambridge.
Kraenzlin, F. (1893). Orchidaceae Africanae VII. Bot.
Jahrb. Syst. 33: 53 – 75.
____ (1898 – 1901). Orchidacearum Genera et Species. 1.
Mayer & Mueller, Berlin.
Lindley, J. (1835). Orchideae Tribe IV. Ophrydeae. Genera
and species of Orchidaceous plants. Ridgeways, London.
Perrier de la Bâthie, H. (1930). Catalogue des Plantes de
Madagascar publié par L'Académie Malgache.
Orchidaceae d'après R. Schlechter. Colonie de Madagascar et Dépendances, Paris.
____ (1931, unpublished). Arch. Bot. Bull. Mens. 5: 36, 48,
72.
____ (1936). Notes sur quelques Habenaria de Madagascar. Bull. Soc. Bot. France 83: 579 – 585.
KEW BULLETIN (2020) 75: 50
____ (1939). Orchidaceae. In: H. Humbert, Flore de
Madagascar. 49e. Famille. – Orchidées. Tananarive
Imprimerie Officielle, Madagascar.
____ (1951) Orchidées de Madagascar et des
Comores. Notul. Syst. (Paris) 14: 138 – 165.
Reichenbach, H. G. f. (1885). Comoren-Orchideen
Herrn Léon Humblot’s. Flora 68: 377 – 382.
____ (1888). Orchideae describuntur. Flora 71: 535 –
544.
Ridley, H. N. (1883). Descriptions and Notes on
new or rare Monocotyledonous Plants from
Madagascar, with one from Angola. J. Linn.
Soc., Bot. 20: 329 – 338.
____ (1885). The Orchids of Madagascar. J. Linn. Soc.,
Bot. 21: 456 – 523.
____ (1886). On Dr. Fox's collection of Orchids from
Madagascar, along with some obtained by the Rev.
R. Baron from the same Island. J. Linn. Soc. Bot. 22:
116 – 127.
Schlechter, R. (1913). Orchidacées de Madagascar.
Orchidaceae Perrieranae Madagascarienses. Ann.
Mus. Colon. Marseille, sér. 3, 1: 148 – 202.
____ (1915). Kritische Aufzählung der bisher von
Madagaskar, den Maskarenen, Komoren und
Seychellen bekantgewordenen Orchidaceen. Beih.
Bot. Centralbl. 33: 390 – 440.
____ (1924). Orchidaceae Perrieranae. Repert. Spec.
Nov. Regni Veg. Beih. 33: 1 – 240.
Szelengowicz, M. & Tamon, J. M. (2013). Les Orchidées
des Mascareignes, Printec. Seychelles.
Page 25 of 25
50
Szlachetko, D. (2005). Habenaria — Beitrage zu einer
Revision der Subtribus (12). Orchidee (Hamburg)
56: 68 – 70.
____ & Kras, M. (2006). Matériaux pour la révision des
Habenariinae (Orchidaceae, Orchidoideae) – 6.
Richardiana 6: 139 – 146.
Thouars, A. du Petit (1822). Histoire particulière des
Plantes Orchidées recueillies sur les trois Îles Australes
d’Afrique. Bertrand, Treuttel & Wurtz, Paris.
Toilliez-Genoud, J. & Bosser, J. (1961). Contribution à
l’étude des Orchidaceae de Madagascar et des
Mascareignes. IV. Adansonia 1: 100 – 105.
____ & ____ (1962). Contribution à l’étude des
Orchidaceae de Madagascar et des Mascareignes.
III. Naturaliste Malgache 13: 25 – 30.
Turland, N. J., Wiersema, J. H., Barrie, F. R., Greuter,
W., Hawksworth, D. L., Herendeen, P. S., Knapp, S.,
Kusber, W.-H., Li, D.-Z., Marhold, K., May, T. W.,
McNeill, J., Monro, A. M., Prado, J., Price, M. J. &
Smith, G. F. (eds) (2018). International Code of
Nomenclature for algae, fungi, and plants
(Shenzhen Code) adopted by the Nineteenth
International Botanical Congress Shenzhen, China,
July 2017. Regnum Veg. 159. Koeltz. Glashütten. DOI
https://doi.org/10.12705/Code.2018.
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