Introduction

The genus Cynorkis Thouars (Orchidaceae: Orchidoideae) comprises around 170 species in total, 120 of which are largely endemic to Madagascar (Hermans et al. 2007), with about 30 endemic to the surrounding islands (Mascarenes, Comores) and fewer than 20 species restricted to mainland Africa (Govaerts et al. 2016). Recently, while exploring an area in Mahajanga province in Northern Madagascar, a team from the Botanical Gardens of the University of Vienna and PBZT Antananarivo discovered six large and attractive new Cynorkis, growing together with other already known species. Only a handful of species have been previously recorded from this region. Earlier collectors, such as Baron and Hildebrandt in the 19th century, very rarely ventured there, while more recent French explorers, particularly Perrier de la Bâthie and Humbert, only visited the area occasionally and not necessarily during the wet and short flowering season when travel is difficult (Dorr 1997 and herbarium material). As part of ongoing research on the orchid flora of Madagascar the material was compared with descriptions, herbarium material and drawings of all the type specimens of the genus to ascertain their novelty. One discovery, C. christae, has even larger flowers than C. gigas Schltr. (Schlechter 1924: 50) which until now was the most impressive species in the genus.

A further four new Cynorkis are newly described from Madagascar, together with a comparison of C. elegans Rchb. f. and C. elata Rolfe and a re-evaluation of the taxonomic status of Arnottia A. Rich. and Physoceras Schltr., both genera now being included in Cynorkis.

The sectional treatment of Cynorkis proposed by Perrier de la Bâthie (1939: 73 – 74) and based on column structure is in need of comprehensive revision and therefore the new species are left unplaced at sectional level; here we indicate the sections of the species that are compared with our new taxa, for reference only. A detailed DNA and morphological analysis of Cynorkis is necessary before a new meaningful classification can be proposed.

IUCN Red List assessments

The conservation status of the new species given in this paper are summaries of the full IUCN Red List assessments for these species which will be completed and submitted for review and publication by IUCN once the species names are validly published and therefore available for assessment. The assessments have been compiled based on current knowledge of these taxa, by two of the authors (LR and LG), who are IUCN Red List assessors, using the IUCN Red List Categories and Criteria (2012).

New Species

Cynorkis aconitiflora Hermans, Andriant. & Sieder sp. nov. Type: Madagascar, Mahajanga prov., road to Bealanana, 13 Jan. 2016, 918 m, Sieder A. & C. & Andriantiana 6964 (Hermans 7983) (holotype WU!, isotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:77164799-1

Erect to semi-erect terrestrial herb, up to 40 cm tall, including the inflorescence. Tuber characteristics not known. Stem up to 4 cm long, 5 – 6 mm in diam., enveloped by a thin brown basal sheath, up to 33 mm long, 7 mm wide, with a pair of almost opposite leaves above. Leaves pale green, more or less spotted with dark red, margins of young leaves purple, ovate-lanceolate, acuminate, up to 15 cm long, 4 cm wide. Inflorescence pale green, glabrous, spotted purple-red longitudinally, emerging from the centre of the leaf petiole, up to 35 cm, c. 4 mm in diam., 2 – 3 lanceolate sheaths enveloping the peduncle, c. 20 mm long; rachis 8 – 12 cm, densely 12 – 26-flowered, unilateral; floral bracts amplexicaul at the base, acuminate, apiculate at the apex, green with a purple margin and tip, 12 – 16 mm long, 2.5 – 3.5 mm wide; pedicel and ovary glabrous, 12 – 14 mm long, 1.5 – 1.7 mm wide, pale green with pale purple bands, curved. Flowers overall c. 11 mm long, 10 mm wide, glabrous, petals and sepals purple, the petals a little paler, lip pale purple with a slightly darker longitudinal central ridge, the base and part of the lateral lobes white, spur paler purple, rostellum white. Dorsal sepal forming a tight hood with the petals, deeply concave, base inflated, the apex slightly recurved, 6.8 – 7.1 mm long, 3.3 – 3.6 mm wide; lateral sepals spreading and somewhat recurved, oblong-obovate 8.9 – 9.2 mm long, 4.9 – 5.2 mm wide. Petals appressed to the inside of the dorsal sepal margins, narrowly oblong-obovate, the margins a little undulate, 7.3 – 7.5 mm long, 2.0 – 2.2 mm wide. Lip 3-lobed, overall 10.4 – 10.7 mm long, 5.3 – 5.9 mm wide, the lateral lobes small, triangular, curved upwards, mid-lobe linguiform, concave with a longitudinal raised central ridge; spur basally infundibuliform and parallel with the ovary then narrowed, somewhat descending and rounded, 8.1 – 8.3 mm long, 1.5 – 1.6 mm in the middle. Column in a horizontal plane 4.1 – 4.9 mm long 3.1 – 3.3 mm wide, with two long rostellum arms surpassing the nose-like mid-lobe by double the length of the rostellum, staminodes short, rounded, verrucose; anther loculi erect; pollinia including the thin caudicle c. 4 mm long. Figs 1 and 2.

Fig. 1
figure 1

Cynorkis aconitiflora. A habit; B inflorescence. photos: Johan Hermans.

Fig. 2
figure 2

Cynorkis aconitiflora. A habit (from photograph); B large inflorescence (from photograph); C flower, lateral view; D flower, face view; E dorsal sepal and petal, lateral view; F dorsal sepal, inner face; G petal, inner face; H lateral sepal, inner face; J labellum; K labellum, transverse section; L column, in situ, lateral view; M column, ¾ dorsal view. From Sieder 6964 (spirit material & field photographs). *approximate scale. drawn by andrew brown.

recognition

The new species is characterised by the two-leaved plant with dark spots on the young leaves, peduncle and long floral bracts. The dense unilateral raceme carries up to 26 flowers. The medium-sized glabrous flowers have a distinct 3-lobed lip with the basal lobes small and triangular, the mid-lobe tongue-shaped with a thick central ridge, the spur which is at first wide and then narrows into a thinner tube. With its secund rachis and 3-lobed lip the new species bears some similarity to Cynorkis lilacina Ridl. (Ridley 1885: 515) in section Hemiperis, from highland Madagascar but it has two larger leaves, longer floral bracts, glabrous rather than hirsute flowers and a lip of different proportions (up to 8 × 7 mm, vs 10 × 6 mm in the new species) with upturned side lobes. It is similar to C. lemurica Bosser (2015: 20) in section Gibbosorchis from the same area but at a higher elevation than the new species. It also differs in its ovate-lanceolate rather than linear leaves, a unilateral rachis, floral bracts almost as long as the ovary, a larger lip (up to 7.5 × 5 mm in C. lemurica vs up to 10.7 × 5.9 mm in the new species), verrucose staminodes and purple rather than whitish-pink flowers.

distribution

So far only known from a small locality in the Antsahabe area in Northern Madagascar.

specimens examined

madagascar. Mahajanga prov., on the road to Bealanana, in the understory between stones, 918 m, 13 Jan. 2016, Sieder A. & C. & Andriantiana 6964 (Hermans 7983) (holotype WU!, isotype K!).

habitat

Terrestrial in the shade of small trees and ferns amongst rocks in seasonally very dry forest remnants within grassland on basement rock, around 900 m.

conservation status

This species is likely to be Critically Endangered (CR), according to the IUCN Red List Categories and Criteria. It is only known from a single individual, at a single unprotected locality, in a forest fragment alongside a road leading to a remote town in the NE of the country — such fragments are known to be particularly at risk from human development. The species is CR based on criterion B2ab(iii) (area of occupancy likely to be less than 10 km2, number of locations = 1, and continuing decline in the quality of habitat inferred).

etymology

The species name refers to the resemblance of flower shape and colour to the Monkshood, Aconitum napellus L. (Ranunculaceae).

Cynorkis christae Hermans, Andriant. & Sieder sp. nov. Type: Madagascar, Mahajanga prov., road to Bealalana, between Antsahabe and Ambatosia, 598 m, 23 Jan. 2016, Sieder A. & C. & Andriantiana 7028 (Hermans 7980) (holotype WU!, isotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:60474976-2

Pendent or arching terrestrial herb, up to 56 cm tall including the inflorescence. With two elongate tubers clustered at the stem base, c. 6 × 2 cm, roots 2 – 3 mm in diam. Stem, combined with the leaf petiole up to 9 cm long, subtended by 1 – 2 funnel-shaped sheaths, with a large solitary arching radical leaf. Leaf glabrous, pale to mid-green, glossy above, lanceolate-elliptic, acuminate and a little narrowed towards the base, obtuse apically, up to 18 cm long, 9 cm wide, probably larger when mature. Inflorescence up to 40 cm long, 2.3 – 3.1 mm in diam., green with a few brown marks towards the base, glabrous, emerging from the base of the unfurling leaf, with a large ovate leaf-like peduncle sheath in the upper third, enveloping the peduncle in the lower half then expanded and acuminate, 6.1 – 7.1 cm long, 2.8 – 3.6 cm wide; with 3 – 4 flowers at the apex, opening in succession; floral bracts green, similar in shape to the peduncle sheath, apiculate, 25 – 29 mm long, 8 – 11 mm wide; pedicel and ovary almost straight, up to 45 mm long, 3 mm wide, hirsute, yellowish-green. Flowers very large for the genus, strongly and pleasantly scented, overall 6 – 6.8 cm long, 4.5 – 6.2 cm wide, not including the spur; sepals greenish-white with the veins green, the interior marked with purple red, the exterior densely papillose, petals white, lip white, spur pale green, column yellowish white, anther loculi pale yellow. Dorsal sepal elliptic-oblong, subacute, concave, together with the petals, forming a hood over the column, 30 – 33 mm long, 15 – 19 mm wide; lateral sepals spreading, oblong-ligulate, somewhat concave, 31 – 33 mm long, 12 – 13 mm wide. Petals lanceolate-ligulate acute, 26 – 28 mm long, 6.0 – 6.5 mm wide, the anterior margin expanded at the base. Lip base elongated and connate to the base of the column, first narrowly angular then expanded and 3-lobed; with two large oblong, spreading lateral lobes, the mid-lobe narrowed at the base, bifid, the divisions oval-obtuse, sometimes with a small appendage in between, disk, with a longitudinal swelling in the middle, overall 45 – 47 mm long, 47 – 49 mm wide; spur at first shortly angular to the lip base then pendent, indistinctly swollen in the lower quarter, then narrowed at the tip, 16 – 19 cm long, 2.8 – 3.1 mm in diam. Column in a horizontal plane 26 – 28 mm long, 10 – 11 mm wide, with two parallel rostellum arms for a quarter its length, mid-lobe forming a nose-like protrusion, staminodes elongate, hidden below the rostellum; pollinia 2, composed of a small disc-like viscidium and a long needle-like stipe up to 27 mm long running in a groove along the column, the pollen masses enclosed in a pair of anther loculae in the base of the hood of the column. Figs 3 and 4.

Fig. 3
figure 3

Cynorkis christae. A habit; B with co-finders Jacky Andriantiana and Christa Sieder; C flowers; D labellum. photos: A, C, D: Johan Hermans, B: ANTON SIEDER.

Fig. 4
figure 4

Cynorkis christae. A habit (from photograph); B flower, ventro-lateral view (from photograph); C flower, face view (from photograph); D flower, lateral view; E dorsal sepal, inner face; F petal, inner face; G lateral sepal, inner face; H labellum; J terminal lobes of another labellum; K column, in situ, lateral view; L underside of column to show (concealed) staminode auricles; M glandular hairs from dorsal sepal. From Sieder 7028/6977, spirit material, field photographs. drawn by andrew brown.

recognition

This very large-flowered species is instantly recognisable by the broad 5 cm white lip and long pendent spur which can be over 19 cm long, the large single leaf and the 3- – 4-flowered inflorescence with a large distinct peduncle sheath in the upper third. It differs from Cynorkis gigas Schltr. (Schlechter 1924: 50), in section Gibbosorchis, in having an unspotted leaf, a longer 40 cm long, 2 – 4-flowered inflorescence, shorter bracts, \( \raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right. \) the length of the ovary, a lip with white rounded side lobes and a 16 – 19 cm long spur swollen in the apical third. C. gigas is the only species with which it could be confused but the differences in morphology and distribution set them well apart, as shown in Table 1.

Table 1. Comparison of Cynorkis christae and C. gigas

distribution

Known from two neighbouring localities in the Mahajanga province of Northern Madagascar.

specimens examined

madagascar. Mahajanga prov., road to Bealalana, between Antsahabe and Ambatosia, 598 m, on wet rocks, 23 Jan. 2016. Sieder A. & C. & Andriantiana 7028 (Hermans 7980) (WU!, K!); between Antsahabe and Ambatosia, on a slope, in pockets of humus on wet rock, 13 Jan. 2016, 633 m, Sieder A. & C. & Andriantiana 6977 (paratype WU!).

habitat

Seasonally dry grassland on basement rock, water seepage areas on basalt rock, in crevices and shallow pockets of humus, around 600 m, in full sun. Growing sympatrically with Cynorkis lentiginosa and C. siederi.

conservation status

This species is likely to be Critically Endangered (CR), according to the IUCN Red List Categories and Criteria. It is known from two small subpopulations a couple of kilometres apart, in unprotected grassland alongside a road leading to a remote town in the NE of the country — such areas are known to be at high risk from human development. The species is CR based on criterion B2ab(iii) (area of occupancy likely to be less than 10 km2, number of locations = 1, and continuing decline in the quality of habitat inferred).

etymology

Named for Christa Sieder, co-finder of the new species.

notes

The species has the largest flowers in the genus Cynorkis, exceeding C. gigas Schltr. (Schlechter 1924: 50) in size, to which it seems to be closely related. C. gigas was not described until 1924: it had been collected earlier but was confused (Hermans & Cribb 2014: 13) with C. uniflora Lindl. (Lindley 1835: 331). It is somewhat astonishing that our new species has remained undiscovered until now, given its large showy flowers. This may be explained by the somewhat remoteness of the area in northern Madagascar where it was found; 19th century exploration was mainly concentrated in the Highlands around missionary posts, whereas prolific French collectors only occasionally ventured to these parts.

Cynorkis elephantina Hermans, Andriant. & Sieder sp. nov. Type: Madagascar, Mahajanga prov., Anjiamangirana, Andeboka near Antsohihy, 319 m, 25 Jan. 2016, Sieder A. & C. & Andriantiana 7035 (Hermans 7984) (holotype WU!, isotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:60474977-2

Erect terrestrial or lithophytic herb, up to 35 cm tall, including the inflorescence. Tubers elongate, clustered, surface velvety, roots a little hairy, fleshy, wiry c. 2 mm in diam. Stem pale green, short, 4 cm long, 8 – 12 mm in diam., enclosed by 1 – 2 leaf-like sheaths and the leaf petiole; 1 – 2 large leaves followed by several leaf-like sheaths above, decreasing in size, small groups of plants often clustered together creating a rosette-like formation. Leaves obliquely lanceolate, narrowed at the base into a short petiole, with a strong mid-vein and 2 – 3 parallel veins, 18 – 23 cm long, 6 – 9 cm wide, leaves and sheaths pale green blotched with dark red, glossy above, paler below. Inflorescence emerging from the developing growth, pale green becoming paler towards the rachis, somewhat ridged, hirsute, flecked with purple-red, up to 29 cm, 1.8 – 3.5 mm in diam., a large lanceolate leaf-like sheath c. 4 cm above the leaf, of a similar colour pattern as the leaf, c. 8 cm long, 3 cm wide, followed by another smaller leaf-like sheath and two or more much smaller sheaths along the peduncle; rachis 8 – 18 cm, lengthening as the flowers open, laxly 15 – 22-flowered; floral bracts pale green blotched purple-red towards the base, amplexicaul at the base for ½ its length, serrate-hirsute at the margin, acuminate, then apiculate at the apex, less than ½ the length of the mature pedicellate ovary 10 – 15 mm long, 2.5 – 3.3 mm wide; pedicel and ovary almost straight, 22 – 27 mm long, 1.8 – 2.3 mm wide, densely hirsute, pale green flecked with purple-red longitudinally. Flowers overall c. 4 cm long, 5.5 cm wide, not including the spur, all segments greenish white, becoming almost pure white when mature, sepals and petals with a few glandular hairs, spotted and flecked on the exterior with dark-purple, auricle on the lateral sepal margin darker, spur purplish-white becoming darker towards the tip, the base streaked longitudinally with purple, column entirely white. Dorsal sepal forming a hood with the petals, strongly gibbose, the apical \( \raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right. \) prominently recurved, 15 – 16.2 mm long, 7 – 7.8 mm wide when flattened; lateral sepals spreading and slightly twisted towards in the apical half, spathulate somewhat concave, with a distinctive auricle on the upper margin towards the tip, 16 – 17 mm long, 6.0 – 6.2 mm wide. Petals appressed to the inside of the dorsal sepal margins, almost scimitar-shaped, curved apically, 14 – 14.5 mm long, 2.1 – 2.2 mm wide. Lip entire, obovate-lanceolate, strongly curved backwards from the middle with the tip almost reaching the base of the ovary, 20.1 – 21.2 mm long, 5 – 5.2 mm wide, concave with a central raised longitudinal swelling; spur down-curved then pendent, lengthening greatly as the flowers develop, subulate towards the end, 70 – 78 mm long, c. 1.5 in diam. in mature flowers. Column in an angular plane 16 – 17 mm long 1.5 – 1.8 mm wide, with two long distinct rostellum arms for almost ½ its length, a rounded swelling half way along their length, a small nose-like mid-lobe, staminodes towards the base, forming triangular auricles, anther loculi elongate, longer than wide, pollinia including the long caudicle c. 15 mm long. Figs 5 and 6.

Fig. 5
figure 5

Cynorkis elephantina. A habit; B inflorescence; C flower. photos: anton sieder.

Fig. 6
figure 6

Cynorkis elephantina. A habit (from photograph); B flower bud in situ, lateral view; C flower, lateral view; D flower, face view (from photograph); E indumentum on peduncle; F indumentum on distal portion of ovary; G dorsal sepal and petal, lateral view; H dorsal sepal, inner face; J petal, inner face; K labellum, flattened, with transverse section to show callus; L lateral sepal, inner face, flattened; M column in situ, lateral view; N underside of column to show staminode auricles; P ovary, transverse section. From Sieder 7035, spirit material. drawn by andrew brown.

recognition

The new species is distinct by the large glossy spotted leaves, large leaf-like sheaths, the entire recurved lip with a long pendent spur which almost quadruples in length as the flower develops, recurved hood, lateral sepals with an auricle towards the apical margin and the long tusk-like rostellum arms. It is a very distinct, large-flowered species that somewhat resembles Cynorkis gibbosa Ridl. (Ridley 1883: 331) in section Gibbosorchis, in its leaf and floral morphology but is distinguished by its white flowers (vs brick red in C. gibbosa) with an entire lip (vs strongly lobed), much longer spur and distinct rostellum arms. Few larger-flowered Cynorkis species have an entire lip.

distribution

Only known from the Antsohihy area in Northern Madagascar.

specimens examined

madagascar. Mahajanga prov., Anjiamangirana, Andeboka, near Antsohihy, 319 m, 25 Jan. 2016, between basalt rock, humus and many geophytes, Sieder A. & C. & Andriantiana 7035 (Hermans 7984) (holotype WU!, isotype K!).

habitat

Seasonally dry forest remnants within grassland on basement rock at c. 300 m altitude. Water seepage areas on basalt rock, or near seasonal streams, in crevices and pockets of humus. In shade and semi-shade, with ferns, Euphorbia guillauminiana Boiteau and E. moratii Rauh.

conservation status

This species is currently considered to be Least Concern (LC), according to the IUCN Red List Categories and Criteria. It is known from many individuals, at a single unprotected locality, in the shade on basaltic rock, but appears to be distant from the nearest human settlements and roads, no obvious imminent threats are apparent. Further research is needed to check the population status, threats and habitat.

etymology

The species name refers to the likeness of the flower to the head of an elephant with the rostellum arms as tusks, ear-like lateral sepals, long trunk-like spur and the lip and dorsal hood resembling the mouth.

Cynorkis jackyi Hermans & Sieder sp. nov. Type: Madagascar, Fianarantsoa prov., Mahavelona, between Manambondro and Manantenina, 38 m, 27 Jan. 2014, Sieder A. & C., Knirsch & Andriantiana 6637 (Hermans 7015) (holotype WU!, isotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:77164800-1

Erect terrestrial herb, up to 21 cm tall, including the inflorescence. Tubers 2, ovoid-ellipsoid, c. 15 mm long, 10 mm wide, with a woolly surface; roots white, fleshy, woolly c. 1.5 mm in diam. Stem partly underground, white, very short, merging into the leaf petiole, with a short brown sheath at the base, c. 10 mm long, 4 – 5 mm in diam., with a pair of sub-opposite leaves. Leaves pale green strongly reticulate with pale veins and cross-bars and sunken darker patches, paler beneath, ovate, acuminate at the tip, almost flat on the ground, with a short petiole, the second leaf generally smaller and sometimes almost sheath-like, 38 – 41 mm long, 24 – 28 mm wide. Inflorescence reddish green to dark brown, a little angular, carrying a few short hairs, emerging from the centre of the leaves, up to 19 cm long, 1.2 – 1.5 mm in diam., with a short apiculate peduncle sheath around the middle, c. 8 × 2 mm; rachis 5 – 7 cm, laxly up to 10-flowered; floral bracts reddish green, amplexicaul at the base, acuminate, apiculate at the apex 2.5 – 3.2 mm long, 0.8 – 1.0 mm wide; pedicel and ovary microscopically verrucose, pale green longitudinally ridged with purple-red, slightly curved, 4 – 5 mm long, 1.0 – 1.3 mm wide. Flowers overall c. 5 mm long, 6 mm wide, all parts more or less fleshy, lateral sepals pale green-yellow, dorsal sepal and petals orange-pink darker around the margin of the petals, lip white, more or less pink in the middle, the basal callus darker pink, pollinarium pink, spur green becoming reddish brown towards the apex, some forms are almost entirely greenish-yellow in all parts. Dorsal sepal ovate, forming a tight hood with the petals, concave, 2.1 – 2.2 mm long, 1.0 – 1.1 mm wide, base inflated, the apex a little recurved; lateral sepals spreading, partly recurved, obovate-oblong 2.9 – 3.1 mm long, 1.5 – 1.6 mm wide. Petals appressed to the dorsal sepal margins, obovate-oblong curved apically, 2.0 – 2.1 mm long, 1.1 – 1.2 mm wide. Lip obscurely 3-lobed, overall 2.5 – 2.7 mm long, 2.0 – 2.2 mm wide, the lateral lobes rounded, the mid-lobe triangular-ligulate, disk raised in the middle, with a distinct rounded callus at the base; spur narrowed at the base becoming inflated in the lower \( \raisebox{1ex}{$2$}\!\left/ \!\raisebox{-1ex}{$3$}\right. \), 3.2 – 3.5 mm long, 0.5 – 0.6 mm diam. in the middle. Column short, in one plane, 1.2 – 1.4 mm long 1.1 – 1.2 mm wide, with two short angular rostellum arms, staminodes rounded and lobule-like, anther loculi distinct, ovate, pollinia on a very short caudicle. Figs 7 and 8.

Fig. 7
figure 7

Cynorkis jackyi in cultivation. A inflorescence; B close-up of flower. photos: anton sieder.

Fig. 8
figure 8

Cynorkis jackyi. A habit; B inflorescence of A; C flower, lateral view; D flower, face view; E dorsal sepal; F petal; G lateral sepal; H labellum; J column, spur and ovary, lateral view; K column ¾ face view. From Sieder 6637 (A – D from photographs, E – K after J. Hermans drawings). Graduated single bar = 2 mm, double bar = 1 cm, graduated double bar = 5 cm). drawn by andrew brown.

recognition

The new species is recognised by the two basal leaves with strongly tessellate venation and the long slender inflorescence with up to 10 small flowers. The lip has a distinct rounded callus at its base and a somewhat swollen spur. It has small flowers somewhat similar to those of species in the genus Benthamia A. Rich. (Richard 1828: 43) but it has ovoid-ellipsoid tubers (vs long and fasciculate in Benthamia), ovate leaves (vs generally narrow in Benthamia), a distinct callus on the base of the lip (not known in Benthamia) and short rostellum caudicles (vs lacking in Benthamia). Reticulate or otherwise patterned basal leaves are not uncommon in Cynorkis and can be found in species like C. tryphioides Schltr. (Schlechter 1913: 155) but with a completely different habit and floral morphology. It is most similar to C. falcata (Frapp. ex Cordem.) Schltr. (Schlechter 1915: 400), endemic to Réunion, with which it shares the size, lip shape and other floral characteristics but it differs in having more than one leaf with distinct net-like venation, is laxly flowered and has a rounded callus at the base of the lip

distribution

So far only known from a small coastal area in Southern Madagascar.

specimens examined

madagascar. Fianarantsoa prov., Mahavelona, between Manambondro and Manantenina, 38 m, 27 Jan. 2014, Sieder A. & C., Knirsch & Andriantiana 6637 (Hermans 7015) (holotype WU!, isotype K!).

habitat

Terrestrial, coastal low elevation forest on white sand, with lichens on the ground.

conservation status

This species is likely to be Critically Endangered (CR), according to the IUCN Red List Categories and Criteria. It is known from few individuals from a remnant of unprotected white sand forest, where it is known that human activity is likely to threaten remaining fragments of forest and species’ survival. The species is CR based on criterion D (fewer than 50 mature individuals), and criterion B2ab(iii) (area of occupancy likely to be less than 10 km2, number of locations = 1, and continuing decline in the quality of habitat inferred).

etymology

The new species is named for Jacky Lucien Andriantiana of the Parc Botanique et Zoologique de Tsimbazaza, Madagascar, recognising his outstanding contribution to the discovery of new orchids in Madagascar.

Cynorkis lentiginosa Hermans, Andriant. & Sieder sp. nov. Type: Madagascar, Mahajanga prov., road to Bealanana, 13 Jan. 2016, 633 m, Sieder A. & C. & Andriantiana 6981 (Hermans 7985) (holotype WU!, isotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:77164801-1

Erect to semi-erect terrestrial or lithophytic herb, up to 19 cm tall, including the inflorescence. Tubers 1 – 2, elliptic, surface woolly; c. 20 mm long, 12 mm wide, roots white, fleshy, glabrous c. 1.8 mm in diam. Stem up to 9 cm long, 4 – 5 mm in diam., white, faintly spotted with red, basal sheath sometimes absent, up to 48 mm long, 16 mm wide, then a 3 – 4 cm bare section to a pair of sub-alternate leaves. Leaves ovate-lanceolate, acuminate, the margins of the mature leaves undulate, up to 11 cm long, 3.5 cm wide, pale green flecked throughout with dark red in a longitudinal series, the margins purple-red, paler below. Inflorescence emerging from the centre of the upper leaf petiole, up to 16 cm long, 1.1 – 1.5 mm in diam., pale green, glabrous, flecked and spotted purple-red throughout but more densely towards the base, a large single lanceolate sheath c. 2 – 3 cm above the leaf, of a similar colour pattern as the leaf, enveloping the inflorescence at the base, 40 – 50 mm long, 12 – 18 mm wide; rachis 4 – 8 cm, laxly 6- to 9-flowered; floral bracts pale green spotted purple-red, amplexicaul at the base, acuminate, apiculate at the apex 8 – 12 mm long, 2.5 – 3.2 mm wide; pedicel and ovary glabrous, pale green flecked with purple-red, slightly curved, 15 – 21 mm long, 1.5 – 1.9 mm wide. Flowers overall c. 25 mm long, 20 mm wide, petals and sepals pale purple becoming green towards the apex, densely spotted with dark purple-red on the exterior, lip lilac fading to white towards the base, pollinarium purplish-pink, spur greenish violet with a purple longitudinal stripe, colours fading to pale-purple to almost white with age. Dorsal sepal forming a hood with the petals, 9.8 – 10.7 mm long, 3.1 – 3.9 mm wide, deeply concave, base inflated, the apex recurved; lateral sepals spreading almost parallel with the lateral lobes of the lip, oblong-obovate curved apically 8 – 9.2 mm long, 4.1 – 4.3 mm wide. Petals appressed to the inside of the dorsal sepal margins, narrowly oblong-obovate curved apically, 6.6 – 6.9 mm long, 1.7 – 1.9 mm wide. Lip 3-lobed, overall 10.1 – 10.4 mm long, 8.3 – 8.7 mm wide, the lateral lobes subrectangular, the mid-lobe linguiform, concave with a central raised ridge; spur narrowed at the base then narrowed into a short appendix at the tip, 8.5 – 9.3 mm long, 1.5 – 1.8 mm in the middle. Column in a horizontal plane 9.7 – 9.9 mm long 5.1 – 5.6 mm wide, with two distinct rostellum arms surpassing the nose-like mid-lobe by c. 3 mm, staminodes short and rounded, anther loculi somewhat verrucose on the anterior margin, covered by the dorsal sepal and petal hood, pollinia including the thin caudicle c. 9 mm long. Figs 9 and 10.

Fig. 9
figure 9

Cynorkis lentiginosa. A inflorescence, lateral view; B flowers, face view. photos: anton sieder.

Fig. 10
figure 10

Cynorkis lentiginosa. A habit (from photograph); B flower, lateral view; C flower, face view; D dorsal sepal and petal, lateral view; E dorsal sepal, inner face; F petal, inner face; G lateral sepal, inner face; H labellum; J column, lateral view. From Sieder 6981, spirit material). drawn by andrew brown.

recognition

The new species is characterised by the distinctive purple-red spotting on the two ovate-lanceolate leaves which have a purple margin, the spotting throughout the plant and flowers, including the sheaths, peduncle, bracts, ovary and exterior of the petals and sepals. All parts are glabrous. The three-lobed angular lip and medium-sized plant, combined with a sausage-shaped spur also set it apart from other Cynorkis. Its lip and spur shape are somewhat similar to those of C. cinnabarina (Rolfe) Hermans & Cribb (2014: 9) but it differs in having pale purple petals and sepals becoming green towards the apex and densely spotted with dark purple-red on the exterior, a lilac lip fading to white towards the base, purplish-pink pollinaria, a greenish violet spur with a purple longitudinal stripe (vs bright orange flowers in C. cinnabarina), ovate-lanceolate leaves (vs lanceolate-linear in C. cinnabarina). More or less dark spotting or flecking on leaves and flowers is not uncommon in other species of Cynorkis (as in C. gibbosa Ridl., for example) but no other species shows this characteristic throughout the plant and flower.

distribution

So far only known from a small area in the Antsahabe area in Northern Madagascar.

specimens examined

madagascar. Mahajanga prov., Antsahabe area, road to Bealanana, on a slope, 633 m, 13 Jan. 2016, Sieder A. & C. & Andriantiana 6981, (Hermans 7985). (holotype WU!, isotype K!).

habitat

In full sun in water seepage areas on basement basalt rock, in crevices and shallow pockets of humus in seasonally very dry grassland, around 600 m, growing sympatrically with Cynorkis christae and C. siederi.

conservation status

This species is likely to be Critically Endangered (CR), according to the IUCN Red List Categories and Criteria. It is known from only a single individual, found in unprotected grassland alongside a road leading to a remote town in the NE of the country — such areas are known to be at high risk from human development. The species is CR based on criterion D (fewer than 50 mature individuals), and criterion B2ab(iii) (area of occupancy likely to be less than 10 km2, number of locations = 1, and continuing decline in the quality of habitat inferred).

etymology

The species name refers to the freckled appearance of the plant and flowers.

Cynorkis mammuthus Hermans & P. J. Cribb sp. nov. Type: Madagascar, Mahajanga prov., road to Antsohihy, near Anjalazala, fl. Feb. 1994, Hermans 2699 (holotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:60474978-2

Plant tubers, roots and foliage not known. Inflorescence total length not known, c. 3 mm in diam.; rachis c. 15 cm, densely racemose with up to 11 flowers; floral bracts funnel-shaped, serrate-hirsute at the margin, acuminate, then apiculate at the apex, about \( \raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right. \) the length of the pedicellate ovary 21 – 25 mm long, 8 – 10 mm wide; pedicel and ovary densely hirsute, almost straight, 60 – 68 mm long, 1.8 – 2.1 mm wide. Flowers overall c. 4 – 5 cm long, 2.5 – 3.5 cm wide, not including the spur. Dorsal sepal 32 – 34 mm long 14 – 16 mm wide when flattened, forming a hood with the petals, gibbose, expanded and hirsute at the base; lateral sepals spreading, oblong-elliptic with a distinctive wing on the upper margin towards the tip, 29 – 30 mm long, 12 – 13 mm wide. Petals appressed to the inside of the dorsal sepal margins, linear-lanceolate, 31 – 33 mm long, 2.5 – 2.8 mm wide. Lip entire, oblanceolate, naviculate, acuminate at the tip, 36 – 38 mm long, 12 – 14 mm wide; spur down-curved then pendent, subulate, 10.5 – 13 cm long, c. 2 mm in diam. Column in an angular plane 33 – 35 mm long 9 – 11 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. 28 mm long. Fig. 11H – T.

Fig. 11
figure 11

A – G Cynorkis mangabensis. A habit (from herbarium sheet); B dorsal sepal; C lateral sepal; D lateral petal; E labellum; F lateral view of column, ovary and spur; G lateral view of column in situ. From Antilahimena 2644; A from herbarium sheet, * approximate scale; B – G after J. Hermans drawings). H – T Cynorkis mammuthus. H rehydrated flower, rather compressed laterally; J glandular hairs from outer surface of dorsal sepal; K indumentum from ovary surface; L dorsal sepal; M petal; N lateral sepal; P labellum; Q column and ovary; R spur; S floral bract; T detail of column. From Hermans 2699; H – K from herbarium sheet, L – T after J. Hermans drawings. drawn by andrew brown.

recognition

The new species is distinct by the raceme of very large flowers for the genus, up to 11 on the raceme, densely hirsute pedicellate ovary and base of flower, the entire lip with a very long pendent spur, lateral sepals with an auricle towards the apical margin and the very long tusk-like rostellum arms. It could be perceived as a peloric form of Cynorkis christae but the longer rostellum arms, lobed lateral sepals and 8- to 11-flowered inflorescence (vs 3 – 4 in C. christae) make it unlikely. It shares its large flower size with C. gigas Schltr. (Schlechter 1924: 50) but has an 8 – 12-flowered inflorescence, an entire vs a distinctly lobed lip, clearly lobed lateral sepals and a longer spur (c. 10 cm in C. gigas vs 13 cm in the new species). The large flower and long spur are reminiscent of C. christae Hermans, Andriant. & Sieder described above, which comes from the same area in Madagascar, but differs in its lip and lateral sepals. In flower morphology the new species is closest to C. elephantina Hermans, Andriant. & Sieder described above, with which it shares the entire lip, the densely hirsute pedicellate ovary, lobed lateral sepals and long rostellum but its sepals and petals are about twice as large, its lip and spur almost a third larger, its floral bracts broad and funnel-shaped (vs shortly amplexicaul in C. elephantina), its hood, formed by the dorsal sepal and petals, is not recurved, its rostellar arms are longer and its spur is up to 13 cm long (vs 7.8 cm).

distribution

Only known from the Antsohihy area in Northern Madagascar.

specimens examined

madagascar. Mahajanga prov., road to Antsohihy, near Anjalazala, fl. Feb. 1994, Hermans 2699 (holotype K!); Befandriana, Feb. 1942, Jard. Bot. Tana 6242 (P!).

habitat

Not known

conservation status

This species is currently considered to be Data Deficient (DD), according to the IUCN Red List Categories and Criteria. It is only known from two individuals, both with incomplete locality information, and one of which is an old specimen from 1942. Further research is needed to check the population status, threats and habitat.

etymology

The species name refers to the likeness of the profile of the flower to the head of a mammoth with the rostellum arms as tusks and long trunk-like spur. It is also larger than its nearest relative Cynorkis elephantina, possibly extinct in the wild and only known from dried specimens.

notes

This enigmatic new species is known from two dried inflorescences only; the first from a herbarium sheet of flowers collected in 1994 as Cynorkis uniflora Lindl. (Lindley 1835: 331) by a Malagasy orchid nursery and a second collection from 1942 in the Paris Herbarium (P), last seen about 10 years ago and currently only known from a photographic record. The latter specimen, originally from the ‘Herbier du Jardin Botanique Tananarivo Madagascar’, was identified as C. gigas Schltr. but annotated by Jean Bosser as ‘not C. gigas, nothing to do with this species’.

Cynorkis mangabensis Hermans & P. J. Cribb sp. nov. Type: Madagascar, Toamasina prov., Maroantsetra, Ambinanitelo, Marovovonana near Ambatofotsy, Mangabe forest, 556 m, 30 Aug. 2004, Antilahimena 2644 (holotype P!, isotype MO).

http://www.ipni.org/urn:lsid:ipni.org:names:77164802-1

Epiphytic herb, up to 48 cm tall including the inflorescence. Tubers unknown, roots, wiry, woolly, c. 1.5 mm in diam. Stem short, merging into the leaf petiole, c. 30 mm long, 3 – 4 mm in diam., with a single leaf. Leaf oblong, acuminate at the tip, narrowed into a 3 – 4 cm petiole at the base, 15 – 17 cm long, 4 – 4.5 cm wide. Inflorescence emerging from the base of the leaf, with a few short hairs towards the rachis, up to 47 cm long, 3 – 4 mm in diam., with a short long peduncle sheath towards the base, c. 45 × 2 – 3 mm, with 2 – 3 shorter sheaths along the stem, c. 20 × 3 mm; rachis 6 – 7 cm, up to 9-flowered; floral bracts acuminate, apiculate at the apex 6 – 7 mm long, 2.1 – 3.0 mm wide; pedicel and ovary hirsute-glandular, slightly curved, 15 – 17 mm long, 1.8 – 2.1 mm wide. Flowers overall c. 20 mm long, 18 mm wide, white, exterior of the sepals hirsute-glandular. Dorsal sepal ovate-lanceolate, forming a hood with the petals, concave, 12 – 13 mm long, 3 – 4 mm wide; lateral sepals, obovate-oblong 11.2 – 11.5 mm long, 5.5 – 5.8 mm wide. Petals asymmetric with the anterior border expanded into a rounded lobe, 14 – 15 mm long, 4 – 5 mm wide. Lip entire, oblong, angular, 14 – 15 mm long, 10.3 – 10.5 mm wide, the margins undulate; spur broad at the base then subulate, curved upwards towards the apex, hirsute underneath the base, c. 9 mm long, 1.2 mm in diam. in the middle. Column short, in one plane, c. 3.5 mm long 3 mm wide, with two short rostellum arms, staminodes rounded, anther loculi ovate-elongate. Fig. 11A – G.

recognition

The new species is typified by its long inflorescence, medium-sized white flowers with an entire lip. It is an epiphyte similar to Cynorkis muscicola Bosser (1969: 346), in section Imerinorchis, in the length of its inflorescence and entire lip but its flowers are twice as large and the lip is oblong and angular rather than linguiform. C. muscicola was found at high altitude in South-Central Madagascar, the new species comes from wet forest in the North-East.

distribution

Only known from the type locality in Mangabe forest in northern Madagascar.

specimens examined

madagascar. Toamasina prov., Maroantsetra, Ambinanitelo, Marovovonana near Ambatofotsy, Mangabe forest, 556 m, 30 Aug. 2004, epiphyte on tree, Antilahimena 2644 (holotype P!, isotype MO).

habitat

Epiphyte in rainforest with high precipitation rates.

conservation status

This species is currently considered to be Data Deficient (DD), according to the IUCN Red List Categories and Criteria. It is only known from the type specimen, which was collected recently (in 2004). It was found within the Makira newly protected area, in dense forest, no obvious imminent threats are apparent. Further research is needed to check the population status, threats and habitat.

etymology

The species name refers to the type locality of the Mangabe forest.

vernacular name

Mantabe: ‘Manta’ is referred to in the literature (e.g. Dandouau 1909) as a Betsimaraka (of Northern Madagascar) dialect name for the Angraecoid orchid Aerangis citrata (Thouars) Schltr., ‘be’ normally refers to ‘large’ or ‘great’.

Cynorkis sanguinolenta Hermans, L. Gaut. & P. J. Cribb. sp. nov. Type: Madagascar, Antsiranana prov., Andrafiamena, forêts aux alentours d’Anjahankely, 7 Jan. 2011, 694 m, Burivalova ZB182 (holotype G!, isotypes K!, MO, P!, TEF).

http://www.ipni.org/urn:lsid:ipni.org:names:77164803-1

Terrestrial or lithophytic herb, up to 35 cm tall including inflorescence, forming small clumps or groups of plants. Tubers and roots not known. Stem short, partly underground, c. 1 – 2 cm long, with a solitary erect to arching radical leaf. Leaf glabrous, the central vein strongly keeled underneath, especially at the base, obtuse apically, up to 18 cm long, 12 mm wide but probably larger when mature, pale to mid-green, paler below, oval-oblong. Inflorescence green, up to 32 cm, 1.7 – 2.0 mm in diam., glabrous, produced from the new developing shoot, within the base of the unfurling leaf, with a peduncle sheath around the middle, c. 3 cm long; rachis secund sub-racemose, up to 14 flowers but generally fewer; floral bracts green sometimes a little hirsute on the exterior towards the apex, ovate-lanceolate, 12 mm long, 2 – 3 mm wide; slightly auriculate at the base, pedicel and ovary with very few scattered hairs, mainly at the apex, yellowish-green, almost straight, up to 55 mm long, 2 mm wide. Flowers variable in size with some forms substantially larger than others, the larger dimensions are measured from living specimens (Sieder et al. 6987), the smaller from the type: overall 26 – 40 mm long, 18 – 33 mm wide, not including the spur; sepals and petals purple, the exterior papillose, lip pure white with the entire disk a distinct carmine continuing into the throat and the spur pinkish-purple, column dark purple with the rostellum white below, anthers darker. Dorsal sepal oblong, deeply concave and, together with the petals, forming a hood over the column, 7.5 – 9.7 mm long 3.4 – 4.1 mm wide, the base inflated; lateral sepals swept backwards against the ovary, concave, oblong to sickle-shaped, 9.1 – 9.6 mm long, 4.1 – 4.2 mm wide. Petals narrowly oblong-obovate, slightly curved apically, appressed to the inside of the dorsal sepal margins forming a narrow rim 9.3 – 9.4 mm long, 1.3 – 1.9 mm wide. Lip somewhat connate to the base of the column, lateral lobes a little smaller than the apical lobes, all deeply serrate around the anterior margins, with the disk slightly raised, overall 18 – 38 mm long, 15 – 27 mm wide; spur almost parallel with the ovary, somewhat sigmoid, subulate towards the end, up to 32 mm long, av. 1.2 mm in diam. Column in an almost horizontal plane 5.9 – 6.1 mm long 4.1 – 4.3 mm wide, with two parallel rostellum arms for almost half its length, mid-lobe forming a nose-like protrusion, staminodes club-shaped, hidden below the rostellum, anther loculi incurved, pollinia including the caudicle c. 5.5 mm long. Figs 12 and 13.

Fig. 12
figure 12

Cynorkis sanguinolenta. A habit, in habitat; B flowers lateral view; C flower, face view, showing labellum. photos: A: ANTON SIEDER, B, C: ZUZANA BURIVALOVA.

Fig. 13
figure 13

Cynorkis sanguinolenta. A habit (from photograph); B habit; C flower, lateral view; D flower, face view; E dorsal sepal and petals, lateral view; F dorsal sepal, inner face; G petal, inner face; H lateral sepal, inner face; J labellum; K column, in situ, lateral view; L column face view; M glands from base of flowers; N surface of inner face of sepal, lower edge; P surface of labellum, proximal region. From Sieder 6987, spirit material, Burivalova 182 K000718773). drawn by andrew brown.

recognition

This large-flowered species is distinct by its large single leaf, tall secund raceme of up to 14 flowers, the proportionally very large white lip with a distinct carmine area towards the base, the deeply serrate anterior margins of the lip and the somewhat sigmoid spur. The new species is allied to Cynorkis fimbriata H. Perrier ex Hermans (in Hermans et al. 2007: 291), section Lowiorchis, which has been found in the same area but whilst both have a typical deeply serrated lip there are considerable differences in both habit and flower: C. fimbriata has a long 5 – 7 cm leaf petiole, a linear lanceolate leaf, large leaf-like lower floral bracts (up to 20 mm long, 10 mm wide), bright purple flowers with the spur inflated in the apical third. C. sanguinolenta has a short 1 – 2 cm leaf petiole, a larger oval-oblong leaf, consistently short amplexicaul floral bracts (12 mm long, 2 – 3 mm wide), larger flower segments, including the lip which is pure white with a carmine base and a slightly sigmoid straight spur. In habit the new species also resembles C. angustipetala Ridl. (Ridley 1885: 514), in section Gibbosorchis, but its rachis is not so densely flowered, its flowers are larger in all their parts and its lip is more deeply serrate at the margins.

distribution

Known from three localities in the Antsiranana province of Northern Madagascar.

specimens examined

madagascar. Antsiranana prov., Andrafiamena, forest near Anjahankely, 7 Jan. 2011, 694 m, grassland on sandstone, Burivalova ZB182 (G, K!, MO, P!, TEF); Ambilobe, Betsiaka, Andasibe forest, wet forest, 505 m, Jan. 2006, Guittou et al. 251, (K, MO); near Ambilobe, 91 m, 15 Jan. 2016, in gullies/crevices, Sieder A. & C. & Andriantiana 6987, (Hermans 7990) (K!, WU!).

habitat

Seasonally dry secondary grasslands on sandstone and white sands and erosion gullies in areas once dominated by semi-deciduous forest, up to 700 m elevation.

conservation status

This species is likely to be Endangered (EN), according to the IUCN Red List Categories and Criteria. It is known from just three localities, only one of which has a provisional status of protection, the area has a lot of human development activity, which could threaten its survival. The localities do encompass a range of habitats, and the specimens were recently collected: 2004, 2011 and 2016. However, based on the current known Extent of Occurrence (approximately 250 km2) and Area of Occupancy (approximately 12 km2), this species would be assessed as Endangered under criterion B1ab(ii,iii)+B2ab(ii,iii) as the extent of occurrence is likely to be less than 5,000 km2, area of occupancy is likely to be less than 500 km2, number of locations less than 5 and continuing decline in the area of occupancy and quality of habitat is inferred).

etymology

The name refers to the red blood-stained appearance of the flower.

Cynorkis siederi Hermans & Andriant. sp. nov. Type: Madagascar, Mahajanga prov., road to Bealanana, 633 m, 13 Jan. 2016, Sieder A. & C. & Andriantiana 6983 (Hermans 7986). (holotype WU!, isotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:60474980-2

Erect terrestrial herb, up to 20 cm high including inflorescence, forming small clumps of several plants. Tubers 1 – 2, elongate, surface woolly, c. 30 mm long, 7 mm wide, roots, fleshy, glabrous c. 2 mm in diam. Stem very short, c. 4 mm in diam., enclosed by a thin greenish-brown scarious sheath, with a single almost erect radical leaf. Leaf and sheaths green, somewhat fleshy, shiny on top, conduplicate, long-lanceolate, acuminate, up to 18 cm long, 12 mm wide. Inflorescence emerging from the leaf axil, up to 19 cm, 1.5 – 1.8 mm in diam., green, glabrous, with a long leaf-like lanceolate-subulate sheath c. 4 cm above the leaf base, 85 – 92 mm long, 3 – 4 mm wide, a second sheath about half way up the inflorescence of the same shape but shorter, 21 – 24 mm long, 2 – 3 mm wide; rachis sub-umbellate, up to 5 flowers; floral bracts green, ovate-lanceolate 12 – 14 mm long, 2.5 – 3 mm wide; pedicel and ovary carrying a few scattered hairs, mainly in the upper half, green, almost straight, up to 25 mm long, 2 mm wide. Flowers overall 18 – 20 mm long, 14 – 16 mm wide, not including the spur, sepals greenish white with the veins green, brownish at the tip, petals white with a pale lilac margin, lip white with two pale lilac lines towards the base, spur white becoming green in the lower half, column greenish-white the pollinia brown, exterior of sepals with a densely pilose indumentum on the exterior. Dorsal sepal forming a hood with the petals, ovate, deeply concave, 7.0 – 8.2 mm long 3.2 – 3.4 mm wide; lateral sepals spreading, oblong-obovate, curved apically, 8.1 – 8.3 mm long, 3.1 – 3.2 mm wide. Petals loosely attached to the dorsal sepal margins, ovate, 7.9 – 8.1 mm long, 3.9 – 4.1 mm wide. Lip with two large and spreading lateral lobes with a gutter-like depression in between, a broadly notched to bilobed central lobe, overall 13.1 – 14.4 mm long, 10.5 – 11.1 mm wide, the lateral lobes almost oval, the margins a little undulate, the lateral lobes much smaller with distinct sinuate to serrate margins; spur pendent then incurved in the lower quarter, subulate towards the end, up to 32 mm long, c. 1 mm in diam. Column in an almost horizontal plane 5.2 – 5.4 mm long 4.5 – 4.8 mm wide, with two short rostellum arms for almost half its length, with a distinct recurved bilobed mid-lobe forming a snout-like protrusion, staminodes below the base of the rostellum arm forming verrucose auricles, anther loculi incurved, pollinia including the caudicle c. 4.5 mm long. Figs 14 and 15.

Fig. 14
figure 14

Cynorkis siederi. photo: anton sieder.

Fig. 15
figure 15

Cynorkis siederi. A habit; B inflorescence (from photograph); C flower, lateral view; D flower, face view; E glandular hairs at apex of ovary; F dorsal sepal and petals, side view; G dorsal sepal, inner face; H petal, inner face; J lateral sepal, inner face; K labellum; L column face view; M column, lateral view; N column, dorsal view. From Sieder 6983 (spirit material & field photographs). drawn by andrew brown.

recognition

The species is distinct by its somewhat fleshy, long lanceolate single leaf and a long thin leaf-like lower peduncle sheath. The medium-sized flowers, which are almost completely white, have a distinct lip with two large rounded lateral lobes and a smaller divided mid-lobe with more or less serrate margin. The snout-like mid-lobe of the rostellum is unusual in Cynorkis. In overall plant habit it resembles C. uniflora Lindl. (Lindley 1835: 331) in section Gibbosorchis, but its flowers have a completely different shape, size and colour. It is probably nearest to C. speciosa Ridl. (Ridley 1886: 122) in section Gibbosorchis, a widespread and variable species, but it has side lobes to the lip that are larger than the divided midlobe (vs almost equal in C. speciosa), a straight and longer spur (vs inflated a little in the middle in C. speciosa), distinctly shaped and coloured mid-lobe of the rostellum. Its lip and spur are similar to those of C. aphylla Schltr. (Schlechter 1913: 150) in section Lowiorchis, but apart from the flower colour (dark pink in C. aphylla) the leaves are not generally developed at flowering time and are broadly oblong (vs lanceolate in C. aphylla).

distribution

Known only from a small area in the Antsahabe area in Northern Madagascar.

specimens examined

madagascar. Mahajanga prov., road to Bealanana, on a slope, amongst rock, 633 m, 13 Jan. 2016, Sieder A. & C. & Andriantiana 6983 (Hermans 7986), (holotype WU!, isotype K!).

habitat

Water seepage areas on basement basalt rock, amongst large rocks in crevices and shallow pockets of humus and other detritus in seasonally very dry forest remnants within grassland, around 600 m, growing sympatrically with Cynorkis christae and C. lentiginosa.

conservation status

This species is likely to be Critically Endangered (CR), according to the IUCN Red List Categories and Criteria. It is only known from a small population of approximately five individuals at a single locality, in unprotected grassland alongside a road leading to a remote town in the NE of the country — such areas are known to be at high risk from human development, including intense, frequent grassland fires. The species is CR based on criterion D (fewer than 50 mature individuals), and criterion B2ab(iii) (area of occupancy likely to be less than 10 km2, number of locations = 1, and continuing decline in the quality of habitat inferred).

etymology

Named for Anton Sieder, of the Botanical Gardens, University of Vienna, Austria who discovered the new species.

Cynorkis syringescens Hermans & P. J. Cribb sp. nov. Type: Madagascar, Fianarantsoa prov., near Mahatsanda, c. 1500 m, Jan. 1996, Hermans 7055 (holotype K!).

http://www.ipni.org/urn:lsid:ipni.org:names:60474981-2

Erect terrestrial herb, up to 28 cm high including the inflorescence, plants generally solitary. Tubers 1 – 2, elongate, surface woolly, c. 25 mm long, 15 mm wide, roots, fleshy, woolly c. 2 mm in diam. Stem very short, c. 20 mm long 10 mm in diam., enclosed by one or two thin scarious brown sheaths, with two or rarely one radical leaves. Leaves pale green, sub-erect, somewhat fleshy, conduplicate when developing, lanceolate-linear, acuminate, up to 20 cm long, 15 – 22 mm wide. Inflorescence emerging from the leaf axil, green, glabrous, up to 25 cm long, 1.8 – 2.1 mm in diam., with a lanceolate-subulate sheath in the middle, 25 – 28 mm long, 5 – 8 mm wide; rachis loosely racemose, up to 14 flowers but generally fewer; floral bracts green, brownish-red in the apical half, ovate-lanceolate 18 – 24 mm long, 5.9 – 7.1 mm wide; pedicel and ovary carrying a few glandular hairs, mainly towards the base, purplish-brown, almost straight, up to 35 mm long, 2 mm wide. Flowers overall c. 30 mm long, 20 mm wide, not including the spur, sepals green with the margins and apex dark reddish-brown, petals greenish-white with the margins mauve, lip mauve with the disk white, spur reddish-brown, column white, staminodes red-purple, anther dark green, base of the sepals with a few apiculate hairs. Dorsal sepal forming a hood with the petals, ovate, deeply concave, 9.0 – 9.2 mm long, 3.8 – 4.0 mm wide; lateral sepals spreading, ovate, 9.8 – 9.9 mm long, 4.4 – 4.5 mm wide. Petals loosely attached to the dorsal sepal margin, lanceolate, 8.9 – 9.1 mm long, 2.5 – 2.6 mm wide. Lip 4-lobed with a narrowed deflexed base and raised disk in between, lobes all of about the same size, ovate truncate, margins undulate and somewhat deflexed, overall 19 – 21 mm long, 19 – 20 mm wide; spur parallel with the ovary, narrowed at the base, then slightly inflated and acuminate towards the end, up to 32 mm long, c. 2 mm in diam. Column angular to almost vertical, 4.0 – 4.2 mm long 3.9 – 4.1 mm wide, with two short parallel rostellum arms for about \( \raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right. \) its length, curved upward, with a nose-like mid-lobe in between, staminodes below the base of the rostellum arm forming verrucose auricles, anther loculi oval, pollinia including the caudicle c. 6 mm long. Figs 16 and 17.

Fig. 16
figure 16

Cynorkis syringescens. A habit, in habitat; B close-up of flower, showing labellum. photos: johan hermans.

Fig. 17
figure 17

Cynorkis syringescens. A habit; B flower, face view (from photograph); C flower, lateral view (from photograph); D floral bract; E dorsal sepal lateral view (collapsed); F dorsal sepal, inner face; G petal, inner face; H lateral sepal, inner face; J labellum; K column, lateral view; L column, lateral view (from J. Hermans drawing); M caudicle and pollinium. From Hermans 7055 (spirit material & field photographs). drawn by andrew brown.

recognition

The species is distinct by its somewhat fleshy, lanceolate-linear leaves; the medium-sized flowers with a 4-lobed lip which is mauve with a white disk, the base and lobes with the margins somewhat deflexed, lobe margins undulate, a spur about as long as the pedicellate ovary and the narrow mid-lobe of the column and slightly curved rostellum arms about \( \raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right. \) of the column length. It is similar in habit to the very widespread and variable Cynorkis speciosa Ridl. (Ridley 1886: 122) in section Gibbosorchis but the flowers are a different shape and colour. The flowers of the new species are generally fewer and smaller, the overall shape of the lip is almost rounded and about 20 mm long × 19 mm wide (vs elliptic and about 27 mm long × 21 mm wide in C. speciosa); the deflexed margins also set it apart. C. syringescens has a mauve blade and a white disk (the lip in C. speciosa can be white, pink or purple but almost invariably has a large distinct rectangle of deep purple or pink on the disk and base). The spur is of a similar size but is more inflated than in C. speciosa. The column and especially the mid-lobe are narrower and not as broad at the apex as in C. speciosa. It is also similar to C. fastigiata Thouars (1822: t3), another common variable and widespread species that grows in the same area; that has similar flower proportions, a white to pale violet-pink lip with a white disk and a darker coloured area in the throat only, the margins are more or less deflexed. The main differences with the new species are the generally single leaf which is ovate-lanceolate rather than lanceolate linear, the smaller size of the flowers and elliptic outline of the lip (average 12 mm long × 9 mm wide).

distribution

Known only from a small highland area in Central Madagascar.

specimens examined

madagascar. Fianarantsoa Prov, near Mahatsanda, Tapia forest, c. 1500 m, Jan. 1996, Hermans 7055 (holotype K!).

habitat

Granite and quartz rocks in pockets of sand, amongst grasses in sub-humid degraded sclerophyllous woodland, amongst Tapia (Uapaca bojeri Baill.), c. 1500 m.

conservation status

This species is likely to be Critically Endangered (CR), according to the IUCN Red List Categories and Criteria. It is only known from a small (albeit persistent) population of 4 – 6 individuals at a single locality, in unprotected Tapia woodland and grassland, near to a major settlement (Mahatsanda) and the Route Nationale 7. Vegetation in these areas is known to be at high risk from human development activity, including intense, frequent grassland fires. The species is CR based on criterion D (fewer than 50 mature individuals), and criterion B1ab(i,ii,iii)+B2ab(i,ii,iii) (extent of occurrence likely to be less than 100km2, area of occupancy likely to be less than 10 km2, number of locations = 1, and continuing decline in the extent of occurrence, area of occupancy, and quality of habitat inferred).

etymology

The epithet refers to its passing resemblance to the lilac (Syringa vulgaris L., Oleaceae) in the colour of the lip and somewhat pyramidal inflorescence.

notes

This interesting new species has been observed in the same locality for several years, including recently in January 2016 when the habitat and surrounding Cynorkis species were looked at in more detail. The species grows together with various colour forms of C. speciosa Ridl. (Ridley 1886: 122), C. gibbosa Ridl. (Ridley 1883: 331) and C. fastigiata Thouars (1822: t3).

Considering its somewhat intermediate morphology between Cynorkis speciosa and C. fastigiata the possibility that it is a natural hybrid between the two should be considered; both grow in the same locality and flower around the same time in January – February. However, as a number of stable colonies of the new species have been recorded and there is little variability in its flowers it seems best to recognise it as a distinct species.

Nomenclatural Notes

Cynorkis elegans

Cynorkis elegans (as Cynosorchis elegans) was described by H. G. Reichenbach f. in the journal ‘Flora’ (Reichenbach 1888a: 150) together with a random group of other orchids that included Cynosorchis lowiana, both cultivated at the Nursery of Messrs Hugh Low & Co. in England. Cynorkis elegans was described in some detail in Latin. At about the same time he also described the same species in the Gardeners’ Chronicle (Reichenbach 1888b: 424) with a similar description in English. The plant flowered at the Low nursery in 1883 and apart from the type there are several detailed sketches by Reichenbach respectively dated July 1882, November 1882 and October 1883, kept at the Vienna Herbarium (W). In addition, there is a watercolour by John Day (in his famous 'Scrapbooks' housed at the Royal Botanic Gardens Kew), presumably of the same Low plant, which includes some details of the flower; dated September 1883 (Fig. 18). Apart from ‘Madagascar’ there is no record of the provenance of Low’s plant. The description and illustrations are clear and depict a short plant with two ornately patterned basal leaves, white flowers with a three-lobed lip, the lateral lobes angular and the mid-lobe smaller and narrower. The lip has deep-purple spots in longitudinal lines, the spur is a little shorter than the pedicel and ovary and the column has a distinct 3-lobed rostellum with divergent arms.

Fig. 18
figure 18

Cynorkis elegans Rchb. f. in John Day's Scrapbook, 1883. Reproduced by kind permission of the Trustees of the Royal Botanic Gardens, Kew.

Later, covering the new collections from Madagascar by George Francis Scott Elliot in the Journal of the Linnean Society, Rolfe (1891: 58) described Cynorkis elata. The type came from Fort Dauphin (Tôlanaro) in Southern Madagascar and specimens are kept at K, BM and P. The type material and description show exactly the same characteristics as C. elegans. Rolfe compared his new species with C. lilacina Ridl. describing it as differing in being a ‘plant with few to 20 flowers’ and ‘quite glabrous’. However, the type material has fewer flowers and a shortly hirsute pedicellate ovary. There is no doubt that C. elegans Rchb. f. and C. elata Rolfe are the same with Reichenbach’s name having precedence.

Cynorkis elegans Rchb. f. (Reichenbach 1888a: 150) & (Reichenbach 1888b: 424). Type: Madagascar, Hort, Messrs. H. Low & Co s.n. (holotype. W, Rchb. Orch. 46502), as Cynosorchis elegans.

Gymnadenia muricata Brogn. ex Kraenzl. (Kraenzlin 1898: 482).

Bicornella elegans (Rchb. f.) Szlach. & Kras (Szlachetko & Kras 2006: 144).

Cynorkis elata Rolfe (1891: 58) synon. nov. Type: Madagascar, Fort-Dauphin, Scott Elliot 2477 (holotype K!; isotype BM!, P!).

specimens examined

madagascar. Mananajary, Geay 7991 (P!); Ambila, S of Tamatave, May 1928, Decary 6427 (P!); Toliara prov., Ambila, Decary, R.6427 (P); Vinanibe forest near Fort Dauphin, April 1947, Humbert 20769 (P!); near St Luce, April 1969, Veyret 1030 (P!); Tulear, Mandena forest, March 1989, Gereau, Rabevohitra, Dumetz 582 (K!, MO, P!, TAN); Tolanaro, Ste Luce, Jan. 1990, Dumetz 1189 (MO, P!); Mananajary prov., coastal zone, March – April 1990, Geay 7992 (P!); Tulear prov., N of Fort Dauphin, St Luce forest, Feb. 1995, Du Puy et al. M847 (K!, P!); Toliara prov., Ampasy, Taolagnaro, Rabenantoandro et al. 275 (MO); Hort., 1996, Hermans 3736 (K!).

Arnottia

Achille Richard (1828: 29) established the genus Arnottia in his work on the flora of Mauritius and Réunion, based on an unspecified Commerson specimen of Arnottia mauritiana A. Rich from Réunion. He illustrated a floral dissection and also cited Amphorkis inermis Thouars. He gave as the main characters distinguishing it from Gymnadenia R. Br. (where most of the African species now in Cynorkis Thouars were then placed) the lip being uppermost in the flower, erect and fused to the base of the lateral sepals and lacking a spur, together with the lateral sepals being winged. Examination of the accompanying drawing of A. mauritiana (Richard 1828: pl.7.1) shows that the latter characteristics are based on confusion in interpretation between the dorsal sepal and the labellum of the dried material.

In effect, the only character separating Arnottia from Cynorkis is the absence of a spur. Spur length in Cynorkis is variable within the genus and within the species. Within the same subtribe the European former genus Aceras R. Br. (Brown 1813: 191), lacks a spur, but was shown by DNA data (Pridgeon et al. 1997: 98) to be nested in the otherwise spurred genus Orchis Tourn. ex L. (von Linné 1753: 939). Within the Orchidaceae of the region, there are several examples, such as Angraecum appendiculatum Frapp. ex Cordem (Frappier 1895: 211) and Angraecum cucullatum Thouars (1822: 48), with populations with not just peloric or partially peloric flowers in which the lip is more or less petal-like but also a spur that is greatly reduced or absent. In addition, flowers with a spur are occasionally found in inflorescences of normally spurless flowers of Arnottia mauritiana (Bernet 2010b: 122 & pers. comm.). A. mauritiana is virtually identical to Cynorkis nutans (Ridl.) H. Perrier (1936: 582) from the Madagascar Highlands but just lacks a spur. The name Arnottia was previously also used by Reinwardt & Hornschuch (1828: 709) for a genus of mosses from Java, presumably also named for the botanist Walker Arnott. As suggested in Genera Orchidacearum (Pridgeon et al. 2001: 276) there is also no genetic evidence for keeping Arnottia separate from Cynorkis. Therefore a number of new combinations need to be made, as shown below.

In addition confusion exists over the identity of Amphorkis inermis Thouars (1822 t.5). In the first text table of the book Thouars separated the spurless species in his genus Amphorkis into Imermamphis (Ophrys) dubia Thouars (1822: t.1) but the corresponding plate b.2-t.5 is captioned Amphorchis inermis. The distribution is said to be Mauritius and Réunion. Thouars’ herbarium sheet at P comprises a specimen plus his original plate of Amphorchis inermis and is labelled ‘O dubia no 4’ and ‘Amphorchis’. The label referring to ‘Thouars no 4’ is either an error as his plate no 4 is Cynorkis squamosa Lindl. with a broad lip (vs very narrow in the specimen) or refers to one of his different numbering systems. Although the leaf and tip of the rachis is missing, the specimen corresponds well with that shown in the plate; it is therefore clear that this is the type specimen of Amphorchis inermis. Thouars’ drawing is somewhat confusing but, on closer examination, the plant and flower detail correspond well with the description and type (assigned below) of Richard’s Arnottia mauritiana with Thouars’ name having priority. It is interesting to note that Richard cites ‘Amphorchis inermis ? Du Petit-Thouars t5?’ before his description of Arnottia mauritiana. The densely hirsute ovary, the oval-oblong lateral sepals and the ligulate-lanceolate lip correspond well with the characteristics of the many specimens of A. mauritiana, the shape of the lip is somewhat variable and can be expanded at the base to almost lobed.

Cynorkis inermis (Thouars) Hermans & P. J. Cribb comb. nov. Type: Mauritius or Réunion, Thouars 4 (holotype P!) (as Amphorchis inermis).

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Amphorkis inermis Thouars, Hist. Orchid. t. 5 (1822).

Ophrys dubia Thouars (1822: t.1).

Amphorchis dubia Thouars (1822: t.1), nom. superfl.

Imermamphis dubia Thouars (1822: t. 1), nom. superfl.

Rodriguezia mascarenensis Spreng. (Sprengel 1826: 719).

Amphorkis nilarmis Steudel (1840: 80).

Arnottia inermis (Thouars) S. Moore in Baker (1877: 339).

Arnottia mauritiana A. Rich. (Richard 1828: 30). Type: Bourbon, Commerson s.n. in Herb. Richard, dated 1771 (lectotype P!, selected here).

There are several specimens in Richard’s herbarium in P; none are specifically noted as being Commerson’s but one specimen is annotated by Richard with his description and is dated 1771, during the period that Commerson was on Bourbon. We therefore select this specimen as the lectotype of Arnottia mauritiana.

Cynorkis imbellis (Frapp. ex Cordem.) Hermans & P. J. Cribb comb. nov. Type: Réunion, St. Pierre, de la plaine des Cafres au coteau maigre, 1500 – 1200 m, Potier s.n. (not found).

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Hemiperis imbellis Frapp. ex Cordem., Fl. Réunion: 244 (Frappier 1895).

Arnottia imbellis (Frapp. ex Cordem.) Schltr. (Schlechter 1915: 398).

Cynorkis simplex (Frapp. ex Cordem.) Hermans & P. J. Cribb comb. nov. Type: Réunion, Richard s.n. (holotype, not found).

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Hemiperis simplex Frapp. ex Cordem., Fl. Réunion: 248 (Frappier 1895).

Arnottia simplex (Frapp. ex Cordem.) Schltr. (Schlechter 1915: 398).

Physoceras

Schlechter (1924: 78) established the genus Physoceras based on four new species collected by Perrier de la Bâthie in Madagascar. The genus currently comprises 12 species and one variety, mainly from Madagascar and two from Réunion with a historical record from Mauritius. They include terrestrial, epiphytic and lithophytic plants that grow in a variety of habitats, the great majority above 1000 m in northern Madagascar. Schlechter distinguished his new genus from Cynorkis on it having a perennial long stem with a single leaf borne towards the middle, a short inflated spur which is first contracted and then expanded and anther canals that are lobed at the end forming a bursicule covering the viscidium. These characters are not always clear or present, especially in the more recently described species, all the characteristics overlap with those in species of Cynorkis. The length of stem and placement of the leaf varies considerably, even within single species of Physoceras: in P. boryanum (A. Rich.) Bosser (1980: 257), for example, the leaf can be in the lower quarter of the stem, in other collections it can be in the upper half. Leaf shape, position and size is very variable in Cynorkis, ranging from basal to several distributed along the stem. A number of Cynorkis, including C. fimbriata H. Perrier ex Hermans (in Hermans et al. 2007: 291), C. lentiginosa Hermans, Andriant. & Sieder described above and C. dens-serpens Hermans & Cribb (2014: 1), also have a leaf positioned high on the stem. The short inflated spur is also not unique to Physoceras with several typical Cynorkis having the same feature, including C. boinana Schltr. (Schlechter 1913: 150) and C. brachystachya Bosser (1980: 260). P. bifurcum H. Perrier (1955: 254) has an expanded bifurcate spur. Some mainland African Cynorkis, including C. uncata (Rolfe) Kraenzl. (Kraenzlin 1902: 53) have a more or less lengthened stem and a short inflated spur.

The bursicule lobes at the apex of the rostellum arms are not evident in some Physoceras: with P. lageniferum H. Perrier (1955: 254) having a gynostemium similar to that of Cynorkis species in section Hemiperis and P. bifurcum that of section Gibbosorchis. Because of the lack of reliable material, no phylogenetic work has been done on this group and this is unlikely to change in the near future, meanwhile the morphological comparison is conclusive.

Considering the overlap of all characteristics between the two genera and the existence of species with intermediate morphology it is proposed to include Physoceras within the genus Cynorkis; therefore the following combinations have to be made:

Cynorkis Thouars (1809: 317).

Physoceras Schltr. (Schlechter 1924: 78), synon. nov. Type species: P. bellum Schltr. (Schlechter 1924: 79) = Cynorkis unguiculata Hermans & P. J. Cribb (lectotype selected in Index Nom. Gen. 64/24066, 1967), synon. nov.

Cynorkis australis (Boiteau) Hermans & P. J. Cribb comb. nov. Type: Madagascar, Ambondrombe, S. Betsileo, 1947, Boiteau — Jard. Bot. Tananarive 4921 (holotype P!).

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Physoceras australe Boiteau, Bull. Trimestriel Acad. Malgache, n.s., 24: 88 (1942).

Cynorkis betsomangensis (Bosser) Hermans & P. J. Cribb comb. nov. Type: Madagascar, Androranga valley, Bemarivo, 1950, Humbert & Capuron 24370 (holotype P!).

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Physoceras betsomangense Bosser, Adansonia 20: 258 (1980).

Cynorkis bifurca (H. Perrier) Hermans & P. J. Cribb comb. nov. Type: Madagascar, NE, eastern summit of Marojejy, W of high Manantenina, 1949, Humbert & Cours 23878 [P00117707] (holotype P!), [P00117708] (isotype P!).

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Physoceras bifurcum H. Perrier, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 254 (1955).

Cynorkis bobyi Hermans & P. J. Cribb nom. nov. Type: Madagascar, S of Mt Tsaratanana, Sambirano valley, 1923, Perrier 15710 [P00117701] (holotype P!); [P00117702] (isotype P!)

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Physoceras perrieri Schltr., Repert. Spec. Nov. Regni Veg. Beih. 33: 82 (Schlechter 1924).

Owing to the earlier use of Cynorkis perrieri Schltr. (Schlechter 1916: 310), this name cannot be used for this species. We therefore propose the epithet bobyi, for the new combination, referring to Henri Perrier de la Bâthie’s dog Boby who accompanied him during his exploration of Madagascar.

Cynorkis boryana (A. Rich.) Lindl. (Lindley 1835: 331). Type: Ile de France (Mauritius), La Pouce, 1825, Bory de St. Vincent s.n. (holotype [P0051595]!)

Gymnadenia boryana A. Rich. (Richard 1828: 26).

Physoceras boryanum (A. Rich.) Bosser (1980: 257) synon. nov.

Cynorkis boryana var. aristei (J.-B. Castillon) Hermans & P. J. Cribb comb. nov. Type: Réunion, Plaine des Palmistes, 2007, J.-B.Castillon 1 (holotype P not found; isotype: TAN).

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Physoceras boryanum var. aristei J.-B.Castillon, Richardiana 11: 14 (2010).

Cynorkis castillonii (P. Bernet) Hermans & P. J. Cribb comb. nov. Type: Réunion, Plaine des Palmistes, 2001, Bernet 3 (holotype P!)

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Physoceras castillonii P. Bernet, Richardiana 10: 145 (2010a, b).

Cynorkis epiphytica (Schltr.) Hermans & P. J. Cribb comb. nov. Type: Madagascar, Mt Tsaratanana, 1922, Perrier 15268 (holotype P!).

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Physoceras epiphyticum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 33: 80 (Schlechter 1924).

Cynorkis hyacinthina Hermans & P. J. Cribb nom. nov. Type: Madagascar, Masoala, 1912, Perrier 11332 (holotype P!).

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Physoceras violaceum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 33: 83 (Schlechter 1924).

Owing to the earlier use of Cynorkis violacea Schltr. (Schlechter 1913: 155), this epithet cannot be used for this species. We therefore propose the epithet hyacinthina, for the new combination, referring to the violet colour of the flowers.

Cynorkis lagenifera (H. Perrier) Hermans & P. J. Cribb comb. nov. Type: Madagascar, Lokoho valley, Mt Beondroka N of Maroambihy, 1949, Humbert 23603 (holotype [P00114060]!); (isotypes [P00114061-3]!)

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Physoceras lageniferum H. Perrier, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 254 (1955).

Cynorkis mesophylla Schltr. (Schlechter 1916: 308). Type: Madagascar, Mt Tsaratanana, 1912, Perrier 11355 (holotype P!).

Physoceras mesophyllum (Schltr.) Schltr. (Schlechter 1924: 82).

Cynorkis rotundifolia (H. Perrier) Hermans & P. J. Cribb comb. nov. Type: Madagascar, Mts between Vohemar & Ambilobe, N of Mt Tsaratanana, 1949, Decary 14701 (holotype P!).

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Physoceras rotundifolium H. Perrier, Notul. Syst. (Paris) 14: 144 (1951).

Cynorkis unguiculata Hermans & P. J. Cribb nom. nov. Type: Madagascar, Centre, Tsaratanana Massif, c. 2000 m, Oct. 1923, Perrier 15267 (holotype [P00115693]!; (isotype [P00115694]!). Madagascar, Centre, Mt Tsaratanana, 1500 m, Jan. 1923, Perrier 15709 (syntype P!).

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Physoceras bellum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 33: 79 (Schlechter 1924).

Owing to the earlier use of Cynorkis bella Schltr. (Schlechter 1924: 42), this epithet cannot be used for this species. We therefore propose the epithet unguiculata, for the new combination, referring to the claw-like spur.