Perrier 1951: 140 Benthamia herminioides subsp. angustifolia H.Perrier Perrier 1934: 37 Revision of the genus Benthamia A. Rich. (Orchidaceae, Orchidioideae, Habenariinae) Hervouet, Jean-Michel Descourvières, Pascal Hermans, Johan Longou, Ludivine Adansonia 2025 3 2025-07-21 47 11 171 249 LHN3 H. Perrier H. Perrier [277,630,1491,1519] Liliopsida Orchidaceae Benthamia Plantae Asparagales 59 228 Tracheophyta species perfecunda  ( Figs 81; 82; 83)     Notulae Systematicae14 (2) (  Perrier 1951: 140).    Benthamia herminioidessubsp. angustifoliaH.Perrier, Bulletin de la Société botanique de France81(  Perrier 1934: 37). — Type:  Madagascar• Haute Matsiatra, mont Tsaratanana; 14°01’17”S, 48°57’60”E; alt.  2600 m;  IV.1924;  H. Perrier16503; holotype: P[ P00094516]!, syn. nov.    TYPE  MATERIAL. —   Madagascar• Sava, sommet oriental du massif de Marojejy; 14°26’57”S, 49°43’57”E; alt.  1830-2135 m;  26.III.1949;  H. Humbert23754: lectotype: P[ P00094564]!, selected by Hermans & Cribb (2021: 76);  isolectotypes: MO[ MO2176327]!,  P[ P00094565, P00094566]!.   ETYMOLOGY. — “ Perfecunda”: Latin for very fertile, alluding to the fact that all flowers produce fruits.   PHENOLOGY. — From January to April.   DISTRIBUTION AND ECOLOGY. — Madagascarand La Réunion. Widespread, from 600 to 2000 m, in medium altitude moist evergreen forest, montane grasslands and ericoid thickets ( Fig. 84).   CONSERVATION. — This species is widespread in La Réunionand therefore has a global conservation status LC (Least Concern). It is Data Deficient in Madagascar(DD) and is currently known in about ten localities in this country, probably because it has been overlooked so far.  ADDITIONAL SPECIMENS EXAMINED. —   Madagascar• sine loc.;  A.M. Homolle1174; P[ P01805136]! •  Alaotra-Mangoro, forêt d’Andasibe sur l’Onive; 18°28’S, 48°28’E;  1200 m;  II.1925;  H. Perrier17089; P[ P00692456]! •  Sava, Vallée inférieure de l’Androranga, massif du Betsomanga; 14°15’30”S, 49°44’00”E; alt.  1300-1350 m;  XI.1950;  H. Humbert et al.24331; P[ P00692279]! •  Diana, massif de Marivorahona; 13°46’40”S, 48°59’30”E; alt.  2000-2244 m;  18-26.III.1951;  H. Humbert et al.25753; P[ P02115322plant on the left]! •  Haute Matsiatra, Andringitra; 22°07’40”S, 46°51’48”E;  IV.1964;  J. Bosser19647; P[ P00692264]! •  Analamanga, Angavokely; 18°55’36”S, 47°45’07”E;  26.II.1988;  B. Petterssonet al. 189; P[ P02115316]! •  Sava, Réserve naturelle de Marojejy; alt.  1900-2133 m;  15.II.1989;  J.S. Miller4164; MO[ MO3022619]! •  Sava, Réserve Naturelle de Marojejy; 14°26’S, 49°13’E; alt.  1900-2133 m;  15.II.1989;  J.S. Miller4168; TAN! •  Sava, Marojejy, au dessus du village Manantenina; 14°26’S, 49°44’E; alt.  1300-1450 m,  III.1990;  A. Randrianasolo111; K!,  MO!, TAN!.   France, La Réunion• Bois de la rivière de l’Est;  2.VI.1851;  L.H. Boivins.n.; P[ P00334799specimen on the right could be  Benthamia spiraloides]! •  Sentier du Bras Cabot, Plaine des Palmistes; 21°06’S, 55°37’E; alt.  1400-1500 m;  VI.1957;  J.Bosser11490and 11491; P[ P00738403, P00738378]! •  Bébour forest; 21°06’40”S, 55°33’47”E; alt.  1300 m;  26.III.1969;  T. Cadet2106; REU[ REU006443]! •  Plateau de Bébour; 21°06’S, 55°33’E;  III.1970;  J. Bosser20068; P[ P00738405]! •  Gîte du volcan; 21°06’36”S, 55°29’46”E;  IV.1970;  J. Bosser20058; P[ P00738404]! •  Sentier du Bras Cabot, îlet de Patience; 21°06’S, 55°37’E;  19.II.1971;  J. Bosser20574; P[ P00738377]! •  Montée de la Plaine des Chicots; 20°58’26”S, 55°25’47”E;  II.1971;  J. Bosser20684; P[ P00738397]! •  Plateau de Bébour; 21°06’S, 55°33’E;  II.1971;  J. Bosser20703; P[ P00738311]! •  Coteau Kerveguen; 21°07’27”S, 55°30’16”E;  II.1971;  J. Bosser20743; P[ P00738385]! •  Piton Taïpoul, Foc-Foc, piton de la Fournaise; 21°17’07”S, 55°42’04”E; alt.  2100 m;  16.IV.1971;  T. Cadet3225; REU[ REU00644, REU017470]! •  Brûlé de Saint-Denis; 20°55’S, 55°26’E;  19.III.1974;  J. Bosser21658; P[ P00738379and spirit]! •  Pied de Piton Mare à Boue, Plaine des Cafres; 21°09’35”S, 55°34’35”E; alt.  1550 m;  30.III.1974;  T. Cadet4615; REU[ REU006439]! •  Plaine des Cafres, base du Piton Mare à Boue; 21°09’35”S, 55°34’35”E;  30.III.1974;  J. Bosser21781; P[ P00738383]! •  Plaine des Cafres, Piton Mare à Boue; 21°09’35”S, 55°34’35”E;  30.III.1974;  J. Bosser21771and 21771bis; P[ P00738384, P00738533]! •  Brûlé de Saint-Denis; 20°55’S, 55°26’E;  2.V.1974;  J. Bosser21925; P[ P00738380and spirit]!;  Brûlé de Saint-Denis; 20°55’S, 55°26’E;  9.V.1974;  J. Bosser21927; P[ P00738382]! •  Descente Plaine des sables à la ravine Langevin; 21°15’17”S, 55°39’45”E;  10.V.1976;  J. Bosser22126; P[ P00738400]! •  Coteau Kervéguen; 21°07’27”S, 55°30’16”E;  16.VI.1976;  J. Bosser22319; P[ P00738396]! •  Sentier du Bras Cabot, îlet de Patience; 21°06’S, 55°37’E;  7.IV.1978;  J. Bosser22465; P[ P00738406]! •  Sentier du Piton des Neiges, Cilaos; 21°07’06”S, 55°29’12”E;  15.IV.1978;  J. Bosser22508; P[ P00738399]! •  Descente Plaine des Sables, Le Tremblet; 21°17’14”S, 55°43’20”E;  20.IV.1978;  J. Bosser22538; P[ P00738398]! •  SF Roche-Écrite; 21°00’55”S, 55°27’39”E; alt.  1320 m;  19.III.1988;  R. LavergneRL1367; REU[ REU007766]! •  Saint-Joseph, Caverne de Cotte; 21°14’16”S, 55°38’18”E; alt.  2240 m;  III.2004;  J. Férardet al. 1043; CBNM!.   NOTES After examining the types, we see no significant difference between  Benthamia herminioidessubsp. angustifoliaand  Benthamia perfecunda. In his publication of  Benthamia perfecundaPerrieronly cited  Benthamia macraand  Benthamia verecundaas similar species. Perrier stated that the leaves of  Benthamia herminioidessubsp. angustifoliaare 5 cmlong, this is a mistake since one leaf of the typeis 11 cmlong, which falls within the  Benthamia perfecundaleaf dimensions. The fact that all flowers are pollinated is observed on all specimens and is a strong indication that the plant is autogamous. This species is very different from  Benthamia herminioides,even though it was initially described as the subspecies angustifolia. Perrier himself says that the stigmas are very different from those of other subspecies of  Benthamia herminioides. It is generally a larger plant, with differently shaped leaves (linear to grasslike vs elliptic-lanceolate to lanceolate for  Benthamia herminioides). The flowers are fleshy,campanulate,and uniformly green (vs with calyx green and corolla white for  Benthamia herminioides). Avery distinctive and unique feature in the genus is the 3-ranked rachis (vs secund). Changing the rank from subspecies to species was therefore necessary. Then the earliest name at the rank of species has priority (article 11.2 of the International Code of Nomenclature), which is  Benthamia perfecunda,  B.herminioidessubsp. angustifoliabecoming a heterotypic synonym. Surprisingly this species has long been overlooked in La Réunion, only Bernet illustrated it as  Benthamiasp. 4( Bernet 2010). Jean Bosser consistently noted on his specimens that there were three helicoidal rows of flowers. See also: Perrier (1939: 38), Cribb & Hermans (2009: 43), Cribb & Hermans (2009: 48), Bernet (2010: 141, as  Benthamiasp. 4), Szelengowicz & Tamon (2013: 245, as  Benthamia herminioidessubsp. arcuata ,photograph on the left only), Hervouet (2018: 171), Szelengowicz & Tamon (2013: 252).   DESCRIPTION Erect terrestrial herb 30-80 cmtall. Tubers 2, roots around 4-6, up to 6 cmlong, 1-2 mmin diameter. Stem cylindrical, 2-4 mmin diameter, covered below by blackish sheaths. Leaves 2-4, cauline in the basal half of the stem or slightly higher, 5-15 cm× 5-12 mm, erect, linear or grass-like, acute, amplexicaul at base, with 7-8 veins. 4-5 sterile bracts in the upper half of the stem. Rachis terminal and dense, 6-15 cmlong, bearing 20-60 flowers, broadly cylindrical but in fact arranged in 3 slightly spiralling rows, 2-5 mmapart. Floral bracts acicular, 3-10 × 1.1-1.8 mm, slightly smaller than the ovary, rarely a little longer. Ovary elongated, a little arcuatereflexed, green, fusiform, at right angle to the rachis at the end of anthesis, 3-8 × 1-2 mm. Flowers not opening fully, campanulate, 2-3 mmlong, nearly aligned with the ovary, green or greenish-yellow; dorsal sepal oblong, 1.5-2.8 × 0.5-1.1 mm, 1-veined; lateral sepals asymmetrically oblong, concave, spreading, 1.5-3.1 × 0.4-1.2 mm; petals broadly oval, 1.2-2.9 × 0.3-2.1 mm, shorter than the lateral sepals, adnate to the dorsal sepal, forming a hood over the lip; lip ovate, trilobed at apex, obtuse, calceolate, fleshy, 1.5-3.2 × 1-3.2 mm, without callus, the lobes reduced to 3 rounded teeth, lateral lobes 0.5 mmlong, central lobe equal or longer, blunt, concave in proximal part, lateral lobes folded upwards, up to the petals, so that a rectangular entrance into the flower is formed; spur 0.4-1.5 × 0.2-0.8 mm, horizontal and parallel to the ovary or touching it, obtuse, cylindrical or slightly flattened, not narrowed at the base, straight or curved along the ovary. Column 0.5-1.1 mmhigh, 0.4-1.2 mmwide, obtuse, anther with an obtuse apiculus; auricles very short, less than a quarter of the length of the anther; rostellum thin, more tri-crenate than trilobed; stigmata distinct, well developed, thick, porrect-claviform, 0.5 mmlong. 5898708689 1949-03-26 P H. Humbert Madagascar 1983 -14.449166 sommet oriental du massif de Marojejy 21 49.732502 Sava 59 228 P00094564 1 lectotype 5898708918 [419,774,1898,1922] 1949-03-26 MO H. Humbert Madagascar 1983 -14.449166 sommet oriental du massif de Marojejy 21 49.732502 Sava 59 228 MO2176327 1 isolectotype 5898709092 [134,407,1926,1950] 1949-03-26 P H. Humbert Madagascar 1983 -14.449166 sommet oriental du massif de Marojejy 21 49.732502 Sava 59 228 P00094565, P00094566 1 isolectotype 5898708798 P A. M. Homolle Madagascar Madagascar 59 228 P01805136 1 5898708507 1925-02 P H. Perrier 1200 -18.466667 Alaotra-Mangoro 1275 48.466667 59 228 P00692456 1 5898708987 1950-11 P H. Humbert et al. 1325 -14.258333 Sava 21 49.733334 59 228 P00692279 1 5898708821 1951-03-18 1951-03-26 1951-03-18 P H. Humbert et al. 2122 -13.777778 Diana 21 48.99167 59 228 P02115322 1 5898708870 1964-04 P J. Bosser 60 229 -22.127777 Haute Matsiatra 21 46.86333 59 228 P00692264 1 5898708692 1988-02-26 P B. Pettersson -18.926666 Analamanga 21 47.751945 60 229 P02115316 1 5898708738 1989-02-15 MO J. S. Miller 2016 Sava 60 229 MO3022619 1 5898708655 1989-02-15 TAN J. S. Miller 2016 -14.433333 Sava 1288 49.216667 60 229 1 5898708815 1990-03 K A. Randrianasolo 1375 -14.433333 Sava 1288 49.733334 60 229 1 5898708874 [218,335,400,424] MO, TAN 1288 60 229 1 5898709033 1851-06-02 P L. H. Boivin France Bois de la riviere de l'Est La Reunion 60 229 P00334799 1 5898708403 1957-06 P J. Bosser Réunion 1450 -21.1 Sentier du Bras Cabot 1265 55.616665 60 229 P00738403, P00738378 1 5898709139 1969-03-26 REU T. Cadet Réunion 1300 -21.11111 Bebour forest 21 55.563053 Bebour forest 60 229 REU006443 1 5898708557 1970-03 P J. Bosser Réunion -21.1 Plateau de Bebour 1265 55.55 Plateau de Bebour 60 229 P00738405 1 5898708779 1970-04 P J. Bosser Réunion -21.11 Gite du volcan 21 55.496113 60 229 P00738404 1 5898708314 1971-02-19 P J. Bosser Réunion -21.1 Sentier du Bras Cabot 1265 55.616665 60 229 P00738377 1 5898708330 1971-02 P J. Bosser Réunion -20.97389 Montee de la Plaine des Chicots 21 55.42972 Montee de la Plaine des Chicots 60 229 P00738397 1 5898708521 1971-02 P J. Bosser Réunion -21.1 Plateau de Bebour 1265 55.55 60 229 P00738311 1 5898708748 1971-02 P J. Bosser Réunion -21.124166 Coteau Kerveguen 21 55.504444 Coteau Kerveguen 60 229 P00738385 1 5898708644 1971-04-16 REU T. Cadet Réunion 2100 -21.285276 Piton Taipoul 21 55.70111 Foc-Foc 60 229 REU00644, REU017470 1 5898708554 1974-03-19 P J. Bosser Réunion -20.916666 Brule de Saint-Denis 1266 55.433334 Brule de Saint-Denis 60 229 P00738379 1 5898708564 1974-03-30 REU T. Cadet Réunion Plaine des Cafres 1550 -21.159721 Pied de Piton Mare a Boue 21 55.57639 60 229 REU006439 1 5898708584 1974-03-30 P J. Bosser Réunion -21.159721 Plaine des Cafres 21 55.57639 60 229 P00738383 1 5898708957 1974-03-30 P J. Bosser Réunion -21.159721 Plaine des Cafres 21 55.57639 60 229 P00738384, P00738533 1 5898708574 1974-05-02 1974-05-09 1974-05-02 P J. Bosser Réunion -20.916666 Brule de Saint-Denis 1266 55.433334 Brule de Saint-Denis 60 229 P00738380, P00738382 1 5898708996 1974-05-02 1974-05-09 1974-05-02 P J. Bosser Réunion -20.916666 Brule de Saint-Denis 1266 55.433334 Brule de Saint-Denis 60 229 P00738380, P00738382 1 5898708480 1976-05-10 P J. Bosser Réunion -21.254723 Descente Plaine des sables a la ravine Langevin 21 55.662502 60 229 P00738400 1 5898709143 1976-06-16 P J. Bosser Réunion -21.124166 Coteau Kerveguen 21 55.504444 60 229 P00738396 1 5898708875 1978-04-07 P J. Bosser Réunion -21.1 Sentier du Bras Cabot 1265 55.616665 60 229 P00738406 1 5898708690 1978-04-15 P J. Bosser Réunion -21.118334 Sentier du Piton des Neiges 21 55.486668 Cilaos 60 229 P00738399 1 5898708791 1978-04-20 P J. Bosser Réunion -21.287222 Le Tremblet 21 55.72222 Descente Plaine des Sables 60 229 P00738398 1 5898708543 1988-03-19 REU R. Lavergne Réunion 1320 -21.015278 SF Roche-Ecrite 21 55.460835 60 229 REU007766 1 5898708722 2004-03 CBNM J. Ferard Réunion 2240 -21.237778 Saint-Joseph 21 55.638336 Caverne de Cotte 60 229 1