Revision of the genus Benthamia A. Rich. (Orchidaceae, Orchidioideae, Habenariinae) Author Hervouet, Jean-Michel Fédération France Orchidées (FFO) and Muséum national d’Histoire naturelle, Direction générale déléguée aux collections, Herbier national, case postale 39, 57 rue Cuvier 75005 Paris (France) jmhervouet @ free. fr jmhervouet@free.fr Author Descourvières, Pascal France Orchidées (FFO), 27 rue Auguste Blanqui, 94400 Vitry-sur-Seine (France) descourvieres. pascal @ neuf. fr descourvieres.pascal@neuf.fr Author Hermans, Johan , Royal Botanic Gardens, Kew, Richmond, Surrey, TW 9 3 AE (United Kingdom) and Core Facility, Botanical Garden, University of Vienna, Rennweg 14, 1030, Vienna (Austria) j. hermans @ kew. org j.hermans@kew.org Author Longou, Ludivine Atelier d'iconographie scientifique, UAR 2047 DoHNÉE, MNHN / CNRS, Muséum national d’histoire naturelle, 57 rue Cuvier, 75005 Paris (France) ludivine. longou @ mnhn. fr ludivine.longou@mnhn.fr text Adansonia 2025 3 2025-07-21 47 11 171 249 https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/adansonia2025v47a11.pdf journal article 313251 10.5252/adansonia2025v47a11 b8e5d9ec-7e27-4bbf-a8a8-6cee2bf7465f 1639-4798 16262489 Benthamia perfecunda H.Perrier ( Figs 81 ; 82 ; 83 ) Notulae Systematicae 14 (2) ( Perrier 1951: 140 ). Benthamia herminioides subsp. angustifolia H.Perrier , Bulletin de la Société botanique de France 81 ( Perrier 1934: 37 ). — Type : Madagascar • Haute Matsiatra , mont Tsaratanana ; 14°01’17”S , 48°57’60”E ; alt. 2600 m ; IV.1924 ; H. Perrier 16503 ; 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. Humbert 23754: 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 . — Madagascar and 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 Réunion and 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. Homolle 1174 ; P [ P01805136 ] ! • Alaotra-Mangoro , forêt d’Andasibe sur l’Onive ; 18°28’S , 48°28’E ; 1200 m ; II.1925 ; H. Perrier 17089 ; 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 [ P02115322 plant on the left] ! • Haute Matsiatra , Andringitra ; 22°07’40”S , 46°51’48”E ; IV.1964 ; J. Bosser 19647 ; P [ P00692264 ] ! • Analamanga , Angavokely ; 18°55’36”S , 47°45’07”E ; 26.II.1988 ; B. Pettersson et al. 189 ; P [ P02115316 ] ! • Sava , Réserve naturelle de Marojejy ; alt. 1900-2133 m ; 15.II.1989 ; J.S. Miller 4164 ; MO [ MO3022619 ] ! • Sava , Réserve Naturelle de Marojejy ; 14°26’S , 49°13’E ; alt. 1900-2133 m ; 15.II.1989 ; J.S. Miller 4168 ; TAN ! • Sava , Marojejy , au dessus du village Manantenina ; 14°26’S , 49°44’E ; alt. 1300-1450 m , III.1990 ; A. Randrianasolo 111 ; K !, MO !, TAN !. France , La Réunion • Bois de la rivière de l’Est ; 2.VI.1851 ; L.H. Boivin s.n. ; P [ P00334799 specimen 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.Bosser 11490 and 11491 ; P [ P00738403 , P00738378 ] ! • Bébour forest ; 21°06’40”S , 55°33’47”E ; alt. 1300 m ; 26.III.1969 ; T. Cadet 2106 ; REU [ REU006443 ] ! • Plateau de Bébour ; 21°06’S , 55°33’E ; III.1970 ; J. Bosser 20068 ; P [ P00738405 ] ! • Gîte du volcan ; 21°06’36”S , 55°29’46”E ; IV.1970 ; J. Bosser 20058 ; P [ P00738404 ] ! • Sentier du Bras Cabot , îlet de Patience ; 21°06’S , 55°37’E ; 19.II.1971 ; J. Bosser 20574 ; P [ P00738377 ] ! • Montée de la Plaine des Chicots ; 20°58’26”S , 55°25’47”E ; II.1971 ; J. Bosser 20684 ; P [ P00738397 ] ! • Plateau de Bébour ; 21°06’S , 55°33’E ; II.1971 ; J. Bosser 20703 ; P [ P00738311 ] ! • Coteau Kerveguen ; 21°07’27”S , 55°30’16”E ; II.1971 ; J. Bosser 20743 ; 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. Cadet 3225 ; REU [ REU00644 , REU017470 ] ! • Brûlé de Saint-Denis ; 20°55’S , 55°26’E ; 19.III.1974 ; J. Bosser 21658 ; P [ P00738379 and 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. Cadet 4615 ; REU [ REU006439 ] ! • Plaine des Cafres , base du Piton Mare à Boue ; 21°09’35”S , 55°34’35”E ; 30.III.1974 ; J. Bosser 21781 ; P [ P00738383 ] ! • Plaine des Cafres , Piton Mare à Boue ; 21°09’35”S , 55°34’35”E ; 30.III.1974 ; J. Bosser 21771 and 21771bis ; P [ P00738384 , P00738533 ] ! • Brûlé de Saint-Denis ; 20°55’S , 55°26’E ; 2.V.1974 ; J. Bosser 21925 ; P [ P00738380 and spirit]!; Brûlé de Saint-Denis ; 20°55’S , 55°26’E ; 9.V.1974 ; J. Bosser 21927 ; P [ P00738382 ] ! • Descente Plaine des sables à la ravine Langevin ; 21°15’17”S , 55°39’45”E ; 10.V.1976 ; J. Bosser 22126 ; P [ P00738400 ] ! • Coteau Kervéguen ; 21°07’27”S , 55°30’16”E ; 16.VI.1976 ; J. Bosser 22319 ; P [ P00738396 ] ! • Sentier du Bras Cabot , îlet de Patience ; 21°06’S , 55°37’E ; 7.IV.1978 ; J. Bosser 22465 ; P [ P00738406 ] ! • Sentier du Piton des Neiges , Cilaos ; 21°07’06”S , 55°29’12”E ; 15.IV.1978 ; J. Bosser 22508 ; P [ P00738399 ] ! • Descente Plaine des Sables , Le Tremblet ; 21°17’14”S , 55°43’20”E ; 20.IV.1978 ; J. Bosser 22538 ; P [ P00738398 ] ! • SF Roche-Écrite ; 21°00’55”S , 55°27’39”E ; alt. 1320 m ; 19.III.1988 ; R. Lavergne RL1367 ; REU [ REU007766 ] ! • Saint-Joseph , Caverne de Cotte ; 21°14’16”S , 55°38’18”E ; alt. 2240 m ; III.2004 ; J. Férard et al. 1043 ; CBNM !. FIG . 83. — Benthamia perfecunda H.Perrier,Bourg-Murat,La Réunion,26 March 2013. Photograph by Jean-Michel Hervouet. NOTES After examining the types , we see no significant difference between Benthamia herminioides subsp. angustifolia and Benthamia perfecunda . In his publication of Benthamia perfecunda Perrier only cited Benthamia macra and Benthamia verecunda as similar species. Perrier stated that the leaves of Benthamia herminioides subsp. angustifolia are 5 cm long, this is a mistake since one leaf of the type is 11 cm long, which falls within the Benthamia perfecunda leaf 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 ). A very 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.herminioides subsp. angustifolia becoming a heterotypic synonym. Surprisingly this species has long been overlooked in La Réunion , only Bernet illustrated it as Benthamia sp. 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 Benthamia sp. 4 ), Szelengowicz & Tamon (2013: 245 , as Benthamia herminioides subsp. arcuata , photograph on the left only), Hervouet (2018: 171) , Szelengowicz & Tamon (2013: 252) . FIG . 84. — Distribution map of Benthamia perfecunda H.Perrier. DESCRIPTION Erect terrestrial herb 30-80 cm tall. Tubers 2, roots around 4-6, up to 6 cm long, 1-2 mm in diameter. Stem cylindrical, 2-4 mm in 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 cm long, bearing 20-60 flowers, broadly cylindrical but in fact arranged in 3 slightly spiralling rows, 2-5 mm apart. 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 mm long, 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 mm long, 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 mm high, 0.4-1.2 mm wide, 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 mm long.