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 erinacea (Cordem.) Hermans & P.J.Cribb ( Figs 33 ; 34 ; 35 ; 36 ) Lankesteriana 21 (2) ( Hermans & Cribb 2021: 74 ). — Habenaria erinacea Cordem ., Flore de l’île de la Réunion : 261 ( Cordemoy 1895 ). — Peristylus erinaceus Frapp. , Catalogue provisoire des orchidées de la Réunion : 9 ( Frappier 1880 ), nomen nudum. Peristylus secundiflorus Boivin ex S. Moore , non Kraenzl., Journal of Botany 5: 293 ( Moore 1876 ) , nomen nudum. Habenaria secundiflora Cordem ., Flore de l’île de la Réunion : 260, 551 ( Cordemoy 1895 ). Benthamia nigrescens Schltr., Beihefte zum Botanischen Centralblatt 34 (2): 301 ( Schlechter 1916 ) . — Type: Madagascar • Vakinankaratra , flanc est du mont Tsiafajavona ; 19°22’S , 47°19’E ; alt. 2000 m ; III.1921 ; H. Perrier 13506 ; holotype: P [ P00094545 ]!. Benthamia nigrescens subsp. borbonica H.Perrier , Bulletin de la Société botanique de France 81: 30 ( Perrier 1934 ) . — Type: France , La Réunion • L.H. Boivin s.n. ; lectotype: P[P00334740]!, here designated; isolectotype: P[P00334741]!. Benthamia nigrescens subsp. decaryana H.Perrier , Bulletin de la Société botanique de France 81 ( Perrier 1934: 30 ). — Type: Madagascar • Diana , Ankaizina ; 14°30’S , 48°55’E ; alt. 1700 m ; 19.IV.1923 ; R. Decary 1982 ; holotype: P [ P00094548 ]!. Benthamia nigrescens subsp. humblotiana H.Perrier , Bulletin de la Société botanique de France 81 (1934). — Type: Madagascar • Diana , Mt. Tsaratanana ; 14°01’17”S , 48°58’00”E ; alt. 2000 m ; IV.1924 ; H. Perrier 16110 ; lectotype: P[P00094552]!, here designated; isolectotype: P[P00094553]!. FIG . 36. — Benthamia erinacea (Cordemoy) Hermans & P.J.Cribb : Montagne d’Ambre, 5 April 2024. Side view of flower showing the spur. Photograph by Jean-Michel Hervouet. FIG . 35. — Benthamia erinacea (Cordemoy) Hermans & P.J.Cribb :Bourg-Murat, La Réunion, 19 April 2018. Photograph by Jean-Michel Hervouet. Benthamia nigrescens subsp. secundiflora H.Perrier , Bulletin de la Société botanique de France 81: 30 ( Perrier 1934 ) . — Based on Peristylus secundiflorus Boivin ex S. Moore. , which is a nomen nudum (see notes below). Benthamia nigrescens subsp. typica H.Perrier , Bulletin de la Société botanique de France 81: 30 ( Perrier 1934 ) . — Type : unknown. A wrong specimen has been cited by Perrier: H. Perrier 8078 , i.e. P00094439, which is Habenaria truncata . By subsp. typica Perrier meant Benthamia nigrescens . Benthamia catatiana H.Perrier , Bulletin de la Société botanique de France 81 ( Perrier 1934: 29 ). — Type: Madagascar • Anosy , Vallée d’Ambolo , au nord de Fort-Dauphin; 1 Jul., L.D.M. Catat 4338 ; lectotype: P[P00094478]!, here designated; isolectotype: P[P00094479]!, syn. nov . Benthamia cuspidata H.Perrier , Bulletin de la Société botanique de France 81: 29 ( Perrier 1934 ) . — Type: Madagascar • Atsinanana , Mananara , sur la côte est; 02.IX.1920 ; R. Decary 79 ; holotype 2-part specimen : P [ P00094493 , P00094494 ]!, syn. nov . Peristylus micranthus A.Rich. & Benthamia micrantha A.Rich. , nomen nudum in sched. , based upon specimens from La Réunion , ex Herb. Richard (K-LINDL!, W!). TYPE MATERIAL . — France , La Réunion • Plaine des Palmistes ; II.1883 ; Cordemoy s.n. Herb. Cordemoy ; lectotype : MARS [ MARS087704 ]!, designated by Hermans & Cribb (2021: 74) . ETYMOLOGY . — From Latin “ erinaceus ”, hedgehog, due to the bristled rachis. PHENOLOGY . — Mostly February to May, but occasionally up to October. DISTRIBUTION AND ECOLOGY . — Madagascar , La Réunion . Widespread but in Madagascar only in the eastern half, from 700 to 2400 m . Epiphytic, mostly in medium altitude moist evergreen forest ( Fig. 37 ). CONSERVATION . —The status of this species is of Least Concern (LC) since it is widespread in Madagascar and La Réunion . It was been deemed Endangered by IUCN in 2015, under the name Benthamia nigrescens , under criteria B2ab(i,ii,iii,iv,v) (https://doi.org/10.2305/ IUCN. UK .2015-2.RLTS.T69222181A69245307.en) but at that time all subspecies were considered separately. ADDITIONAL SPECIMENS EXAMINED . — Madagascar • Sine loc. ; L. Humblot s.n. ; P [ P00094551 ] ! • Diana , île de Nosy Be , dans les plaines ; 11.IV.1841 ; A. Pervillé 801 ; P [ P00094549 , P00094550 ] ! • Balfour s.n. ; 1875; K ! • Alaotra-Mangoro , massif de l’Andrangovalo au sud-est du lac Alaotra ( réserve de Zahamena ); 17°40’S , 48°45’E ; alt. 1200 m ; X.1937 ; H. Humbert et al. 17783 ; P [ P00094546 ] ! • Haute Matsiatra , Ambalavao ; 21.III.1953 ; H. Randriamiera RN5581 ; P [ P00692313 , P00692498 ] ! • Vakinankaratra , Manjakatompo ; alt. 1700-1800 m ; V.1960 ; J. Bosser 14443 ; TAN ! • Analamanga , lake Mantasoa ; 19°00’45”S , 47°50’17”E ; VIII.1962 ; J. Bosser 16153 ; P [ P00692431 ]! photograph n°721 by Jean Bosser • Alaotra-Mangoro , route de Lakato ; 19°11’30”S , 48°26’00”E ; VII.1963 ; J. Bosser 17176 ; P [ P00692238 ] ! • Diana , Montagne d’Ambre ; VI.1970 ; J. Bosser 20377 ; P [ P00692267 ] ! • Diana , Tsaratanana massif ; alt. 1800-2000; 8.V.1974 ; A. Gentry 11589 ; MO [ MO3248452 ] ! • Diana , Ambahatra ; 14°00’31”S , 48°25’49”E ; III.1999 ; L. Gautier et al. LG3647 ; G ! • Alaotra-Mangoro , road to Lakato ; 19°03’S , 48°21’E ; alt. 1041 m ; II.2000 ; J. Hermans 5561 ; K ! • Analamanga , Tampoketsa d’Ankazobe ; 18°12’56”S , 47°17’17”E ; 2000; J. Hermans 5187; K ! • Alaotra-Mangoro , Maromizaha ; 19.VIII.2022 ; L.R. Rajaovelona et al. RJL1421 ; K [ K001522014 ]!, TAN !. France , La Réunion • Sine loc. ; Cordemoy s.n. ; MARS [ MARS087702 , MARS087703 , MARS087704 !, MARS087706 ! and two specimens without number] • sine loc. ; A. Delteil s.n. ; P [ P00334755 ]!, BR [ BR0000021833151 ] ! • sine loc. ; A.N. Desvaux 259 ; P [ P00334756 ] ! • sine loc. ; L.H. Boivin 1068 ; W ! • sine loc. ; L.H. Boivin s.n. ; P [ P00334747 , P00334748 , P00334750 ] ! • sine loc. ; L.H. Boivin s.n. ; P [ P00738485 ]! among Cynorkis citrata • sine loc. ; L.H. Boivin s.n. ; P [ P00738486 ] ! • sine loc. ; C. Richard 501 and 501bis ; P [ P00334743 , P00334742 ] ! • sine loc. ; C. Richard s.n. ; P [ P00334744 ] ! • sine loc. ; C. Richard 387 ; P [ P00334745 !, P [ P00334746 ] ! • sine loc. ; VII.1837 ; Gaudichaud s.n. ; P [ P00334753 ] ! • sine loc. ; 1837; Gaudichaud s.n. ; G ! • II.1847 ; L.H. Boivin 1063 ; P [ P00738489 !, P00334751 !, P00334752 ] ! • sine loc. ; 1849; Giraudy s.n. ; P [ P00738478 ] ! • Plaine des Cafres ; 21°09’S , 55°34’E ; 6.VI.1851 ; L.H. Boivin s.n. ; P [ P00334749 ] ! • sine loc. ; 1867; Herb. A. Richard. R19909 ; W !, drawing of flower by Rchb.f., as Herminium spirale • sine loc. ; 26.II.1875 ; G. de l’Isle 69 ; P [ P00334754 ] ! • sine loc. ; 1932; s.c. ; s.n. ; P [ P00334757 ] ! • Bébour forest , Bélouve ; 21°07’58”S , 55°34’23”E ; alt. 1400 m ; IV.1956 ; J. Bosser 9464 ; P [ P00738461 ] ! • Bébour forest , Bélouve ; 21°07’58”S , 55°34’23”E ; alt. 1500 m ; IV.1956 ; J. Bosser 9465 ; P [ P00738463 , P00738465 ]!, MAU [ MAU0001989 ] ) • Hauts de Maka ; alt. 1600 m ; V.1957 ; J. Bosser 11762 ; P [ P00738468 ] ! • Brûlé de Saint-Denis ; 20°55’S , 55°26’E ; alt. 1200 m ; V.1957 ; J. Bosser 11842 ; P [ P00738462 ] ! • Grande Montée ; 20°56’S , 55°31’E ; alt. 1600 m ; V.1964 ; Staub 11123 ; MAU [ MAU0001991 ] ! • Saint-Denis , Hauts de Brûlé , Roche écrite path , Mamode forest camp; 20°55’S , 55°26’E ; alt. 1200 m ; III.1966 ; T. Cadet 417, Herb.Maur. n°12073 ; REU [ REU006408 ]!, MAU [ MAU0001990 ] ! • Bébour forest ; 21°06’40”S , 55°33’47”E ; alt. 1300 m ; 26.III.1969 ; T. Cadet 2018 ; REU [ REU006409 , REU006411 ] ! • Mafate , Petite Mavis , Sentier Augustave ; 21°02’34”S , 55°26’43”E ; alt. 1100 m ; 1.II.1970 ; R. Lavergne RL1792 ; REU [ REU007779 ] ! • Col de la Plaine des Palmistes , Plaine des Cafres ; 21°09’S , 55°34’E ; III.1970 ; J. Bosser 20057 ; P [ P00738474 ] ! • Plateau de Bébour ; 21°06’S , 55°33’E ; IV.1970 ; J. Bosser 20066 ; P [ P00738494 ] ! • Petite Plaine des Palmistes , berge de la Ravine Sèche ; alt. 1100 m ; 12.IV.1970 ; T. Cadet 2298 ; REU [ REU006410 , REU006412 ] ! • Plateau de Bébour ; 21°06’S , 55°33’E ; II.1971 ; J. Bosser 20694 ; P [ P00738470 ] ! • Plateau de Bébour ; 21°06’S , 55°33’E ; II.1971 ; J. Bosser 20708 ; P [ P00738473 ] ! • Plaine des Affouches ; 20°58’50”S , 55°25’00”E ; 13.II.1971 ; J. Bosser 20513 ; P [ P00738475 ! and spirit] • Saint-Philippe , Hauts de Basse Vallée ; alt. 1200 m ; 15.II.1971 ; T. Cadet 3043 ; REU [ REU006413 ] ! • Notre-Dame de la Paix , Plaine des Cafres ; 21°09’S , 55°34’E ; alt. 1600 m ; 18.II.1971 ; T. Cadet 3073 ; REU [ REU006414 ] ! • Sentier du Bras Cabot , îlet de Patience ; 21°06’S , 55°37’E ; 19.II.1971 ; J. Bosser 20573 ; P [ P00738479 ] ! • Forêt de Bébour ; 21°05’37”S , 55°36’43”E ; alt. 1300 m ; III.1973 ; D. Lorence 36 ; K !, MO ! • Forêt de Bébour ; 21°05’37”S , 55°36’43”E ; alt. 1300 m ; III.1973 ; D. Lorence R15 ; P [ P00738454 ]!, K !, MO ! • Forêt de Bébour ; 21°07’33”S , 55°34’21”E ; alt. 1300 m ; 26.III.1973 ; D. Lorence R17 ; K !, MO ! • Hauts de Saint-Benoit , forêt dans l’encaissement de la Rivière des Marsouins ; 21°05’37”S , 55°36’43”E ; alt. 350 m ; 18.IX.1973 ; T. Cadet 4429 ; REU [ REU006415 ] ! • Basse Vallée ; 3.III.1974 ; J. Bosser 21810 ; P [ P00738469 ] ! • Grand Matarum , cirque de Cilaos ; 21°07’19”S , 55°28’40”E ; 23.III.1974 ; J.Bosser 21698 ; P [ P00738471 , spirit]!, K ! • Grand Matarum , cirque de Cilaos ; 23.III.1974 ; J. Bosser 21698 bis ; P [ P00738477 ] ! • Sentier du morne de Fourche , cirque de Salazie ; 21°03’49”S , 55°26’38”E ; 28.III.1974 ; J. Bosser 21748 ; P [ P00738472 ] ! • Brûlé de Saint-Denis ; 20°55’S , 55°26’E ; 19.III.1974 ; J. Bosser 21657 ; P [ P00738467 ] ! • Bébour , Bélouve ; 21°06’40”S , 55°33’47”E ; alt. 1350-1450 m ; I.1975 ; L. Bernardi 15119 ; G ! • Bébour , Cassé de Takamaka ; 21°05’S , 55°38’E ; 1350- 1450m ; I.1975 ; L. Bernardi 15144 ; G ! • Salazie , on path Terre Plate-Cap des Anglais ; alt. 1450 m ; 7.IV.1976 ; T. Cadet 5322 ; REU [ REU006416 ] ! • Cirque de Mafate , sentier du bélier à Aurère ; 21°02’24”S , 55°26’29”E ; 14.V.1976 ; J. Bosser 22154 ; P [ P00738457 ] ! • Forêt de Bébour , 21°06’40”S , 55°33’47”E ; 3.III.1978 ; J. Bosser 22355 ; P [ P00738460 ] ! • Hauts de Sainte-Rose ; 21°09’S , 55°45’E ; 18.IV.1978 ; J. Bosser 22515 ; P [ P00738466 ] ! • Rivière des Marsouins ; 21°05’37”S , 55°36’43”E ; 29.IV.1980 ; F. Billiet 886 ; BR [ BR00000215511000 ] ! • Bébour , Coteau des Calumets ; 21°03’40”S , 55°25’40”E ; alt. 1500 m ; 6.IV.1984 ; T. Cadet 6576bis ; REU [ REU006417 ] ! • sine loc. ; X.1986 ; J. Dupont s.n. ; P ! spirit only • Sentier du Piton par la Plaine des Cafres ; 21°09’S , 55°34’E ; alt. 1700 m ; 24.IV.1988 ; J. Dupont s.n., P [ P00738356 ] ! • Plaine des Palmistes ; alt. 800 m ; II.2002 ; T. Pailler TP40 ; REU ! • Takamaka ; 21°05’S , 55°38’E ; alt. 1375 m ; 14.II.2003 ; Hoarau et al. 12 ; REU [ REU006407 ] ! • Tévelave ; 21°12’11”S , 55°21’51”E ; 1.III.2003 ; T. Pailler et al.TP161 ; REU [ REU006418 , REU006419 ] ! • L’Étang-Salé , Source Nouvelle Bras Sec , Chemin Canal ; 21°13’32”S , 55°22’09”E ; alt. 1250 m ; X.2003 ; V. Grondin et al. 838 ; CBNM ! • La Plaine des Palmistes , Piton Bébour path; 21°07’41”S , 55°33’43”E ; alt. 1310 m ; II.2004 ; J. Férard et al. 994 ; CBNM ! • La Plaine des Palmistes , La Tanguière ; alt. 750 m ; III.2004 ; V. Grondin et al. 1222 ; CBNM ! • Saint-Philippe , Piton Mare d’Arzule ; 21°19’00”S , 55°46’00”E ; alt. 700 m ; III.2004 ; C. Fontaine et al. 1127 ; CBNM ! • Plaine des Palmistes , Sentier vers Piton des Cabris ; 21°09’24”S , 55°39’03”E ; alt. 1200 m ; 21.V.2008 ; F. Martos FM328 and FM329 ; REU [ REU007870 , REU007872 ] ! • Plaine des Palmistes , Gros Piton Rond ; 21°07’41”S , 55°33’43”E ; alt. 1420 m ; 2008; F. Martos FM407 ; REU [ REU007869 ] ! • Cilaos , Mare à Joseph vers Kerveguen ; 21°07’27”S , 55°30’16”E ; alt. 1200 m ; 24.V.2009 ; F.Martos FM697 , FM698 and FM699; REU [ REU007696 , REU007695 , REU007692 ] !. NOTES In the protologue of Benthamia catatiana , Perrier states that this species is close to B. nigrescens (synonym of B. erinacea ), with the following differences: lip calceolate and attenuate but with apex obtuse, petals 5.5-6 mm longer than sepals 4.5 mm , anther shortly apiculate, and ovary short, less than 3 mm . However, examination of a rehydrated flower from the type specimen ( Catat 4338 ) shows no significant difference with flowers from specimens of B. erinacea , it is well within the variation range of B. erinacea . The cuspidate leaves of Benthamia cuspidata can be found also in Benthamia nigrescens subsp. humblotiana . Perrier gave specimen “ Boivin (sans n°) ” as type of Benthamia nigrescens subsp. borbonica . However there are in P two such specimens L . H . Boivin s.n. bearing his handwriting “ Benthamia nigrescens subsp. borbonica H.Perr. ”, without mention of which one is the type. We have chosen P 00334740 as the lectotype , with more individuals and more variations between them, P 00334741 being an isolectotype . Peristylus secundiflorus is a nomen nudum which was probably meant for specimens L . H . Boivin 1063 , according to handwritings on sheets P 00738489, P 00334751 and P 00334752, but it was never validly published. For Benthamia nigrescens subsp. humblotiana , Perrier gave no specific type but mentioned Humblot’s specimens ( sine loc. and s.n. ), from the north-east of Madagascar , and his own gathering H. Perrier 16110 from Mt. Tsaratanana, which has two sheets. We have chosen sheet P00094552 of H. Perrier 16110 as lectotype because it includes a dissected flower. For Benthamia catatiana H.Perrier , there are two sheets of the specimen cited in the protologue, L.D.M. Catat 4338 in P. We choose P00094478 as the lectotype , P00094479, a dissected flower, being an isolectotype . The watercolour 55 by Eudoxie de Cordemoy in MAU has been designated as a paratype by Hermans & Cribb (2021: 74) . FIG . 37. — Distribution maps of Benthamia erinacea (Cordemoy) Hermans & P.J.Cribb. See also: Schlechter (1930 : t.11 n°43), Perrier (1939: 25 , as Benthamia catatiana ), Perrier(1939: 26 , as Benthamia cuspidata ), Perrier (1939: 27) , Cadet (1989 : plate 23), Cribb & Hermans (2009: 43 , as Benthamia catatiana ), Cribb & Hermans (2009: 44) , Cribb&Hermans (2009:48 ,as Benthamia cuspidata ), Bernet (2010: 134) , Bosser & Lecoufle (2011: 178) , Pailler et al. (2013: 29) , Szelengowicz &Tamon (2013:242 , as Benthamia cuspidata ), Szelengowicz & Tamon (2013: 249) , Hervouet (2018: 170) , Pailler & Henze (2020: 75) , Hermans & Cribb (2023: 112) . DESCRIPTION An erect epiphyte or lithophytic herb 8-45 cm tall. Tubers several, clustered, fleshy and elongate, roots wiry. Stem 1.4- 4 mm in diameter, bearing two lanceolate, acuminate sterile bracts. Leaves cauline, 2-5, narrowly elliptic, oblong-elliptic or elliptic-lanceolate, acute or cuspidate, 4-15 × 1-4.5 cm , sheathing at the base, the upper two bract-like. Rachis erect, dense, often somewhat secund, narrowly cylindrical, 5 to 25 cm long. Floral bracts single-veined, lanceolate, very acute or acuminate, 4-20 mm × 1-4.5 mm , variable in size but generally longer than the ovary, especially the lower ones. Ovary 3-9.5 × 1-3 mm . Flowers c. 5 mm across, off-white or yellow to yellowish green, the petals and lip often more yellow; dorsal sepal single-veined, ligulate-lanceolate, acute, 2.8-6.3 × 0.7-1.3 mm ; lateral sepals single-veined, narrowly lanceolate, keeled, 2.9-6.5 × 1-2.1 mm ; petals triangular-ligulate, obtuse, 3-5.9 × 0.6-2 mm ; lip concave at the base, ligulate, entire or slightly trilobed, without callus, obtuse, 3-6.5 × 1-3.3 mm ; spur scrotiform, a little broader than long, 0.9-1.6 × 0.5-1 mm . Column c. 1-1.2 × 0.4-1 mm , anther subglobose, obtuse to shortly apiculate, auricles linear, subfalcate, longer than the anther; rostellum short, the midlobe obtuse, lateral lobes very small. Capsule less than 10 mm long.