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]!.

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).

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.