Benthamia africana (Lindl.) Hermans

(Figs 4; 5; 6)

Kew Bulletin 69 (2)-9517: 10 (Hermans & Cribb 2014). — Spiranthes africana Lindl., Botanical Register 10: t. 823 (Lindley 1824). — Satyrium spirale Thouars, Histoire particulière des plantes orchidées recueillies sur les trois îles australes d’Afrique (Thouars 1822: t.9), nom. illeg.— Habenaria spiralis A.Rich., Mémoires de la Société d’Histoire Naturelle de Paris, tome IV (Richard 1828: 21), nom. illeg. — Herminium spirale (Thouars) Rchb.f., Bonplandia 3: 213 (1855). — Peristylus spiralis (Thouars) S.Moore, in Baker, Flora of Mauritius and the Seychelles (Moore 1877: 335), nom. illeg.

Habenaria minutiflora Ridl., Journal of the Linnean Society, Botany 21: 503 (Ridley 1885). — Benthamia minutiflora (Ridl.) Schltr ., Repertorium Specierum Novarum Regni Vegetabilis 33: 29 (Schlechter 1924). —

Type: Madagascar • Est Imerina, Andrangoloaka; 19°02’S, 47°55’E; XI.1880; J.M. Hildebrandt 3729; holotype: BM!; isotypes: P [P00094580, P00094581]!, K [K000415552]!, AMES [AMES00099954]!, M [M0103630]!, G [G00190645, G00300208]!, HBG [HBG500730]!, WU!, Z [Z000064380]!, BM [BM000034593]!, GOET [GOET008457]!, W!, FR [FR0036981]!.

Benthamia procera Schltr., Beihefte zum Botanischen Centralblatt 34 (2): 301 (Schlechter 1916). — Type: Madagascar • Alaotra-Mangoro; alt. 800 m; 11.X.1912; H. Perrier 11380; holotype: P [P00738317]!; isotype: P [P00094573]!, syn. nov.

TYPE MATERIAL. — Sine loc.; L.M.A. du Petit-Thouars 5 (or 9 in BM); holotype: P [P00738391]!; isotype: BM!

ETYMOLOGY. — Of Africa.

PHENOLOGY. — Mostly from April to November.

DISTRIBUTION AND ECOLOGY. — Madagascar, Mauritius, La Réunion, possibly Comoros. From sea level to 1500 m, in lowland and medium altitude moist evergreen forest. Widespread in Madagascar in the eastern part of the country. It grows on dry banks as well as in damp places (Fig. 7).

CONSERVATION. — It is a very widespread and locally common species, of Least Concern so far (LC) in Madagascar and La Réunion. It was considered Endangered on Mauritius in 1994 (Strahm: app. 14) and should be looked for in the Comoros.

ADDITIONAL SPECIMENS EXAMINED. — Comoros • Grande Comore; 14.VI.1899; Pobeguin s.n.; P [P00209897]!. With doubts, as already noticed by Perrier de la Bâthie, the specimen does not allow identification at species level. The flowers are in bud and are indeed reminiscent of Benthamia africana .

Madagascar • Nord Betsileo; I.1881; Hildebrandt 3975a; W [R14939] !• Analamanga, rochers de la Mandraka; 2.X.1912; H. Poisson 602; P [P00094607] ! • Atsinanana, Andovoranto, district d’Anivorano; 18°53’S, 48°34’E; alt. 380 m; 14.X.1912; R. Viguier et al. 687 and 687a; P [P00334796, P00094608] ! • Atsinanana, Andovoranto, district d’Anivorano; 18°53’S, 48°34’E; 15.X.1912; alt. 800 m; R. Viguier et al. 727; P [P00334727] ! • Atsinanana, Ranolalina; 18°00’54”S, 49°00’09”E; 20.IX.1920; R. Decary 82; P [P00094586] ! • Alaotra-Mangoro, district d’Ambatondrazaka; 17.VII.1921; R. Decary 295; P [P00094590] ! • Analamanga, Betatao; 18°12’41”S, 47°52’41”E; alt. 1300 m; XI.1922; H. Perrier 14962; P [P00094605] ! • Atsimoatsinana, Vondrozo; 22°49’S, 47°20’E; 4.IX.1926; R. Decary 5089; P [P00738390] ! • Atsimo-atsinanana, Vondrozo; 5.IX.1926; R. Decary 5191; P [P00094593] ! • Analamanga, Mandraka, est de Tananarive; 18°55’31”S, 47°56’20”E; XI.1927; H. Perrier 18006; P [P00094538] ! • Alaotra-Mangoro, entre Moramanga et Nosibe; alt. 400-700 m, X.1927; H. Perrier 18027; P [P00094572] ! • Centre, rocailles humides du Tampoketsa du haut Bemarivo; alt. 1000 m; 1928; H. Perrier 1878; P [P00094602], P [P00094603], P [P00094604] ! • Alaotra-Mangoro, Analamazaotra; 18°56’05”S, 48°24’43”E; X.1936; P. Boiteau 2175; P [P00692295]!, TAN ! • Alaotra-Mangoro, district d’Ambatondrazaka, Sasamanga Onibe; 800 m; XI.1938; G. Cours 1039; P [P00094589] ! • Alaotra-Mangoro, sud de Moramanga; 1.IX.1942; R. Decary 18163; P [P00094594] ! • Alaotra-Mangoro, sud de Moramanga; 2.IX.1942; R. Decary 18172; P [P00094595]!, TAN!, K [K000718260] ! • Alaotra-Mangoro, district de Moramanga, Lakato; 19°11’30”S, 48°26’00”E; 2.IX.1942; R. Decary 18240; P [P00094596]!, K [K000718261]!, TAN ! • Alaotra-Mangoro, Anosibe, sud de Moramanga; 19°25’29”S, 48°12’37”E; 6.IX.1942; R. Decary 18385; P [P00094597, P02115324]!, TAN, G, K [K000718263] ! • Alaotra-Mangoro, Périnet; 18°56’S, 48°25’E; 4.IX.1951; R. Benoist 1114; P [P00692221, P00692222] ! • Haute Matsiatra, Anjamana Ambalarondra; 18°28’08”S, 49°00’02”E; 1954; Vigreux 15341; P [P00692311] ! • Atsinanana, Ambodiriana Canton, Mariandrano; 18°27’06”S, 48°58’37”E; X.1955; J. de la Croix Rakoto 7694RN; G!, TAN!, K [K000718258]!, P [P01805121, P01805145]!, WAG [WAG.1924443] !) • Alaotra-Mangoro, Nombe, Moramanga; 2.VIII.1956; P. Soga 8012; P [P00692317] ! • Analanjirofo, Sahatavy, district de Vavatenina; 17°28’22”S, 49°12’02”E; 17.IX.1960; Laibosaka 11346; P [P00692319] ! • Alaotra-Mangoro, route de Moramanga, Anosibe; VIII.1962; J. Bosser 16179; P [P00692232, P00692233] ! • Mountain forest; alt. 600 m; VII.1963; W. Rauh R10032; HEID [HEID789485, HEID789486, spirit 250342] ! • Alaotra-Mangoro, route Moramanga-Anosibe; VIII.1963; J. Bosser 18534; P [P00692245] ! • Atsinanana, Tamatave, near road; IX.1969; J. Stewart 1122; TAN ! • Alaotra-Mangoro, forêt d’Analamazaotra; 18°56’05”S, 48°24’43”E; 4.XI.1970; Keraudren-Aymonin 25358; P [P00326136] ! • Toamasina, in a marsh 4 km south of Mahavelona; 17°43’S, 49°30’E; alt. 0 m; 16.X.1986; B. Pettersson et al. 11; UPS!, K ! • Alaotra-Mangoro, Andasibe, près d’Antanambe; 1.XII.1989; P. Morat 8523; P [P00094530] ! • Vatovavy-Fitovinany, Vohipeno; 22°21’13”S, 47°50’25”E; X.1990; Beaujard 368; P [P00323116, P00323117] ! • Atsinanana, east coast, Betampona Réserve Naturelle Intégrale, 40 km NW of Toamasina; 17°51’S, 49°12’E-17°55’S, 49°15’E; alt. 275-650 m; IX.1993; B. Lewis et al. 612; P [P00755163]!, K!, MO • Antsiranana, district d’Andapa, RNI de Marojejy, à 10 km au nord-ouest du village de Manantenina; 14°26’S, 49°46’E; alt. 750 m; 19.X.1996; M. Rakotondrainibe 3416; P [P01778631] ! • Alaotra-Mangoro, Andasibe, forest around Vakona Lodge; 18°53’57”S, 48°26’38”E; alt. 1101 m; I.2000; J. Hermans 6565; K ! • Haute Matsiatra, Ranomafana by the Namorona falls; V.2001; 21°14.70’S, 47°23.80’E; alt. 1135 m; J. Hermans 5590; K ! • Alaotra-Mangoro, Moramanga, Lakato; 19°04’29”S, 48°21’59”E; alt. 1002 m; XI.2002; G. Fischer et al. FS282; WU [WU084252] ! • Alaotra-Mangoro, Ambatovy; 18°53’S, 48°34’E; 3.VIII.2005; C. Rakotovao 1930; P [P02091797]!, K!, MO, TAN ! • Alaotra-Mangoro, Ambatovy; 18°50’51”S, 48°18’31”E; 22.VIII.2005; H. Razanantsoa et al. 359; P [P02091796]!, K [K000718736]!, MO, TAN ! • Alaotra-Mangoro, Andasibe, forest around Vakona Lodge; 18°53’38”S, 48°26’36”E; alt. 1060 m; I.2007; J. Hermans 6760; K ! • Alaotra-Mangoro, Andasibe, forest around Vakona Lodge; 30.IX.2016; J. Hermans 8025; K !.

Mauritius • sine loc.; L. H. Boivin s.n.; P [P00738537]! (plant on the right only) • Au-dessous du plateau du piton du Pouce; 20°11’44”S, 57°31’20”E; X.1849; L.H. Boivin s.n.; P [P00738420] ! • Forêt de Curepipe; 20°19’40”S, 57°32’14”E; 20.IX.1851; L.H. Boivin s.n.; P [P00738392]!; [very linear leaves, possibly hybrid with Benthamia spiralis A.Rich. ] • VIII.1862; P.B. Ayres s.n.; P [P00738413]! (specimens on the left and on the right only), K ! • In high and damp places; 1864; L. Bouton s.n.; K ! • Grand Bassin, above Black Gorge; 20°25’05”S, 57°29’30”E; IV.1953; Cooper s.n.; K![spirit collection 29789] • Mare Longue; 20°22’36”S, 57°27’22”E; alt. 300 m; VII.1961; H. St John 26479; G ! • Cascade; 20°22’22”S, 57°37’37”E; 9.IV.1974; J. Bosser 21830; P [P00738521] ! • Pétrin; 20°24’31”S, 57°28’19”E; 9.IV.1974; J. Bosser 21827; P [P00738519, spirit] ! • Plaine Champagne; 20°25’35”S, 57°23’30”E; 3.VI.1976; J. Bosser 22262; P [P00738375] ! • Les Mares, track from Plaine Paule to Mt. Cocotte; 20°26’09”S, 57°28’11”E; alt. 650 m; 13.V.1979; D. Lorence 2627; P [P00738374]!, K !. France, La Réunion • J.B. Potier s.n.; P [P00738526] ! • L.H. Boivin s.n.; P [P00738394] ! • L.H. Boivin s.n.; P [P00738414] ! • L.H. Boivin s.n.; P [P02115323] ! • L.H. Boivin s.n.; P [P00738530] ! • Les Hauts du Breton; L.H. Boivin 1064; P [P00738531] ! • L.H. Boivin s.n.; P [P00738393] ! • sine loc.; 1846-1852; L.H. Boivin s.n.; G! Herb. Delessert • 1848; L.H. Boivin 1064; P [P00738423, P00738525] ! • 1862; Vieillard et al. s.n.; P [P00738495] ! • Montagne du Brûlé; 1862; Anonymous s.n.; P [P00738388] ! • 8 km SSW Saint-Denis; alt. 1090 m; VII.1967; Gill 728; K ! • Saint-Philippe, Mare Longue; 21°21’16”S, 55°44’22”E; 25.IX.1968; T. Cadet 1625; REU [REU017476] ! • Salazie plain; 21°06’S, 55°32’E; alt. 1500 m; 7.III.1971; T. Cadet 3135; REU [REU017472] ! • Takamaka, Saint-Benoit; 21°05’S, 55°38’E; alt. 600 m; 21.IV.1971; T. Cadet 3236; P [P00738416, spirit]!, REU [REU017473] ! • Grand Brûlé; 21°14’50”S, 55°47’50”E; alt. 120 m; VIII.1972; T. Cadet 3753; REU [REU017475] ! • Takamaka; 21°05’S, 55°38’E; alt. 600 m; VII.1972; F. Friedmann 1695; P [P00738419] ! • La Montagne; 20°54’S, 55°24’E; alt. 650 m; 12.VII.1972; T. Cadet 3657; REU [REU006431, REU017474] ! • Grand Brûlé; 21°14’50”S, 55°47’50”E; 30.VIII.1972; T. Cadet 3756; P [P00738415], REU [REU006432] ! • Plate Découverte, Hauts de Saint-André; 20°59’33”S, 55°34’24”E; alt. 750 m; 12.VI.1973; T. Cadet 4268; REU [REU006433], P [P00738412] ! • Basse Vallée; 27.V.1974; J.Bosser 22068; P [P00738534] ! • Les Trois Bras, Hauts de Sainte Anne; 21°09’08”S, 55°43’43”E; 30.V.1974; J.Bosser 22082; P, not found, anther drawn by Szlachetko in (Szlachetko & Rutkowski 2000: 141) under the name Benthamia spiralis, tentatively put here but not seen • Les Trois Bras, Hauts de Sainte Anne; 21°09’08”S, 55°43’43”E; 30.V.1974; J.Bosser 22083; P [P00738309, P00738310] ! • Les Trois Bras, Hauts de Sainte Anne; 21°09’08”S, 55°43’43”E; 30.V.1974; J. Bosser 22084; P [P00738325, P00738326] ! • Hauts de Sainte Anne, left bank of the Rivière de l’est; 21°07’S, 55°43’E; alt. 300-1100 m; 30.V.1974; T. Cadet 4710; REU [REU006434, REU006435] ! • Bord de la RN3, au dessous de la Plaine des Palmistes; 21°06’58”S, 55°38’56”E; alt. 800 m; 4.VI.1974; T. Cadet 4716; P [P00738395]!, REU [REU006436, REU006437 ! • Sentier de l’îlet à Guillaume; 20°57’S, 55°24’E; 11.V.1976; J. Bosser 22292; P [P00738318, P00738319] ! • Basse Vallée; 21°21’16”S, 55°44’22”E; 24.V.1976; J.Bosser 22202; P [P00738320, P00738321] ! • Basse Vallée; 21°21’16”S, 55°44’22”E; 24.V.1976; J. Bosser 22268; P [P00738497] ! • Plaine des Palmistes; 21°06’58”S, 55°38’56”E; 8.VI.1976; J. Bosser 22223; P [P00738323, P00738324 ! • Plaine des Palmistes; 8.VI.1976; J. Bosser 22223bis; P [P00738368, spirit] ! • Hauts de Sainte-Marie; 20°58’S, 55°31’E; alt. 1250 m; 18.VI.1976; T. Cadet 5519; REU [REU006438] ! • Hauts du Tremblet; 21°17’43”S, 55°45’34”E; 20.IV.1978; J.Bosser 22531; P [P00738314, P00738315] ! • Basse-Vallée; 21°21’16”S, 55°44’22”E; alt. 500 m; 13.VII.1979; J. Stewart 2127; K ! • Sentier de l’îlet Patience; 21°06’S, 55°37’E; alt. 1100 m; 22.IX.2002; T. Pailler TP48 & TP48bis; REU [REU007939, REU006427] ! • Forêt de Bélouve; alt. 1560 m; 13.V.2003; M. Pignal & C. Lartigau-Roussin 2086; P [P00340567] ! • La Plaine des Palmistes, La Tanguière; alt. 800 m; IV.2004; V. Grondin et al. 1179; CBNM ! • La Plaine des Palmistes; alt. 850 m; VI.2004; V. Grondin et al. 1187; CBNM ! • Sainte-Rose, Piton Crac; 21°13’38”S, 55°45’20”E; alt. 1000 m; V.2008; Fontaine et al. 3374; CBNM !. Mascareignes • Sine loc.; McGregor s.n.; BM [BM 000034596]! • Moore s.n.; BM [BM000034598]! • L.M.A. du Petit-Thouars 1011; BM [BM000034597] !.

NOTES

There has been much confusion over the identity of the species. In La Réunion, several other species have long been included in the species concept, in Madagascar it also encompassed at least three other species, and in Mauritius it is thought to be sympatric with Benthamia spiralis A.Rich. (see the notes under that entry). Even with these clarifications Benthamia africana certainly remains a species complex, and further field observations and DNA analysis may show differences that remained unnoticed in dried specimens. We have kept the holotype as commonly accepted, the only Thouars specimen known of Benthamia africana . However, the illustration given by Thouars (1822, plate 9) shows leaves adpressed on the ground, which is the case for some plants but not all.

Benthamia procera is included here as a synonym, only the lip midlobe length seems to be the distinctive feature, but the variations of the lip midlobe length on the type itself, from 1 mm to 1.6 mm, which is in the variation range of Benthamia africana, show that it is preferable to consider the type of Benthamia procera as Benthamia africana with an unusually long lip.

See also: Schlechter (1930: t.12 n°47), Perrier (1939: 31, as Benthamia spiralis A.Rich.), Perrier (1939: 32), Cribb & Hermans (2009: 47, as Benthamia spiralis (Thouars) A.Rich), Bernet (2010: 136, as Benthamia spiralis (Thouars) A.Rich.), Szelengowicz & Tamon (2013: 253, as Benthamia nigrescens Schltr.), Szelengowicz & Tamon (2013: 255, as Benthamia verecunda Schltr.), Hervouet (2018: 157), Pailler & Henze (2020: 72), Hermans & Cribb (2023: 101, except figure 58 p. 105 which is Benthamia mascula Hervouet & Descourv., sp. nov.).

DESCRIPTION

Slender terrestrial herb, 20-62 (108) cm tall.Tubers 1-2, ovoid, 20-35 × 4-5 mm, roots fleshy and woolly, up to 65 mm long. Stem cylindrical, 1-5 mm in diameter, with 2 to 3 sheaths. Leaves 3-5, basal or cauline in the lower part of inflorescence, narrowly ligulate or linear-lanceolate, acute, 6-20 × 0.5-2 cm, shortly petiolate, higher on the stem grading into small sheaths. Rachis many-flowered, 6-38 cm long, cylindrical. Floral bracts linear, acuminate 4-12 mm × 0.8-3 mm. Ovary suberect, 4.5-9 × 1-1.9 mm. Flowers very small, 2-5 mm in diameter, secund, spreading, with greenish sepals, green or yellow petals and lip; dorsal sepal cucullate, elliptic, obtuse, 1.8-4.2 × 0.6-1.1 mm; lateral sepals obliquely oblong-ovate, obtuse, 1.8-3.8 × 0.5-1.1 mm, concave at the apex; petals porrect, recurved in apical half, lanceolate, subacute, 1.8-3.8 × 0.5- 1.2 mm; lip trilobed, 1.8-3.1 × 1-2 mm, recurved, lateral lobes 0.2-0.4 mm, obtuse, midlobe fleshy, much longer than the lateral lobes, 0.8-1.3(1.6) mm; spur scrotiform to subrectangular, 0.5-1 to 0.4-0.9 mm across. Column 0.6-1.1 × 0.5-1 mm, auricles short, anther retuse or obtusely apiculate; rostellum trilobed, the three lobes subequal, 0.1-0.2 mm long, obtuse, hardly visible.