Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae)

2025  47  11 adansonia Adansonia est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Adansonia is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-8571/ ISSN (électronique / electronic) : 1639-4798 Directeur De la publication / Publication director: Gilles Bloch Président du Muséum national d’Histoire naturelle réDacteur en chef / editor-in-chief : Thierry Deroin réDacteurs / editors : Porter P. Lowry II ; Zachary S. Rogers ; Mathieu Gardère assistant De réDaction / assistant editor : Emmanuel Côtez ([email protected]) Mise en page / Page layout : Emmanuel Côtez coMité scientifique / scientific board : F. Blasco (CNRS, Toulouse) M. W. Callmander (Conservatoire et Jardin botaniques de Genève) J. A. Doyle (University of California, Davis) P. K. Endress (Institute of Systematic Botany, Zürich) P. Feldmann (Cirad, Montpellier) L. Gautier (Conservatoire et Jardin botaniques de Genève) F. Ghahremaninejad (Kharazmi University, Téhéran) K. Iwatsuki (Museum of Nature and Human Activities, Hyogo) A. A. Khapugin (Tyumen State University, Russia) J.-Y. Lesouef (Conservatoire botanique de Brest) P. Morat (Muséum national d’Histoire naturelle, Paris) J. Munzinger (Institut de Recherche pour le Développement, Montpellier) S. E. Rakotoarisoa (Millenium Seed Bank, Royal Botanic Gardens Kew, Madagascar Conservation Centre, Antananarivo) P. H. Raven (Missouri Botanical Garden, St. Louis) G. Tohmé (Conseil national de la Recherche scientifique Liban, Beyrouth) J. G. West (Australian National Herbarium, Canberra) J. R. Wood (Oxford) couverture / cover : Réalisée à partir des Figures de l’article/Made from the Figures of the article. Adansonia est indexé dans / Adansonia is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Adansonia est distribué en version électronique par / Adansonia is distributed electronically by: – BioOne® (http://www.bioone.org) 171 ADANSONIA, sér. 3 • 2025 • 47 (11) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com Jean-Michel HERVOUET 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) [email protected] Pascal DESCOURVIÈRES Fédération France Orchidées (FFO), 27 rue Auguste Blanqui, 94400 Vitry-sur-Seine (France) [email protected] Johan HERMANS Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE (United Kingdom) and Core Facility, Botanical Garden, University of Vienna, Rennweg 14, 1030, Vienna (Austria) [email protected] Ludivine LONGOU Atelier d'iconographie scientifique, UAR 2047 DoHNÉE, MNHN/CNRS, Muséum national d’histoire naturelle, 57 rue Cuvier, 75005 Paris (France) [email protected] Submitted on 3 September 2024 | accepted on 29 December 2024 | published on 21 July 2025 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) Hervouet J.-M., Descourvières P., Hermans J. & Longou L. 2025. — Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae). Adansonia, sér. 3, 47 (11): 171-249. https://doi.org/10.5252/adansonia2025v47a11. http://adansonia.com/47/11 ABSTRACT A revision of the genus Benthamia A.Rich is presented, based on herbarium material, literature and field observations. A taxonomic history of the genus is given. Thirty-one species are recognized and described. Seven new species are described: Benthamia litoralis Hervouet& Descourv., sp.nov., Benthamia decaryana Descourv.& Hervouet, sp.nov., Benthamia lakatoensis Descourv.& Hervouet, sp.nov., Benthamia mascula Hervouet& Descourv., sp.nov., Benthamia trifida Hermans& Hervouet, sp.nov., Benthamia corona Hermans& Hervouet, sp.nov., Benthamia vulcanorum Hervouet& Descourv., sp.nov. A new form is established: Benthamiaerinacea (Cordemoy) Hermans& P.J.Cribb f. reticulata Hervouet& Descourv., f.nov. One subspecies is raised to species level: Benthamia herminioides subsp. arcuata H.Perrier becomes Benthamia arcuata (H.Perrier) Descourv.& Hervouet, comb. nov., stat. nov., seven taxa are reduced to synonymy: Benthamia herminioides subsp. angustifolia H.Perrier is a synonym of Benthamia perfecunda H.Perrier, Benthamia verecunda Schltr. and Benthamia nigrovaginata H.Perrier are synonyms of Benthamia elata Schltr., Benthamia exilis var. tenuissima Schltr. is a synonym of Benthamia exilis Schltr., Benthamia herminioides subsp. intermedia is a synonym of Benthamia herminioides Schltr., Benthamia nivea subsp. parviflora H.Perrier is a synonym of Benthamia nivea Schltr., Benthamia perularioides Schltr. is a synonym of Benthamia rostrata Schltr. Two species remain undescribed because they are known only from photographs, four specimens are discussed for their specific interest and are potentially new species but this remains to be confirmed when more data becomes available. Twelve taxa are lectotypified: B.calceolata H.Perrier, B.elata Schltr., B.exilis var. tenuissima Schltr., B.nigrovaginata KEY WORDS Orchidaceae, Madagascar, Mascarenes, La Réunion, Mauritius, Comoros, Seychelles, conservation, new synonyms, new status, lectotypifications, epitypifications, new forma, new species. 172 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. INTRODUCTION The need for a revision of The genus Benthamia In 2024 the genus Benthamia comprised 32 species, five subspecies and one variety (Plants Of the World Online (POWO): https://powo.science.kew.org, on November the 4th) from the West Indian Ocean. We give in the next section, for reference, a detailed history of the genus, from 1828 on, leading to the current state of the art at the beginning of this study. It will be seen that species currently recognised as Benthamia have been previously classified in Cynorkis, Habenaria, Hemiperis, Herminium, Holothrix, Peristylus, Platanthera and Rolfeella. This complicated history and a morphological examination showing a large diversity suggest that Benthamia as currently circumscribed is not monophyletic. A majority of species have a scrotiform or very small spur and show an affinity with the type of the genus Benthamia latisatis (Thouars) Bytebier, they are mostly terrestrial but not exclusively. Others, mostly epiphytes, have tubular flowers. One species, Benthamia calceolata, stands apart with a calceolate lip. Finally, a number of species have cylindrical spurs and are reminiscent of Cynorkis or Habenaria. Ngugi etal. (2020) published the first phylogenetic study showing that Benthamia is polyphyletic. The study included five species of Benthamia. Two species with long spurs, namely Benthamia monophylla (under the name B.bathieana) and B.rostrata (under the name B.perularioides), appeared to be embedded in Cynorkis. Three species with short spurs, namely Benthamia africana (under the names B.spiralis and B.sp.), B.erinacea (under the name B.nigrescens) and B.flavida, appeared to be close to the genus Tylostigma, and closer to Habenaria than to Cynorkis. We will come back to this study and the possible MOTS CLÉS Orchidaceae, Madagascar, Mascareignes, Île de La Réunion, Île Maurice, Comores, Seychelles, conservation, synonymes nouveaux, statut nouveau, lectotypifications, épitypifications, formes nouvelles, espèces nouvelles. H.Perrier, B.verecunda Schltr., B.nigrescens subsp. borbonica H.Perrier, B.nigrescens subsp. humblotiana, B.catatiana, B.humbertii H.Perrier, B.melanopoda Schltr., B.perularioides Schltr., B.praecox Schltr. Three species, Benthamia dauphinensis (Rolfe) Schltr., Benthamia macra Schltr. and Benthamia misera (Ridl.) Schltr., are considered ambiguous or insufficiently known. Benthamia longecalceata H.Perrier, is excluded, being a synonym of Cynorkis falcata (Frapp.) Schltr. Benthamia micrantha (Frapp. ex Cordem.) Hermans& Hervouet (nom. illeg.) is excluded, being Cynorkis micrantha (Frapp. ex Cordem.) Schltr. The relationship with other genera is investigated and several groups of similar species within the genus Benthamia are proposed as a starting point for a subgeneric classification. RÉSUMÉ Révision du genre Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae). Cet article présente une révision du genre Benthamia A.Rich, fondée sur des observations de terrain et des spécimens d’herbier. L’histoire du genre est présentée. 31 espèces sont reconnues. Sept nouvelles espèces sont décrites : Benthamia litoralis Hervouet& Descourv., sp.nov., Benthamia decaryana Descourv.& Hervouet, sp.nov., Benthamia lakatoensis Descourv.& Hervouet, sp.nov., Benthamia mascula Hervouet& Descourv., sp.nov., Benthamia trifida Hermans& Hervouet, sp.nov., Benthamia corona Hermans& Hervouet, sp.nov., Benthamia vulcanorum Hervouet& Descourv., sp.nov. Une nouvelle forme est décrite : Benthamiaerinacea (Cordemoy) Hermans& P.J.Cribb f. reticulata Hervouet& Descourv., f. nov. Une sous-espèce est élevée au rang d’espèce : Benthamia herminioides subsp. arcuata H.Perrier devient Benthamia arcuata (H.Perrier) Descourv.& Hervouet, comb. nov., stat. nov., sept taxons sont placés en synonymes d’autres espèces : Benthamia herminioides subsp. angustifolia H.Perrier est synonyme de Benthamia perfecunda H.Perrier, Benthamia verecunda Schltr. et Benthamia nigrovaginata H.Perrier comme synonymes de Benthamia elata Schltr., Benthamia exilis var. tenuissima Schltr. comme synonyme de Benthamia exilis Schltr., Benthamia herminioides subsp. intermedia comme synonyme de Benthamia herminioides Schltr., Benthamia nivea subsp. parviflora H.Perrier comme synonyme de Benthamia nivea Schltr., Benthamia perularioides Schltr. comme synonyme de Benthamia rostrata Schltr. Deux espèces dont on ne possède que des photos sont mentionnées mais non décrites. L’attention est attirée sur quatre spécimens intéressants et qui pourraient se révéler être des nouvelles espèces, insuffisamment documentées pour l’instant. Douze taxons sont lectotypifiés: B.calceolata H.Perrier, B.elata Schltr., B.exilis var. tenuissima, B.nigrovaginata H.Perrier, B.verecunda Schltr., B.nigrescens subsp. borbonica H.Perrier, B.nigrescens subsp. humblotiana, B.catatiana, B.humbertii H.Perrier, B.melanopoda Schltr., B.perularioides Schltr., B.praecox Schltr. Trois espèces sont considérées comme ambiguës ou insuffisamment connues : Benthamia dauphinensis (Rolfe) Schltr., Benthamia macra Schltr. et Benthamia misera (Ridl.) Schltr. Benthamia longecalceata H.Perrier, est exclue car synonyme de Cynorkis falcata (Frapp.) Schltr. Benthamia micrantha (Frapp. ex Cordem.) Hermans& Hervouet (nom. illeg.) est exclue, étant Cynorkis micrantha (Frapp. ex Cordem.) Schltr. Enfin les relations avec d’autres genres sont envisagées et différents groupes d’espèces semblables au sein du genre Benthamia sont donnés comme base d’une future subdivision du genre. 173 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) groups of species within the genus Benthamia in the discussion, once nomenclatural issues have been solved. Other molecular studies, with more species, will help characterise the genus Benthamia, but a key issue appears to be the correct identification of specimens. As is the case with many difficult genera, a large number of Benthamia specimens remain unidentified in herbaria, many are incorrectly identified, others are misfiled under other genera, and a number are labelled Benthamia but are not. A complete revision is therefore necessary. Molecular work can help, but ultimately taxa will need to be circumscribed on the basis of morphology, it thus appeared necessary to identify morphologically the existing specimens, thereby avoiding compromising further DNA analysis by misidentifications and this is the goal of the present study. The remaining taxonomical problems can then be addressed by later studies. hisTory of The genus Benthamia The genus was created in 1828 by Achille Richard (Richard 1828: 37) for two species, Benthamia latifolia A.Rich., a new combination for Satyrium latifolium Thouars, and Benthamia spiralis A.Rich. Richard placed his new genus in the tribe Ophrydeae. Lindley (1835: 297) considered the genus Benthamia to be a synonym of the genus Peristylus. In Ridley’s work on orchids of Madagascar (Ridley 1885), Benthamia was merged into Habenaria, with, in a section Peristylus, Habenaria spiralis, Habenaria minutiflora Ridl., which was later reduced into the synonymy of the former, and Habenaria misera Ridl. The year after (Ridley 1886) he described Holothrix glaberrima, which Perrier moved to Benthamia (Perrier 1934). In Kraenzlin’s major work (Kraenzlin 1899) Benthamia was not included, members of the genus were scattered e.g. in Herminium and Platanthera. Schlechter (1915) published a critical evaluation of orchids in Madagascar, Mascarenes, the Comoros and Seychelles, in which all Benthamia species appeared as Habenaria or Cynorkis. Schlechter (1916) after studying the specimens collected in Madagascar by Perrier de la Bâthie, described Benthamia nigrescens, B.procera, B.flavida, B.perrieri (which, in 1925, he considered a synonym of B.flavida), and B.praecox. He also mentioned Benthamia latifolia A.Rich. from Mt. Ibity in Madagascar, which he later described as the new species Benthamia bathieana Schltr. In 1918 he added Benthamia elata (Schlechter 1918). In the year before his death, Schlechter (1924), created Benthamia bathieana, B.dauphinensis (based on Habenaria dauphinensis Rolfe), B.exilis and its variety tenuissima, B.herminioides, B.macra, B.melanopoda, B.misera (based on Habenaria misera Ridl.), B.monophylla, B.nivea, B.perularioides, B.rostrata (a new name for his Platanthera madagascariensis published in 1916, which was a nom. illeg.) and B.verecunda. He moved three species, Holothrix madagascariensis Rolfe, Habenaria minutiflora Ridl. and Habenaria misera Ridl, to the genus Benthamia. After Schlechter’s death, Henri Perrier de la Bâthie (Perrier 1934) revised and published a monography of the genus, with a key. He included twenty-five species and created Benthamia calceolata, B.catatiana, B.cuspidata, B.nigrovaginata and B.humbertii. He also added four sub-species to Benthamia nigrescens: subsp. borbonica, subsp. decaryana, subsp. humblotiana, subsp. secundiflora, three sub-species to Benthamia herminioides: subsp. angustifolia, subsp. arcuata and subsp. intermedia, and a sub-species to Benthamia nivea: subsp. parviflora. Subsequently, he placed Holothrix glaberrima Ridl. and also Habenaria cinnabarina Rolfe in the genus Benthamia, with Benthamia flavida Schltr. as a new synonym, but this was contradicted in 2014 by J. Hermans& P.J. Cribb (Hermans& Cribb 2014) who placed Habenaria cinnabarina in Cynorkis. Perrier recognised Benthamia minutiflora (Ridl.) Schltr. as a synonym of the former Satyrium spirale published by Thouars in 1822. Based on plants collected by Henri Humbert and others in Madagascar, Perrier (Perrier 1951) created four more species: Benthamia leandriana (later renamed Cynorkis tryphioides var. leandriana by Jean Bosser in1969, later becoming Cynorkis leandriana (Perrier) Hervouet in 2022), Benthamia longecalceata (here considered to be Cynorkis falcata (Frapp.) Schltr. from La Réunion), Benthamia majoriflora and B.perfecunda. Dressler (1990: 191) placed the genus Benthamia in the subtribe Habenariinae, together with Peristylus and Tylostigma. In 1998, Garay& Romero (1998) considered Satyrium latifolium Thouars to be illegitimate, being a name already used by Linnaeus, and thus created Benthamia chlorantha (Spreng.) Garay& Romero, based on Habenaria chlorantha Spreng. (1826). In the year 2000, Szlachetko& Rutkowski (2000) transferred Oligophyton drummondii Linder& Williamson to Benthamia and created Benthamia drummondii (Linder& Williamson) Szlach.& Rutk., although this is generally not accepted. Cribb, in Genera Orchidacearum vol. 2 (Pridgeon etal. 2001: 261), recognises about 25 species of Benthamia, endemic to Madagascar and the Mascarenes. In 2014, J. Hermans (in Hermans& Cribb 2014) argued that Thouars’s name Satyrium spirale is illegitimate as it had been used already by Swartz in 1788, for a plant from Jamaica. Lindley (1824: t. 823) consequently described Spiranthes africana based upon Thouars’s Satyrium spirale and his epithet now prevails. In 2014, Hervouet etal. (2014) published two new species, B.boiteaui and B.bosseri, found on the Ambondrombe mountain in Madagascar, the type material for both having been collected by Pierre Boiteau in April 1941. Chase etal. (2015), in their updated classification of Orchidaceae, lumped Orchidinae and Habenariinae in one subtribe. However, Ngugi etal. (2020), including 368 samples representing 278 species of Orchideae, showed that a subdivision in three subtribes, Bartholininae, Orchidinae and Habenariinae, is perfectly justified, and that all Benthamia (and all Cynorkis but two) belong to Habenariinae. In 2021, J. Hermans and P.J. Cribb (Hermans& Cribb 2021), in preparation for the publication of the «Flore des Mascareignes» volume on Orchidaceae, created Benthamia erinacea (Cordemoy) Hermans& P.J.Cribb, based on Habenaria erinacea Cordemoy, preceding Benthamia nigrescens Schltr. by 21 years, the latter hence a new synonym. In addition, all the former subspecies of Benthamia nigrescens were merged into this taxon. Habenaria spiraloides Cordem., described in 1895, was also recognised as a Benthamia under the name Benthamia spiraloides (Cordem.) Hermans& P.J.Cribb. Finally, in 2025 Bytebier coined the new name Benthamia latisatis (Thouars) Bytebier for the type of the genus, based on the Thoursian name Satorkis latisatis, replacing Benthamia chlorantha (Spreng.) Garay& Romero. 174 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. Key To The genus Benthamia a.rich. 1. Spur elongated, cylindrical .......................................................................................................................... 2 — Spur short, scrotiform ................................................................................................................................. 7 2. Leaves reduced to sheaths or not developed at flowering .............................................................................. 3 — Leaves developed at flowering ...................................................................................................................... 4 3. Lip clearly trilobed ................................................................................................. Benthamia praecox Schltr. — Lip obscurely trilobed ................................................................................................ Benthamia exilis Schltr. 4. Leaf single ................................................................................................................................................... 5 — More than one leaf ...................................................................................................................................... 6 5. Lip trilobed ....................................................................................... Benthamia glaberrima (Ridl.) H.Perrier — Lip entire ......................................................................................................... Benthamia monophylla Schltr. 6. Leaves 2, rarely 3 .................................................................................................... Benthamia rostrata Schltr. — Leaves 3 to 6 ...................................................................................................... Benthamia bathieana Schltr. 7. Spur scrotiform or dorso-ventrally flattened ................................................................................................ 8 — Spur a simple hollow at the base of the lip ........................................................ Benthamia boiteaui Hervouet 8. Plants epiphytic ........................................................................................................................................... 9 — Plants terrestrial ......................................................................................................................................... 14 9. Plants more than 45cm tall ......................................................................... Benthamia majoriflora H.Perrier — Plant less than 45cm tall ........................................................................................................................... 10 10. Lip entire or obscurely trilobed .................................... Benthamia erinacea (Cordem.) Hermans& P.J.Cribb — Lip distinctly trilobed ................................................................................................................................ 11 11. Lobes of lip truncate .................................................. Benthamia lakatoensis Descourv.& Hervouet, sp.nov. — Lobes of lip not truncate ........................................................................................................................... 12 12. Sepals green, petals white ................................................... Benthamia trifida Hermans& Hervouet, sp.nov. — Sepals white or off-white ........................................................................................................................... 13 13. Flowers pure white .................................................................................................... Benthamia nivea Schltr. — Flowers off-white ................................................................ Benthamia corona Hermans& Hervouet, sp.nov. 14. Petals white ............................................................................................................................................... 15 — Petals green or yellow ................................................................................................................................ 16 15. Basal leaf solitary ............................................................................................ Benthamia herminioides Schltr. — Basal leaves several ........................................................................ Benthamia madagascariensis (Rolfe) Schltr. 16. Leaf solitary ............................................................................................................................................... 17 — Leaves several ............................................................................................................................................ 19 17. Lip calceolate .................................................................................................. Benthamia calceolata H.Perrier — Lip not calceolate ...................................................................................................................................... 18 18. Leaf linear ................................................................. Benthamia spiraloides (Cordem.) Hermans& P.J.Cribb — Leaf elliptic or elliptic-lanceolate ..... Benthamia arcuata (H.Perrier) Descourv.& Hervouet, comb. nov., stat. nov. 19. Leaves near middle of stem ........................................................................................................................ 20 — Leaves basal ............................................................................................................................................... 24 20. Leaves 5 or more .................................................................................... Benthamia spiralis (A.Rich.) A.Rich. — Leaves fewer than 5 ................................................................................................................................... 21 21. Flowers arranged in 3 rows ............................................................................ Benthamia perfecunda H.Perrier — Flowers not arranged in 3 rows .................................................................................................................. 22 22. Lip with prominent callus ............................................................................... Benthamia melanopoda Schltr. — Lip lacking a prominent callus .................................................................................................................. 23 175 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) MATERIAL AND METHODS Acronyms of herbaria follow Index Herbariorum (http:// sweetgum.nybg.org/science/ih/). This study is mainly based on approximately 580 herbarium specimens stored at the Muséum national d’histoire naturelle (MNHN, herbarium P), but also using more than 450 specimens at AMES, BM, BR, CBNM, G, GOET, HEID, K, L, M, MO, R, REU, TAN, WU, herbaria visited by Johan Hermans and by Jean-Michel Hervouet for REU and TAN. This was combined with extensive field observations by two of the authors (Johan Hermans and Jean-Michel Hervouet), either in type localities of previously described species, or elsewhere. The descriptions are therefore based on both in situ observations and on herbarium material. Flowers were rehydrated for analysis under a binocular microscope and were drawn by either Johan Hermans (244 drawings) or Pascal Descourvières (87 drawings). The 87 original drawings by the latter author will be deposited at P. Maps used as a background for the distribution maps have been downloaded from the following websites: http://www.fao.org/geonetwork/srv/en (land cover of Madagascar, dated 2009) https://www.data.gouv.fr/fr/datasets/carte-des-departements-2 (La Réunion island) https://gadm.org (Mauritius). Similar data can be found at http://www.diva-gis.org. The locality coordinates are either from original collectors, from field observations by the authors, using a Garmin etrex 10 GPS with a WGS 84 datum, from Google Earth or from the Tropicos website gazetteer to Malagasy botanical collecting localities: http://legacy.tropicos.org/projectwebportal. aspx?pagename=mad_gazetteer&projectid=17 All specimen localities were double-checked with Google Earth (https://earth.google.com). A relational database was built using the database management system 4D version 17.1 (https://www.4d.com), storing a total of about 500 observations or localities. The Opensource Geographical Information System QGIS version 2.18.10 (www.qgis.org) was used to compose the final images. For the plant identification terminology, we used Harris& Harris (2000). The plant size given in the descriptions includes the inflorescence, it will not be repeated. To simplify and avoid confusion with the roots we call tubers what is, strictly speaking with orchids, tuberous roots. Following all previous descriptions of Benthamia species, the peduncle is included in the description of the stem, as there is no clear transition between the two in the genus, see e.g. Benthamia elata Schltr, with leaves up to the rachis. As it is a common character to all species it will not be repeated that the ovary is pedicellate. For classifying the vegetation of Madagascar, we use the terminology introduced by Gautier etal. (2018: table 23). Conservation status assessments are given following the IUCN Red List categories and criteria, version 3.1 (IUCN 2012). They are mainly based on the number of localities known, the area of occupancy (AOO) and the extent of occupancy (EOO). When available, the official IUCN assessment is also given with a reference, it is taken from https://www.iucnredlist.org/. The differences between these two assessments are most likely due to the fact that many Benthamia specimens were not or not correctly identified in herbaria, some species appearing now more widespread than initially thought. 23. Spur ventrally flattened .................................................................................. Benthamia vulcanorum sp.nov. — Spur scrotiform .............................................................. Benthamia mascula Hervouet& Descourv., sp.nov. 24. Inflorescence arcuate ........................................................................................... Benthamia bosseri Hervouet — Inflorescence erect ..................................................................................................................................... 25 25. Lip lateral lobes very short ......................................................................................................................... 30 — Lip clearly trilobed .................................................................................................................................... 26 26. Leaves more than 4 times longer than wide ............................................................................................... 27 — Leaves less than 4 times longer than wide .................................................................................................. 29 27. Leaves erect and regularly alternate .......................................................................... Benthamia flavida Schltr. — Leaves not alternate ................................................................................................................................... 28 28. Leaves basal, rachis often less than a third of plant height ....... Benthamia litoralis Hervouet& Descourv., sp.nov. — Leaves up to rachis, rachis often half of plant height ................................................... Benthamia elata Schltr. 29. Leaves 5 or more ............................................................................................ Benthamia humbertii H.Perrier — Leaves fewer than 5 ................................ Benthamia latisatis (Thouars) Bytebier. (La Réunion and Mauritius) 30. Leaves more than 5cm long, basal or cauline ....................................... Benthamia africana (Lindl.) Hermans — Leaves less than 5cm long, basal ....................................................................... Benthamia decaryana sp.nov. Key To The genus Benthamia a.rich. (continuation) 176 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. TAXONOMIC TREATMENT Family o rchidaceae Juss. Subfamilyo rchidoideae A.A. Eaton Tribe Orchideae (Dressler & Dodson) Verm. Subtribe Habenariinae sensu Ngugi etal. (2020) Genus Benthamia A.Rich. (Figs 1; 2) Monographie des Orchidées des Îles de France et Bourbon (Richard 1828: 37), non Lindl. (w). Rolfeella Schltr., Repertorium Specierum Novarum Regni Vegetabilis 33: 18 (Schlechter 1924). — Type species: Benthamia glaberrima (Ridl.) Schltr. T ype species .— Benthamia latisatis (Thouars) Bytebier (as Benthamia latifolia A.Rich.). disTribuTion and ecology. — The genus Benthamia is widespread in Madagascar, La Réunion and Mauritius, with possibly one species (probably B.africana (Lindl.) Hermans) on the Comoros (Fig. 3). It ranges from sea level to high altitudes (2600m). It is generally found in open places, both dry and marshy. e Tymology . — Named for the English botanist George Bentham (1800-1884), who together with Sir Joseph Hooker produced Genera Plantarum (1865-1883). d escripTion Terrestrial or less commonly epiphytic (5 species) sympodial glabrous herbs with 2-7 swollen generally glabrous tubers, which are in fact tuberous roots. Stem erect or arcuate, sometimes with sheaths only (one species), or with 1-2 radical sheathing leaves, or with 2-10 cauline leaves. Leaves lanceolate, ovate or elliptic, sometimes not developed at anthesis (3 species). Rachis terminal, erect or rarely curved, fewto densely many-flowered. Floral bracts small, usually as long or longer than the ovary. Ovary pedicellate, fusiform or oblong, generally twisted (except Benthamia bosseri). Flowers small, resupinate (except one species with non-resupinate and pendent flowers), arranged in a narrow rachis, white, green or yellow; sepals and petals free or almost free, entire; labellum entire or trilobed; spur cylindrical (nine species) or short and scrotiform, entire or bilobed at apex, or a small hollow only in one species (B.boiteaui). Column short and massive, subterete, erect; anther erect to slightly deflexed, placed above the rostellum, apiculate or not, bilocular, locules parallel, without anther canals; pollinia two, ellipsoid, granular, 2 caudiculae rudimentary, with two viscidia, detachable, flat, ovate or subelliptic; rostellum small, tridentate or trilobedat the anterior edge; stigma always very short, cushion-shaped, with short convex lobes; auricles two, generally conspicuous, subspathulate or falciform, sometimes lacerated, on lateral sides of column. Fruit a dehiscent capsule, with floral parts persistent on the fruit. 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• AlaotraMangoro; alt. 800m; 11.X.1912; H. Perrier 11380; holotype: P[P00738317]!; isotype: P[P00094573]!, syn. nov. T ype 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. d isTribuTion 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). c onservaTion . — 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. a ddiTional 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]!• AlaotraMangoro, 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]!• 177 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) A B C E D fig. 1. — Benthamia latisatis (Thouars) Bytebier, specimen P00334779, showing the different parts of the plant: A, rachis; B, sterile bract; C, stem (including the peduncle); D, leaf; E, sheath. Photograph by Jean-Michel Hervouet. 184 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. diameter, with a short apiculus at apex; rostellum trilobed, short, with midlobe rounded; auriclesstrap-shaped, half the column length. BenthamiabathieanaSchltr. (Figs 11; 12) Repertorium Specierum Novarum Regni Vegetabilis 33 (Schlechter 1924: 25). Benthamia latifolia Schltr., Beihefte zum Botanischen Centralblatt 34(2) (Schlechter 1916: 303), nom. illeg. non A.Rich.— Type: Madagascar• Vakinankaratra, Mt. Ibity; II.1914; H. Perrier s.n.; holotype: B, could not be traced, possibly destroyed during WWII. Type maTerial.— Madagascar• Vakinankaratra, Mt. Ibity; 20°06’25”S, 46°59’51”E; alt. 2000m; IV.1921; H. Perrier 13582; holotype: P[P00094469]!; isotype: P[P00094470]!. eTymology. — Named for the French botanist Henri Perrier de la Bâthie, who collected the type specimen on Mt. Ibity. phenology. — February to April. disTribuTion and ecology. — Madagascar. Only recorded from the centre: from Mt. Ibity near Antsirabe and in the Itremo massif near Ambatofinandrahana, amongst rocks, in montane grasslands. The altitude range is 1700-2100m (Fig. 13). conservaTion. — This species is only known with certainty from Mt. Ibity and the Ambatofinandrahana area (including Itremo National Park). This is fewer than five localities, with an AOO less than 500km² and a quality of habitat in continuing decline due to bush fires, it should be considered Endangered (EN) according to criterion B2. a ddiTional specimens examined . — Madagascar• Amoron’i Mania, environs d’Ambatofinandrahana; 20°40’11”S, 47°07’19”E; alt. 1600-1800m; 23.II.1938; R. Decary 13230; P[P00094473]!• Amoron’i Mania, Itremo, 20°34’12”S, 46°34’55”E; IV.1964; J. Bosser 19588; P[P00094471, P00094472]!, TAN!• Vakinankaratra, Mt. Kiboy, Ibity massif; 20°3.66’S, 47°0.01’E; V.2001; J. Hermans 121; K!. noTes The identity of this species has only recently been correctly understood. It can easily be confused with Benthamia rostrata in photographs, though it is a much larger plant. Several Benthamia rostrata specimens have been labelled Benthamia bathieana in the literature and herbaria. Apart from the plant size it differs from Benthamia rostrata by the leaves which are closer together and wider, and its denser inflorescence. The fig. 12 . — Benthamia bathieana Schltr., Mount Ibity, Madagascar, 1 April 2018. Photograph by Jean-Michel Hervouet. fig. 11 . — Benthamia bathieana Schltr., Mount Ibity, Madagascar, 1 April 2018. Photograph by Jean-Michel Hervouet. 185 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) flowers differ from B.rostrata by the petals and sepals twice as long, the side-lobes of the lip which are not equilateral vsequilateral in B.rostrata, the shorter spur, and the paleyellow colour of the flowers (vs. greenish in B.rostrata). The gathering H. Perrier 13582has two sheets in P, but one of them (P00094469) is clearly annotated in Perrier’s handwriting that it is the material that was sent to Schlechter for the description, and can therefore be considered the holotype, with P00094470 being an isotype. See also: Schlechter (1930: t.9 n°36), Perrier (1939: 23), Cribb& Hermans (2009: 42, photograph is Benthamia rostrata Schltr). descripTion Robust terrestrial herb up to 70cm tall. Tubers 20-30, narrowly fusiform, extended by a root up to 12cm long. Stem 10-15mm in diameter, with base enclosed by sheaths, with leaves higher up. Leaves 5-6, big, oval or elliptic, the middle ones up to 16× 7.5cm, the upper ones becoming gradually smaller. Rachis rigid, elongate, 22-30× 1.8cm, densely 40-80-flowered. Floral bracts lanceolate, very acuminate, the lower ones longer than the ovary, 5-10× 2mm. Ovary 6-10× 1.8mm long. Flower butter yellow; sepals oblong, obtuse, 1-veined, dorsal sepal 3.5-3.9× 1-2mm long, concave and erect; lateral sepals 4-4.5× 1.2-2mm; petals oval-obtuse, 2-veined, 4× 2-2.8mm, a little expanded at the front edge towards the base. Lip 4-6× 2.2-3mm, without callus, broadly angular or wedge-shaped at the base, clearly trilobed from the middle; lateral lobes flat, triangular-falciform, midlobe linear-ligulate, somewhat obtuse and fleshy, upcurved at the apex; spur pendent, cylindrical, 3.6-4.5× 0.5-0.7mm long. Column 1.2-2× 1mm, anther oval-obtuse; auricles 2-3 times shorter than the anther. Benthamia boiteaui Hervouet (Figs 14; 15) Adansonia, sér. 3, 36 (2): 213 (Hervouet etal. 2014). T ype maTerial . — Madagascar• Haute Matsiatra, Ambondrombe; 21°52’47”S, 47°15’33”E; alt. 1900m; 11.IV.1941; P. Boiteau 4621; holotype: P[P00692438]!. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 13. — Distribution map of Benthamia bathieana Schltr. fig. 14. — Benthamia boiteaui Hervouet, Mount Ambondrombe, Madagascar, 9 March 2009. Photograph by Jean-Michel Hervouet. 186 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. eTymology. — Dedicated to Pierre Boiteau (1911-1980), collector of the type and former director of the Tsimbazaza botanical and zoological garden in Antananarivo. phenology. — March to April. disTribuTion and ecology. — Madagascar. In Haute Matsiatra and Amoron’i Mania regions. Always found in damp places in montane grasslands or marshes, from1000 to 1900m. During the present study a specimen has been identified from Antoetra, near Ambositra, it remains however rather rare (Fig. 16). conservaTion. — Now known from three different locations, each in very limited areas giving an AOO of less than 500km², in habitats of montane grasslands threatened by bush fires, indicating a conservation status as Endangered (EN) according to criterion B2. addiTional specimens examined. — Madagascar• Haute Matsiatra, Ambondrombe; 21°52’47”S, 47°15’33”E; alt. 1900m; 11.IV.1941; P. Boiteau 4621; P[P00692439, P00692440]!• Haute Matsiatra, Ambondrombe; alt. 1700m; 11.IV.1941; P. Boiteau 4637; P[P00692439, P00692440]!• Haute Matsiatra, Ambondrombe; alt. 1800m; 12.IV.1941; P. Boiteau 4687; P[P00692303]!• Amoron’i Mania, Antoetra, piste du Vohibe, sud d’Ambositra; 20°46’27”S, 47°19’16”E; V.1969; P. Morat 3148; P[P00692448, P00692449]!• Haute Matsiatra, parc de Ranomafana; 21°11’S, 47°25’E; alt. 1150m; 10.III.1995; Turk etal. 722; K!, K[spirit 72273.000], MO!, TAN!• Haute Matsiatra, Ambondrombe; alt. 1900m; 1999; J. Hermans 4403 ex Malala orchidées; K!. noTes This species shares some similarities with the genus Tylostigma, e.g. the spur is similar to that of Tylostigma nigrescens Schltr., but the lip lacks a transverse lamellate callus and is typical for Benthamia. See also: Hervouet (2018: 158). descripTion Erect terrestrial herb 27-38cm tall. Tubers and roots not seen. Stem cylindrical, 2mm in diameter, dull pale green, with one or two sheaths above the ground followed, in the first 2/3rd, by 2-3 leaves, and then in the upper part 1-2 sterile bracts, linear, acute, 15-30× 2-3mm. Leaves amplexicaul, flexible, linear and acute, 50-130× 5-8mm. Rachis 5-8cm long, terminal, erect, with 15-33 flowers. Floral bracts acicular, fig. 15. — Benthamia boiteaui Hervouet, Mount Ambondrombe, Madagascar, 9 March 2009. Photograph by Jean-Michel Hervouet. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 16. — Distribution map of Benthamia boiteaui Hervouet. 187 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) longer than ovary, 4-11× 1.5-2mm. Ovary 4.8-7mm× 1.93mm. Flowers 6mm deep, 4.5mm high, 5mm wide, bright white; dorsal sepal elliptic, obtuse with a slightly rounded and concave apex, 3-6mm× 1.2-3mm; lateral sepals elliptic, 3.1-5.5× 1.2-3mm, with a visible vein, dorsally slightly keeled, ventrally slightly concave, apex rounded, sub-acute, margin smooth; petals partly overlapping and together with the dorsal sepal forming a hood over the column and lip, elliptic, 3-4.1× 1.8-2mm, with a visible vein, apex rounded, sub-acute, margin smooth or slightly irregular; lip trilobed, 2.5-5× 1.6-3mm, lateral lobes acute, 0.5mm long, central lobe fleshy, 1.0-1.3mm long; spur a shallow hollow only, very short and obtuse, 0.1-0.3× 1mm, adpressed to the ovary. Column apiculate, 1-1.5× 1-1.2mm, lateral auricles large, suborbicular; rostellum trilobed, with midlobe apiculate; masullae very small, less than 0.12mm in diameter. Benthamia bosseri Hervouet (Figs 17; 18) Adansonia, sér. 3 36 (2): 217 (Hervouet etal. 2014). T ype maTerial . — Madagascar• Haute Matsiatra, Ambondrombe; 21°52’47”S, 47°15’33”E; alt. 1700m; 11.IV.1941; P. Boiteau 4634; holotype: P[P00692301]!. eTymology. — Dedicated to Jean Bosser (1922-2013), who collected a specimen of the species and was instrumental in the advancement of our current knowledge of the orchids of Madagascar and the Mascarenes. phenology. — March and April. disTribuTion and ecology. — Madagascar. First described from the Haute Matsiatra region in Madagascar, on Ambondrombe and Andringitra, in montane grasslands among rocks. Since 2014 it has fig. 17 . — Benthamia bosseri Hervouet: Andringitra massif, Madagascar, 7 April 2018. Photograph by Jean-Michel Hervouet. fig. 18. — Benthamia bosseri Hervouet: Mount Ambondrombe, Madagascar, 9 March 2009. Photograph by Jean-Michel Hervouet. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 19. — Distribution map of Benthamia bosseri Hervouet. 188 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. also been found around Antoetra in Amoron’i Mania region (first author’s observation and photographs, without specimen). The altitudinal range is 1600-2100m (Fig. 19). conservaTion. — Although found in two national parks, it is only known from three locations with an AOO less than 500km², in habitats threatened by fires coming up from prairies around the mountains. This species can therefore be considered Endangered (EN) according to criterion B2. addiTional specimens examined. — Madagascar• Haute Matsiatra, Ambondrombe; 21°52’47”S, 47°15’33”E; alt. 1700m; 11.IV.1941; P. Boiteau 4634; P[P00692265, P00692299, P00692300, P00692302]!• Haute Matsiatra, Parc National de l’Andringitra; 22°07’40”S, 46°51’48”E; IV.1964; J. Bosser 19648; P[P00692265]!. noTes The flowers remain fresh long after pollination, they are probably autogamous. See also: Hervouet (2018: 159). descripTion Terrestrial herb 21-71cm long. Tubers and roots not seen. Stem erect, cylindrical, 2-5mm in diameter, green or purple (on the same plant), with leaves in the lower part, followed by 1-2 sheaths and 2-3 sterile bracts in the upper part. Leaves 2-3, amplexicaul, thin, flexible, linear, acute, 50-140× 5-10mm; sterile bracts linear, acute, 13-25× 2-3mm. Rachis 7-19cm long, arcuate, bearing 20-50 flowers. Floral bracts decurrent, acicular, reddish, as long as ovary + flower, 7-12× 2mm. Ovary not twisted, 3-10mm long, 1-2mm in diameter, up to 3mm when ripe, greenish or reddish. Flowers secund, not resupinate, drooping, yellow and reddish; sepals subsimilar, linear, concave, 3-3.5× 1mm, with a central vein, slightly keeled dorsally, reddish; petals linear, similar to sepals but slightly narrower, slightly fleshy at apex, 3-3.5× 0.8-1.1mm, a little narrower at the base, yellow; lip trilobed, 3-4× 0.82mm, with a longitudinal rounded callus prolonged into a thickened midlobe, 1mm long, lateral lobes a little shorter than midlobe, thickened; spur very short, 1-1.5mm, round and flattened, bilobed when dry. Column terete, very short, 0.9-1mm, anther apiculate; auricles relict. Benthamia calceolata H.Perrier (Figs 20; 21; 22) Bulletin de la Société botanique de France 81 (Perrier 1934: 32). Type maTerial. — Madagascar• Sava, Andratamarina, near Mananara; 16°17’49”S, 49°43’37”E; 20.VIII.1920; R. Decary 24; lectotype: P[P00094475]!, here designated; isolectotype: P[P00094476]!, only a flower dissection of the former. eTymology. — From Latin “calceolatus”: slipper-shaped, referring to the shape of the lip. phenology. — From June to November. disTribuTion and ecology. — Madagascar. Only known from four locations, all near the east coast, almost from north to south. In humid places, in riparian Pandanus forest or lowland moist evergreen forest. Altitude from 100 to 500m (Fig. 23). conservaTion. — Known so far from only four localities but the very large EOO suggests that it has been largely overlooked due to its diminutive habit. IUCN quoted it EN in 2015, under criteria B2ab(ii,iii,iv,v) (https://doi.org/10.2305/IUCN.UK.2017-2.RLTS. T70102664A70132920.en). The only recent observation has been in the Ambodiriana forest, which has an area of 240ha. As fewer than 50 individuals have been counted, considering only the recently observed living specimens would indicate a Critically endangered species (CR) status under criterion D. addiTional specimens examined. — Madagascar• Sava, Manantenina; 24°16’49”S, 47°19’10”E; 10.VI.1925; R. Decary 3877; P[P00094477]!• Sava, Masoala Peninsula, west of village of Andrem - bona; 15°17’25”S, 50°13’10”E; alt. 470m; 19.XI.1993, K.Lance 39; K!. fig. 20. — Benthamia calceolata H.Perrier: Ambodiriana forest, Manompana, Madagascar, 18 October 2022. Photograph by Véronique Lavergne. fig. 21. — Benthamia calceolata H.Perrier: Ambodiriana forest, Manompana, Madagascar, 18 October 2022. Photograph by Véronique Lavergne. 189 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) noTes This species differs from all the others in the genus by the pouched lip. It has recently been seen in October 2022 in the Ambodiriana forest on the east coast of Madagascar, by Véronique Lavergne, who provided us with the first photographs of the species. The fasciculate habit observed on the type was not confirmed in recent observations. There are two sheets of the R. Decary 24 gathering, we chose P00094475 as the lectotype, the isolectotype P00094476 be - ing a dissected flower. See also: Perrier (1939: 28), Cribb& Hermans (2009: 43). descripTion Erect terrestrial herb, 30-60cm tall. Tubers 5-8, elongated, up to 3cm, cylindrical or fusiform, ending in a root. Stem erect, 2-3mm in diameter, with a basal leaf and then 6-8 scale-like leaves, slowly turning into very acute floral bracts with linear blade. Leaf 1, oblong-lanceolate, c.14× 3.2cm, acutely narrowed at both ends. Rachis slender, lax, narrow, 17× 1cm, bearing 30-45 flowers. Floral bracts lanceolate, very acute, in the middle of the rachis slightly longer than half the length of the ovary. Ovary 4.5-6× 1.3mm. Flowers 3mm without the ovary, greenish yellow; all sepals 1-veined and with an obscure dorsal keel; dorsal sepal oval, obtuse or with apex subacute, 1.5-2.2× 0.8-1mm; lateral sepals oval-obtuse, 1.5-2.2× 0.81mm; petals much thicker, very concave, obovate, 1.4-2× 0.7-0.8mm, a little cucullate and very rounded at the tip; lip entire or very obscurely trilobed at apex, 1.8× 1-1.1mm, slipper shaped and very short, a little verrucose on the outside, with apex truncated; spur scrotiform, bilobed, as wide at the tip as long, 1-1.1× 0.8-1mm, contracted at the base, with a clear deep furrow at the tip. Column sub-terete, very different from others in the genus, 0.7-1× 0.5-1mm, with apex digitate, 0.3mm long, slightly canaliculate in its internal part, overhanging the rostellum; rostellum trilobed with an erect midlobe around 0.3mm long and two short lateral lobes; auricles basal, strap-like, nearly reaching the anther margins. Benthamia corona Hermans& Hervouet, sp.nov. (Figs 24; 25; 26) diagnosis. — Benthamia corona Hermans& Hervouet, sp.nov. is distinct by its epiphytic habit, narrowly elliptic leaves, long sheaths fig. 22. — Benthamia calceolata H.Perrier: Ambodiriana forest, Manompana, Madagascar, 18 October 2022. Photograph by Véronique Lavergne. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 23. — Distribution map of Benthamia calceolata H.Perrier. 190 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. decreasing in length as they become floral bracts, its loosely spiral rachis, the yellowish-white flowers (large for the genus), with a long corolla tube, its distinctly trilobed lip with triangular lobes and an incurved midlobe, and its anther that lacks a distinct midlobe. It shares a similar habit and spur with Benthamia erinacea but has narrower leaves, a less dense rachis and a trilobed lip (vsentire). It is most similar to Benthamia nivea and Benthamia lakatoensis Descourv.& Hervouet, sp.nov. with which it shares the epiphytic habit, the sheath-floral bract arrangement, relatively large pale flowers and long corolla tube but differs by its narrowly elliptic (vselliptic to ovate) leaves, flowers that are yellowish-white (vspure white) and about 1/3rd smaller, the lobes of the lip that are triangular attenuate (vsobtuse to acute), its bi-globose flattened (vsglobose-saccate) spur and its anther lacking a distinct midlobe. Type maTerial. — Madagascar• Analamanga, Anjozorobe area, Hort. Malala Orchidées; 2001; J. Hermans 5183; holotype: K!. eTymology. — The species epithet refers to the crown-like appearance of the flowers and its morphological resemblance to, and commemorating the coronavirus (COVID-19), an infectious disease caused by the SARS-CoV-2 virus, which resulted in a world pandemic from 2019 to 2022. phenology. — Flowering time is not known. disTribuTion and ecology. — Madagascar. Only known from Tsaratanana in Diana region and from Anjozorobe area in Analamanga region. Altitude and habitat are not known, it is thought to be from medium altitude moist evergreen forest (Fig. 25). conservaTion. — The extent of occurrence (EOO) of Benthamia corona Hermans& Hervouet, sp.nov. cannot be estimated since it is only known from one locality and from cultivated material but given the limited collection it is estimated to be in Category CR (Critically Endangered) under criterion D if we estimate that there are fewer than 50 individuals in the wild. a ddiTional specimens examined . — Madagascar• Diana, Tsaratanana; 14°01’17”S, 48°58’00”E; 1950; R. Paulian Jard. Bot. 786; P[P00692271]!. descripTion Slender epiphytic herb, 5.5-12cm tall. Tubers 1-2, ovoid to fusiform, woolly, 22-32× 5-10mm, roots wiry, glabrous, 0.6– 0.8mm in diameter. Stem 3-5mm in diameter, with 1-2 thin basal enveloping sheaths, then leaves, then 3-5 sheaths, 7-15× 2-5mm, decreasing in size until they merge with the floral bracts. Leaves 2-3, cauline with the base enveloping the stem for 5-11mm, erect to arching, narrowly elliptic, attenuate, 3.5-11.5× 0.4-0.8cm, petiolate, almost flat, pale to dark fig. 24. — Benthamia corona Hermans & Hervouet, sp. nov., Hort. Malala Orchidées, 2001. Photograph by Johan Hermans. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 25 . — Distribution map of Benthamia corona Hermans & Hervouet, sp. nov. 191 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) A C E F G B H D fig. 26. — Benthamia corona Hermans & Hervouet, sp. nov.: A, floral bract; B, flower side view and front view; C, dorsal sepal, petal and lateral sepal; D, lip and spur from above; E, ovary and flower with sepals and petals removed side view and view from above; F, column from above and below; G, plant and inflorescence; H, inflorescence. Drawn by Deborah Lambkin. From Hermans 5183 and photographs. Scale bars: A, B, 3 mm; C-E, 2.5 cm; F, 1 mm; G, 1.5 cm; H, 1 cm. 192 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. green. Rachis up to 10cm long, erect, pale green, loose in the upper half, with up to 30 spirally arranged flowers, developing acropetally with the lower flowers fertilized before the apical ones open. Floral bracts narrowly lanceolate, acuminate, somewhat incurved, decreasing considerably in size towards the apex with the basal ones more than twice the length of the ovary, the apical ones a little shorter, 6-13.5× 1.1-1.8mm, green. Ovary fusiform, at first erect then divergent, with three longitudinal ridges, 4.5-6.3× 1.6-1.9mm, pale green. Flowers large for the genus, fleshy, c.3× 3.5× 5mm long, corolla fused-tubular in the basal half; dorsal sepal recurved, narrowly elliptic, acute, 3.5-4.2× 0.9-1.2mm, greenish-white, green at the base; lateral sepals spreading to recurved, narrowly elliptic, margins involute, acute, 4.9-5.3× 1.6-1.9mm, greenish-white, green at the base; petals spreading, recurved, elliptic, the apex and anterior margin, thickened, erose, 2.8-3.6× 1-1.5mm, creamy-white; lip distinctly trilobed, 4.3-4.6× 3.9-4.4mm, creamy-white, with lateral lobes decurved, triangular and attenuate and midlobe with the apex distinctly incurved; spur bi-globose, laterally flattened, 0.9-1.2× 0.9-1.3mm, pale green. Column 0.8-0.9× 0.9-1mm, anther equally bilobed; auricles falcate, scabrid. Benthamia decaryana Descourv.& Hervouet, sp.nov. (Figs 27; 28; 29) diagnosis. — This species has broadly similar flowers to those of Benthamia africana but the habit is smaller (largest plant known 35cm high vs62cm). The most distinctive difference lies in the fact that leaves are all basal, less than 5cm long, conduplicate, coriaceous, generally with undulate margins and arcuate, at least when dried (vsleaves linear or ligulate, 6-18cm long, remaining flat when dried). The rostellum has a very wide and large midlobe, 0.4-0.5mm long, protruding and pointing downwards (vsmidlobe 0.1-0.2mm). T ype maTerial . — Madagascar• Atsimo-Atsinanana, Midongy du sud; 23°35’S, 47°01’E; 23.VIII.1926; R. Decary 5138; holotype: P[P00094591]!; isotypes: P[P00094592]!, K!, TAN!. e Tymology . — Named for Raymond Decary (1891-1973), co - lonial administrator in Madagascar. He undertook seven scientific missions from 1922 to 1944, collecting, among many other things, 40000 specimens of plants. phenology. — September. disTribuTion and ecology. — Madagascar. Known only from the Atsimo-Atsinanana region, therefore the south-east, in forest on laterite soil. A humid roadside is mentioned on the type specimen (Fig. 28). conservaTion. — This species has been collected only twice and has not been recorded since 1926. However, the locus classicus, South Midongy, is not very accessible. With only two known locations and the continuing decline of the forests in the region it must be considered Endangered (EN) under criterion B1. addiTional specimens examined. — Madagascar• Atsimo-Atsinanana, Vondrozo; 22°49’14”S, 47°19’10”E; 5.IX.1926; R. Decary 5195; P[P00094587]!. auricle rostellum fig. 27. — Benthamia decaryana Descourv. & Hervouet, sp. nov.: drawing of the column by Pascal Descourvières, showing the auricle and the rostellum. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 28 . — Distribution map of Benthamia decaryana Descourv. & Hervouet, sp. nov. 193 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) noTes This small species has a strikingly distinctive rostellum compared with other Benthamia species. The most distinctive difference is however the short basal conduplicate leaves, which are unique among other Benthamia. fig. 29. — Benthamia decaryana Descourv. & Hervouet, sp. nov.: drawing by Ludivine Longou, after dried specimens: A, habit; B, side view of part of the rachis. Scale bars: A, 10 mm; B, 5 mm. AB 200 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. eTymology. — From Latin “reticulatus”, veined by a network. phenology. — Same as typical form. d isTribuTion and ecology . — Madagascar. Potentially within the distribution range of Benthamia erinacea in Madagascar, in the same habitat. conservaTion. — Very little is known so far on this form. Its conservation is certainly linked to that of the typical form, as it probably appears in its populations. It is provisionally quoted Data Deficient (DD). noTes Specimens with reticulated or white-marbled leaves have been recorded by Jean Bosser (P[P00692431]) and Johan Hermans, they are frequent among populations in Andasibe region. descripTion As Benthamia erinacea, except that the leaves are finely reticulated in white. Benthamia exilis Schltr. (Fig. 40) Repertorium Specierum Novarum Regni Vegetabilis 33: 26 (Schlechter 1924). Benthamiaexilisvar. tenuissima Schltr., Repertorium Specierum Novarum Regni Vegetabilis 33 (Schlechter 1924: 26).— Type: Madagascar• Vakinankaratra, Mt. Tsiafajavona; 19°21’25”S, 47°18’28”E; alt. 2400m; III.1921; H. Perrier 13522; lectotype: P[P00094501]!, designated here; isolectotype: P[P00094502]!, syn. nov. Type maTerial. — Madagascar• Haute Matsiatra, massif d’Andringitra; 22°07’28”S, 46°52’07”E; alt. 2200m; IV.1921; H. Perrier 13652; lectotype: P[P00094498]!; isolectotype: P[P00094499]!, designated by Hermans etal. (2007: 70). eTymology. — From the Latin “exilis”: slender, meagre, thin. phenology. — January to April. disTribuTion and ecology. — Madagascar. So far only known from Mt. Tsiafajavona in the Vakinankaratra region and the Andringitra massif in Haute Matsiatra region, in montane grasslands, at elevations ranging from 1700 to 2400m (Fig. 41). conservaTion. — Known only from two localities, Mt. Tsiafajavona and the Andringitra massif, this species has been seldom collected fig. 38. — Benthamia erinacea (Cordemoy) Hermans & P.J.Cribb: Iaroka forest near Andasibe, 11 July 2023. Typical form and f. reticulata f. nov. mixed. Photograph by Jean-Michel Hervouet. fig. 39. — Benthamia erinacea (Cordemoy) Hermans & P.J.Cribb f. reticulata f. nov.: forêt de Iaroka, 11 July 2023. Reticulated leaf. Photograph by JeanMichel Hervouet. fig. 40. — Benthamia exilis Schltr., Andringitra Massif, Madagascar, 12 March 2009. Photograph by Jean-Michel Hervouet. 201 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) and is considered Endangered (EN), according to criterion B2, with fewer than five known localities and threatened habitat. a ddiTional specimens examined . — Madagascar• Haute Matsiatra, massif d’Andringitra; alt. 2200m; II.1922; H. Perrier 14390; P[P00094500]!• Haute Matsiatra, massif d’Andringitra; alt. 2200m; II.1922; H. Perrier 14579; P[P00094503]! for Benthamia exilis var. tenuissima• Haute Matsiatra, Andringitra; 2001; J. Hermans 2063; K!. noTes The specimens available show a variation of habit from 12cm to about 50cm high and a great variation of rachis length. Considering this variability we include Benthamia exilis var. tenuissima Schltr. in the synonymy of B.exilis. For this variety Schlechter cited Perrier 13522 for the type, but there are in P two specimens with this collection number. We have chosen P00094501 as lectotype because Perrier indicated on it that it was sent to Schlechter in April 1921. Plants referred to var. tenuissima in Hervouet (2018: 162) are Cynorkis micrantha (Frapp. ex Cordem.) Schltr. In the same page 162 the specimen cited Perrier 74579 should read Perrier 14579. An undescribed species close to B.exilis has been observed on the summit of Ambondrombe, but with a much longer midlobe of the lip. No herbarium specimen has been located. See also the notes under Benthamia micrantha in excluded species. See also: Schlechter (1930: t.10 n°38), Perrier (1939: 20); Cribb& Hermans (2009: 43), Hervouet (2018: 162, the bottom photograph is Cynorkis micrantha). descripTion Very slender terrestrial herb, 13-50cm tall. Tubers oblong and smooth. Stem filiform, 0.8-1.5mm in diameter, covered by 4-7 distant sheaths with blade acute. Leaves absent. Rachis 8× 1-1.2cm, rather dense, many-flowered. Floral bracts lanceolate, almost as long as the flowers, 2.8-5× 1-1.2mm. Ovary fusiform-cylindrical, smooth, 4-5× 0.8mm. Flowers yellowish or greenish, sometimes with brown sepals; sepals oblong, somewhat obtuse, 2-3.5× 0.8-1.5mm, 3-veined, the dorsal sepal concave, the laterals oblique; petals oblong, obtuse, 1.9-3.1× 0.9-2mm, 3-veined and slightly expanded at the anterior edge in the lower half; lip narrowly oval in outline, 3-4.1× 1.6-2.5mm, almost orbicular and concave in its basal half, narrowly lanceolate and obscurely trilobed in the apical half, midlobe about 1.8mm; spur with a wide base, narrowly cylindrical towards the apex, 1.8-3× 0.3-0.4mm long and bifid at the tip. Anther obtuse and short, 0.6-1.2× 0.5-0.8mm; auricles erect, linear, curved in a scythe shape and taller than the anther. Benthamia flavida Schltr. (Figs 42; 43) Beihefte zum Botanischen Centralblatt 34(2) (Schlechter 1916: 302). Benthamia perrieriSchltr., Beihefte zum Botanischen Centralblatt 34(2) (Schlechter 1916: 302). Type: Madagascar• Vakinankaratra, Antsirabe, marais tourbeux; alt. 2000m; II.1914; H. Perrier 8081; holotype: P[P00094490]!. Benthamia cinnabarinasensu H.Perrier, Bulletin de la Société botanique de France 81(Perrier 1934: 38). Perrier cited a wrong type: Trabonjy; V.1880; J.M. Hildebrandt 3450; P[P00094480]!, K!, which is indeed Benthamia flavida, but the basionym Habenaria cinnabarina Rolfe is referable to Cynorkis cinnabarina (Rolfe) Hermans& P.J.Cribb; the type is in K, it is a specimen cultivated by Lewis& Co in England in 1893. T ype maTerial . — Madagascar• Vakinankaratra, Cime du Famoizankova, au nord d’Antsirabe; 19°39’32”S, 47°03’48”E; alt. 2200m; V.1912; H. Perrier 11386; holotype: P[P00094481]!. eTymology. — Derived from Latin «flavus», yellow, referring to the yellowish colour of the flower. phenology. — January to May. disTribuTion and ecology. — Madagascar. Except for one locality in the north, mainly in montane grasslands in the central plateaux, in dry and open places, but sometimes also in wet areas, from 1200m to 2200m (Fig. 44). conservaTion. — This plant is very common in Madagascar and can therefore be considered of Least Concern (LC). It was considered NT by IUCN in 2008, under the name Benthamia cinnabarina. See https://doi.org/10.2305/IUCN.UK.2013-1.RLTS. T44392752A44475517.en. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 41. — Distribution map of Benthamia exilis Schltr. 202 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. addiTional specimens examined. — Madagascar• Sine loc.; R. Baron 5193; P[P00094484]!, K!• sine loc.; R. Baron 5245; P[P00094485]!• sine loc.; Bojer etal. s.n.; P[P00094486]!• Nord Betsileo; I.1881; J.M. Hildebrandt 3975; W[R14942, R8645]!, WU, G, M, BM[BM000034589]!• Central, North Betsileo; I.1881; comm. C. Rensch, J.M. Hildebrandt s.n.; K!• Vakinankaratra, mont Ibity au sud d’Antsirabe; 20°05’44”S, 47°00’12”E; alt. 2000m; II.1914; H. Perrier 8097; P[P00094487, P00094488]!• Haute Matsiatra, massif d’Andringitra; alt. 2000m; IV.1921; H. Perrier 13614; P[P00094492]!, K!• Haute Matsiatra, massif d’Andringitra; alt. 2200m; II.1922; H. Perrier 14578; P[P00094482, P00094491]!• Amoron’i Mania, environs d’Ambatofinandrahana; alt. 1600-1800m; 19.II.1938; R. Decary 13074; P[P00094489]!• Sofia, de la haute Maevarano au bas Sambirano par Bealanana et la haute Sandrakoto; alt. 1400-1700m; II.1951; H. Humbert etal. 25402; P[P00692280]!, TAN!, K!• Diana, Betainkankana, Ankaizina; 14°37’S, 48°43’E; V.1952; J. Bosser 2560; P[P00094483, P00692223]!• Amoron’i Mania, montagnes à l’ouest d’Itremo; 20°34’12”S, 46°34’55”E; 1955; H. Humbert 30046; P[P00692281]!, another specimen Humbert 30046 is Tylostigma nigrescens Schltr. or Tylostigma perrieri Schltr.• Amoron’i Mania, montagnes à l’ouest d’Itremo; 20°34’12”S, 46°34’55”E; 1955; H. Humbert 30050; P[P00692283]!, TAN!• Amoron’i Mania, montagnes à l’ouest d’Itremo; 20°34’12”S, 46°34’55”E; alt. 1500-1700m; 1955; H. Humbert 30055; P[P00692284 second from left only, P00692285 left only, P00692287]!, MO[3248465]!• Haute Matsiatra, Sendrisoa, Ambalavao, Antanifotsy; 22°13’S, 46°55’E; 5.II.1955; C. Rakotovao 7303; P[P00692316]!• Amoron’i Mania, route d’Ambositra à Ambatofinandrahana; 20°40’03”S, 47°08’03”E; III.1960; M. Keraudren 148; P[P00692288 plant in the middle is Benthamia glaberrima]!• Amoron’i Mania, Soanieranana, col d’Antoetra, district d’Ambositra; 20°46’27”S, 47°19’16”E; 21.III.1960; M. Peltier 2229; P[P00692293, P00692294]!• Amoron’i Mania, Ambositra, Anjoma; III.1960; J.& M. Peltier 2173; P[P01805103]!, TAN! plant on the left• Amoron’i Mania, route d’Ambatofinandrahana, à 10km d’Ivato; 20°40’03”S, 47°08’03”E; II.1963; F. Chabonis s.n.; P[P00692269]!• Atsimo-Andrefana, environs de Betroka, région de Tuléar; 23°16’20”S, 46°05’58”E; II.1963; J. Bosser 17387; P[P00692239]!• PK300, RN7; IV.1963; J. Bosser 16815; P[P00692237]!• PK300, RN7; VIII.1963; J. Bosser 17553; P[P00692240]!• Haute Matsiatra, Andringitra; IV.1964; J. Bosser 19508; P[P00692254]!• Amoron’i Mania, massif de l’Itremo; 20°34’12”S, 46°34’55”E; IV.1964; J. Bosser 19596; P[P00692256, P00692257]!• Amoron’i Mania, 10km route Ivato, route d’Ambatofinandrahana; 20°40’03”S, 47°08’03”E; IV.1964; J. Bosser 19599; P[P00692260]! plant on the left, the two others are Tylostigma• Haute Matsiatra, 25km SW of Fianarantsoa; 5.III.1969; W. Rauh 22258; HEID[HEID740739, HEID740740, HEID740741, HEID740742, HEID740743, specimens in alcohol]!• Vakinankaratra, Mt. Ibity; 6.V.1970; J. Bosser 20258; P[P00692266]!• Amoron’i Mania, Morondava Ambatofinandrahana road, Col d’Itremo; 20˚40’S, 46˚35’E; alt. 1600m; III.1971; D.J. Mabberley 763; K!, TAN!• Amoron’i Mania, near Morondava-Ambatofinandrahana road RN35, Col d’Itremo; 20°40’S, 46°35’E; III.1971; D.J. Mabberley 770; K!• Analamanga, Angavokely; 5.III.1988; B. Pettersson etal. 222; P[P02115318]!, K[spirit K55125.000]!• Analamanga, Angavokely; 22.III.1988; B. Pettersson etal. 282; fig. 42 . — Benthamia flavida Schltr., Andringitra Massif, Madagascar, 13 March 2009. Photograph by Jean-Michel Hervouet. fig. 43 . — Benthamia flavida Schltr.: Mount Ibity, Madagascar, 13 March 2015. Photograph by Jean-Michel Hervouet. 203 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) P[P02115317]!• Amoron’i Mania, Antoetra village; 20°46’27”S, 47°19’16”E; alt. 1710m; V.1993; C.H. Jongkind etal. 870; K[K000624472]!, BR[BR0000005242719]!, MO!, TAN!• Fianarantsoa province, SW of Ivato, off RN7, about 9km on Ambatofinandrahana road; 20°40’03”S, 47°08’03”E; 1994; J. Hermans 3519; K!• Amoron’i Mania, Itremo, près du col; 20°34’28”S, 46°34’57”E; alt. 1710m; 19.IV.1995; D.J. Du Puy M912; P[P00059608]!, K!, K[spirit SPC-72244.000], TAN!• Highlands, cultivated plant, found in bog; 1995; J. Hermans 2740; K!• Amoron’i Mania, Ambatofinandrahana; 1998; J. Hermans 5047; K!• Haute Matsiatra, Andringitra foothills, near the village of Soavihiny; 22°06’08”S, 46°47’04”E; V.2001; J. Hermans 5240; K!• Haute Matsiatra, Andringitra foothills, near village of Soavihiny; 22°04’54”S, 46°46’28”E; alt. 1506m; V.2001; J. Hermans 5245; K!• Amoron’i Mania, road towards Ambatofinandrahana, Inselberg a few km before Anjomam Ankona; 20°40’11”S, 47°07’19”E; IV.2001; alt. 1548m; J. Hermans 5256; K!• Amoron’i Mania, near Ambatofinandrahana; 20°40’03”S, 47°08’03”E; alt. 1500m; 2001; J. Hermans 3381; K!• Vakinankaratra, Antsirabe, Mt. Ibity; 20°05’07”S, 47°01’06”E; alt. 2010m; II.2003; T. Andriamihajarivo 95; P[P00692192]!, MO[ MO3022646 ]!, TAN!• Haute Matsiatra, Ambalavao, réserve d’Anja; 21°50’10”S, 46°50’58”E; alt. 1171m; 22.III.2006; J.N. Labat 3656; P[P00533700]!• Vakinankaratra, Mt. Ibity, Ankobohy; 20°04’19”S, 47°00’33”E; II.2007; T. Ranarivelo etal. 463; K!• Ihorombe, commune Iarintsena, Anja, mont Amboalandy; 21°51’08”S, 46°50’18”E; 1173m; 16.III.2010; Rajaonary etal. 70; P[P06773265]!• Haute Matsiatra, Ambalavao, Iarintsana, Anja; 21°50’44”S, 46°49’55”E; alt. 1275m; 16.III.2010; Razafindraibe etal. 290; P[P06796928]!, TAN!• Ihorombe, inselberg Bonnet du pape, fokontany Ifanadana, commune Zazafotsy, district Ihosy, RN7; 22°00’23”S, 46°21’42”E; III.2010, L. Ramandimbisoa etal. 119; P[P06773267]! and samples in alcohol• Amoron’i Mania, Ivato, Inselberg à 11.7km d’Ivato vers la route d’Ambatofinandrahana; 20°39’55”S, 47°07’49”E; alt. 1542m; 24.III.2010; Rakotoarivelo etal. 292; P[P06797815]!• Amoron’i Mania, Ambatofinandrahana, Itremo; 20°34’36”S, 46°35’06”E; alt. 1556m; 3.V.2010; Ralimanana RLI1450; K[K000751569]!, P[P00939974]!, TAN!• Amoron’i Mania, Ambatofinandrahana, Itremo, Ambatoantrano; 20°33’55”S, 46°34’54”E; 4.V.2010; S.E. Rakotoarisoa etal. 695; K[K000664251]!, P[P00939969]!, MO!, TAN!, TEF!, SNGF!• Analamanga, Carion, inselberg Maharidaza; 47°43’39”S, 18°53’30”E; alt. 1501m; 6.III.2011; N. Ravololomanana 222; P[P06796775]!, MO[ MO3022632 ]!, TAN!• Amoron’i Mania, Antoetra, on inselberg; 20°46’27”S, 47°19’16”E; 14.II.2015; alt. 1676m; A.& C. Sieder etal. L6803; WU! spirit material• Amoron’i Mania, Antoetra, inselberg; 20°46’27”S, 47°19’16”E; V.2018; J. Hermans 8255; K!. noTes The leaves are generally basal and alternate, but this is not the case for all specimens, e.g. Bosser 2560 (P) from the north, which is in a disjunct area. This needs to be clarified from fresh material since plants with unusual patterns of leaves often occur in marshy soils (e.g. in Ambondrombe and Itremo). Specimen H. Perrier 113 cited by Schlechter in the protologue is an alternative numbering of H. Perrier 11386, which was sent to Schlechter in Berlin and returned, so there is no doubt that this is the holotype. See also: Schlechter (1930: t.10 n°39), Perrier (1939: 39, as Benthamia cinnabarina), Cribb& Hermans (2009: 46, as Benthamia cinnabarina, photograph is Benthamia africana), Bosser& Lecoufle (2011: 175, as Benthamia cinnabarina), Hervouet (2018: 163). descripTion Erect terrestrial herb, 15-35cm tall. Tubers many, oblong. Stem 1.5-3mm in diameter, with 4-5 leaves, basal or up to the middle, a little higher carrying a few sheaths gradually decreasing in size. Leaves, erect, alternate, linear or lanceolatelinear, 6-12cm× 5-12mm, acute. Rachis 5-16cm long, several-flowered, but not very densely flowered, interrupted and spiral. Floral bracts narrow, lanceolate-linear, normally longer than the ovary, 6.5-12× 1.1-2.5mm. Ovary 2.5-7.5× 0.8-2mm. Flowers small, 4mm without the ovary; sepals narrowly oblong, obtuse, 3-veined, dorsal sepal 3.5-4.9× 0.6-1.5mm, lateral sepals 2.7-4.8× 0.3-1.2mm; petals of the same length as sepals, wider, 3-5× 0.9-1.5mm, thickened and opaque in the upper half, a little expanded towards the middle of the front edge; lip oblong, 3.6-5.1× 1.9-3.6mm, trilobed, in the upper half, the lateral lobes very obtuse and about 2/3rd the size of the midlobe which is wider and around 1.5mm long; spur scrotiform and a little flattened, as broad as high, 0.8-1.2× 0.4-1.2mm. Column 0.7-1.2× 0.7-1mm, anther shortly apiculate; auricles broadly spathulate, slightly shorter than the anther; rostellum with two very small lateral lobes and a larger obtuse midlobe, 0.1-0.2mm long, upwards. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 44. — Distribution map of Benthamia flavida Schltr. 204 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. Benthamia glaberrima (Ridl.) H.Perrier (Fig. 45) Bulletin de la Société botanique de France 81 (Perrier 1934: 28).— Holothrix glaberrimaRidl., Journal of the Linnean Society, Botany, vol.22: 125 (Ridley 1886); Perrier (1939: 22). — Peristylus glaberrima(Ridl.) Rolfe, Orchid Review 7 (73): 4 (Rolfe 1899).— Platanthera glaberrima(Ridl.) Kraenzl., Orchidacearum Genera et Species, vol.1: 609 (Kraenzlin 1899).— Habenaria glaberrima(Ridl.) Schltr., Beihefte zum Botanischen Centralblatt 33: 405 (Schlechter 1915).— Rolfeella glaberrima(Ridl.) Schltr., Repertorium Specierum Novarum Regni Vegetabilis 33: 18 (Schlechter 1924). Habenaria deceptrix Baron ex Ridley, nomen nudum in Baron’s “Compendium des plantes Malgaches”. Revue de Madagascar, vol.3, no.10: 722 (Baron 1901). T ype maTerial . — Madagascar• Analamanga, Ambatovory; 18°54’S, 47°42’E; III.1882; J.T. Fox 23; holotype: K[K000415546]!. e Tymology . — From Latin “glaber”, glabrous, with superlative ending “rima” thus completely smooth, without hairs. phenology. — January to May. d isTribuTion and ecology . — Restricted to the central mountains and plateaux of Madagascar, from 1500 to 2250m, in dry montane grasslands, also rocky outcrops and sometimes humid places (Fig. 46). conservaTion. — Widespread in Madagascar, this species is considered of Least Concern (LC). However, it was considered Near Threatened (NT) by IUCN in 2015 (https://doi.org/10.2305/ IUCN.UK.2015-2.RLTS.T68002690A68015027.en). addiTional specimens examined. — Madagascar• Imerina; R. Baron 849; BM!• sine loc.; Deans Cowan 23; BM!, B!, M000032247!• sine loc.; B. Pettersson etal. 221; K! flowers in spirit [K55124.000]• Montagnes d’Amoron Kay, est de l’Imerina; alt. 1600-1900m; H. Rusillon 166; G!• Ambatovory; III.1886; J.T. Fox 23; K! ex LDS• north west, IX.1887; R. Baron 5240; K! mounted with G.F. Scott Elliot 1944 and Baron 849• Near Mahabo; 19°07’S, 46°58’E; received IV.1890; G.F. Scott Elliot 1944; K!• Vakinankaratra, cime du Famoizankova près d’Antsirabe; 19°39’32”S, 47°03’48”E; alt. 2200m; V.1912; H. Perrier 11385; P[P00094511]!• Vakinankaratra, pente du mont Ibity au sud d’Antsirabe; alt. 1800-2200m; 20°05’44”S, 47°00’12”E; II.1914; H. Perrier 8087; P[P00094510 the plant on the right is Benthamia rostrata]!• Haute Matsiatra, massif d’Andringitra; alt. 2000m; I.1922; H. Perrier 14326; P[P00094512]!• Vakinankaratra, flanc Est du Tsiafajavona; 19°19’58”S, 47°16’08”E; alt. 2000m; 1928; H. Perrier 13518; P[P00094508, P00094509]!• Analamanga, massif de l’Angavo près d’Ankazobe; 18°14’S, 47°02’E; 10.III.1930; R. Decary 7325; P[P00094505]!• Analamanga, Tampoketsa au nord d’Ankazobe; 18°12’56”S, 47°17’17”E; 12.III.1930; R. Decary 7453; P[P00094504]!, TAN• Analamanga, massif de l’Angavo près d’Ankazobe; 18°14’S, 47°02’E; 16.III.1930; R. Decary 7600; P[P00094506]!• Amoron’i Mania, montagnes à l’ouest d’Itremo; 20°34’12”S, 46°34’55”E; alt. 1500-1700m; 1955; H. Humbert 30055; P[P00692284 all but second specimen]!, P[P00692285 specimen on the right only]!, P[P00692286]!, TAN!• Vakinankaratra, environs de Faratsiho (face ouest de l’Ankaratra); 19°24’S, 46°57’E; alt. 1800m; II.1957; J. Bosser 10864; P[P00094507]!, TAN!• Analamanga, Iarinandriana, 40km from Antananarivo; 19°10’S, 47°30’E; IV.1957; J. Bosser 11061; TAN!• Vakinankaratra, massif de l’Ibity; 20°10’S, 47°01’E; 19.III.1960; M. Peltier 2112; P[P00692291]!• Amoron’i Mania, Ambositra, Anjoma; III.1960; J.& M. Peltier 2173; TAN! plant on the right• Amoron’i Mania, route Ambositra à Ambatofinandrahana; 20°40’03”S, 47°08’03”E; III.1960; M. Keraudren 148; P[P00692288, plant in the middle only]!• Haute Matsiatra, route Ambositra à Ambatofiterahana à 99km de l’embranchement d’Ivato; 20°40’03”S, 47°08’03”E; III.1960; M. Keraudren 157; P[P00692289]!• Vakinankaratra, col des Tapias, 45km avant Ambositra; 20°14’34”S, 47°06’04”E; IV.1964; J. Bosser 19597; P[P00692258]!• Haute Matsiatra, Andringitra Forest Reserve, along path to Pic Boby; 22°11’S, 46°55’E; alt. 1900-2050m; 23.III.1989; P. Goldblatt etal. 8965; K!, MO[MO3011473]!• Vakinankaratra, Ankaratra Massif W of Ambatolampy; 19°20’S, 47°16’E; III.1989; P. Goldblatt etal. 9011; K!, MO[MO4314444]!• Analamanga, c.85km in direction Ankazobe; 18°15’59”S, 47°10’53”E;.XII.1998; W. Rauh R21598; HEID[HEID744389]!• Vakinankaratra, Ibity, Mt. Kiboy; 20°05’44”S, 47°00’12”E; alt. 1600m; 1999; J. Hermans 2410; K!• Haute Matsiatra, Ambondrombe; alt. 2000m; 1999; J. Hermans 4406; K!• Amoron’i Mania, Ambatofinandrahana, W of Itremo; 20°34’12”S, 46°34’55”E; 2002; J. Hermans 3382; K!• Vakinanakaratra, Ibity Massif; 20°04’10”S, 47°00’16”E; alt. 1700m; II.2003; G.E. Schatz etal. 4097; TAN!, MO!• Vakinanakaratra, Ibity Massif; alt. 1700m; 17.II.2003; G.E. Schatz etal. 4139; MO[MO3022642]!• Haute Matsiatra, Andringitra; alt. 1987m; I.2005; G. Fischer etal. FS2031-2005; WU0848511!• Vakinankaratra, Mt. Ibity, Ankobohy; 20°04’30”S, 47°00’47”E; II.2007; T.Ranarivelo etal. 465; TAN!, K!• fig. 45. — Benthamia glaberrima (Ridl.) H.Perrier: Mount Ibity, Madagascar, 5 March 2009. Photograph by Jean-Michel Hervouet. 205 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) Amoron’i Mania, Ambatofinandrahana, Itremo massif, about 25km from Itremo village; 20°33’42”S, 46°33’49”E; alt. 1647m; II.2010; L.R. Rajaovelona etal. RJL145; K!, TAN!• Haute Matsiatra, RN7 PK420.7, bifurcation à droite vers la commune d’Andoharanomaitso; 21°30’07”S, 46°58’43”E; alt. 1293m; 14.III.2010; L. Ramandimbisoa etal. 58; P[P06773266]!. noTes The species can be easily recognised in herbaria by its basal leaves followed by a series of prominent long acuminate sheaths, then by its dense rachis and also by its long and filiform spur. See also: Cribb& Hermans (2009: 40), Bosser& Lecoufle (2011: 176), Hervouet (2018: 164). descripTion Erect terrestrial herb, 30-60cm tall. Tubers 4-5, bunched, oblong-fusiform; Stem 2-3mm in diameter, with a single radical sheathing leaf, higher covered by 6-11 acuminate sheaths 20-35mm long. Leaf narrowly lanceolate-acute, 8.516× 0.9-1.4cm, very gradually narrowed towards the base, 5-veined. Rachis very dense, with 20-55 flowers, unilateral, 5-19cm long. Floral bracts lanceolate, longly acuminate, 10-20× 2.5-5mm, normally longer than the flower. Ovary twisted, 5.5-10× 1-2.5mm. Flowers small, sulphur yellow; all sepals narrowly ligulate, a little fused to the petals at the base, dorsal sepal 5.1-7.5× 0.6-1.3mm, lateral sepals 4.5-8× 0.5-1.8mm; petals lanceolate, 5.2-8× 0.6-2.1mm, slightly longer than the sepals; lip oval in outline, 5.2-7.5× 3-4.5mm, obviously trilobed, a little fused with the sepals at the extreme base; midlobe oval-acute, the lateral lobes much smaller; spur filiform, pendent, 7.2-10× 0.5-1.2mm, a little longer than the ovary. Column straight, 1.1-2× 0.9-1.5mm; anther obtuse or subapiculate; auricles small, rounded, pollinia claviform with a very short caudicula; rostellum small, tridentate. Benthamia herminioides Schltr. (Figs 47; 48; 49) Repertorium Specierum Novarum Regni Vegetabilis 33 (Schlechter 1924: 27). Benthamiaherminioidessubsp.intermediaH.Perrier, Bulletin de la Société botanique de France 81 (Perrier 1934: 37). — Type: Madagascar• Diana, Mt. Tsaratanana; 14°01’17”S, 48°58’00”E; alt. 2000m; IV.1924; H. Perrier 16479; holotype P[P00094515]!, syn.nov. Type maTerial. — Madagascar• Vakinankaratra, mont Ibity au sud d’Antsirabe; 20°06’25”S, 46°59’51”E; alt. 2000m,.II.1914; H. Perrier 8103 (XIX); holotype: P[P00094513]!. eTymology. — Named for its similarity with species in the genus Herminium, according to Schlechter. phenology. — February to April. d isTribuTion and ecology . — Madagascar. Mainly in Analamanga, Alaotra-Mangoro, Haute Matsiatra and Amoron’i Mania regions, but with one locality in Sava region, Mt. Marojejy. In montane grasslands or marshes from 1000 to 2100m (Fig. 50). Benthamia herminioides (subsp. herminioides) and Benthamia herminioides subsp. intermedia were listed as occurring in La Réunion by Szelengowicz& Tamon (2013: 244-245). However, there is no voucher for either and the photographs of Benthamia herminioides subsp. intermedia show a young Benthamia latisatis. conservaTion. — This species seems widespread in Madagascar but is only known from fewer than 10 localities which do not cover more than 2 000km², with a decline of the habitat quality, it is therefore considered Vulnerable (VU) according to criterion B2. a ddiTional specimens examined . — Madagascar• Sava, sommet oriental du massif de Marojejy; III-IV.1939; H. Humbert etal. 23880; P[P00334789]!• Haute Matsiatra, Ambondrombe; 21°52’30”S, 47°15’37”E; alt. 1500m; 11.IV.1941; P. Boiteau 4840; P[P00692304]!• Sava, pentes orientales du massif de Marojejy; 24.III.1949; alt. 1000m; H. Humbert 23638; P[P00334791]!• Sava, sommet oriental du massif de Marojejy; 14°26’57”S, 49°43’57”E; 26.III-2.IV.1949; H. Humbert etal. 23879; P[P00334790]!, G!• Sava, sommet oriental du massif du Marojejy; 14°25’S, 49°43’E; alt. 1850-2137m; 17-20.XII.1949; H. Humbert 22717; P[P00692985, P00692988, P01778630]!, G!, K!• Atsinanana, Brickaville; VI.1959; W. Rauh 158; HEID[HEID713727]!• Haute Matsiatra, Andringitra; IV.1964, J. Bosser 19643; P[P00692263]!• Analamanga, SE Carion, near the granite dome of Angavokely; alt. 1750-1780m; 27.II.1988; B. Pettersson etal. 203; P[P02115315 flowers in bud]!, K[spirit K55118.000]!• Sava, Réserve naturelle de Marojejy, the trail to the summit of Marojejy Est; 14°26’S, 49°13’E; alt. 1900-2133m; 15.II.1989; J.S. Miller etal. 4150; P[P00754986]!, 45°E 15°S 20° 25° 50° 88,427 200 km fig. 46. — Distribution map of Benthamia glaberrima (Ridl.) H.Perrier. 206 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. MO[MO3022650]!, TAN!• Sava, Réserve naturelle de Marojejy; alt. 1900-2133m; 15.II.1989; J.S. Miller etal. 4179; MO[MO3022625]!• Haute Matsiatra, Ranomafana National Park, parcelle n°3; alt. 1374m; 21°20’03”S, 47°24’49”E; III.1992; S.T. Malcomber etal. 1364; P[P00754904]!, K, MO[MO3022611]!, TAN!• Haute Matsiatra, Ranomafana National Park, vicinity of Ampasina; 21°11’S, 47°25’E; 10.III.1995; D. Turk etal. 727; P[P00094582]!, K!, K[spirit K72250.000]!, WAG[WAG.1924442]!• Haute Matsiatra, Ambondrombe; alt. 1875m; 9.III.2009; photos Jean-Michel Hervouet• Alaotra-Mangoro, Ambatovy; alt. 1120m; 25.III.2011; T. Stévart etal. 4115; BRLU!, MO[MO3060441]!, K!, TAN!, spirit• Amoron’i Mania, Antoetra, on Inselberg; 20°46’27”S, 47°19’16”E; alt. 1676m; II.2015; A.& C. Sieder etal. L6811& J. Hermans 8076; K!, WU!, and spirit material• Alaotra-Mangoro, Andasibe, Iaroka forest; 29.IX.2019; J. Hermans 835;5 K!. noTes The flowers are amongst the smallest in the genus. They generally have a green calyx and white corolla, but the petals can be also greenish (original subsp. intermedia) or sulphur yellow (Marojejy and Ambondrombe, with plants with white corolla also present). The differences given by Perrier to establish the subspecies intermedia fall within the variability of the species fig. 48. — Benthamia herminioides Schltr.: Mount Ibity, Madagascar, 1 April 2018. Photograph by Jean-Michel Hervouet. fig. 47. — Benthamia herminioides Schltr.: Mount Ibity, Madagascar, 1 April 2018. Photograph by Jean-Michel Hervouet. fig. 49 . — Benthamia herminioides Schltr.: Mount Ambondrombe, Madagascar, 11 April 2018. Yellow flowers. Photograph by Jean-Michel Hervouet. 207 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) itself, but he noted on the herbarium sheet that the flowers are greenish. Perrier also refers to Benthamia herminioides subsp. typica H.Perrier (Perrier 1934: 36). This name is not validly published (article 24.3 of the International Code of Nomenclature). With subsp. typica Perrier was referring to the autonym name, i.e. subsp. herminioides. See also: Schlechter (1930: t.10 n°40), Perrier (1939: 36), Cribb& Hermans (2009: 48), Hervouet (2018: 165). descripTion Small or slender terrestrial herb, 20-70cm tall. Tubers 2-3, 15-25× 3-4mm. Stem green, 1.5-3mm in diameter, with 1-2 tubular sheaths at the base, 15-32cm× 3mm, and leaves in the lower half. Leaves 2-3, elliptic-lanceolate or lanceolate, 3-10× 1.5-2.5cm, shortly acuminate, very gradually and slightly narrowed towards the base. Rachis 5-25cm, laxly 15-45 flowered, flowers fairly close together on stem, 3-6mm apart, unilateral. Floral bracts lanceolate-acuminate, 2.2-6.5× 0.7-1.5mm, variable, generally about the same length as the ovary but sometimes shorter or longer, folded around the base of the ovary and apiculate. Ovary 2.5-5× 0.9-1.5mm, barrelshaped with 3 distinct ridges. Flowers with green calyx, and petals generally white, sometimes yellow or greenish, small. Overall flower size 3mm wide, 2mm high, 2.5mm deep; sepals oblong, obtuse, 1-veined, 1.8-2.75× 0.4-1mm, dorsal sepal forming a hood over the column together with the petals, lateral sepals concave or navicular, spreading and joined slightly at the base with the dorsal sepal and petals; petals 1-veined very obtuse, gradually thickened towards the apex, 1.7-2× 0.6-1mm, a little shorter than the sepals; lip rhomboid in outline, 1.1-2.7× 0.9-1.2mm, base a little concave, trilobed in the upper third, the midlobe triangular-acute, thick, twice as long as the lateral lobes which are obtuse and thickened at the apex, erect to inwards folding, embedded into the petals; spur very short, subglobose, saccate or a little claviform, 0.6-0.8× 0.3-0.8. Column 0.5-0.7× 0.5-0.7mm, anther apiculate; auricles reaching half the anther length or more, slightly spathulate; rostellum with the midlobe triangularacute; stigmata in a cushion, free only at the apex; pollen grains minute, around 0.2mm. Benthamia humbertii H.Perrier (Figs 51; 52) Bulletin de la Société botanique de France 81: 35 (Perrier 1934). Type maTerial. — Madagascar• Ihorombe, pic d’Ivohibe; 22°30’25”S, 46°58’15”E; alt. 1500-2000m; 5-11.XI.1924; H.Humbert 3287; lectotype: P[P00094517]!, here designated; isolectotype: P[P00094518]!, B[B100365917]!, one of the few specimens saved from the destruction of the Berlin herbarium at the end of second World War, G!, K[K000415555]!, TAN[TAN000409]!. e Tymology . — Named for Henri Humbert (1887-1967) who collected the type material. phenology. — October and November. disTribuTion and ecology. — Madagascar. In the southern half of the country, in montane grasslands from 1100 to 2000m (Fig. 53). conservaTion. — So far only known from fewer than 10 localities only, with an AOO less than 2000km², and habitat threatened by frequent bushfires, it is considered Vulnerable (VU), despite a very large EOO according to criterion B2. It was considered EN by IUCN in 2015 (https://doi.org/10.2305/IUCN.UK.2015-2.RLTS. T70102654A70155404.en). addiTional specimens examined. — Madagascar• Haute Matsiatra, Andringitra, Andrianony, cirque de Manjarivolo; alt. 1800m; 1.XI.1970; J.L. Guillaumet 3451; TAN!• Anosy, massif du Beampingaratra, col de Vohipaha; 24°32’S, 46°53’E; alt. 11001400; 16.XI.1928; H. Humbert 6658; syntypes: P[P00094519, P00094520]!• Atsimo-Atsinanana, bassin de l’Itomampy, mont Papanga près de Befotaka; 23°51’S, 46°57’E; alt. 1300-1700m; 2-3. XII.1928; H. Humbert 6906 bis; syntype: P[P00094521]!• Haute Matsiatra, Andrambovato, près de Fianarantsoa; 21°31’S, 47°25’E; XI.1963; J. Bosser 18306; P[P00692242, P00692243]!• Amoron’i Mania, Antoetra; 20°46’27”S, 47°19’16”E; photographs by Anton Sieder• Haute Matsiatra, Ankazomivady; 20°46’22”S, 47°10’48”E; 1721m; 12.VIII.2017; photographs by Hasimbola Rakotobe• Haute Matsiatra, Ambondrombe; 21°52’30”S, 47°15’37”E; alt. 1875m; 29.X.2022; photographs by Jean-Michel Hervouet. 45°E 15°S 20° 25° 50° 88,427 200 km fig. 50. — Distribution map of Benthamia herminioides Schltr. 208 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. noTes One of the largest Benthamia species in Madagascar. It closely resembles B.latisatis from the Mascarenes but is easily distinguished by the leaves (alternate amplexicaul, 6-8cm long vs12-30cm long, higher on the stem), by the sepals and petals which are 1-veined (3-veined in B.latisatis), the bigger lip with a crest, the non-apiculate anther and the indented rostellum in the middle. See also: Perrier (1939: 36), Cribb& Hermans (2009: 49), Bosser& Lecoufle (2011: 177, the photograph shows Benthamia elata). descripTion Robust terrestrial herb, 50-70cm tall. Tubers many, fusiform. Stem thick, 5-10mm in diameter, carrying only 2-3 very lax sheaths in its lower part, of which the 2 outer ones are black, then 4-8 developed alternate amplexicaul leaves above the middle and higher again a few short sheaths becoming gradually similar to the floral bracts. Leaves rather thick, broadly oval-lanceolate, 6-8× 2-4cm, acuminate-acute. Rachis elongate, 15-18cm, cylindrical, 2-2.4cm in diameter, dense, with 35-100 flowers. Floral bracts narrowly lanceolate, concave at base, acuminate-acute, 1-veined, 10-14× 3mm, the lower ones a little longer than the ovary and flower, the upper ones about as long. Ovary twisted, 7-20× 1.5-3mm, reddish. Flowers greenish yellow, thick and short, 4-5mm without the ovary; sepals 1-veined, dorsal sepal broadly elliptic, 4-5× 1.7-2mm, lateral sepals concave, elliptic to subtriangular or even navicular, very obtuse to subacute, 4.2-4.5× 1.7-1.8mm; petals elliptic, asymmetrical, 3.5-4× 1.5-2mm; lip broadly obovate, 4.5-5× 3-4.5mm, thick, trilobed in the apex, the lateral lobes short, triangular with obtuse apex, 0.40.5mm long, the midlobe truncate, obtuse, 1-1.5× 1mm, thickened; spur scrotiform, compressed, wider than long, 1-2× 1.8-2mm. Column 1-1.2× 1-1.2mm, anther obtuse, not or hardly apiculate; auricles slightly spathulate, equalling half the length of the anther or less; rostellum trilobed, with fig. 51 . — Benthamia humbertii H.Perrier: Mount Ambondrombe, Madagascar, 11 April 2018. Photograph by Jean-Michel Hervouet. fig. 52 . — Benthamia humbertii H.Perrier: Mount Ambondrombe, Madagascar, 29 October 2022. Photograph by Jean-Michel Hervouet. 209 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) two small horizontal lateral lobes, midlobe toothed, short and thick, slightly emarginate in the middle; stigma short, forming two distinct small, rounded masses. Benthamia lakatoensis Descourv.& Hervouet, sp.nov. (Figs 54; 55; 56; 57) diagnosis. — Similar to Benthamia nivea but differing by the shape of the leaves, elliptic with apex rounded (vselliptic with apex acute in Benthamia nivea), and in the smaller flowers, 4-6mm (vs5-10mm for Benthamia nivea), forming a straight tube in their lower third (vsforming a curved tube), with petals not or hardly asymmetrical (vscompletely asymmetrical), lateral lobes of lip divergent (vsparallel when flattened), base of lip narrow, hardly wider than the midlobe (vsbase of lip twice wider than the midlobe), the dorsal sepal and petals ending in a single vertical plane in fresh specimens (vsflowers not opening widely). The auricles are nearly twice as long and the column is much less apiculate, the spur is half the length. The flowering season is August-November (vsFebruary-April). T ype maTerial . — Madagascar• Alaotra-Mangoro, route de Lakato; 19°11’30”S, 48°26’00”E; XI.1963; J. Bosser 18340; holotype: P[P00094562]!. eTymology. — Referring to «Lakato», a village near Andasibe in the Alaotra-Mangoro region, where the type of the plant was first found. phenology. — August to November. disTribuTion and ecology. — Madagascar. Only few plants have been recorded so far, from c.725 to 1200m, in medium altitude moist evergreen forest, from north to south. The type was collected along the road between the Route Nationale 2 and Lakato, a region that has suffered from severe deforestation in the recent years, it is very unlikely that it still exists there. A specimen has been located in Zahamena forest reserve, another one in Ranomafana national park, and more recently another from Tsaratanana area (Fig. 56). conservaTion. — Apart from the specimens found recently, this species has been collected in two localities only, one of them (road to Lakato) being heavily deforested. It can be considered Critically endangered (CR) according to criterion B2, with AOO less than 10km², severely fragmented and in continuing decline. addiTional specimens examined. — Madagascar• AlaotraMangoro, massif de l’Andrangovalo au sud-est du lac Alaotra, réserve de Zahamena; 17°40’S, 48°45’E; alt. 1200m; X.1937; 45°E 15°S 20° 25° 50° 88,427 200 km fig. 53. — Distribution map of Benthamia humbertii H.Perrier. fig. 54 . — Benthamia lakatoensis Descourv. & Hervouet, sp. nov., Lakato road, Andasibe region, Madagascar, September 1963. Photograph by Jean Bosser. 216 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. 49: 23 (1899).— Peristylus madagascariensis(Rolfe) Rolfe, Orchid Review 7 (73): 4 (Rolfe 1899).— Platanthera madagascariensis(Rolfe) Kraenzl., Orchidacearum Genera et Species1: 609 (Kraenzlin 1899). Peristylus macropetalusFinet, Notulae Systematicae 2: 24 (Finet 1911).— Type: sine loc.; culture in P; XII.1910, M.F. Geay s.n.; holotype: P[P00334758]!. A B D C fig. 63. — Benthamia litoralis Hervouet & Descourv., sp. nov.: drawing by Marie-Aimée Allard: A, habit; B, side view of flower and ovary; C, front view of flower; D, part of rachis with swollen fruits. Scale bars: A, 1 cm; B, C, 1 mm; D, 0.5 cm. 217 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) T ype maTerial . — Madagascar• Anosy, near Fort-Dauphin; 25°02’S, 46°59’E; G.F. Scott Elliot 2643; lectotype: K!, selected by Hermans etal. (2007: 73); isolectotype: P[P00094529]!. eTymology. — “Madagascariensis”, from Madagascar. phenology. — February to August. disTribuTion and ecology. — Madagascar. Only in the Anosy region in the north of Fort-Dauphin, except for one possibly dubious record from the south of Antananarivo. In coastal marshes (Fig. 65). conservaTion. — This species is widespread in marshland in the Fort-Dauphin region but is threatened by the mining activities in the coastal sands of this region, so it should be considered now as Near Threatened (NT). addiTional specimens examined. — Madagascar• Atsimo-Atsinanana, plains near Vaingandrano; 23°20’50”S, 47°36’50”E; 6.III; G.F. Scott Elliot 2257; K[K000415554], BM[BM000076235]!• sine loc.; 1897; L.D.M. Catat 4326; P[P00334762, P00334763, P00334764]!• Anosy, région de Fort-Dauphin; 1897; G. Paroisse 28; P[P00334765, P00334766]!• Analamanga, damp area near Manjakandriana; 18°54’37”S, 47°48’29”E; VI.1905; C.d’Alleizette 7066b; L[L1487718]!• Anosy, Manantenina; 10.VI.1926; R. Decary 3876; P[P00334770]!• Anosy, Fort-Dauphin; 24.VI.1926; R. Decary 4087; P[P00334773]!• Anosy, Fort-Dauphin; 26.VI.1926; R. Decary 3982; P[P00334771, P00334772]!• Anosy, Fort-Dauphin; 1.V.1932; R. Decary 9793; P[P00334774]!• Anosy, FortDauphin, Pic Saint-Louis; alt. 300m; 3.VII.1932; R. Decary 9973; P[P00334776]!, BM[BM000034591!, BM000034592]!, K!, TAN!• Anosy, Fort-Dauphin; 30.VIII.1932; R. Decary 9841; P[P00334775]!• Anosy, Soanierana, district de Fort-Dauphin; 25°00’S, 46°53’E; 16.X.1932; R. Decary 10776; P[P00334777]!• Anosy, Fort-Dauphin; 21.VIII.1937; Poisson 2635; TAN!• Anosy, Fort-Dauphin; 6.VI.1939; R. Decary 14129; P[P00334778]!• Anosy, vallée du Mandrare, affluent de la Manampanihy, montagne au sud de Tanandava; 24°26’00”S, 47°00’00”E; alt. 850m; 11-16.III.1947; H. Humbert 20555; P[P00334759]!• Anosy, bassin de la Manampanihy, mont Vohimavo au nord d’Ampasimena; 24°22’S, 47°10’E; alt. 830m; 27-28.III.1947; H. Humbert 20720; P[P00334760, P00334761]!, K[K000718265]!, TAN!, G!• Anosy, RN 11, Canton Enomihila, district de Manantenina; 24°17’S, 47°19’E; 28.V.1953; D. Ramarokoto 5432RN; P[P00334767, P00326093]!• Anosy, Fort-Dauphin; 10.IV.1956; Raonarivelo 85111; P[P00692460]!• Anosy, Fort-Dauphin, Pic Saint-Louis trail; 25°00’29”S, 46°57’42”E; 12.V.1968; D. Seligson 594; P[P00334768 noted ”purple flowers” so questionable]!• Anosy, route de Manantenina, environs de Fort-Dauphin; 4.IV.1969; Y. Veyret 1041; P[P00334769]!• Anosy, Fort Dauphin, Pic Saint-Louis; VII.1969; F. Friedmann 181; TAN!• Anosy, Sainte-Luce; 23.IV.1969; W. Rauh R22146; HEID[HEID740744, HEID740745, HEID740746]!• AtsimoAndrefana, Tuléar, route entre Mahatalaky et Manafiafy, 2km à l’ouest du village d’Ambandrika; 24°46’S, 47°12’E; alt. 0-10m; 13.VI.1994; A. Randrianasolo 367; P[P02103231]!, K!, MO[MO3022641]!, TAN!• Anosy, Fort-Dauphin area; 24°47’30”S, 47°05’00”E; 1999; J. Hermans 4242; K!• Anosy, near Mahatalaky, Tolagnaro province; 2001; J. Hermans 5351; K!• Anosy, Iabakoho, Antsotso Avaratra, Ivohibe; 24°34’10”S, 47°12’26”E; alt. 105m; V.2006; C. Birkinshaw etal. 1639; K[K000395392]!, P[P00593236]!, MO!, TAN!• Anosy, Mahatalaky; alt. 10m; 5.VIII.2012; N. Rakotonirina 783; MO[MO3022627]!. noTes According to Rolfe, Holothrix madagascariensis (Rolfe 1893: 55) has the habit of Habenaria glaberrima Ridley but is readily distinguished by the laxer rachis with larger flowers, the much larger lateral lobes of the lip, and the very short saccate spur. In Rolfe’s publication the species is illustrated on plate XII, which is mislabelled Habenaria elliotii. The specimen C.d’Alleizette 7066b in L (L1487718) was collected close to Antananarivo, very far from the distribution range given by other records. We consider the locality dubious. Note that there is also a second specimen C.d’Alleizette 7066b in L (L1487725), of Benthamia monophylla. A specimen in P (D. Seligson 594) has a note saying that the flowers are purple. This is questionable since all specimens seen in the wild so far have white flowers. See also: Cribb& Hermans (2009: 47), Bosser& Lecoufle (2011: 178), Hervouet (2018: 166). descripTion Erect terrestrial herb, 25-50cm tall. Tubers 1-2, ovoid, roots wiry, c.1mm in diameter, up to 20cm long. Stem 2-5mm in diameter, with leaves on the lower half, higher up with 3-4 narrow bladed sheaths. Leaves 3-6, linear, 8-15cm× 3-5mm. Rachis 5-10cm, with 15-40 flowers rather densely distributed. Floral bracts narrow, very acute, 7-10× 1-2mm. Ovary fusiform, straight, 7-12× 1-2mm. Flowers forming a green tube at the base, secund, white; sepals 1-veined, clearly linear, dorsal sepal 5-10.5× 0.6-2.2mm, the lateral sepals a little shorter, 5-10× 0.92mm, adnate to the petals and to the lip; petals longer and wider than sepals, 5.2-10× 0.3-2mm, oblanceolatelinear, rounded at the tip, 1-veined; lip 8-10× 3-7mm, narrow, divided into three lobes in the upper quarter; lobes equal, c.3mm long, 1-1.5mm wide; spur very shortly saccate, 0.8-1.1× 0.6-1mm, as wide at the base as high. Column 1.2-2.8× 1-2mm, anther apiculate; auricles broadly oblong, as long as the anther; rostellum with the median tooth twice as long as the lateral, obtuse and gutter-shaped at the apex. table 1. — Differences between Benthamia africana, B. elata and B. litoralis sp. nov. Benthamia africana Benthamia elata Benthamia litoralis sp. nov. Leaves linear-lanceolate, basal or cauline in lower part of inflorescence Leaves lanceolate, nearly up to the rachis Leaves linear strap-like, mostly basal, leaving a long gap between leaves and rachis Rachis very long, 6-38 cm Rachis very long, 20-40 cm Rachis short, 4.5-17 cm Plant medium or large (20-62 cm) Plant large (25-100 cm) Plant small (20-41 cm) Flowers small (2-5 mm) Flowers large (5 mm) Flowers small (3 mm) Lip lateral lobes very small and obtuse Lip lateral lobes from 1/3 to size of central lobe Lip lateral lobes acute, less than 1/3 of central lobe Sheaths 2-3 below the leaves Sheaths 2 below the leaves Sheaths absent below the leaves 218 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. Benthamia majoriflora H.Perrier (Fig. 66; 67) Notulae Systematicae 14 (2): 140 (Perrier 1951). T ype maTerial . — Madagascar• Sava, pentes occidentales du massif de Marojejy, district d’Andapa; 14°26’05”S, 49°45’38”E; 28.XI06.XII.1948; H. Humbert etal. 22224; holotype: P[P00094531]!. eTymology. — From Latin “major”, greater, and “florus”, flowered. phenology. — From October to December. disTribuTion and ecology. — Madagascar. So far only known in the slopes of Mt. Marojejy in the north-east, in Sava region. Epiphytic, often on Ravenea palms, from 500m to 1500m (Fig. 68). Specimen Iharivolana etal. IH021 in Geneva (G[G00409570]) is possibly a withered Benthamia majoriflora from Sorata forest, Amboriala in the district of Vohemar, which would be the first record outside the Mt. Marojejy area. conservaTion. — If we exclude the specimen recently collected in the district of Vohémar, this species has been collected from a single locality along the path to the summit of Mt. Marojejy, it can be considered Critically endangered (CR) according to criterion B2, AOO less than 10km², a single location and quality of habitat on decline. addiTional specimens examined. — Madagascar• Sava, massif du Marojejy; alt. 1300m; XI.1972; P. Morat 4009; P[P00692450, P00692451]!• Sava, massif du Marojejy; alt. 700-1150m; 8.X.1988; J.S. Miller 3494; MO[MO3022624]!• Sava, massif du Marojejy; alt. 1000-1200m; 25.XI.1989; J.S. Miller 4593; MO[MO3022635]!• Sava, NW of Manantenina; 1995; J. Hermans 2751; K!• Sava, Réserve Naturelle Intégrale de Marojejy; 14°26’12”S, 49°44’30”E; alt. 1300m; 26.X.1996; M. Rakotondrainibe 3518; P[P06796735]!• Sava, Marojejy massif; alt. 1400m; 1999; J. Hermans 4408; K!• Sava, Marojejy massif; alt. 1400m; 1999; J. Hermans 5089; K!• Sava, Marojejy National Park, camp n°3 and camp n°2; 14°25’09”S, 49°44’46”E-14°25’24”S, 49°45’29”E; alt. 1280-1350m; XI.2003; H.H. Schmidt etal. 4313; K! spirit material, MO• Sava, Marojejy, between encampment Marojejya& Propithecus; 14°26’15”S, 49°45’16”E; alt. 1106m; XI.2010; L.R. Rajaovelona etal. RJL281; K[K000664873]!, TAN!. noTes A very distinctive, large epiphytic plant with white flowers. Flowers and habit are similar to those of Benthamia humbertii or B.latisatis, both terrestrial. fig. 64. — Benthamia madagascariensis (Rolfe) Schltr., Sainte-Luce Forest, near Fort-Dauphin, Madagascar, 25 February 2001. Photograph by Jean-Michel Hervouet. 45°E 15°S 20° 25° 50° 88,427 fig. 65. — Distribution map of Benthamia madagascariensis (Rolfe) Schltr. 219 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) fig. 66. — Benthamia majoriflora H.Perrier, Mount Marojejy, Madagascar, 28 October 2005. Photograph by Jean-Michel Hervouet. fig. 67. — Benthamia majoriflora H.Perrier: Mount Marojejy, Madagascar, 28 October 2005. Photograph by Jean-Michel Hervouet. 220 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. See also: Cribb& Hermans (2009: 49), Hervouet (2018: 167). descripTion Erect robust epiphytic herb, 60-100cm tall, turning black when dry, yellowish-green when alive. Tubers in bundles, very fleshy, up to 5, roots not seen. Stem 5-11mm in diameter, the lower part covered with dark red sheaths. Leaves 1-5, acutely lanceolate, 22× 4cm, in lower half or up to the rachis. Rachis 10-18cm long, 2.5-3cm wide, very dense, bearing 40-100 flowers. Floral bracts narrow, 7× 2mm, much shorter than the ovary. Ovary 12-13× 2-3mm long. Flowers 5mm long, white or very light green; sepals narrowly elliptic, obtusely narrowed, dorsal sepal 5-5.2× 1.2-2mm, lateral sepals 4-5× 1.3-2.1mm; petals asymmetric, 4.5-4.9× 1.8-2.7mm, lower margins widely expanded, base 2.5mm high; lip widely ventricose, concave, 5-6× 3.1-4.5mm, obscurely 4-veined, apex trilobed, lobes narrow, acute, midlobe thicker, with a thick callus; spur purse-shaped, 1-1.5× 0.7-1mm. Column 1-2.5× 1-1.5mm; anther acute at the top; auricles prominent, nearly as wide as the anther is high. Benthamia mascula Hervouet& Descourv., sp.nov. (Figs 69; 70; 71; 72) diagnosis. — Similar to Benthamia africana but differing by the clearly trilobed lip (vsobscurely trilobed), the absence of a callus on the lip, the larger scrotiform spur and the 2-3 leaves in the middle of the stem (vsbasal or in the lowest part of the stem), it differs from Benthamia perfecunda by the secund rachis (vsin three ranks), flowers that are not autogamous (vsautogamous), smaller leaves and a lax rachis (vsdense). Type maTerial. — France, La Réunion• Sentier du Bras Cabot, îlet de Patience; 21°06’S, 55°37’E; 7.IV.1978; J. Bosser 22460; holotype: P[P00738370]!. e Tymology . — Referring to the proportionally large scrotumshaped spur. phenology. — March to May. disTribuTion and ecology. — La Réunion and Madagascar. Widespread in La Réunion and so far only known from photographs from Ambondrombe in Madagascar, in montane grassland. From 1600 to 2300m (Fig. 73). conservaTion. — This species is not very rare in La Réunion, and has largely been overlooked, it is found in protected areas in La Réunion. It is thus considered of Least Concern (LC). In Madagascar it is provisionally considered Data Deficient (DD). addiTional specimens examined. — Madagascar• Haute Matsiatra, summit of Ambondrombe; 21°52’30”S, 47°15’37”E; alt. 1875m; 10.IV.2018; photograph by Jean-Michel Hervouet (Fig. 69). France, La Réunion• Massif de la Fournaise, Ramon crater area; 21°17’41”S, 55°42’22”E; alt. 2000m; 15.II.1971; T. Cadet 3045 and 3046; REU[REU006428, REU006429]!• Plaine des Salazes; 21°06’S, 55°32’E; alt. 2500m; 3.III.1971; T. Cadet 3134; REU[REU017471]!• Salazie plain; 21°06’S, 55°32’E; alt. 1500m; 7.III.1971; T. Cadet 3136; REU[REU006430]!• Brûlé de Saint-Denis; 20°55’S, 55°26’E; 45°E 15°S 20° 25° 50° 88,427 fig. 68. — Distribution map of Benthamia majoriflora H.Perrier. fig. 69. — Benthamia mascula Hervouet & Descourv., sp. nov., Mount Ambondrombe, Madagascar, 10 April 2018. Photograph by Jean-Michel Hervouet. 221 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) 2.V.1974; J. Bosser 21924; P[P00738536]!• Takamaka; 21°05’S, 55°38’E; 24.V.1974; J. Bosser 22050; P[P00738381]!• Sentier du Bras Cabot, îlet de Patience; 21°06’S, 55°37’E; 7.IV.1978; J. Bosser 22472; P[P00738367]!• Hauts de Sainte-Rose; 21°09’S, 55°45’E; 18.IV.1978; J. Bosser 22521; P[P00738517]!• Morne des patates à Durand; alt. 1000-1100m; 20°56’S, 55°27’E; 5.V.1978; J. Bosser 22448; P[P00738376]!• Plaine des Fougères; 21°00’15”S, 55°29’21”E; alt. 1200m; 28.V.2003; T. Pailler TP79 and TP79.2; REU[REU006442, REU007937 except plant on the right]!• L’Étang-Salé, source of La Nouvelle Bras Sec; alt. 1300m; II.2004; V. Grondin etal. 1014; CBNM!• Plaine des Palmistes, sentier Piton Bébour; 21°07’41”S, 55°33’43”E; 26.II.2004; J. Férard etal. 1009; CBNM!• Piton Rouge, Plaine des Cafres; 21°09’S, 55°34’E; alt. 1900m; 9.III.2004; T. Pailler TP162 and TP163; REU[REU006440, REU006424]!• Route du Volcan, vers Le Piton de l’Eau; 21°11’02”S, 55°40’28”E; alt. 2100-2200m; 15.III.2007; F. Martos FM88 and FM89; REU[REU007631, REU007632]!• Plaine des Palmistes, Piton des Cabris; 21°09’24”S, 55°39’03”E; alt. 1200m; 21.V.2008; F. Martos FM318 and FM319; REU[REU007893, REU007902]!• Route du Volcan, vers Le Piton de l’Eau; 21°11’02”S, 55°40’28”E; 21.V.2008; F. Martos FM348 and FM349; REU[REU007892, REU007901]!. noTes The species also seems to be in Madagascar, according to photographs from Ambondrombe, however no herbarium specimen from this island has been located so far. The position of the leaves on the stem is similar to Benthamia macra Schltr. from Madagascar but the latter has a smaller spur, different bracts (acicular and shorter than ovary in Benthamia mascula sp.nov., lanceolate acuminate and longer than ovary in Benthamia macra), the flowers are smaller (lip at most 2.5mm long vs4mm for Benthamia macra) and the rachis is denser. It is apparent on some specimens that not all the flowers are pollinated and some remain fresh late in the season, while others are fruiting on the same rachis, it is thus probably allogamous. See also: Bernet (2010: 139, as Benthamia sp. 2), Szelengowicz& Tamon (2013: 245, as Benthamia herminioides subsp. arcuata H.Perrier, photograph on the right only). descripTion Erect terrestrial herb, 25-60cm tall. Tubers 2, digitate, up to 25mm long, roots thick, starting from the junction between stem and tubers. Stem 2-5mm in diameter, with 2 basal sheaths, then leaves in the middle, then 3-4(7) acicular sterile bracts, 1-2cm long. Leaves 2-3, cauline, narrowly ovate or elliptic-lanceolate, acuminate, erect, 4-8cm× 4-12mm for the larger. Rachis unilateral, lax, 5-13cm long, with 5-10mm between successive flowers, bearing7-30(50) flowers. Floral bract nearly as long or ¾ of the length of the ovary. Ovary4-8mm long, at right angle to the rachis when mature. Flowers greenish, partly closed except the spreading lateral sepals, around 2-3mm long; dorsal sepal elliptic, concave, 1-veined, apex rounded, 1.8-2× 0.8-1.1mm; lateral sepals obliquely elliptic, concave, 1-veined, apex subacute or fig. 70 . — Benthamia mascula Hervouet & Descourv., sp. nov., Le Brûlé, La Réunion, 18 April 2018. Photograph by Jean-Michel Hervouet. fig. 71 . — Benthamia mascula Hervouet & Descourv., sp. nov., Le Brûlé, La Réunion, 18 April 2018. Photograph by Jean-Michel Hervouet. 222 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. obtuse, 1.6-2.3× 1-1.2mm; petals more or less obliquely elliptic, 1-veined, apex subacute, 1.8-2× 1mm, forming a hood with the dorsal sepal. Lip 1.5-2.5× 2-3mm, trilobed in upper third, with thick lobes, without callus, smooth and glossy, canaliculate, with the lateral lobes slightly shorter then midlobe, forming with the perianth a narrow entrance into the spur; spur markedly bilobed, scrotiform, 1mm long, nearly as wide as long, narrowed at base. Column subterete, 0.6mm long, with two basal auricles shorter than the column, rectangular, with distal margin irregular. AB C D fig. 72. — Benthamia mascula Hervouet & Descourv., sp. nov., drawing by Ludivine Longou, after photographs and dried specimens: A, habit; B, front view of flower; C, side view of flower and ovary; D, part of rachis with fresh flowers and fruits. Scale bars: A, 10 mm; B, 0.6 mm; C, 0.5 mm; D, 2 mm. 223 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) Benthamia melanopodaSchltr. (Fig. 74) Repertorium Specierum Novarum Regni Vegetabilis 33: 29 (Schlechter 1924). Type maTerial. — Madagascar• Vakinankaratra, flanc est du mont Tsiafajavona; 19°22’S, 47°19’E; alt. 2000m; III.1921; H. Perrier 13543; lectotype: P[P00094534]!, here designated; isolectotype: P[P00094535 and flowers in spirit]!. eTymology. — From the Greek “melano”, black, and “poda”, foot, due to the black sheaths at the base of the plant. phenology. — March to April. disTribuTion and ecology. — Madagascar. From Analamanga south to Anosy region. Found in montane grassland, but also in marshes at Ambohitantely in Analamanga region, from 1500m to 2000m (Fig. 75). conservaTion. — This species is currently known from fewer than five locations, with an AOO less than 500km² and a habitat threatened by bushfires, it is therefore considered as Endangered (EN) according to criterion B2. addiTional specimens examined. — Madagascar• Haute Matsiatra, au pied de l’Ambondrombe; 21°52’30”S, 47°15’37”E; alt. 1200m; 10.IV.1941; P. Boiteau 4559; P[P00692298]!• Anosy, sommet de la Menakompy à l’est d’Ampandrandava; 24°07’S, 45°48’E; alt. 1100m; IV.1943; A. Seyrig 658; P[P00094533]!; det. Benthamia macra by Perrier)• Vakinankaratra, Manjakatompo; 1.V.1951; R. Benoist 909; P[P00692220]!• Haute Matsiatra, Andringitra, versant du Vohidray; alt. 1500m; 13 .I.1971; J.L. Guillaumet 3760; TAN!. noTes The only Benthamia with basal lobes of the lip that can be longer than the midlobe. The very prominent callus at the apex of the lip is also very distinctive. The species was recorded from La Réunion by Szelengowicz& Tamon (2013: 247). The text refers to the Madagascan species but the photograph probably shows a withered Benthamia perfecunda. See also: Schlechter (1930: t.11 n°41), Perrier (1939: 38), Cribb& Hermans (2009: 47), Hervouet (2018: 168). descripTion Erect slender terrestrial herb, 60-78cm tall. Tubers 2, fusiform, elongated, ending in a root, roots 7-8, 1-2mm in diameter. Stem 3-6mm in diameter, with the base en45°E 15°S 20° 25° 50° Madagascar Mauritius La Réunion 20 km La Réunion 20 km La Réunion 88,427 203,665 mm fig. 73. — Distribution maps of Benthamia mascula Hervouet & Descourv., sp. nov. 224 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. veloped by 2-3 black sheaths, in the middle with leaves, higher up with several sheaths with their blade decreasing in size with height. Leaves 3-4, 10-12× 0.8-1cm, grass-like, clasping the stem. Rachis cylindrical, dense, bearing 36-90 flowers, 10-15× 1.5cm. Floral bracts narrowly lanceolateacuminate, 7-18× 1-2mm, the lower ones up to three times longer than the flower. Ovary fusiform-cylindrical, twisted, 3.5-7× 2-4mm. Flowers butter yellow, 2-3× 2.53mm without the ovary; sepals oblong, obtuse, slightly mucronate, 3-veined, dorsal sepal 2-2.6× 1-1.4mm, lateral sepals 2-2.6× 1-1.4mm, slightly asymmetrical; petals oval, obtuse, with one main vein and two smaller, 2-2.5× 1.2-1.5mm, almost as long as the sepals, but a little wider; lip broadly oval, 2-3× 2-2.5mm, concave, trilobed in the upper third; lobes oval-triangular obtuse, almost equal, the lateral lobes a little thickened and narrowed towards the tip, 0.3-0.7mm long, sometimes larger than the midlobe, the midlobe very fleshy, with a very prominent central callus at the tip; spur very short, 0.8-1× 0.5-0.7mm, oblongly saccate, obtuse, thickened at the base. Column subterete, short, 0.6-0.7× 0.5-0.8mm, anther not apiculate, obtuse to hardly apiculate; auricles rounded, crenulate, almost half the length of the anther or smaller; rostellum trilobed, lateral lobes short, horizontal, midlobe toothed, erect, 0.10.2mm long. Capsules 6× 3mm, developing before the apical flowers have opened. Benthamia monophylla Schltr. (Figs 76; 77) Repertorium Specierum Novarum Regni Vegetabilis 33: 30 (Schlechter 1924). Type maTerial. — Madagascar• Haute Matsiatra, massif d’Andringitra; alt. 2200-2600m; III.1922; H. Perrier 14387; holotype: P[P00094540]!; isotype: P[P00094541]!. eTymology. — “Monophylla”, in Greek with a single leaf. phenology. — January to April. fig. 74 . — Benthamia melanopoda Schltr.: Ambohitantely, Madagascar, 10 March 2015. Photograph by Jean-Michel Hervouet. 45°E 15°S 20° 25° 50° 88,427 fig. 75. — Distribution map of Benthamia melanopoda Schltr. 225 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) disTribuTion and ecology. — Madagascar. A high elevation plant, from 1600 to 2600m, found in the southern half of the country, mostly from Andringitra massif (Haute Matsiatra region) to Andohahela in the south (Anosy region), in humid mountain grasslands or small patches of vegetation on rocks (Fig. 78). c onservaTion . — This species has a large EOO and is locally abundant, but the AOO is probably less than 2000km². It was quoted NT by IUCN in 2016 (https://doi.org/10.2305/IUCN. UK.2018-1.RLTS.T44393380A69785668.en). addiTional specimens examined. — Madagascar• Analamanga, near Mandraka; 18°55’30”S, 47°56’20”E; XII.1905; C.d’Alleizette 7066b; L[L1487725]!• Haute Matsiatra, pic Boby, Andringitra; 22°11’S, 46°53’E; 1946; A.M. Homolle 1192; P[P02115319, P02115320]!• sine loc.; 1946; A.M. Homolle 1263; P[P00692277]!• sine loc.; 1946; A.M. Homolle 1277; P[P00692278]!• Anosy, massif de l’Andohahela; 24°36’S, 46°42’E; alt. 1800-1975m; I.1934; H. Humbert 13608; P[P0009454, P00094543, P00094544]!, G!, TAN!• Haute Matsiatra, canton Sendrisoa, district Ambalavao; 22°13’S, 46°55’E; 17 .I.1958; C. Rakotovao 9938RN; P[P00692318, P00692461]!• Haute Matsiatra, 30km NW of Fianarantsoa; VIII.1963; W. Rauh 10613; HEID[HEID2500341]!• Haute Matsiatra, pente du Vohidray, Andringitra; 13.I.1971; J.L. Guillaumet 3761; TAN!• Anosy, massif de l’Andohahela, RN11; I.1974; P. Morat 4389; P[P00692454, P00692455]!• Haute Matsiatra, Andringitra reserve; 22˚10’S, 46˚56’E; alt. 1975m; 13 .I.1987; M. Nicoll 262; K[K000453691, K000453692]!, MO[MO3022623]!, TAN!• Haute Matsiatra, Andringitra, near to Pic Boby; II.2007; T. Ranarivelo etal. 484; K!, TAN!• Haute Matsiatra, Andringitra; D. RobertsDLR119& DLR120; K!• Haute Matsiatra, Andringitra; D. RobertsDLR125; K[spirit 76706]!. noTes Plants sometimes have two basal leaves (e.g. on Ambondrombe). The flowers have a lip that is very similar to the petals and are reminiscent of some Holothrix in South Africa. It can form huge colonies, e.g. on Andringitra. With its elongate spur, this species is not a typical Benthamia with a short spur like the type of the genus, Benthamia latisatis. Perhaps it belongs to a different genus. The specimen H. Perrier 14387has two sheets in P, but Perrier’s handwriting on specimen P00094540 clearly shows that it is the sheet sent to Berlin for the description by Schlechter, it is therefore the holotype. See also: Schlechter (1930: t.11 n°42), Perrier (1939: 24), Perrier (1939: 30), Cribb& Hermans (2009: 42), Cribb& Hermans (2009: 46), Hervouet (2018: 169). fig. 76. — Benthamia monophylla Schltr.: Andringitra massif, Madagascar, 7 April 2018. Photograph by Jean-Michel Hervouet. fig. 77 . — Benthamia monophylla Schltr., Andringitra massif, Madagascar, 7 April 2018. Photograph by Jean-Michel Hervouet. 232 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. nondum evolutis”). This is questionable since all specimens in P have leaves, only one specimen at K has no leaves. Schlechter doubted that this species would belong to the genus Benthamia, because of the long spur and the short staminodes (here auricles). See also: Schlechter (1930: t.12 n°46), Perrier (1939: 20), Cribb& Hermans (2009: 43). descripTion Erect slender terrestrial herb, 10-40cm tall. Tubers 5-7, stalked, oblong-claviform, downy, roots numerous. Stem 1.5-3mm in diameter, with 2-3 basal amplexicaul sheaths, 20-35× 6-10mm, a tuft of leaves and a few bract-like sheaths higher up. Leaves 5-9, erect, acuminate-aristate, 40-70× 2-5mm. Rachis rather dense, bearing 25-45 flowers, elongated, 8-23cm long, unilateral. Floral bracts lanceolate-linear, 4-7× 1-1.6mm, very acute, a little longer than the ovary. Ovary 4-8× 1-2mm, a little curved. Flowers small, greenishbrown; sepals elliptic-oblong, obtuse, dorsal sepal 2-2.9× 0.9-1.3mm, lateral sepals 2.2-3.1× 0.9-1.3mm; petals as long, 2-3× 0.9-1.4mm, lanceolate-ligulate obtuse, a little expanded towards the base of the front edge; lip broadly cuneate, 3-4× 1.6-2.5mm, trilobed above the middle, lobes not equal, ligulate-obtuse, the middle one thick, around 1.8mm long, the laterals divergent, around 1mm long; spur narrowly cylindrical or conical, somewhat obtuse, 2-3× 0.3-0.8mm. Anther subglobose; auricles hardly visible at the base of column, suborbicular, twice as short as the anther. Benthamia rostrata Schltr. (Fig. 87) Repertorium Specierum Novarum Regni Vegetabilis 33: 33 (Schlechter 1924).— Platanthera madagascariensissensu Schltr., Beihefte zum Botanischen Centralblatt 33: 299 (Schlechter 1916), non Kraenzl. (Kraenzlin 1899: 609). BenthamiaperularioidesSchltr., Repertorium Specierum Novarum Regni Vegetabilis 33: 32 (Schlechter 1924). — Type: Madagascar• Vakinankaratra, mont Tsiafajavona; 19°19’58”S, 47°16’08”E; alt.2500m; III.1921; H. Perrier 13523; lectotype: P[P00094567]!, here designated; isolectotype: P[P00094568]!, syn. nov. Type maTerial. — Madagascar• Vakinankaratra, mont Ibity au sud d’Antsirabe; alt. 2000m; II.1914; H. Perrier 8101; holotype: P[P00094574]!; isotype: P[P00094575]!. eTymology. — From the Latin “rostratus”, beaked, referring to the midlobe of the lip. phenology. — January to April. disTribuTion and ecology. — Madagascar. Widespread in montane grasslands from 1400 to 2370m. Can also grow on bare laterite soils (Fig. 88). conservaTion. — The conservation status of this species is of Least Concern (LC). It has the largest number of specimens of the genus in herbaria and is still widespread in Madagascar. It has however been assessed EN by IUCN in 2015 (https://doi.org/10.2305/IUCN. UK.2015-2.RLTS.T70102386A70124752.en). addiTional specimens examined. — Madagascar• sine loc.; Baron 849; P[P00094569, as Rolfeella glaberrima, on the same sheet as Dr. Fox 23 but different]!, K[K000415545]!• sine loc.; R. Baron 5252; P[P00094570]!• Ankaratra; R. Capuron 759; P[P00692268]!• sine loc.; A.M. Homolle 1112; P[P00692276]!• Ihorombe, plaine de la Menarahaka, entre Ihosy et Ivohibe; 22°27’S, 46°15’E; B. Descoings 983; P[P00692270]!• Ankaratra; G.F. ScottElliot 2096; BM[BM000034590]!• Ankaratra; G.F. Scott-Elliot 1982; P[P00334731]!• Central Madagascar; R. Baron 2271; K!• sine loc.; R. Bonet s.n.; P[P00326135]!• Analamanga, Tampoketsa d’Ankazobe; E. Ursch 1019; TAN!, P[P00692272]!• Analamanga, Ambatovory; 18°54’S, 47°42’E; III.1883; Dr. Fox 24; K! ex LDS• Centre; 1887; C.M. Le Myre de Vilers s.n.; P[P00094474]!• Analamanga, Ambohimanga; 18°45’42”S, 47°33’49”E; 12.VI.1905; C.d’Alleizette s.n.; L[L1487720]!• Analamanga, environs de Nanisana près de Tananarive; 18°54’S, 47°42’E; 20.I.1906; L. Rotereau s.n.; P[P06796776]!• Analamanga, Nanisana; 18°54’S, 47°42’E; I.1906; C.d’Alleizette 977M; P[P00334732]!• Vakinankaratra, Manandona; alt. 1400m; IV.1912; H. Perrier 8024; P[P00094576]!• Vakinankaratra, mont Ibity; 20°04’03”S, 47°00’04”E; alt. 2000m; II.1921; H. Perrier 13581; P[P00094578]!• Analamanga, Ambohimanga; 18°45’42”S, 47°33’49”E; 27.III.1921; R. Decary 302; P[P00334738]!• Analamanga, Ambohimanga; 18°45’42”S, 47°33’49”E; IV.1921; R. Decary 52; P[P00334739]!• Haute Matsiatra, massif d’Andringitra; alt. 2000m; II.1923; H. Perrier 14431; P[P00094579]!• Diana, Ankaizina; alt. 1500m; 19.IV.1923; R. Decary 1918; P[P00334737]!• Diana, Anfig. 87. — Benthamia rostrata Schltr.: Mount Ibity, Madagascar, 1 April 2018. Photograph by Jean-Michel Hervouet. 233 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) kaizina; alt. 1500m; 19.IV.1923; R. Decary 1936; P[P00334736]!• Diana, Ankaizina; 18°54’S, 47°42’E; alt. 1700m; 19.IV.1923; R. Decary 1997; P[P00334735]!• Vakinankaratra, entre Ambatolampy et Antsirabe; alt. 1600m; II.1927; H. Perrier 17906; P[P00334733, P00334734]!• Haute Matsiatra, Andringitra; 26.II.1938; P. Boiteau 3135; P[P00692296, P00692297]!• Haute Matsiatra, au sommet de l’Ambondrombe; 21°52’30”S, 47°15’37”E; alt. 2000m; 11.IV.1941; P. Boiteau 4611; P[P00692437]!• Analamanga, Ambohitantely; 18°05’S, 47°16’E; 24.II.1945; A.M. Homolle s.n.; P[P00692274]! the four plants on the left are Tylostigma nigrescens Schltr. or Tylostigma perrieri Schltr.• Analamanga, Tampoketsa d’Ankazobe; 18°13’S, 47°17’E; 7.II.1948; P. Boiteau 6725; TAN!• Diana, massif de Marivorahona; alt. 2000-2244m; 18-26.III.1951; H. Humbert etal. 25753; P[P02115322 plant on the right only, P02115321, plant on the right only]!• Alaotra-Mangoro, Ambatosoratra, Atozaka, RN3; 17°38’30”S, 48°50’00”E; 20.III.1953; ResNat 6055; P[P00692314]!• Haute Matsiatra, RN5, Canton Sendrisoa, District Ambalavao; 27.II.1954; R. Rakoto 6484; P[P00692315]!• Vakinankaratra, route d’Ambatolampy, Faratsiho; 19°23’S, 47°16’E; alt. 1800; VIII.1955; J. Bosser 10880; P[P00692227]!• Vakinankaratra, Ambohimandroro, Ampatolampy; XI.1955; J. Bosser 9206; TAN!• Vakinankaratra, Nanokely, Ankaratra; 19°32’S, 47°03’E; alt. 2000m; II.1957; J. Bosser 10878; P[P00692226]!• Vakinankaratra, Nanokely (district de Faratsiho); 19°32’S, 47°03’E; 26.II.1960; M. Peltier 1915; P[P00692290]!• Analamanga, Soavina PK 22 route de Tamatave; 18°52’S, 47°40’E; III.1960; J. Bosser 13532; P[P00692228]!• Amoron’i Mania, Fiadanana, Ilaka, Ambositra; 20°34’30”S, 46°57’30”E; 20.III.1960; M. Peltier 2149; P[P00692292]!• Analamanga, vallée de Soavina, PK 23 route de Tamatave; II.1961; J. Bosser 14863; TAN!• Vakinankaratra, Manjakatompo; 19°21’25”S, 47°18’28”E; III.1961; J. Bosser 15024; P[P00692229]!• Analamanga, lake Mantasoa; 19°00’45”S, 47°50’17”E; IX.1962; J. Bosser 16144; P[P00692230]!• Vakinankaratra, Nanokely, Ankaratra; 19°32’S, 47°03’E; II.1963; J. Bosser 17573; P[P00692241]!• Route de Majunga; III.1963; J. Bosser 16783; P[P00692235]!• Vakinankaratra, route d’Ambatolampy; 19°23’S, 47°16’E; III.1963; J. Bosser 16797; P[P00692236]!• Vakinankaratra, route Ambatolampy-Faratsiho; II.1964; J. Bosser 19264; P[P00692246, P00692247]!• Haute Matsiatra, sentier du pic Boby, Andringitra; 22°11’S, 46°53’E; IV.1964; J. Bosser 19478; P[P00692248, P00692249]!• Haute Matsiatra, Andringitra; 22°13’S, 46°55’E; IV.1964; J. Bosser 19492; P[P00692251]!• Haute Matsiatra, Andringitra; IV.1964; J. Bosser 19506; P[P00692252, P00692253]!• Vakinankaratra, Col des Tapias, 45km avant Ambositra; 20°14’34”S, 47°06’04”E; IV.1964; J. Bosser 19598; P[P00692259]!• sine loc.; 28.III.1969; anonym; HEID[HEID250340]!• Vakinankaratra, Mt. Ibity western slope; 20°05’44”S, 47°00’12”E; alt. 1900m; III.1972; G. Cremers 2037; TAN!• Vakinankaratra, south of Antsirabe, Mt. Ibity; 20°10’S, 47°03’E; alt. 1300-1800m; III.1985; L.C. Barnett etal. 491; TAN!, MO[MO3022638]!• Haute Matsiatra, Andringitra Forest Reserve, along path to Pic Boby; 22°11’S, 46°55’E; alt. 1900-2050m; III.1989; P. Goldblatt etal. 8964; P[P00692435]!, MO[MO4314431]!, TAN!, WAG[WAG.1130546]!• Vakinankaratra, Ankaratra Massif, W of Ambatolampy; 19°20’S, 47°16’E; alt. 2350m; III.1989; P. Goldblatt etal. 9012; MO[MO4314448]!, TAN!• Amoron’i Mania, on hillside near Talavina, PK 204 from Antananarivo to Ambositra; 20°10’42”S, 47°05’24”E; alt. 12301330m; V.1993; C.H. Jongkind etal. 829; K!, MO[MO3022616]!, P[P00692179]!, TAN!• Vakinankaratra, Col Tapia, 36km S of Antsirabe; 20°10’41”S, 47°05’16”E; alt. 1400m; IV.1995; D. DuPuy etal. M906; K!, P[P00059605]!, MO!, WAG!, TAN!• Analamanga, Tampoketsa d’Ankazobe; 17°52’07”S, 47°03’39”E; 18.III.1999; J.N. Labat 2951; P[P00160016, P00160017]!, G!, K!, MO!, TAN!• Between Antananarivo and Ambositra, Tapia forest; 2000; J. Hermans 4393; K!• Vakinankaratra, between Antsirabe and Ambositra, on RN7, about 3km before Col Tapia; alt. 1440m; IV.2001; J. Hermans 5251; K!• Vakinankaratra, between Antsirabe and Ambositra, on RN7, about at Col Tapia; 20°14’25”S, 47°06’12”E; alt. 1740m; V.2001; J. Hermans 5205; K! with spirit• Amoron’i Mania, Ambositra, Ivato, road to Antoetra; 20°45’43”S, 47°19’31”E; alt. 1701m; 25 .I.2007; A. Sieder FS4188 and J. Hermans 8086; WU[WU0063773, spirit material]!• Amoron’i Mania, Ambatofinandrahana, 25km W of Itremo village; 20°34’12”S, 46°34’55”E; alt. 1647m; 21.II.2010; L. Rajavelona etal. RJL148; K[K664548]!, TAN!• Amoron’i Mania, Antoetra, on inselberg; 20°46’27”S, 47°19’16”E; II.2015; alt. 1676m; A.& C. Sieder etal. 6801; WU! spirit material• N4, 126km north of Tana, inselberg; II.2017, A.& C. Sieder etal. 7309; WU! flower in alcohol• Vakinankaratra, between Antsirabe and Ambositra, on RN7, Col Tapia; IV.2018; J. Hermans 8241; K!• Analamanga, Andakana RN4; 18°15’58”S, 47°10’51”E; 19.III.2019; alt. 1461m; A.& C. Sieder etal. 7792; WU!• Analamanga, RN4, c.12km after Ankazobe; alt. 1461m; III.2019; Sieder A.& C. etal., L7787; WU!. noTes A very widespread species, with a variable spur length. Any differences between Benthamia rostrata and B.perularioides do not hold when a large number of specimens are examined. Moreover, there are no distinct distribution zones as suggested by Perrier (Perrier 1939: 24, 25), with Benthamia perularioides north of Ankaratra and Benthamia rostrata south of Ankaratra. This species was also often misidentified as Benthamia bathieana, which added to the confusion. 45°E 15°S 20° 25° 50° 88,427 fig. 88. — Distribution map of Benthamia rostrata Schltr. 234 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. Benthamia perularioides and B.rostrata were described in the same publication by Schlechter. The rules of nomenclature allow a choice of which name to prioritise. We have chosen B.rostrata because this name is more widely used. The type indicated by Schlechter for his Platanthera madagascariensis is “Perrier de la Bathie XXI, β“. He copied this on specimen P00094574 which has handwritten XXI and B together with the number 8101. Schlechter (1924: 33) gave “H. Perrier 8101 (XXI B)” when he coined the name Benthamia rostrata. The specimen P00094574 was thus sent to Berlin and returned to Paris, so there is no ambiguity that it is the holotype. For Benthamia perularioides Schlechter cited Perrier 13523 for the type, but there are in P two specimens with this collection number. We have chosen P00094567 as lectotype because Perrier indicated on it that it was sent to Schlechter in April 1921. This species, which has an elongated spur, is different from the type of the genus Benthamia and other species with a scrotiform or short spur. It is perhaps misplaced in Benthamia. As a matter of fact, specimens identified as Benthamia perularioides in (Ngugi etal. 2020) were found to be embedded in Cynorkis. At the beginning of the flowering time the spur is enveloped by the bracts, as is also observed in some African Habenaria. See also: Schlechter (1930: t.12 n°45, as Benthamia perularioides), Perrier (1939: 24, as Benthamia perularioides), Perrier (1939: 25), Cribb& Hermans (2009: 42, as Benthamia perularioides), Cribb& Hermans (2009: 43), Bosser& Lecoufle (2011: 180), Hervouet (2018: 172). descripTion Erect slender terrestrial herb, 22-70cm tall. Tubers 4-11, clustered, fusiform, slender and white, 5-8mm in diameter, 20-60mm long, roots up to 8, wiry, white, 1-2mm in diameter. Stem 2-4mm in diameter, with the base covered by sheaths 25-30mm long, with above slightly fleshy leaves and higher up 7-10 lanceolate sheaths, gradually turning into floral bracts. Leaves 2-3, more or less narrowly oblong, 7-20× 1.54cm, shortly acuminate or mucronate. Rachis rigid, erect, 16-25× 1.2cm, densely many-flowered, somewhat unilateral with 30-50 flowers. Floral bracts lanceolate, very acuminate, erectly spreading, 5-15× 1.5-3mm, embedding the ovary, the lower ones longer than the ovary. Ovary twisted, 5-10mm long, diameter 0.6-1.6mm, the basal part fused to the base of the floral bract. Flowers resupinate, c.7.5mm and 4mm deep, olive green turning yellowish; sepals narrowly oblong, obtuse, slightly navicular, the dorsal obscurely 3-veined, erect, 1.8-3.2× 0.7-1.6mm, the laterals single-veined, spreading, 2.3-4× 0.7-1.9mm; petals narrowly oval-obtuse, obscurely 2-veined, a little expanded at the anterior margin towards the base, 1.7-3.2× 0.9-1.6mm, forming a hood with the dorsal sepal; lip a little longer than the sepals, 2.8-4× 1-2.3mm, rhomboid in outline, tri-lobed, sometimes geniculate between the spur and the lobes, the central lobe tongue-shaped with apex curving up, obtuse, 1.5× 0.5mm, the lateral lobes triangular, obtuse, 0.5× 0.7mm; spur subcylindrical, trumpet-shaped at first then becoming thinner, pendent or trapped into the bract, narrowed from the base to the obtuse apex, whitish green, as long as the sepals, 2.3-6× 0.3-0.6mm. Anther not apiculate, suborbicular, 1-2.2× 0.8-1.3mm, with two distinct chambers; auricles very rounded, thick, almost a quarter the length of the anther; lateral lobes of the rostellum obsolete, the middle one obtuse, pointing backwards. Sweetly scented in the evening. Benthamia spiralis A.Rich. Monographie des Orchidées des Îles de France et Bourbon: 39 (Richard 1828).— Habenaria dissimulata Schltr., Beihefte zum Botanischen Centralblatt 33: 404 (Schlechter 1915).— Benthamiaspiralisvar.dissimulata(Schltr.) H.Perrier, Bulletin de la Société botanique de France 81: 34 (Perrier 1934). Type maTerial. — Mauritius• sine loc.; J. Néraud 8; holotype: P[P00738316]!. eTymology. — From the Latin “spiralis”, spiral, referring to the arrangement of the rachis. phenology. — Mainly May to October, but flowering time rarely noted on specimens. disTribuTion and ecology. — Mauritius. Habitat unknown, except the vicinity of a marsh mentioned for one specimen (Fig. 89). conservaTion. — This species is at least Critically Endangered (CR), with an area of occupancy severely fragmented and less than 10km² and an inferred continuing decline (criterion B2ab). The most recent collection, by Francis Friedmann, dates back to 1971, there is thus no recent proof that it is not extinct. a ddiTional specimens examined . — Mauritius• sine loc.; J.B.G.M. Bory R43052; W!• sine loc.; L.H. Boivin s.n.; P[P00738422]!• sine loc.; L.H. Boivin s.n.; P[P00738537! plant in the middle only]• sine loc.• L.H. Boivin s.n.; P[P00738421]!• sine loc.; L.H. Boivin s.n.; P[P00738538]!• sine loc.; 1831; L. Bouton s.n.; P[P00738527]!• sine loc.; 1846-1848; L.H. Boivin s.n.; W!• Marais contigu au chemin de Chirac près du point le plus élevé; X.1849; L.H. Boivin s.n.; P[P00738389]!• sine loc.; VIII.1862; P.B. Ayres s.n.; P[P00738413 plant in the middle only]!• Pétrin; 20°24’31”S, 57°28’19”E; V.1971; F. Friedman 1162; P[P00738417, P02088511]!. noTes When Achille Richard described this species in 1828, it was for him clearly distinct from Satyrium spirale Thouars. The latter species appears on page21 of the same publication under the nameHabenaria spiralis (Thouars) A.Rich. Moreover, he wrote on the Néraud 8 type specimen: «non in P. Th.», meaning that it was not a Du Petit-Thouars species. The name was validly published since no one had used the name Benthamia spiralis before. When Schlechter wanted to transfer Benthamia spiralis A.Rich. to Habenaria in 1915, he was hindered by the already existing Habenaria spiralis(Thouars) A.Rich. which has Satyrium spirale Thouars as basionym. He therefore created a new name: Habenaria dissimulata Schltr. Many subsequent authors have mistakenly considered Benthamia spiralis A.Rich as a new name for Satyrium spirale Thouars. To add to the confusion the original Thouars name Satyrium spirale was illegitimate because it was already used by Swartz in 1788, for what is now Spiranthes torta (Thunb.) Garay& H.R.Sweet from Central America. For this reason, 235 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) Lindley applied the new name Spiranthes africana in 1824, later changed to Benthamia africana (Lindl.) Hermans in 2014. Examination of the types proves them to be different, we have thus Benthamia spiralis A. Rich, endemic to Mauritius, as a different species from Benthamia africana, from Madagascar and the Mascarenes. That Benthamia spiralis A.Rich. was a different species or variety was also the interpretation of Perrier de la Bâthie who wrote (Perrier 1934: 33): « on doit le distinguer comme variété et certains botanistes pourraient même le considérer comme une espèce distincte» («it must be considered as a distinct variety and some botanists could even consider it as a distinct species»), «avec un port plus robuste et des feuilles plus grandes, plus larges et plus nombreuses» («with a more robust habit, larger, wider and more numerous leaves »), but he mistakenly named it Benthamia spiralis (Thouars) A.Rich. var. dissimulata (Schltr.) H.Perrier and among other specimens he cited one from La Réunion island (Boivin 1064, P[P00738525]!, which is for us merely Benthamia africana. This species differs from Benthamia africana mainly by the more numerous cauline regularly alternate linear leaves, borne up to the base of the rachis, and also by the longer lateral lobes of the lip, the petals with fleshy tips and revolute margins, and the much longer floral bracts, up to twice the length of the ovary. Both plants are growing together and field observations are necessary to know to which extent they hybridize. descripTion Erect terrestrial herb 42-66cm tall. Tubers 2, fusiform, 4× 0.8cm, roots 2-6, up to 6cm long, 1-2mm in diameter. Stem with the base covered by 1-2 sheaths, 3-5mm in diameter. Leaves 5-10, erect, linear or strap-like, alternate subdistichous, acute and glabrous, 8-19cm× 4-11mm, regularly spaced (internode 2-3cm), up to the rachis. Rachis terminal (4)7-17(23)cm, bearing 27-45 flowers roughly arranged in a spiral, 4-6mm apart. Floral bracts linear-lanceolate, acute, enveloping the ovary at the basis, 8-15mm long, up to twice the length of the ovary. Ovary fusiform, twisted, glabrous, slightly curved at the apex, almost vertical, 5-8mm long. Flowers resupinate, 2.5-4.5mm long, green; sepals 1-veined; dorsal sepal elliptic-oblong, 2-3× 0.8-1mm; lateral sepals similar or slightly longer, 2.1-3× 0.9-1mm; petals oblong-triangular, thickened at the apex, with slightly revolute margins, 2.2-2.6× 0.8-1.2mm; lip 2-3× 1.2mm, concave in the proximal part, 45°E 15°S 20° 25° 50° Madagascar Mauritius La Réunion 40 km Mauritius M aur iti u s 88,427 203,665 mm fig. 89. — Distribution map of Benthamia spiralis A.Rich. 236 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. subcanaliculate, clearly trilobed at apex, like a trident, with a thickened midlobe 0.5-1mm long, obtuse, lateral lobes acute, 0.3-0.5mm long; spur 0.8-1mm, slightly bilobed, slightly contracted at the base. Column subterete, 0.5-1× 0.8-1mm, with an obtuse to subacute nose-shaped apiculus 0.2-0.3mm long, with lateral broadly rectangular auricles, with irregular margins; rostellum trilobed, hardly visible in dried material. Benthamia spiraloides (Cordem.) Hermans& P.J.Cribb (Figs 90; 91; 92) Lankesteriana 21 (2): 76 (Hermans& Cribb 2021).— Habenaria spiraloidesCordem., Flore de l’île de la Réunion (Cordemoy 1895: 551). — Type: France, La Réunion• Plaine des Palmistes, Cordemoy s.n.; holotype: [not found] • Plaine des Palmistes, chemin vers l’îlet Patience; 21°06’S, 55°37’E; 1100m; IV.2002; T. Pailler 48; neotype: REU[REU007939]!, designated by Hermans& Cribb (2021: 76). eTymology. — Named for its similarity to what was once called Satyrium spirale Thouars, now Benthamia africana. phenology. — July to March. disTribuTion and ecology. — La Réunion and Madagascar. Terrestrial or epiphytic, in moss, in semi-shade. In Madagascar observed thriving on rotten Eucalyptus stumps, or on the ground in pine plantations. Between 1000 and 1900m (Fig. 93). c onservaTion . — This species has been provisionally quoted Vulnerable (VU) in La Réunion (Hermans etal., 2023), where less than ten locations are known and its semi-shaded habitat among mossy rocks is threatened (criteria B1ab and B2ab). In Madagascar it is present in a number of reserves like Angavokely and Manongarivo and has probably been overlooked in a number of places. It can thrive in pine or Eucalyptus plantations and is therefore of Least Concern (LC). a ddiTional specimens examined . — Madagascar• Diana, massif de Manongarivo; 14°00’00”S, 48°23’30”E; alt. 1200m; IV.1909; H. Perrier 1944 bis; P[P00094536, P00094537]!• Analamanga, Manjakandriana; 18°54’S, 47°48’E; VIII.1955; alt. 1200m; J. Bosser 8292; P[P00692224]!• Analamanga, Lake Mantasoa; 19°00’45”S, 47°50’17”E; IX.1962; J. Bosser 16416; P[P00692234]!• Analamanga, Ankeramadinika, Ambatoloana; VI.1963; J. Bosser 16161; P[P00692231]!• Alaotra-Mangoro, route de Lakato; 19°11’30”S, 48°26’00”E; VIII.1963; J. Bosser 18523; P[P00692244]!• N25, peu après le PK15 en venant de la N7; 17.VII.1969; Y. Veyret 1114; P[P00692305, P00692306, P00692307]!• Analamanga, Angavokely; 18°55’36”S, 47°45’07”E; 2.VI.1970; Y. Veyret 1303; P[P05096694]!• Analamanga, Angavokely; 18°55’36”S, 47°45’07”E; 27.VII.1970; Y. Veyret 1330; P[P05096692, P05096693, P05096695]!• Diana, Réserve spéciale de Manongarivo, chemin d’Ambalafary au vallon supérieur de la Bekolosy; 14°02’S, 48°18’E; alt. 840m; 16.V.1995; L. Gautier etal. LG2668; P[P00098891]!, K!, G!, WAG[WAG.0388478]!• Alaotra-Mangoro, Ambatovy; 18°50’51”S, 48°18’31”E; 21.X.2005; H. Razanantsoa etal. 579; P[P02091795]!• Alaotra-Mangoro, Andasibe, near Mantadia forest; 1.X.2016; J. Hermans 8038; K!• Alaotra-Mangoro, Moramanga, Maromizaha; alt. 1080m; 18.VIII.2022; L.R. Rajaovelona 1402; K[K001522047]!, TAN!. France, La Réunion• Sur les montagnes; C. Richard 697; P[P00738528]!• sine loc.; L.H. Boivin s.n.; P[P00738369]!• sine loc.; J. Renz 13027; RENZ!• sine loc.; 1837; C. Richard 397; P[P00738529]!• sine loc.; 1867; Herb. A. Richard R19908; W!• Takamaka; 21°05’S, 55°38’E; 24.V.1974; J. Bosser 22049; P[P00738522]!• Bélouve forest; alt. 1500m; V.2003; M. Pignal etal. 2079; P[P00340568]!, K!, MO!• Plaine des fougères; 21°00’15”S, 55°29’21”E; alt. 1200m; 28.V.2003; T. Pailler TP78; REU[REU006450, REU007938]!• Piton rouge, Plaine des Cafres; 21°09’S, 55°34’E; 1900m; 9.III.2004; T.Pailler TP164; REU!• Bébour, route forestière de Takamaka, sentier îlet à bananes; 21°05’54”S, 55°35’35”E; alt. 1200m; 16.VI.2004; J. Fournel JF110; REU[REU010244, REU010245, REU010246]!• L’étang salé; 21°13’32”S, 55°22’09”E; alt. 1000m; 5 .VI.2005; V. Grondin 1796; CBNM!• Plaine des Fougères, sentier vers Be Massoune; 21°00’15”S, 55°29’21”E; alt. 1380m; 29.IX.2005; J. Fournel JF306; REU[REU010337]!• Piton Marmite, Salazie; alt. 1850m; 31.VII.2006; J. Fournel JF405; REU[REU010388]!• Bébour, sentier la rivière; 21°06’40”S, 55°33’47”E; alt. 1300-1350m; 7.VI.2007; F. Martos FM128 and FM129; REU[REU007895, REU007897]!• Dimitile, sentier Bayonne; 21°13’10”S, 55°27’45”E; 15 .VI.2008; F. Martos FM362 and FM363; REU[REU007896, REU007898]!• Dimitile, sentier Bayonne; 21°13’10”S, 55°27’45”E; 25.IV.2009; F. Martos FM623; REU[REU007754]!• sine loc.; XI.2017; ML04; CBNM!• sine loc.; IX.2019; ML02; CBNM!• Bébour forest; 21°06’40”S, 55°33’47”E; 15.XI.2019; J. Hermans 8365; K!. fig. 90 . — Benthamia spiraloides (Cordem.) Hermans & P.J.Cribb, Ialatsara forest, Madagascar, 1 August 2016. Photograph by Jean-Michel Hervouet. 237 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) noTes This species has long been overlooked in Madagascar, where it has been confused with Benthamia africana and Benthamia misera. Before the present study many specimens in herbaria were unidentified. See also: Bernet (2010: 138, as Benthamia sp. 1), Szelengowicz& Tamon (2013: 243, as “Benthamia dauphinensis H.Perrier”), Hervouet (2018: 160, as Benthamia dauphinensis (Rolfe) Schltr.), Hermans& Cribb (2023: 122). descripTion A slender epiphytic or terrestrial glabrous herb, 20-45cm tall. Tubers 2-3, ovoid, roots fleshy and woolly. Stem c.1mm in diameter with 2 to 3 short basal sheaths. Leaves 1-3, cauline, narrowly ligulate or linear-lanceolate, acute, 7-14× 0.5-1.5cm, shortly petiolate, the second leaf c.2-3cm above the basal and shorter. Rachis spiral, 5-12cm long, lax to dense, with 15-27 flowers, with 2 to 4more or less leaf-like sheaths, often purplered. Floral bracts linear, acuminate-filiform 4-11mm× 1-3mm, brownish-red. Ovary disproportionately large, at first erect then divergent, with three longitudinal ridges, 5-11× 1-2.7mm, generally brownish purple. Flowers 2-4mm in diameter, corolla almost tubular, generally with reddish-brown sepals, yellow petals, lip and spur; dorsal sepal cucullate, 3-veined, elliptic, obtuse, 1.5-3.2× 0.6-1.4mm; lateral sepals hardly spreading, forming a tube with the dorsal, 3-veined, obliquely oblong-ovate, obtuse, 1.3-3.1× 0.8-3.1mm, concave at the apex; petals lanceolate, subacute, 1.7-3.1× 0.6-1.2mm; lip barely trilobed, 1.8-3.1× 0.9-1.8mm, lateral lobes basal, small, rounded, midlobe thick, fleshy, about as long as the lateral lobes, the apex often incurved; spur scrotiform to subrectangular, often sub-bilobed-sacculate, the base funnel-shaped, 0.5-1× 0.5-0.7mm, twice as long as wide. Column with short auricles, less than 1/3 the size of the anther, anther retuse or obtusely apiculate. Benthamia trifida Hermans& Hervouet, sp.nov. (Figs 94; 95; 96) diagnosis. — Benthamia trifida Hermans& Hervouet, sp.nov. shares the fleshy pale petals and lip with contrasting green sepals with Benthamia herminioides but that species is terrestrial or lithophytic, has a much shorter leaf at anthesis and a more densely flowered rachis. It shares the habit of Benthamia spiraloides (Cordem.) Hermans& P.J.Cribb but in that species the leaves are narrower and borne higher on the stem, the lateral sepals are not spreading-reflexed and the spur is scrotiform (vs flattened). It is also similar to the widespread terrestrial or lithophyte Benthamia africana (Lindl.) Hermans but that species has more leaves (3 to 5 vs 1 to 3) which are narrower and more attenuate, a denser rachis and a more distinctly trilobed lip. fig. 91. — Benthamia spiraloides (Cordem.) Hermans & P.J.Cribb: Bras-Cabot, Bébour forest, La Réunion, 3 November 2021. Photograph by Jean-Michel Hervouet. fig. 92 . — Benthamia spiraloides (Cordem.) Hermans & P.J.Cribb, Ialatsara forest, Madagascar, 1 August 2016. Photograph by Jean-Michel Hervouet. 238 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. Type maTerial. — Madagascar• Alaotra-Mangoro, Andasibe area; 19°00’40”S, 48°27’24”E; alt. 1150m; V.2001; J. Hermans 8224; holotype: K!. eTymology. — The species epithet trifida, refers to the distinctive three-pointed calyx and corolla. phenology. — Flowering from March to May. disTribuTion and ecology. — Madagascar. From Andasibe region (Alaotra-Mangoro region) down to Ranomafana in the south (Haute Matsiatra region), so far known from only two localities. Altitudinal range 950 to 1200m, medium altitude moist evergreen forest (Fig. 97). conservaTion. — The AOO of this species is less than 500km², the two locations known are a national park that suffered poaching of orchids during the Covid years, Ranomafana N.P., and a forest which is not yet an official reserve and is threatened by logging, the Iaroka forest near Andasibe. This species is thus considered Endangered (EN) according to criterion B2. addiTional specimens examined. — Madagascar• Alaotra-Manoro, Andasibe area; 19°00’40”S, 48°27’24”E; alt. 1100m; V.2001; J. Hermans 8231; K!• Haute Matsiatra, Ranomafana N.P.; 21°14’22”S, 47°23’41”E; alt. 1025m; 20.III.2015; photographic records by Jean-Michel Hervouet (Figs 94; 95)• Alaotra-Mangoro, Andasibe area; alt. 1200m; 5.V.2023.; photographic records by J. Hermans. noTes This species is very variable in size and even very small plants can produce flowers. descripTion Slender epiphytic herb, 4-25cm tall. Tubers 2-3, ovoid, woolly, 10-25× 3-8mm, roots wiry, woolly, 0.6-1.4mm in diameter. Stem 3-5mm in diameter, with 2 to 3 short basal enveloping sheaths. Leaves cauline, generally 1 but up to 3 in older plants, erect to arcuate, narrowly ligulate or linear-lanceolate, acute, 3-12.5× 0.5-1.5cm, shortly petiolate, the midvein slightly raised beneath, pale to dark green. Rachis more or less spiral, erect, secund, green, with 1 to 2 sheaths towards the base, loosely flowered in the upper half, with up to 45 flowers but generally fewer. Floral bracts narrowly lanceolate, acuminate, about as long as the ovary, 3-5.6× 1.2-1.8mm, green. Ovary disproportionately large, at first erect then divergent, with three longitudinal ridges, 3.5-6.5× 1-1.4mm, pale green. Flowers fleshy, c.3× 4mm, corolla almost tubular at the base, sepals pale green, petals and lip often pale pink but sometimes pale yellowish-green; dorsal sepal cucullate, narrowly elliptic, obtuse, arcuate over the column, 1.5-2.2× 0.4-0.9mm; lateral 45°E 15°S 20° 25° 50° Madagascar Mauritius La Réunion 20 km La Réunion 20 km La Réunion 88,427 203,665 mm fig. 93. — Distribution map of Benthamia spiraloides (Cordem.) Hermans & P.J.Cribb. 239 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) sepals spreading to recurved, narrowly elliptic, concave, obtuse, 1.9-2.1× 0.8-0.9mm; petals, ovate, subacute, the apex and anterior margin thickened and more or less lobed, 1.82× 1-1.2mm. Lip obtusely trilobed, 1.5-1.9× 1.1-1.7mm; lateral lobes thickened towards the apex, rounded; midlobe thick, fleshy, the apex curved; spur subrectangular, flattened, sometimes sub-bilobed, 0.8-1.3× 0.8-0.9mm, green. Column 0.6-0.9× 0.6-0.8mm, with prominent suborbicular auricles, anther curved, obtusely apiculate. Benthamia vulcanorum Hervouet& Descourv., sp.nov. (Figs 98; 99; 100) diagnosis. — Similar to Benthamia africana but differing in the number and position of leaves (2-3 leaves in the middle of the stem vs3-5 basal and cauline leaves in lower part of the stem), in the non-resupinate flowers, that are half closed and facing upwards (vsresupinate, opened and horizontal), the differently coloured flower parts (yellow tips of petals and lip, yellow spur, vsuniform green for Benthamia africana) and the flattened spur (vsscrotiform). Similar to Benthamia perfecunda but flowers not in three rows, the ovary is vertical (vsperpendicular to the rachis in Benthamia perfecunda), and the number and the position of the leaves are different (2-4 cauline leaves in lower part of the stem for Benthamia perfecunda). T ype maTerial . — France, La Réunion• sine loc.; 14.II.1969; T.Cadet 1963; holotype: P[P00738518]!. eTymology. — The species range lies in high altitude zones between the Fournaise volcano and the Piton des Neiges, hence the name vulcanorum, “of the volcanoes”. phenology. — From March to April, at the end of the rainy season. disTribuTion and ecology. — La Réunion. Collected between 1700 and 2300m, growing on bare soil made of volcanic ash (Fig. 101). c onservaTion . — This species is known from more than ten localities but its AOO is very restricted, less than 20km², it could be threatened by the consequences of a volcanic eruption throwing ashes, it must therefore be considered Vulnerable (VU) according to criterion D2. addiTional specimens examined. — France, La Réunion• Coteau Kervéguen; 21°07’27”S, 55°30’16”E; II.1971; J. Bosser 20737, 20740, 20744 and 20745; P[P00738532, and spirit, P00738371, P00738372, P00738524]!• Sentier du Côteau Maigre, Plaine des Cafres; 21°08’08”S, 55°31’40”E; II.1971; J. Bosser 20479; P[P00738373]!• Sentier du Côteau Maigre, Plaine des Cafres; 21°08’08”S, 55°31’40”E; II.1971; J. Bosser 20481; P[P00738523 and spirit]!• Piton Mare à boue, Plaine des Cafres; 21°07’56”S, 55°30’45”E; 30.III.1974; J. Bosser 21774; P[P00738520]!• Base du Piton Mare à Boue, Plaine des Cafres; 21°09’35”S, 55°34’35”E; 30.III.1974; J. Bosser 21783; P[P00738535]!• Sentier du Piton des Neiges, Cilaos; 21°07’06”S, 55°29’12”E; alt. 1700-1800m; 14.IV.1978; J. Bosser 22506; P[P00738312, P00738313]!• Descente Plaine des sables le Tremblet; 21°17’14”S, 55°43’20”E; alt. 2000m; 20.IV.1978; J. Bosser 22540; P[P00738401]!• sine loc.; alt. 2300m; 1987; J. Renz 14774; RENZ!• Sentier du Piton fig. 94 . — Benthamia trifida Hermans & Hervouet, sp. nov., Ranomafana, Madagascar, 20 March 2015. Photograph by Jean-Michel Hervouet. fig. 95 . — Benthamia trifida Hermans & Hervouet, sp. nov., Ranomafana, Madagascar, 20 March 2015. Photograph by Jean-Michel Hervouet. 240 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. des Neiges par la Plaine des Cafres; 21°07’57”S, 55°32’36”E; alt. 1700m; 24.IV.1988; J. Dupont 1; P[P00738359 and spirit]!• Sommet Kervéguen; 21°07’27”S, 55°30’16”E; alt. 2300m; 24.V.2000; F. Martos FM706; REU[REU007702]!• Route forestière du volcan; 21°11’40”S, 55°38’36”E; alt. 2200m; 1.III.2003; T. Pailler etal. TP70; REU[REU006420]!• Bébour, Piton des Neiges to Bélouve path; alt. 1900m; 5.III.2003; V. Hoarau etal. 13 and 14; REU[REU007941, REU006426] !• Saint-Joseph, Caverne de AF H G B C D E fig. 96. — Benthamia trifida Hermans & Hervouet, sp. nov.: A, dorsal sepal, lateral sepal and petal; B, flower, side view with floral bract and front view; C, lip, spur and column, side and front view; D, column; E, flower with lip removed, front view; F, plant and inflorescence; G, tuber, plant and inflorescence; H, anther. Drawn by Deborah Lambkin. From Hermans 8224, Hermans 8231, and photographs. Scale bars: A, B, E, 2 mm; C, 1.5 mm; D, 1 mm; F, 3 cm; G, 2 cm; H, 0.5 mm. 241 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) Cotte; alt. 2240m; 21°14’16”S, 55°38’18”E; III.2004; J. Férard etal. 1041 and 1042; CBNM!• Saint-Joseph, Oratoire SainteThérèse; 21°12’37”S, 55°39’15”E; 2300m; II.2005; V. Grondin etal. 1564; CBNM!• Maïdo, sentier du Grand Bénard; 21°04’09”S, 55°23’16”E; 21.III.2007; V. Lecomte s.n.; REU[REU005135]!• Route du Volcan, vers Le Piton de l’Eau; 21°11’02”S, 55°40’28”E; 21.V.2008; F. Martos FM340 and FM341; REU[REU007894, REU007903]!• Puy Ramond; 21°17’41”S, 55°42’22”E; 22.II.2020; V. Lavergne; photographs. noTes Although long overlooked, this species has distinctive features, such as the flat spur, the erect flowers and the leaves similar to those of the local Tylostigma, which shares the same altitude and habitat close to the top of the volcanoes Fournaise and Piton des Neiges. It was noted and briefly described by Bernet (Bernet 2010) who provisionally labelled it Benthamia sp. 3. It is likely that Jean Bosser also noticed this plant as he systematically noted on his specimens that the lip and petals were green with yellow tips. Another distinctive feature to look for is the erect ovary often prolonged by erect half-closed flowers. It is surprising that among 20 specimens found none were collected before 1969. However, it is unlikely that this is due to a recent arrival, since the plant is endemic to La Réunion. Given the number of very similar specimens and the absence of would-be parents in the same habitat, it is unlikely that this species is a hybrid. In the Benthamia africana complex in Madagascar, some plants (e.g. observation of first author on Sainte-Marie Island) may have two-colour erect flowers with a flattened spur, but in this case all the leaves are basal, as leaves in the drawing by Thouars (Thouars 1822: t.9). There is also in the spirit collection in P a sample corresponding to the type P00738518. See also: Bernet (2010: 140, as Benthamia sp. 3), Szelengowicz& Tamon (2013: 248, as Benthamia misera). descripTion Erect terrestrial herb 25-80cm tall. Tubers 2-4, fusiform, 2× 0.8cm, roots 2-6, up to 7cm long, 1-3mm in diameter. Stem cylindrical, 1.2-3mm in diameter, with leaves in the 45°E 15°S 20° 25° 50° 88,427 fig. 97. — Distribution map of Benthamia trifida Hermans & Hervouet, sp. nov. fig. 98 . — Benthamia vulcanorum Hervouet & Descourv., sp. nov., Le Tremblet, La Réunion, 20 April 2018. Photograph by Jean-Michel Hervouet. 248 ADANSONIA, sér. 3 • 2025 • 47 (11) Hervouet J.-M. et al. We thank Pr. Thierry Pailler (Université de La Réunion) for showing the first author Benthamia specimens of the La Réunion Herbarium and fruitful exchanges, especially on the former Benthamia micrantha, and for sending spirit material then donated to MNHN. Thanks to Jacques Fournel for providing information on his specimens in La Réunion Herbarium. We acknowledge the important findings of Véronique Lavergne in Ambodiriana forest in Madagascar, of Benthamia calceolata, Benthamia elata and Benthamia litoralis sp.nov. She was also generous in providing photographs and duly documented notes. In situ observations are of utmost importance in a revision and we waermly thank Laurent Catan, Charles Robert and Jean-Maurice Tamon for fruitful fieldtrips in Mascarene Islands. REFERENCES baron r. 1901. — Compendium des plantes Malgaches. Revue de Madagascar 3 (10): 722. b enKe m. 2004. — La Réunion des orchidées sauvages. Éditions Mabé, 504 p. b erneT p. 2010. — Orchidées de la Réunion. Naturae Amici éditions, Entre-Deux, 368 p. b osser J. 1969. — Contribution à l’étude des Orchidaceae de Madagascar. X. Adansonia série 2, 9 (3): 343-359. https://www. biodiversitylibrary.org/page/59837149 bosser J. 2015. — Description de trois Cynorkis Thouars nouveaux (Orchidaceae) du nord de Madagascar. Adansonia, sér.3, 37(1): 19-23. https://doi.org/10.5252/a2015n1a3. https://sciencepress. mnhn.fr/fr/periodiques/adansonia/37/1/description-de-troiscynorkis-thouars-nouveaux-orchidaceae-du-nord-de-madagascar bosser J. & lecoufle m. 2011. — Les orchidées de Madagascar. Biotope, Mèze, 496 p. (Collection Parthénope). buTzin f. 1981. — Typenstudien im Berliner Orchideen-Herbar: Diverse markierte Typen. Willdenowia 11: 119-121. https://www. jstor.org/stable/3995795 byTebier b. 2025. — The correct name for the type species of the genus Benthamia (Habenariinae, Orchidaceae). Candollea 80: 1-5. https://doi.org/10.15553/c2025v801a1 c adeT J. 1989. — Joyaux de nos forêts : les orchidées de la Réunion. Nouvelle Imprimerie Dyonisienne, Saint-Denis-de-La-Réunion, 179 p. chase m. W., cameron K. m., freudensTein J. v., pridgeon a. m., salazar g., van den berg c. & schuiTeman a. 2015.— An updated classification of Orchidaceae. Botanical Journal of the Linnean Society, Botany, vol. 177: 151-174. https:// doi.org/10.1111/boj.12234 cordemoy de J. 1895. — Flore de l’île de la Réunion. Imprimerie de J. Tastu, Paris, 574 p. https://www.biodiversitylibrary.org/ page/21709771 cribb p. J. & hermans J. 2009. — Field Guide to the Orchids of Madagascar. Royal Botanical Gardens, Kew, 456 p. dorr l. J. 1997. — Plant Collectors in Madagascar and the Comoro Islands. Royal Botanic Gardens, Kew, 524 p. d ressler r. l. 1981. — The Orchids, Natural History and Classification. Cambridge, Massachusetts & London, 332 p. https://doi. org/10.2307/1219717 du puy d., cribb p., bosser J., hermans c. & hermans J. 1999.— The Orchids of Madagascar. Royal Botanic Gardens, Kew, 376 p. durand T. & schinz h. 1892. — Conspectus Florae Africae. Vol.5. Jardin botanique de l’état, Bruxelles, 289p. https://www.biodiversitylibrary.org/page/29402748 fineT a. 1911. — Orchidées nouvelles ou peu connues. Notulae Systematicae 2: 23-25. https://www.biodiversitylibrary.org/ page/6677717 frappier c. de monbenoisT 1880. — Orchidées de l’île de la Réunion, catalogue des espèces indigènes découvertes jusqu’à ce jour. Gabriel et Gaston Lahuppe, Saint-Denis, La Réunion, 16p. https://www.biodiversitylibrary.org/page/516129 gauTier l., Tahinarivony J. a., ranirison p. & Wohlhauser s. 2018.— Vegetation, in goodman et al. (eds), The Terrestrial Protected Areas of Madagascar, their History, Description and Biota. Volume 1, chapter 7. Association Vahatra, Antananarivo: 207-242. g aray l. & r omero g. a. 1998. — Schedulae Orchidum. Harvard Papers of Botany 3: 53. https://www.jstor.org/stable/41761555 h arris g. h. & h arris m. W. 2000. — Plant Identification Terminol - ogy, an Illustrated Glossary. Second edition. Spring Lake, Utah, 206p. hermans J. & cribb p. J. 2014. — New species and new names in Malagasy orchids (Orchidaceae). Kew Bulletin 69 (2)-9517. https://doi.org/10.1007/s12225-014-9517-X hermans J. & cribb p. J. 2021. — New combinations and other taxonomic changes for the forthcoming “Flore des mascareignes” orchidaceae accounts. Lankesteriana 21 (2): 61-137 (2020) 75: 50 1-25. https://doi.org/10.15517/lank.v21i2.47156 hermans J. & cribb p. J. 2023. — Flore des Mascareignes. Vol.170. Orchidées, vol. 1. IRD Éditions, Marseille, 340 p. hermans J., hermans c., du puy d., cribb p. J. & bosser J. 2007.— Orchids of Madagascar. Second edition. Royal Botanic Gardens, Kew, 398 p. hermans J., andrianTiana J. l., sieder a., Kiehn m., cribb p. J., raJavelona l. & gardiner l. m. 2017. — New species and nomenclatural changes in Cynorkis (Orchidaceae: Orchidoideae) from Madagascar and the Mascarenes. Kew Bulletin 72: 38 1-38. https://doi.org/10.1007/s12225-017-9715-4 hermans J., raJaovelona l. r., cribb p., hervoueT J.-m., sieder a. & andrianTiana J. 2020. — New species and nomenclatural changes in Cynorkis (Orchidaceae) from Madagascar, the Comoros and the Mascarenes. Kew Bulletin 75: 50 1-25. https://doi.org/10.1007/s12225-020-09904-5 h ervoueT J.-m. 2018. — À la recherche des orchidées de Madagascar. Sur les traces d’Henri Perrier de la Bâthie. Biotope, Mèze, 504p. (Collection Parthénope). hervoueT J.-m. & hermans J. 2022. — Deux nouvelles espèces du genre Cynorkis Thouars (Orchidaceae, Orchidioideae) à Madagascar, et une nouvelle combinaison pour C.tryphioides var. leandriana. Adansonia, sér. 3, 44 (9): 63-79. https://doi. org/10.5252/adansonia2022v44a9. http://adansonia.com/44/9 hervoueT J.-m., descourvieres p., reJo fienena f. & ranaivo J. 2014. — Two new species of the genus Benthamia A.Rich. (Orchidaceae) from Madagascar, B.boiteaui Hervouet, sp.nov. and B.bosseri Hervouet, sp.nov. Adansonia, sér. 3, 36 (2): 209219. https://doi.org/10.5252/a2014n2a4. https://sciencepress. mnhn.fr/fr/periodiques/adansonia/36/2/deux-nouvelles-especesdu-genre-benthamia-rich-orchidaceae-madagascar-b-boiteauihervouet-sp-nov-et-b-bosseri-hervouet-sp-nov iucn 2012. — IUCN Red List Categories and Criteria. Version 3.1. Second edition. Gland, Switzerland and Cambridge, UK. IUCN. iv+32p. https://iucnredlist.org/resources/categories-and-criteria Jin W.-T., schuiTeman a., chase m. W., li J.-W., chung s.- W., hsu T.-c. & Jin x.-h. 2017. — Phylogenetics of subtribe Orchidinae s.l. (Orchidaceae; Orchidoideae) based on seven markers (plastid matK, psaB, rbcL, trnL-F, trnH-psba, and nuclear nrITS, Xdh): Implications for generic delimitation. BMC Plant Biology17: 222. https://doi.org/10.1186/s12870-017-1160-x Kraenzlin f. 1899. — Orchidacearum Genera et Species, vol. I.Berlin, Mayer& Müller,986 p. https://www.biodiversitylibrary. org/page/10350481 leWis b. a., phillipson p. b., andrianarisaTa m., rahaJasoa g., raKoTomalaza p. J., randriambololona m. & mcdonagh J. 1996. — A study of the botanical structure, composition, and diversity of the eastern slopes of the Réserve Naturelle Intégrale d’Andringitra, Madagascar. Fieldiana 85: 24-71. https://www. biodiversitylibrary.org/page/2855999 249 Revision of the genus Benthamia A.Rich. (Orchidaceae, Orchidioideae, Habenariinae) ADANSONIA, sér. 3 • 2025 • 47 (11) linder h. p. & Williamson p. 1986. — Notes on the Orchids of Southern Tropical Africa. II. Oligophyton drummon - dii, gen. et sp.nov. Kew Bulletin 41 (2): 313-317. https://doi. org/10.2307/4102936 lindley J. 1824. — Edwards’s Botanical Register. Vol.10. London. lindley J. 1832. — Herminium cordatum, sub t. 1499. Edwards’s Botanical Register 18. London. lindley J. 1833. — Ornamental flower-garden and shrubbery. Edwards’s Botanical Register 19. https://www.biodiversitylibrary. org/page/239468 lindley J. 1835. — The Genera and Species of Orchidaceous Plants. Ridgways, London. https://www.biodiversitylibrary. org/page/393248 m oore s. 1876. — On the orchids collected at the Island of Bourbon, during the transit of Venus expedition, by Dr. I. B. Balfour. The Journal of Botany, British & Foreign, vol. 5, new series (vol.14 entire work): 292-293. https://www.biodiversitylibrary. org/page/516129 moore s. 1877. — Orchideae, in baKer J. G. (ed.), Flora of Mauritius and the Seychelles. London, L. Reeve: 327-364. https://www. biodiversitylibrary.org/page/54531716 n gugi g., l e p échon T., m arTos f., p ailler T., b ellsTedT d. u. & byTebier b. 2020. — Phylogenetic relationships amongst the African genera of subtribe Orchidinae s.l. (Orchidaceae; Orchideae): Implications for subtribal and generic delimitations. Molecular Phylogenetics and Evolution 153: 106946. https://doi. org/10.1016/j.ympev.2020.106946 pailler T. & henze f. 2020. — Orchidées de la Réunion. Orphie, 209p. pailler T., Tournebize r. & henze f. 2013. — Guide des Orchidées de la Réunion. Université de la Réunion, 209p. pailler T., gauche m., da silva d. & besse p. 2024.— Position taxonomique de Hemiperis micrantha (Frapp. Ex Cordem.) Schltr. (Orchidaceae; Orchidioideae; Habenariinae), par séquençage des genes ITS et matK. Adansonia, sér. 3, 46 (14): 149-155. https://doi.org/10.5252/adansonia2024v46a14. http:// adansonia.com/46/14 perrier de la bâThie h. 1934. — Le genre Benthamia (Orchidées). Bulletin de la Société botanique de France 81: 25-38. https://doi. org/10.1080/00378941.1934.10833927 perrier de la bâThie h. 1939. — 49.Orchidaceae, in humberT h. (ed.), Flore de Madagascar et des Comores. Volume 1. Antananarivo, Madagascar: 16-40. https://www.biodiversitylibrary. org/page/8099159 perrier de la bâThie h. 1951. — Orchidées de Madagascar et des Comores. Nouvelles observations. Notulae Systematicae 14 (2): 138-165. https://www.biodiversitylibrary.org/page/8067581 p ridgeon a. m., c ribb p. J., c hase m. W. & r asmussen f. n. 2001.— Genera Orchidacearum. Volume 2. Orchidoideae (Part1). Oxford University Press, Oxford, 416 p. richard a. 1828. — Monographie des Orchidées des Îles de France et Bourbon. Mémoires de la Société d’Histoire naturelle de Paris, tome IV: 1-70. https://www.biodiversitylibrary.org/page/4191509 ridley h. n. 1885. — The orchids of Madagascar. The Journal of the Linnean Society, Botany 21: 456-522. https://www.biodiversitylibrary.org/page/176990 r idley h. n. 1886. — On Dr. Fox’s collection of orchids from Madagascar. The Journal of the Linnean Society, Botany 22: 116127. https://www.biodiversitylibrary.org/page/228351 rolfe r. 1891. — New and little known Madagascar plants, collected and enumerated by G.F. Scott-Elliot, Orchideae. Journal of the Linnean Society, Botany 29: 50-59. https://www.biodiversitylibrary.org/item/8386 rolfe r. 1893. — New and little-known Madagascan plants, collected and enumerated by G.F. Scott-Elliot. Orchidaceae. The Journal of the Linnean Society, Botany 29: 50-59. https://www. biodiversitylibrary.org/page/227088 rolfe r. a. 1899. — Peristylis glaberrimus and P.madagascariensis. Orchid Review 7 (73): 4 (Jan. 1899). https://www.biodiversitylibrary.org/page/38921231 schlechTer r. 1915. — Kritische Aufzählung der bisher von Madagaskar, den Maskarenen, Komoren und Seychellen bekanntgewordenen Orchidaceen. Beihefte zum Botanischen Centralblatt 33: 390-440. https://www.biodiversitylibrary.org/page/59629600 s chlechTer r. 1916. — Orchidaceae Perrierianae (collection secunda). Beihefte zum Botanischen Centralblatt 34 (2): 294-341. https://www.biodiversitylibrary.org/page/59635226 schlechTer r. 1918. — Orchidaceae novae et criticae, Additamenta ad Orchideologiam madagascarensem. Repertorium specierum novarum regni vegetabilis 15: 324. https://www.biodiversitylibrary. org/page/244429 schlechTer r. 1924. — Orchidaceae Perrierianae. Ein Beitrag zur Orchideenkunde der Insel Madagaskar. Repertorium specierum novarum regni vegetabilis 33: 1-391. https://www.biodiversitylibrary. org/page/57726481 schlechTer r. 1930. — Blütenanalysen neuer Orchideen. Repertorium specierum novarum regni vegetabilis 68: 1-126. s prengel K. 1826. — Systema Vegetabilium. Vol.3. Librariae Dieterichianae, Gottingae, 936p. https://doi.org/10.5962/bhl.title.822 sTrahm W. 1994. — The Conservation and Restoration of the Flora of Mauritius and Rodrigues. Ph.D. thesis, Department of Botany, University of Reading. szelengoWicz m. & Tamon J.-m. 2013. — Les orchidées des Mascareignes. Printec Press Holdings, 427p. szlacheTKo d. l. 2004. — Habenariinae (Orchidaceae, Orchidoideae) – Beiträge zu einer Revision des Subtribus (9). Die Orchidee (Hamburg) 55 (3): 315-316. szlacheTKo d. l. 2005. — Habenariinae (Orchidaceae, Orchidoideae) – Beiträge zu einer Revision des Subtribus (12). Die Orchidee (Hamburg) 56 (1): 68-70. szlacheTKo d. l. & Kras m. 2006. — Matériaux pour la révision des Habenariinae (Orchidaceae, Orchidoideae). Richardiana 6: 138-146. https://www.biodiversitylibrary.org/partpdf/272877 szlacheTKo d. l. & ruTKoWsKi p. 2000. — Gynostema Orchidalium I. Acta Botanica Fennica 169: 1-380. Thouars a.-a. 1809. — Extrait de trois mémoires lus à la première classe de l’institut, sur l’histoire des plantes orchidées des îles australes d’Afrique. Nouveau bulletin des sciences de la société philomatique de Paris 1: 314. https://www.biodiversitylibrary.org/page/4123300 Thouars a.-a. 1822. — Histoire particulière des plantes orchidées recueillies sur les trois îles australes d’Afrique. Réimpression de Earl M. Coleman, 1979, New-York. https://www.biodiversitylibrary.org/page/516129 Submitted on 3 September 2024; accepted on 29 December 2024; published on 21 July 2025.