Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region, with the description of a new species from Mayotte
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Muñoz, Iris Montero, Levin, Geoffrey A., Cardiel, José María (2023): Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region, with the description of a new species from Mayotte. Adansonia (3) 45 (26): 395-496, DOI: 10.5252/adansonia2023v45a26, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/adansonia2023v45a26.pdf
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395 ADANSONIA, sér. 3 • 2023 • 45 (26) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com KEY WORDS , Acalypha, Madagascar, Comoros, Seychelles, Mascarene Islands, brachyblast, lectotypifications, new synonyms, new combination, new species. Iris MONTERO MUÑOZ Departamento de Biología, Facultad de Ciencias, Universidad Autónoma de Madrid, Campus de Cantoblanco, Postal Code 28049, Madrid (Spain) Current address: Real Jardín Botánico de Madrid (RJB), CSIC, Plaza Murillo 2, 28014, Madrid (Spain) [email protected] [email protected] (corresponding author) Geoffrey A. LEVIN Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816 South Oak Street, Champaign, Illinois 61920 (United States) Current Address: Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 (Canada) José María CARDIEL Departamento de Biología, Facultad de Ciencias, Universidad Autónoma de Madrid, Campus de Cantoblanco, Postal Code 28049, Madrid (Spain) and Instituto de Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid, Campus de Cantoblanco, Postal Code 28049, Madrid (Spain) Submitted on 23 May 2022 | accepted on 9 March 2023 | published on 27 November 2023 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region, with the description of a new species from Mayotte Montero Muñoz I., Levin G. A. & Cardiel J. M. 2023. — Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region, with the description of a new species from Mayotte. Adansonia, sér. 3, 45 (26): 395-496. https://doi.org/10.5252/adansonia2023v45a26. http://adansonia.com/45/26 ABSTRACT This work presents a monographic revision of the genus Acalypha L. (Euphorbiaceae) from the Western Indian Ocean Region, including Madagascar, the Comoros Archipelago, the Mascarene Islands, the Seychelles Archipelago, and the Scattered Islands. Forty-nine species are recognized, with 42 native and seven probably introduced; 39 are found in Madagascar, eight in the Comoros, eight in the Mascarene Islands, four in the Seychelles, and one in the Scattered Islands. For each species, we provide a description, information about distribution and habitat, a distribution map, and a preliminary conservation assessment. Original illustrations and the first identification key for Acalypha species of this region are also included. Lectotypes are designated for 10 names. Acalypha crateriana (Coode) I.Montero& Cardiel, comb. nov. is proposed as a new combination and Acalypha berryi I.Montero, Cardiel& G.A.Levin, sp.nov., a new species from Mayotte, is described and illustrated.
396 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. INTRODUCTION Acalypha L. is the type genus of Euphorbiaceae subfamily Acalyphoideae Beilschm. With about 500 species, Acalypha is the third most species-rich genus of Euphorbiaceae Juss. [sensu stricto, since Phyllanthus L. (sensu lato) includes >600 spp], after Euphorbia L. (Riina etal. 2013) and Croton L. (Berry etal. 2005). Acalypha is considered a natural and wellcharacterized genus (Nowicke& Takahashi 2002). Molecular phylogenetic studies strongly support its monophyly (Levin etal. 2005; Sagun etal. 2010; Levin etal. 2022). Acalypha is morphologically characterized by three synapomorphies associated with wind pollination: elongate, pendent, twisted anther sacs; small, brevicolporate, almost smooth pollen grains; and highly laciniate stigmas (Nowicke& Takahashi 2002; Sagun etal. 2006, Levin etal. 2022). It also appears that epidermal crystals are a synapomorphy of the genus (Cardiel etal. 2020). The most recent proposal for an infrageneric classification of the genus (Levin etal. 2022) includes four subgenera: three already recognised in the previous classification by Pax& Hoffmann (1924) – Acalypha, Androcephala Pax& K.Hoffm., and Linostachys (Klotzsch ex Schltdl.) Pax& K.Hoffm.– and a fourth one, Hypandrae (Müll.Arg.) Hurus, proposed by Levin and colleagues based on molecular data. Acalypha species are mainly small trees and shrubs and, to a lesser extent, herbs. Its distribution is mainly pantropical, although some herbaceous species reach temperate areas. They range from tropical humid forests to sub-desert zones, and from sea level to about 4000m altitude in tropical mountains. The greatest diversity appears in tropical America, with around 250 species (Cardiel etal. 2022b), followed by continental Africa, with around 65 species (Cardiel& Montero Muñoz 2018), and the islands of the Western Indian Ocean Region, the subject of this monograph, where we recognize 49 species. According to the biogeographic system of Brummitt (2001), the Western Indian Ocean Region (hereafter WIOR) includes Madagascar, the Comoros Archipelago (Union of Comoros and the French Department of Mayotte), the Mascarene Islands (Mauritius and the French Department of La Réunion), the Seychelles Archipelago, and the Scattered Islands (Fig. 1). This area belongs to the Malagasy region of the Paleotropical kingdom, according to Takhtajan (1986). Taxonomic hisTory of AcAlyphA of Wior The first species of Acalypha from WIOR, A.filiformis Poir. and A.venosa Poir., both from Mauritius, were described by Poiret (1804). Willdenow (1805) soon added another species from Mauritius, A.integrifolia Willd. Bojer’s Hortus Mauritianus (Bojer 1837), which included the first catalogue of the native species of Mauritius, listed eight Acalypha species, two of them newly named (A.lantanaefolia Bojer and A.tomentosa Bojer), but unfortunately Bojer’s designations are both nomina nuda. One of the 19th century botanists who contributed the most to the study of Acalypha in WIOR was Henri Baillon. He initially listed two species from Madagascar (A. reticulata (Poir.) Müll.Arg. and A.salviifolia Baill.) and three from the Mascarene Islands (A.marginata (Poir.) Spreng., A.colorata (Poir.) Spreng., and A.arborea Comm. in Poir.) (Baillon 1858). Soon thereafter, he listed nine species from Madagascar (A.acuminata Vahl ex Baill., A.emirnensis Baill., A.goudotiana Baill., A.gracilipes Baill., A.pervilleana Baill., A.rottleroides Baill., A.salviifolia, A.spachiana Baill., and A.urophylla Boivin ex Baill.), four species from the Comoros (A.chibomboa Baill., A.codonocalyx Baill., A.ovalifolia Baill., and A.richardiana Baill.), and three species from the Mascarene Islands (A.arborea, A.commersoniana Baill., and A.marginata); 11 of these species were newly described (Baillon, 1861). Much later, Baillon’s (1891) contribution to Histoire naturelle des Plantes volume III (Atlas) of Grandidier’s Histoire physique, naturelle et politique de Madagascar (Baillon 1891) included excellent plates, made by the illustrator André Revillon d’Apreval, of ten Acalypha species, five of them new to science (A.diminuta Baill., A.humblotiana Baill., A.leptomyura Baill., A.madreporica Baill., and A.polynema Baill.). It should be noted that only plates were published in this work, with no accompanying descriptions or additional information, but under the International Code of Nomenclature (Turland etal. 2018), these are sufficient to validate Baillon’s new names. Finally, Baillon (1892, 1895a, 1895b) listed 32 Acalypha species from MOTS CLÉS Euphorbiaceae, Acalypha, Madagascar, Comores, Seychelles, Mascareignes, brachyblaste, lectotypifications, synonymes nouveaux, combinaison nouvelle, espèce nouvelle. RÉSUMÉ Révision monographique d’Acalypha L. (Euphorbiaceae) de la région de l’Ouest de l’Océan Indien, avec la description d’une espèce nouvelle endémique de Mayotte. Ce travail présente une révision monographique du genre Acalypha L. (Euphorbiaceae) de la région de l’océan Indien occidental, comprenant Madagascar, les îles Comores, les îles Mascareignes, l’archipel des Seychelles et les îles Éparses. Quarante-neuf espèces sont reconnues, dont 42 indigènes et sept probablement introduites; 39 se trouvent à Madagascar, sept aux Comores, huit aux Mascareignes, quatre aux Seychelles et une aux îles Éparses. Pour chaque espèce, nous fournissons une description, des informations sur la distribution et l’habitat, une carte de distribution et une évaluation préliminaire du statut de conservation. Des illustrations originales et la première clé d’identification des espèces d’Acalypha de cette région sont également incluses. Les lectotypes sont désignés pour 10 noms. Acalypha crateriana (Coode) I.Montero& Cardiel, comb. nov. est proposée comme nouvelle combinaison et Acalypha berryi I.Montero, Cardiel& G.A.Levin, sp.nov., une nouvelle espèce de Mayotte, est décrite et illustrée.
397 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) Madagascar, of which five were described for the first time (A.bakeriana Baill., A.hildebrandtii Baill., A.meiodonta Baill., A.leonii Baill., and A.vulneraria Baill.); he also provided descriptions of some of the species illustrated in the 1891 atlas. The treatment of Acalypha in de Candolle’s Prodromus (Müller Argoviensis 1865, 1866) included 11 Acalypha species from WIOR, six from Madagascar (A.emirnensis, A.reticulata, A.rottleroides, A.spachiana Baill., A.spiciflora Burm.f. and, as a novelty, A.fasciculata Müll.Arg.), three from the Comoros (A.chibomboa, A.codonocalyx, and A.richardiana), and two from the Mascarene Islands (A.commersoniana and A.marginata). Müller Argoviensis (1882) described one more species from Madagascar, A.buchenavii Müll.Arg. Other 19 th century authors also treated Acalypha from the WIOR. Baker (1877) listed five Acalypha species from Mauritius and the Seychelles and provided an identification key. Baker also described four new species from Madagascar (A.baronii Baker, A.hologyna Baker, A.lyallii Baker, and A.radula Baker) (Baker 1883, 1884). Strangely, Baron (1889) included only two Acalypha species (A.radula and A.urophylla) from Madagascar. Pax (1894) described four new species from Madagascar and the Comoros (A.juliflora Pax, A.comorensis Pax, A.urophylla Pax, and A.squarrosa Pax). Cordemoy (1895) wrote the first flora of La Réunion, in which five Acalypha species were included (A.indica L., A.poiretii Spreng., A.reticulata, A.colorata, and A.marginata). Early in the 20th century, Palacký (1907) compiled a list of 44 previously described Acalypha species from Madagascar. Voeltzkow (1917) included seven Acalypha species from the Comoros. Hemsley (1919), in his flora of Aldabra and nearby islands, included two species of Acalypha endemic to Seychelles (A.claoxyloides Hutch. and A.fryeri Hutch.), both previously described by Hutchinson (1918). The last worldwide treatment of the genus, in Engler’s Das Pflanzenreich (Pax& Hoffmann, 1924), included 31 Acalypha species from WIOR, only one of which was newly described (A.madagascariensis Pax& K.Hoffm.). Leandri (1942) made an important contribution to our understanding of Acalypha in WIOR. In a thorough taxonomic review of Acalypha in Madagasar, he treated 22 species and 14 varieties, of which eight species were newly described (A.andringitrensis Leandri, A.boinensis Leandri, A.decaryana Leandri, A.gagnepainii Leandri, A.humbertii Leandri, A.lepidopagensis Leandri, A.linearifolia Leandri, and A.perrieri Leandri). Leandri (1942) also proposed many new synonyms and provided illustrations and an identification key. Shortly after, he cited seven Acalypha species collected by Lam and Meeuse (Leandri, 1948). Coode (1978, 1979, 1982) reviewed Acalypha for the Mascarene Islands. He included five species, three of them introduced, and proposed some new subspecies and varieties. Robertson (1989) recognized five Acalypha species from the SeyWestern Indian Ocean Region SEYCHELLES Aldabra Group Alphonse Group Farquhar Group Agaléga Tromelin Island Cargados Carajos Shoals Rodrigues Mauritius La Réunion Madagascar Europa Island 1000 km 20°S 0° 0° 60°E 50°E 1600 km Bassas da India Juan de Nova Comoros Archipelago Glorioso Islands Southern Coral Group Amirante Islands Granitic group N N fig. 1. — Map of the Western Indian Ocean Region (WIOR).
398 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. chelles. Several thematic works on ethnobotany, conservation, etc. (e.g.: Heckel 1903; Jenkins 1987, 1990; Goodman 1996; Schatz 2001; Seebaluck etal. 2015; Seebaluck-Sandorama, 2018), cited Acalypha species from WIOR, although they do not provide relevant taxonomic information. Due to the insufficient and outdated state of knowledge of Acalypha in this region, we started a monographic study of Acalypha species from WIOR in 2017, as a part of the PhD dissertation of the first author (Montero Muñoz 2021). The first step was a nomenclatural review of the genus in this region (Montero Muñoz etal. 2018a). That was followed by the publication of 12 new species from the region (Montero Muñoz etal. 2018b, 2020a, b, 2022). MATERIAL AND METHODS The present work is based mainly on the study of 2 312 herbarium specimens of Acalypha stored in the following herbaria: A, B, BM, BREM, BRNU, C, CAN, CAS, E, FR, G, GB, GDC, GH, JE, K, L, LD, LE, LMU, M, MA, MAO, MAU, MAUAM, MO, MPU, NY, P, PRE, S, TAN, TEF, TUB, UPS, US, W, and WAG (acronyms according to Thiers 2022). They correspond to 1 646 collections from WIOR. Of them, 1 117 collections (1 597 specimens) are from Madagascar, 91 (140 specimens) from the Comoros, 356 (463 specimens) from the Mascarene Islands, 81 (111 specimens) from Seychelles, and one (one specimen) from the Scattered Islands. We have also reviewed numerous collections from continental Africa and other regions of the Paleotropics to improve our understanding of the diversity and evolution of the genus. Most of the specimens were studied in situ, either in the different herbaria or through loans. We also consulted digitized collections accessible through JSTOR Global Plants (https://plants.jstor.org) and other virtual herbaria, especially to review type collections. The morphological terms used follow those proposed by Harris& Harris (2001) and Ellis etal. (2009). The micromorphological analysis of the epidermal surface was carried out using scanning electron microscopy (SEM). For this purpose, we selected 52 specimens from different herbaria (BREM, COL, G, GH, K, MO, P, SEL, U, and W) corresponding to 30 native and five introduced species of our study area (Appendix I). For the analysis of the foliar surface, we used a scalpel to extract two approximately 3mm2 fragments from each specimen. We also examined bracts and flowers using a fragment or the entire organ. The samples were placed on SEM-specific slides and then metallized with a 15 nm gold layer, using a Quorum Q150T-S metallizer. For the observation and analysis of the samples, we used a HITACHI S-3000N electron microscope with X-ray detector (INCAx-sight, Oxford Instruments), which enables energy dispersive X-ray (EDX) analysis. The information obtained was incorporated into databases (nomenclature, collections, bibliography, morphology, distribution and habitat, images, etc.) using Microsoft Access 2016 © (Redmond, Washington, United States). These databases are integrated and managed through an online information system named Acalypha Taxonomic Information System (Cardiel etal. 2022b). This website aims to collect all the taxonomic and biogeographic information of Acalypha worldwide in open access, and it is integrated into global biodiversity networks through GBIF. The system allows detailed consultation of all the studied specimens and the available images of them; it also generates distribution maps. Most collections studied lacked geographical coordinates (81.7%) so we georeferenced them using GeoLocate v.3.22 (Ríos& Bart 2010) and Google Earth. For the older collections (from the 18th and 19th centuries) we use the “Gazetteer to Malagasy Botanical Collecting Localities” from the Missouri Botanical Garden (Schatz& Lescot 2003), which includes many old names for localities in Madagascar. When localities were ambiguous or imprecise, we assigned generic country or island coordinates. Distribution maps were prepared using QGIS Desktop 3.2.2. (QGIS Development Team 2018). The layers of the administrative boundaries of all the archipelagos of our study area were obtained from DIVAGIS 7.5 (Hijmans etal. 2012; https://www.diva-gis.org/), and the altitude layers from SRTM Tile Grabber (Watkins 2020; https://dwtkns.com/srtm/). Preliminary conservation assessments are based on the IUCN Red List Categories and Criteria (IUCN 2017). Area of occupancy (AOO) and extent of occurrence (EOO) were calculated with GeoCAT, a geospatial conservation assessment tool (Bachman etal. 2011; http://geocat.kew.org/), using a 2× 2km grid cell size as recommended by IUCN (2012, 2017). The nomenclature is based primarily on our previous work (Montero Muñoz etal. 2018a). The genus description is based on the WIOR species. The accepted species are arranged first by subgenera, and then alphabetically arranged followed by their author and place of publication. We then list, as appropriate, the basionym, homotypic synonyms in chronological order, and heterotypic synonyms in chronological order, following the same format as for the accepted name. Types are cited for all valid taxa, indicating the type locality, date of collection, collector, collection number, and herbarium/a where they are deposited, including the herbarium barcode if available. Missing information is indicated as s.loc. (without locality), n.d. (undated), and s.n. (without number). If the locality information is ambiguous or imprecise, we indicate it in quotation marks as it appears in the protologue. We provide the bibliographic sources for illustrations of the species, or the number of the figure included in this work. Descriptions are based on the studied specimens and follow the same sequence. When necessary, we use round brackets (parentheses) to indicate uncommon variation in morphological ranges and square brackets to indicate rare variation. The etymology of the specific epithet of each accepted name is also provided. For each species, we report its general geographic distribution and its distribution in WIOR. For Madagascar, we also indicate the provinces for which there are collection records; for the other archipelagos, we report the island/s where the species occurs. We also describe the habitat, according to the vegetation typology of Gautier etal. (2018), and the altitudinal range. This information is based exclusively on the collections studied.
399 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) Preliminary conservation assessments of each species are provided, indicating the IUCN category, the justification, and the criteria and sub-criteria followed. We cite chronologically the publications (with page numbers) in which the species are cited for WIOR; after that, we report the total number of collections examined, identifying each one with the primary col - AB CD F E GH fig. 2. — Trichomes: A, B, simple trichomes: A, lower leaf surface of Acalypha decaryana Leandri (P. Rakotomalaza 211); B, lower leaf surface of A. diminuta Baill. (H. Ralimanana 239); C, D, stellate and fasciculate trichomes; upper leaf surface of A. radula Baker (J. M. Hildebrandt 3865 and Service Forestier s.n.); E, simple and glandular trichomes, lower leaf surface of A. decaryana Leandri (Rakotomalaza 211); F, G, sessile glandular trichomes, lower leaf surface of A. diminuta Baill. (H. Ralimanana 239); H, pedunculate glandular trichomes, lower leaf surface of A. pervilleana Baill. (A. Mocquerys 159). Scale bars: A, E, 200 µm; B, 20 µm; C, G, 100 µm; D, H, 50 µm; F, 1 mm.
400 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. lector’s name and number; we also cite the herbarium barcode (when available). For full information about each collection, readers can consult the Acalypha Taxonomic Information System website (http://www.acalypha.es/secc/specimens.asp, using the information provided in the material examined section to search for the collection. Finally, in the notes section, we indicate any additional taxonomic or nomenclatural clarification, or other additional relevant information. DescripTive morphology Habit Most native Acalypha species from WIOR are shrubs or small trees to six meters high. We only found two herbs or subshrubs: A.leandrii I.Montero& Cardiel and A.rabesahalana I.Montero& Cardiel. Deciduous species predominate; only A.andringitrensis, A.berryi I.Montero, Cardiel& G.A.Levin, sp. nov., A.chibomboa, A.leonii, A.linearifolia, A.radula, and probably A.leandrii and A.vulneraria, are evergreen species. For ten species (A.crateriana (Coode) I.Montero& Cardiel, comb. nov., A.filiformis, A.gracilipes, A.integrifolia, A.isaloensis I.Montero& Cardiel, A.magistri I.Montero& Cardiel, A.marginata, A.pervilleana, A.richardiana, and A.urophylla Boivin ex Baill.) we do not have enough information to know if they are deciduous or evergreen. Branching is usually lax, with thin and slender branches, sometimes divaricate, and with many lenticels. Some speAB CD fig. 3. — Pedunculate glandular trichomes: A, lower leaf surface of Acalypha levinii I.Montero & Cardiel (P. Phillipson 2503); B, mature female bract of A. radula Baker (Service Forestier s.n.); C, mature female bract of A. rottleroides Baill. (L. Nusbaumer LN2629); D, mature female bract of A. vulneraria Baill. (F. Randriatafika 812). Scale bars: 0.5 mm.
401 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) AB CD EF fig. 4. — Sessile glandular trichomes: A, B, lower leaf surface of Acalypha decaryana Leandri (P. De Block 2383 and P. Rakotomalaza 211); C, lower leaf surface of A. diminuta Baill. (S. Wohlhauser SW310); D, lower leaf surface of A. emirnensis Baill. (L. Gautier LG3889); E, mature female bract of A. claoxyloides Hutch. (F. R. Fosberg 48724); F, mature female bract of A. emirnensis Baill. (L. Gautier LG3889). Scale bars: 0.5 mm.
408 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. Epidermal crystals Epidermal crystals in Acalypha were recently studied by Car - diel etal. (2020), who identified them as a new taxonomic trait of the genus. In that study, four types of crystals were recognized: prismatic crystals, styloids, stellate crystals, and druses. In the WIOR species, we found three types (prismatic, stellate, and druses) in 30 of the 34 analyzed species (Table 1); they appear mainly on the upper and AB CD E G F H fig. 10. — Druses (Dc): A, B, lower leaf surface of Acalypha diminuta Baill. (H. Ralimanana 239); C, D, female flower and ovary surface of A. diminuta Baill. (H. Ralimanana 239); E, ovary surface of A. fimbriata Schumach. & Thonn. (C. C. H. Jongkind 2277); F, sepal of the female flower of A. nusbaumeri I.Montero & Cardiel (L. N. Nusbaumer LN1169); G, H, lower leaf surface of A. vulneraria Baill. (F. Randriatafika 2314). Scale bars: A, B, E, 20 µm; C, 1 mm; D, H, 100 µm; F, 10 µm; G, 50 µm.
409 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) lower leaf surfaces, and often also on the bracts and flowers. Prismatic crystals (Fig. 9) can appear solitary or grouped, and we frequently observed two overlapping perpendicular crystals. These crystals appear on both the upper and lower leaf surfaces of most of the species studied, and on the female bracts of A.mayottensis. Druses can appear solitary or grouped (Fig. 10). We have found them in nine species, on upper and lower leaf surfaces, bracts, and flowers. Stellate crystals (Fig.11A-C) are very infrequent, appearing mainly in some South American species (Cardiel etal. 2020). We have not found them in native species from WIOR, but we have observed them in the male flowers of A. fimbriata Schumach.& Thonn., a species native to continental Africa and introduced in Madagascar. Due to insufficient sampling thus far, the systematic importance of crystals across Acalypha and among WIOR species requires further evaluation. In addition to the crystals described by Cardiel etal. (2020), we have recently identified another crystal form found on the leaf epidermis, which we have provisionally named “granules”. They have a rounded and smooth appearance and are c.5µm in diameter (Fig.11D-F). We have found them only on the foliar surface of three native species, A.claoxyloides, A.emirnensis, and A.mayottensis (Table 1). We still do not have conclusive results regarding their nature and composition. Epicuticular waxes The leaf surface of most Acalypha species studied is covered by epicuticular waxes of different shapes and densities. Following the classification of Barthlott etal. (1998), we found two basic types of waxes: layers and crystalloids (Table 2). Layers are continuous and homogeneous sheets AB C E D F fig. 11. — A-C, Stellate crystals (Cc), male bud surface of Acalypha fimbriata Schumach. & Thonn. (C. C. H. Jongkind 2277); D-F, granules (Ec): D, lower leaf surface of A. claoxyloides Hutch. (F. R. Fosberg 49627); E, lower leaf surface of A. emirnensis Baill., the white rectangle shows a single granule (L. Gautier LG3889); F, lower leaf surface of A. mayottensis I.Montero & Cardiel (J.-N. Labat et al. 3268). Scale bars: A, 100 µm; B, 50 µm; C, 30 µm; D, F, 10 µm; E, 200 µm.
410 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. B D E A F G C H fig. 12. — A, B, Epicuticular waxes, layers: A, Acalypha claoxyloides Hutch. (F. R. Fosberg 49627); B, A. radula Baker (J. M. Hildebrandt 3865); C-H, crystalloids; C, granules in A. filiformis Poir. (M. J. E. Coode 5062); D, granules in A. pervilleana Baill. (A. Mocquerys 159); E, Platelets in A. lamiana (Leandri) I.Montero & Cardiel (H. J. Lam 6127); F, Platelets in A. richardiana Baill. (J. M. Hildebrandt 1661); G, Plates in A. emirnensis Baill. (S. Rakotonandrasana 1279); H, Plates in A. urophylla Boivin ex Baill. (J. M. Hildebrandt 2914). Scale bars: A, H, 20 µm; B, D, F, 10 µm; C, E, 5 µm; G, 2 µm.
411 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) that cover the entire surface; they vary in thicknesses and can appear folded or fractured (Fig.12A, B). Crystalloids form a discontinuous cover, are grouped or solitary, have different shapes and sizes, and can appear together with other types of waxes (Barthlott etal. 1998). In the WIOR species we have found three types of crystalloids: granules, platelets, and plates. Granules are crystalloids with irregular shapes, but usually isodiametric and rounded (Fig.12C, D). Platelets and plates are flat, plate-shaped crystalloids; they are the most common crystalloids in plants (Barthlott etal. 1998). Platelets have irregular margins (Fig.12E, F), whereas plates are polygonal and have entire margins (Fig.12G, H). We do not yet know what, if any, taxonomic significance epicuticular waxes have in Acalypha. TAXONOMIC TREATMENT Family euphorbiaceae Juss. Genus Acalypha L. Species Plantarum: 1003 (Linnaeus 1753); Genera Plantarum, ed. 5: 436 (Linnaeus 1754).— Type: Acalypha virginica L. (lectotype designated by Small 1913: 457).— An Illustrated Flora of the Northern United States, ed. 2., 2: 457. Mercuriastrum Heist. ex Fabr., Enumeratio Methodica Plantarum: 202 (Fabricius 1759).— Type: not designated. Cupameni Adans., Familles des Plantes (Adanson), 2: 356 (Adanson 1763), nom. illeg. et superfl.— Type: not designated. Caturus L., Mantissa Plantarum. Generum Editionis vi et Specierum Editionis II: 19 (Linnaeus1767).— Galurus Spreng., Anleitung zur Kenntniss der Gewachse, Zweite 364 (Sprengel 1817) nom. illeg.— Type: Caturus spiciflora L. [syn. of Acalypha hispida Burm.f.]. Usteria Dennst., Schlüssel Hortus Malabaricus: 31 (Dennstaedt 1818); non Usteria Willd., nec Usteria Medikus, nec Usteria Cav.— Type: not designated. table 1. — Crystals found in the studied species of Acalypha. Abbreviations: FC, foliar crystals; EC, extra-foliar crystals; Ac, prismatic crystals; Cc, stellate crystals; Dc, druses; Ec, granules; 0, crystals not found; –, material not examined). Subgenera Species FC EC Androcephala A. diminuta Baill. Ac, Dc Dc Hypandrae A. baretiae I.Montero & Cardiel Ac 0 Acalypha A. andringitrensis Leandri Dc – A. ankaranensis I.Montero & Cardiel Ac – A. chibomboa Baill. Ac – A. claoxyloides Hutch. Ac, Ec – A. crateriana (Coode) I.Montero & Cardiel, comb. nov. Ac – A. decaryana Leandri Ac – A. emirnensis Baill. Ac, Dc, Ec – A. filiformis Poir. Ac – A. fimbriata Schumach. &Thonn. Ac, Dc Cc, Dc A. hispida Burm.f. Ac – A. indica L. Dc – A. lamiana (Leandri) I.Montero & Cardiel 0 – A. leandrii I.Montero & Cardiel Ac – A. leptomyura Baill. 0 Ac A. linearifolia Leandri 0 – A. marginata (Poir.) Spreng. Ac – A. mayottensis I.Montero & Cardiel Ac, Ec Ac A. menavody (Leandri) I.Montero & Cardiel Ac – A. nusbaumeri I.Montero & Cardiel. Ac – A. perrieri Leandri 0 – A. perrieri aff. Leandri Ac – A. pervilleana Baill. Ac, Dc – A. poiretii Spreng. Ac – A. rabesahalana I.Montero & Cardiel Ac Ac A. radula Baker Dc – A. richardiana Baill. 0 – A. rottleroides Baill. Ac – A. indet. 1 Ac, Dc – A. indet. 2 Ac Dc A. tremula I.Montero & Cardiel Ac – A. urophylla Boivin ex Baill. Ac – A. vulneraria Baill. Ac, Dc – A. wilkesiana Müll.Arg. Ac – table 2. — Epicuticular waxes (EW) found in the studied species of Acalypha. Abbreviations: L, layers; P, platelets; Pl, plates; G, granules; 0, epicuticular waxes not seen. Subgenera Species EW Androcephala A. diminuta Baill. L Hypandrae A. baretiae I.Montero & Cardiel L Acalypha A. andringitrensis Leandri L A. ankaranensis I.Montero & Cardiel G A. chibomboa Baill. G A. claoxyloides Hutch. L A. crateriana (Coode) I.Montero & Cardiel, comb. nov. G A. decaryana Leandri L A. emirnensis Baill. G, Pl A. filiformis Poir. G A. fimbriata Schumach. &Thonn. L A. hispida Burm.f. L, P A. indica L. L, P A. lamiana (Leandri) I.Montero & Cardiel P A. leandrii I.Montero & Cardiel G A. leptomyura Baill. G A. linearifolia Leandri - A. marginata (Poir.) Spreng. L A. mayottensis I.Montero & Cardiel. L A. menavody (Leandri) I.Montero & Cardiel L A. nusbaumeri I.Montero & Cardiel. L A. perrieri Leandri - A. perrieri aff. Leandri L, Pl A. pervilleana Baill. L A. poiretii Spreng. L A. rabesahalana I.Montero & Cardiel L A. radula Baker L A. richardiana Baill. P A. rottleroides Baill. L, Pl A. indet. 1 L, P A. indet. 2 L A. tremula I.Montero & Cardiel G A. urophylla Boivin ex Baill. L, Pl A. vulneraria Baill. L A. wilkesiana Müll.Arg. L, P
412 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. Cupamenis Raf., Sylva Telluriana. Mantis Synopt. New genera and species of trees and shrubs of North America: 67 (Rafinesque 1838), non Cupameni Adans.— Type: Acalypha indica L. Acalyphes Hassk., Catalogus Plantarum in Horto Botanico Bogoriensi Cultarum Alter. Bataviae: 235 (Hasskarl 1844), orth. var. Linostachys Klotzsch ex Schltdl., Linnaea 19: 235 (Schlechtendal 1847).— Type: Linostachys padifolia Schltdl. [syn. of Acalypha schlechtendaliana Müll.Arg.] Odonteilema Turcz., Bulletin de la Société impériale des Naturalistes de Moscou 21: 587 (Turchaninov 1848).— Type: Odonteilema clausseni Turcz. [syn. of Acalypha clausseni (Turcz.) Baill.]. Paracelsea Zoll., Natuurkundig Tijdschrift voor Nederlandsch-Indië, 14(4): 171 (Zollinger 1857), nom. illeg. Calyptrospatha Klotzsch ex Baill., Étude générale du Groupe des Euphorbiacées: 440 (Baillon 1858).— Type: Calyptrospatha pubiflora Klotzsch [syn. of Acalypha pubiflora (Klotzsch) Baill.]. Gymnalypha Griseb., Bonplandia (Hannover) 6: 2 (Grisebach 1858).— Type: Gymnalypha jacquinii Griseb., nom. illeg. [syn. of Acalypha villosa Jacq.]. Corythea S.Watson, Proceedings of the American Academy of Arts and Sciences, Boston 22: 451 (Watson 1887).— Type: Corythea filipes S.Watson [syn. of Acalypha filipes (S.Watson) McVaugh]. Ricinocarpus Burm. ex Kuntze, Revisio Generum Plantarum 2: 615 (Kuntze 1891), nom. illeg., non Ricinocarpos Desf.— Type: Acalypha lanceolata Willd. Schizogyne Ehrenb. ex Pax& K.Hoffm., Das Pflanzenreich (Engler), IV, 147, XVI: 178 (Pax& Hoffmann 1924), nom. illeg., non Schizogyne Cass.— Type: Schizogyne ciliata Ehrenb. [syn. of Acalypha ciliata Forssk.]. Acalyphopsis Pax& K.Hoffm., Das Pflanzenreich (Engler), IV, 147, XVI: 178 (Pax& Hoffmann 1924).— Type: Acalyphopsis celebica Pax& K.Hoffm., nom. dub. [syn. of Acalypha hoffmanniana Hurus.]. DisTribuTion, habiTaT, anD classificaTion.— A pantropical genus of about 500 species, with centers of diversity in Mexico and east tropical Africa, and including 49 species in WIOR (42 native and 7 probably introduced). Acalypha is distributed in all the archipelagos in the WIOR and grows in a wide variety of habitats. Following the subgeneric classification of Levin etal. (2022), subg. Androcephala Pax& K.Hoffm. is represented by one species (A.diminuta), subg. Hypandrae (Müll.Arg.) Hurus. by one species (A.baretiae), and subg. Acalypha by 47 species. The fourth subgenus, Linostachys (Klotzsch ex Schltdl.) Pax& K.Hoffm., does not occur in WIOR. Madagascar is the only island in WIOR where three subgenera are present; other regions of WIOR are home only to species in subg. Acalypha. The species in subg. Acalypha found in WIOR do not represent a monophyletic group (Montero Muñoz 2021; Levin etal. 2022). DescripTion Small trees, shrubs, or rarely herbs, evergreen or deciduous, monoecious or rarely dioecious. Branches usually slender, rarely thorny or with brachyblasts. Indumentum velutinous, pubescent or sparsely hairy, rarely absent (glabrous); trichomes simple (long or short, erect, curved, or appressed, sometimes hyaline), glandular (long, short, or resinous flattened/rounded), rarely stellate. Axillary buds naked or perulate, then with two perules that are imbricate or valvate or four perules that are overlapping (superposed), chartaceous or membranous. Stipules usually caducous, linear to triangular or ovate-lanceolate. Leaves alternate, undivided, usually petiolate and stipulate. Leaf blades entire or more commonly dentate or crenate-serrate, rarely lobed; venation actinodromous or pinnate. Stipels sometimes present at apex of petiole or leaf base, minute, glandular, filiform or triangular-lanceolate. Inflorescences unisexual or bisexual, axillary, terminal or both, usually spiciform, less commonly racemose or glomerate. Male inflorescences always axillary, spiciform, densely or laxly flowered, with several flowers at each node, subtended by a minute bract. Female inflorescences usually spiciform, sometimes glomerulate, bracteate, with 1-3 flowers per node; bracts minute at anthesis, usually accrescent and foliaceous in the fruit, sometimes non-accrescent, margin entire, crenate, or dentate, sometimes with two bracteoles at base. Bisexual inflorescences usually spiciform, sometimes racemose, usually with female flowers below and male above (androgynous), rarely with male flowers below and female above (gynecandrous). Flowers unisexual, apetalous, disc absent. Male flowers inconspicuous, short-pedicellate; calyx 4-lobed, stamens 8, thecae 2, pendulous and vermiform at anthesis; pollen small, oblate-spheroidal to suboblate, apertures 3-8, brevicolporate; pistillode absent. Female flowers usually sessile or subsessile, rarely pedicellate; calyx of 3[-4] or sometimes 5 small sepals, distinct or connate at base; ovary with [2]-3 carpels, surface smooth, papillose, or echinate, usually pubescent to densely hispid, sometimes glabrous; styles [2]-3, usually reddish, distinct or connate at base, each divided in several filiform segments; stigmas smooth; ovules 1 per locule. Allomorphic female flowers sometimes present, solitary, at apex of the inflorescences or axillary; ebracteate, long-pedicellate or subsessile, calyx as in the normal female flowers, ovary 1-2[-3] lobed, distally fimbriate or papillose, styles 1-2[-3], subbasal (rarely terminal). Fruits from normal flowers capsular, [2]-3 lobed, soon dehiscing septicidally into 2-3 bivalved cocci; usually surrounded by the accrescent female bract; surface smooth, papillose, or echinate, usually pubescent to hispid, sometimes glabrous; those from allomorphic flowers indehiscent or schizocarps with indehiscent mericarps, usually distally fimbriate or sharply papillate. Seeds usually piriform, sometimes subglobose or globose, minutely foveolate or foveolate, caruncle usually very reduced. Taxonomic resulTs We found a total of 163 names associated with Acalypha of WIOR (Appendix II) published prior to the beginning of our work, of which we accept 36 names (including three new combinations, one made in this paper), 92 are considered as synonyms (7 for the first time here), and 2 are excluded from the genus. The remaining names are invalid or excluded names. Finally, in previous works, we have described 12 new Acalypha species (Montero Muñoz etal. 2018b, 2020a, 2020b, 2022, and another one in this paper).
413 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) Key To The species of AcAlyphA of WesTern inDian ocean region (Wior) 1. Female flowers pedicellate, calyx with 5 sepals; female bracts not enlarging in fruit (subg. Androcephala and Hypandrae) .................................................................................................................................................. 2 — Female flowers sessile, calyx with 3[4] sepals; female bracts enlarging in fruit, except in A.crateriana comb.nov., A.hispida, and A.marginata (subg. Acalypha) .............................................................................................. 3 2. Inflorescences gynecandrous; male segment 2-3(-3.5) cm long, flowers glomerate .......................................... ..................................................................................................................... A.baretiae I.Montero& Cardiel — Inflorescences androgynous; male segment to 0.5cm long, flowers in a racemose to subumbelliform cluster ...... ........................................................................................................................................... A.diminuta Baill. 3. Inflorescences all unisexual .......................................................................................................................... 4 — Inflorescences mostly androgynous (sometimes with the male segment caducous), rarely male inflorescences and/or solitary axillary female bracts .......................................................................................................... 21 4. Female inflorescences sessile or subsessile, with 1 bract that enlarges in fruit, or flowers glomerate and bracts not enlarging ............................................................................................................................................... 5 — Female inflorescences pedunculate (peduncle more than 5mm long, usually more than 10mm long), with more than 2 bracts, usually many, rarely 1 ............................................................................................................ 9 5. Leaf blades linear, c.0.2cm wide .................................................................................. A.linearifolia Leandri — Leaf blades not linear, 1-7(-8) cm wide ....................................................................................................... 6 6. Petioles (2-)3-4.5cm long; leaf blades membranous, with pocket-shaped domatia on the lower surface ......... ................................................................................................................... A.isaloensis I.Montero& Cardiel — Petioles 0.5-2.5(-3) cm long; leaf blades chartaceous or coriaceous (the mature leaves), without domatia .... 7 7. Leaf blades reddish on lower surface, without discolorous margin; stipules triangular-lanceolate, with two translucent lateral wings; female inflorescences with 1 bract enlarging in fruit ........................ A.integrifolia Willd. — Leaf blades greenish on lower surface, usually with discolorous margin; stipules linear-lanceolate, without wings; female inflorescences in glomerules of 3-4 flowers with bracts not enlarging in fruit .................................... 8 8. Leaf blades to 10cm long, elliptic to obovate, with reddish margin ................................................................ ................................................................................. A.crateriana (Coode) I.Montero& Cardiel, comb.nov. — Leaf blades to 21cm long, elliptic-lanceolate, with usually pale margin .......................................................... ........................................................................................................................... A.marginata (Poir.) Spreng. 9. Female inflorescences with (1-)2-4 bracts, to 6 bracts in A.chibomboa ...................................................... 10 — Female inflorescences with more than (4-)9 bracts ..................................................................................... 13 10. Leaf blades 1.5-2.5(-4)[-7.5] cm long, elliptic to oblong, sometimes obovate ................................................. ...................................................................................................................................... A.decaryana Leandri — Leaf blades (5-)7-12(-18)[-26.5] cm long, elliptic-lanceolate to ovate-lanceolate, sometimes obovate-lanceolate ............................................................................................................................................................ 11 11. Female bracts to 5mm long, orbicular, denticulate ........................................... A.chibomboa Baill. (in part) — Female bracts to 9-15mm long, orbicular-reniform, dentate ..................................................................... 12 12. Leaf blades elliptic-lanceolate to obovate-lanceolate; base acute to cuneate ........................ A.emirnensis Baill. — Leaf blades ovate-lanceolate; base rounded to subcordate ......................................... A.lepidopagensis Leandri 13. Female bracts with only simple trichomes ................................................................................................. 14 — Female bracts with simple and glandular trichomes or only with glandular trichomes ............................... 16 14. Female inflorescences subterminal, laxly flowered, c.3cm long ............ A.rabesahalana I.Montero& Cardiel — Female inflorescences axillary, densely flowered, 10-40cm long ................................................................ 15 15. Leaf blades not variegated; female inflorescences up to 40cm long, pendulous; female bracts not increasing in fruit ....................................................................................................................................... A.hispida L. — Leaf blades variegated; female inflorescences up to 10cm long, more or less erect; female bracts increasing in fruit ...................................................................................................................... A.wilkesiana Müll.Arg. 16. Leaf blades bullate ..................................................................................................................................... 17 — Leaf blades not bullate ............................................................................................................................... 18
414 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. 17. Leaf blades ovate-lanceolate; stipules to 13mm long, elliptic to ovate-elliptic ........ A.andringitrensis Leandri — Leaf blades narrowly triangular-lanceolate to linear-lanceolate; stipules to 8mm long, triangular-lanceolate .............................................................................................................................................. A.radula Baker 18. Leaf blades with simple and glandular trichomes ................................................ A.rottleroides Baill. (in part) — Leaf blades with only simple trichomes ..................................................................................................... 19 19. Leaf blades 2-6(-13) cm long; female bracts dentate ............................................. A.spachiana Baill. (in part) — Leaf blades (3.5-)7-17cm long; female bracts entire ................................................................................. 20 20. Petioles (2-)3-8cm long; leaf blades broadly ovate-lanceolate, base rounded to cordate; female inflorescences to 8.5cm long .................................................................................................. A.leandrii I.Montero& Cardiel — Petioles (1-)2-3cm long; leaf blades ovate-lanceolate to oblong-lanceolate, base rounded to truncate; female inflorescences to 13cm long .............................................................................................. A.vulneraria Baill. 21. Androgynous inflorescences with inconspicuous male segment (3.5-15mm long) ..................................... 22 — Androgynous inflorescences with conspicuous male segment (20-90mm long) ......................................... 31 22. Annual or perennial herbs [allochthonous species] .................................................................................... 23 — Shrubs [native species] ............................................................................................................................... 27 23. Female bracts with only simple trichomes; margin crenate-denticulate ........................................ A.indica L. — Female bracts with simple and glandular trichomes; margin dentate .......................................................... 24 24. Female bracts with c.23 teeth, teeth falcate .............................................. A.fimbriata Schumach.& Thonn. — Female bracts with 7-15 teeth, teeth straight ............................................................................................. 25 25. Female bracts 2-5, central tooth prominent .............................................................. A.bailloniana Müll.Arg. — Female bracts 7-15, central tooth not prominent ....................................................................................... 26 26. Male inflorescence segment c.2mm long; female bracts 12-15, to 2× 4mm, teeth 9-11, broadly triangular ............................................................................. A.lanceolata Willd. var. glandulosa (Müll.Arg.) Radcl.-Sm. — Male inflorescence segment to 10mm long; female bracts 16-20, to 5× 6mm, teeth 7-8, narrowly triangular .................................................................................................................................. A.poiretii Spreng. 27. Leaf blades and/or female bracts with glandular trichomes; androgynous inflorescences with male segment deciduous (inflorescences may appear to be female) ...................................................................................... 28 — Leaf blades and female bracts with only simple trichomes; androgynous inflorescences with male segment persistent ........................................................................................................................................................ 29 28. Leaf blades with glandular trichomes; androgynous inflorescences axillary; female bracts subentire to crenate ........................................................................................................................... A.rottleroides Baill. (in part) — Leaf blades without glandular trichomes; androgynous inflorescences terminal (on lateral branches, sometimes appearing leaf-opposed); female bracts dentate .................................................... A.spachiana Baill. (in part) 29. Androgynous inflorescences to 9cm long, with 4-15 female bracts ................................. A.boinensis Leandri — Androgynous inflorescences 1-2cm long, with 1 female bract ................................................................... 30 30. Female bracts to 5× 9mm, entire; bracteoles absent; inflorescences with male segment to 3.5mm ................ ............................................................................................................. A.gillespieae G.A.Levin& I.Montero — Female bracts to 2× 2.5mm, crenate; bracteoles linear-lanceolate; inflorescences with male segment to 7mm ............................................................................................................... A.nusbaumeri I.Montero& Cardiel 31. Androgynous inflorescences with 1 female bract at base; flattened resinous glands absent .......................... 32 — Androgynous inflorescences with 2 or more female bracts at base, sometimes 1 in A.claoxyloides and A. medibracteata; flattened resinous glands present or absent (if androgynous inflorescences with 1 female bract, then flattened resinous glands present) .............................................................................................................. 46 32. Solitary axillary female bracts absent; androgynous inflorescences sessile or pedunculate, peduncle more than (2.5-)5mm long ....................................................................................................................................... 33 — Solitary axillary female bracts present; androgynous inflorescences usually sessile or subsessile, peduncle to 3mm long (to 10mm in A.filiformis) ................................................................................................................. 38 33. Androgynous inflorescences to 7cm long, sessile or subsessile (peduncle to 2.5mm in A.richardiana); axillary buds to 1mm long with perules chartaceous or membranous ................................................................... 34 — Androgynous inflorescences to 3(-6) cm long, with peduncle 5-15mm long; axillary buds to 2-4mm long with perules membranous ................................................................................................................................. 36
415 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) 34. Leaf blades broadly ovate to subtriangular; capsules to 3mm diameter with long projections to 3.5mm long ..................................................................................................................... A.tremula I.Montero& Cardiel — Leaf blades elliptic to ovate-lanceolate; capsules to 2mm diameter without long projections (only with papillae to 0.5mm long) ........................................................................................................................................ 35 35. Branches erect, densely pubescent; petioles 2-6(-9) cm long, no canaliculate, densely pubescent ................... .......................................................................................................................... A.richardiana Baill. (in part) — Branches flexuous, glabrous; petioles (0.4-)0.7-1.4 (-1.7) cm long, canaliculate, glabrous ............................... .............................................................................................................................. A.gracilipes Baill. (in part) 36. Female bracts to 8mm wide, dentate to subentire; axillary buds subfusiform ................ A.humbertii Leandri — Female bracts to 12mm wide, entire; axillary buds spherical ..................................................................... 37 37. Leaf blades 5-10cm long, acuminate to subacuminate .............. A.mayottensis I.Montero& Cardiel (in part) — Leaf blades 3.5-6(-8.5) cm long, abruptly acuminate ......................................................... A.perrieri Leandri 38. Leaf blades (6-)7-17(-22) cm long; branches never flexuous ...................................................................... 39 — Leaf blades (1.5-)2.5-7.5(-10) cm long (rarely to 11cm long in A.urophylla), if longer than 7cm long, then branches usually flexuous .......................................................................................................................... 41 39. Leaf blades broadly ovate-lanceolate, base cordate .................. A.ankaranensis I.Montero& Cardiel (in part) — Leaf blades ovate-lanceolate to elliptic-lanceolate, base rounded to obtuse ................................................. 40 40. Female bracts usually to 10mm long (sometimes to 20mm), margin dentate with or without prominent central tooth .................................................................................................................. A.pervilleana Baill. (in part) — Female bracts to 7mm long, margin subentire, without prominent central tooth ....................................... ....................................................................................................................... A.richardiana Baill. (in part) 41. Solitary female bracts long-pedunculate .............................................................................. A.filiformis Poir. — Solitary female bracts sessile ...................................................................................................................... 42 42. Branches glabrous; mature female bracts to 4× 3.5mm, bracteoles present; leaf blade apices obtuse, lower surface glabrous; male bracts glabrous; androgynous inflorescences to 2cm long ................................................. ........................................................................................ A.berryi I.Montero, Cardiel& G.A.Levin, sp.nov. — Branches hairy, glabrescent; mature female bracts to 5-10× 5-12mm, bracteoles absent; leaf blade apices subacuminate to acuminate, lower surface hairy, at least on veins, sometimes glabrescent; male bracts sparsely hairy; androgynous inflorescences to 4-6cm long ............................................................................................... 43 43. Leaf blades (1.5-)2.5-6.5cm long, ovate-lanceolate to subtriangular, acumen rounded .............................. 44 — Leaf blades (4-)5-7.5(-11) cm long, ovate to elliptic-lanceolate, acumen acute or mucronate ..................... 45 44. Leaf blades (1.5-)2.5-3.5(-4.5) cm long; lower leaf surface and female bracts with only simple trichomes; female bracts with margin entire to crenate ................................................................................ A.leptomyura Baill. — Leaf blades (3.5-)4-6.5cm long; lower leaf surface and female bracts with simple and minute glandular trichomes; female bracts with margin dentate .................................................... A.levinii I.Montero& Cardiel (in part) 45. Branches straight; female bracts glabrous with reddish margin ...... A.lamiana (Leandri) I.Montero& Cardiel — Branches usually flexuous; female bracts pubescent without reddish margin ........ A.urophylla Boivin ex Baill. 46. Leaf blades and female bracts with flattened resinous glands; ovary and capsule smooth. ............................47 — Leaf blades and female bracts without resinous glands; ovary and capsule papillose to erinaceous. .............. 49 47. Androgynous inflorescences with 2-6 female bracts; female bracts c.5mm long ............... A.chibomboa Baill. — Androgynous inflorescences with 1-2(-3) female bracts; female bracts 10-12mm long .............................. 48 48. Leaf blades 3.5-5(-7) cm long, linear-lanceolate to oblong-lanceolate ...... A.medibracteata Radcl.-Sm.& Govaerts — Leaf blades (5-)6-11(-14) cm long, ovate-lanceolate to elliptic-lanceolate ..................... A.claoxyloides Hutch. 49. Inflorescences androgynous and male ........................................................................................................ 51 — All the inflorescences androgynous ............................................................................................................ 55 51. Androgynous inflorescences with peduncle 10-15mm long; female bracts with rounded teeth ................. 52 — Androgynous inflorescences sessile; female bracts with acute teeth ............................................................. 53 52. Leaf blades 1.8-2.6(-3) cm long, subrhomboidal ..................................... A.cardielii I.Montero& G.A.Levin — Leaf blades 7-10(-12.5) cm long, elliptic-lanceolate to subobovate ........................................... A.leonii Baill.
416 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. Acalypha subgen. Androcephala Pax& K.Hoffm. Das Pflanzenreich (Engler), 147, 16 (Heft 85): 21 (Pax& Hoffmann 1924).— Type: Acalypha diminuta Baill. DescripTion Inflorescences terminal, androgynous; female segment racemose or paniculate; male segment racemose to subumbelliform (in WIOR species). Female bracts minute, non-accrescent in fruit. Female flowers pedicellate; sepals 5; ovary 3-lobed, styles 3. 1. Acalypha diminuta Baill. Histoire physique, naturelle et politique de Madagascar, Atlas, 2, fasc.27: t. 194 (Baillon 1891).— Type: pl. 194 in Baillon (1891), holotype. iconography.— Baillon (1891: pl. 194); Leandri (1942: 254); Fig. 28A. e Tymology .— The specific epithet probably refers to the small size of the male segment of the inflorescences. DisTribuTion anD habiTaT.— Endemic to Madagascar (Diana, Sofia, Boeny, Melaky, Menabe, Atsimo-Andrefana, Androy, and Alaotra-Mangoro [only one collection]). Dry deciduous forest, spiny thickets; sometimes in riparian forest. On Tertiary limestone, sandstone, and unconsolidated sand. Altitudinal range (40-) 50-850 (-1450) m (Fig. 13). p reliminary conservaTion assessmenT .— The EOO of Acalypha diminuta is estimated to be 310 217km2 and its AOO 104km2, which is less than the 500km 2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in the western dry deciduous forests. Most of these forests are highly degraded (40%) and the major threat to the dry deciduous forests is slash-and-burn clearing for grazing and agricultural lands. Grasslands and pastures have replaced the forests in the western slope and the central highlands (Moat& Smith 2007; Gautier etal. 2018). Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha diminuta is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for listing it under the Endangered category. maTerial examineD.— 44 collections. Madagascar. Appert, O.P. 31 (P[P04779831]), 41 (P[P00360914]); Bosser, M.J. 13885 (P[P04779814, P04779816]), 13993 (P[P04779821, P04779824]); Capuron, R. 20581SF (K, P[P05547246]); Cours, G. 1819 (P[P05547244]); d’Alleizette, Ch. s.n. (L[L0241302]); Decary, R. 16089 (K, P[P05547245]), 16122 (P[P05547250]); Gillespie, L. 4034 (US[US01287336]), 10704 (MO); Grevé, H. 223 (P[P00360915]); Hladik, A. AH7158 (P[P00780280, P00780618]); Humbert, H. 11237 (B[B100009884], P[P00508516, P00360918]), 11288 (P[P00360919]), 11335 (BR[BR0000005045020], NY, P[P00508515, P00360917]), 11506 (P[P00360916, P00508517]); Jard. Bot. Tananarive 5749 (P[P00360923]); Keraudren-Aymonin, M. 25933 (P[P00360920]); Leandri, J. 489 (K, P[P05547061]), 491(US[US01287337]), 3533(P[P05547060]), 3569 (B[B100480050], G, NY, P[P05547243]), 3620 (G, P[P05547242]); Lorence, D. 2084 (K, P[P00360921]); Peltier, J. 1432 (P[P05547022]); Perrier de la Bâthie, H. 379 (P[P05577069, P05577067]), 9632 (P[P00508520, P00508521, P00360922]), 9816 (P[P00508519, P00360924, P00360925]), 9818 (P[P00360499, P00360500, P00508518]), 9819 (P[P05547062]); Phillipson, P.B. 2431 (MO, P[P00360497]), 2750 (K, MO, P[P00360498]); Pichon, C. CP113 (P[P00780436]); Rakotozafy, A. 319 (P[P05547065]); Ralimanana, H. 239 (G[G00405396], DBEV, K, MO[MO-2965792], P[P05481882], TAN); Richard, M. 173 (K), 547 (K); Service Forestier Madagascar 16576-SF (P[P00360912, P00508522, P00508523]), 27956-SF (P[P00360913]); Seyrig, A. 254 (P[P05547066]), 254B (P[P05547064]), 254 C (P[P05547063]); Wohlhauser, S. SW 310 (G[G00405345], P[P04779528]). 53. Leaf blades 6-16(-21) cm long, apex long-acuminate to caudate; some androgynous inflorescences terminal ........................................................................................................................... A.pervilleana Baill. (in part) — Leaf blades (4-)5-7.5cm long, apex shortly acuminate; all androgynous inflorescences axillary ................. 54 54. Petioles (0.4-)0.7-1.4cm long; leaf blades elliptic-lanceolate; capsules to 2mm diameter, without long projections (only with papillae to 0.5mm long) ............................................................. A.gracilipes Baill. (in part) — Petioles (3-)4-6cm long, leaf blades broadly ovate to subtriangular; capsule to 3mm diameter with long projections to 3.5mm long ...................................................................... A.tremula I.Montero& Cardiel (in part) 55. Androgynous inflorescences sessile or subsessile ......................................................................................... 56 — Androgynous inflorescences pedunculate (peduncle more than 15mm long) ............................................ 57 56. Leaf blades (6-)7-9cm long, broadly ovate-lanceolate, subcordate to cordate at the base; female bracts with acute to subacute teeth, with sessile glands at apex .......................... A.ankaranensis I.Montero& Cardiel (in part) — Leaf blades (3.5-)4-6.5cm long, ovate-lanceolate to elliptic-lanceolate, rounded to subcordate at the base; female bracts with rounded teeth without glands ......................................... A.levinii I.Montero& Cardiel (in part) 57. Female bracts c.5mm long ......................................................................... A.magistri I.Montero& Cardiel — Female bracts 7-19mm long ..................................................................................................................... 58 58. Leaf blades broadly ovate-lanceolate; androgynous inflorescences with peduncle to 45mm long .................................................................................................. A.menavody (Leandri) I.Montero& Cardiel — Leaf blades ovate-lanceolate to elliptic-lanceolate; androgynous inflorescences with peduncle to 15mm long .... 59 59. Leaf blades 2.5-5.5cm long; female bracts to 10× 11mm, with discolorous margin ..................................... ................................................................................................................. A.burmanii I.Montero& Cardiel — Leaf blades (3-)5-10cm long; female bracts to 19× 21mm, without discolorous margin .............................. .................................................................................................. A.mayottensis I.Montero& Cardiel (in part)
417 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) references.— Baillon (1895b: 1197); Palacký (1907: 25); Pax& Hoffmann (1924: 21); Leandri (1935: 42); Leandri (1942: 253); Govaerts etal. (2000: 60); Sagun etal. (2006: 124); Montero Muñoz etal. (2018a: 95). DescripTion Shrubs, deciduous, to 3m tall, monoecious. Branches laxly pubescent with simple, short, curved trichomes and simple, erect trichomes, glabrescent when mature; brachyblasts present. Axillary buds subspherical, to 1.2× 1mm, perulate, perules 2, imbricate, chartaceous, brownish, glabrous. Stipules to 3mm long, lanceolate to linear-lanceolate, ciliate with some sparse, simple, short trichomes. Petioles 0.2-0.3cm long, indumentum similar to that on young branches. Leaf blades 3-5(-5.5)× 1.42cm, usually elliptic to elliptic-lanceolate, sometimes narrowly elliptic-lanceolate, membranous; base rounded to cordate; apex obtuse and mucronate; margin entire to crenate towards apex, teeth rounded; upper surface subglabrous, with some sparse, simple trichomes on veins; lower surface glabrous, with flattened resinous glands; venation pinnate, secondary veins 5-7 per side. Stipels absent. Inflorescences racemose, androgynous, terminal, to 5cm long, mostly female with short male segment; peduncle filiform, to 30mm long, appressed-pubescent. Female segment to 2cm long; bracts 1-7, not enlarging in fruit, to 1mm long, lanceolate, sparsely hairy; bracteoles absent. Male segment persistent, to 0.5cm long; flowers in racemose to subumbelliform cluster; bracts to 0.8mm long, triangularlanceolate, sparsely hairy with simple, arachnoid trichomes. Male flowers: pedicel to 0.8 mm long, sparsely hairy; buds to 1 mm diameter, sparsely hairy with simple, arachnoid trichomes. Female flowers 1 per bract, pedicellate; pedicel to 7mm long, indumentum similar to that on peduncle; sepals 5, to 0.8mm long, triangular, ciliate with simple trichomes to 0.5mm long; ovary c.1 mm diameter, 3-lobed, papillose, surface glabrous but with flattened resinous glands; styles 3, to 2.5mm long, distinct, glabrous, each divided into c.10 segments. Allomorphic flowers not seen. Capsules to 4mm diameter, papillose, glabrous, with papillae more or less capitate, surface glabrous but with flattened resinous glands. Seeds c.2 mm diameter, globose, minutely foveolate. noTes 1)Acalypha diminuta was first illustrated, without description, in Baillon’s Histoire Naturelle des Plantes (Baillon 1891). The first description of this species, based only on Baillon’s illustration, appears in Engler’s Pflanzenreich (Pax& Hoffmann, 1924); and 2) this is one of only two species included in Acalypha subgen. Androcephala Pax& K.Hoffm.; the other is A.gillmanii Radcl.-Sm. of eastern Africa (Levin etal. 2022). Acalypha subgen. Hypandrae (Müll.Arg.) Hurus. Journal of the Faculty of Science, University of Tokyo, Section III. Botany 6: 297 (Hurusawa 1954).— Acalypha subser. Hypandrae Müll. Arg., Linnaea 34: 9 (Müller Argoviensis 1865).— Type: Acalypha sonderiana Müll.Arg. (lectotype designated by Levin etal. 2022). DescripTion Inflorescences axillary, gynecandrous, racemose or paniculate. Female bracts minute, non-accrescent in fruit. Female flowers pedicellate; sepals 5; ovary 2-lobed, styles 2. 2. Acalypha baretiae I.Montero& Cardiel South African Journal of Botany 119: 420 (Montero Muñoz etal. 2018b).— Type: Madagascar. Toliara. Morondava, 10km de Morondava, route de Malaimbandy, 337m, 20°23’33”S, 44°56’57”E, X.1963, M. J. Bosser 18124 (holo-, P[P04779795]).— Paratypes: Madagascar, Toliara: Atsimo-Andrefana region. Beroroha, fokontany Betorabato. Village le plus proche Ambalamanga. Forêt Akolitsika (rivière), 276m, 21°40’22”S, 44°59’36”E, 18.I.2011. R. Razakamalala 6064 (MO[MO-3025026], P[P01152840], TAN). iconography.— Montero Muñoz etal. (2018b). eTymology.— The epithet honors Jeanne Baret (1740-1807), the first woman to circumnavigate the globe. Disguised as a man and calling herself Jean Baret, she served as naturalist Philibert Commerson’s valet and botanical assistant on Louis Antoine de Bougainville’s expedition from 1766 to 1769, the first French circumnavigational voyage. She visited Madagascar with Commerson in 1770-1772, 45°E 15°S 20° 25° 300 km 50° Acalypha diminuta fig. 13. — Distribution map of Acalypha diminuta Baill. in Madagascar.
424 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. 7. Acalypha boinensis Leandri Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 268 (Leandri 1942).— Type: Madagascar. Prov. Mahajanga, Ambongo et Boïna, bassin moyen du Bemarivo, 200m, XI.1919, H. Perrier de la Bâthie 9823 (lecto-, designated by Montero Muñoz etal. (2018a: 93): P[P00508582]).— Former syntypes: Madagascar. Prov. Mahajanga, Maintirano, s.d., R. Decary 8216 (P[P00508588]); Maromandia (Andranosamontana), s.d., R. Decary 1045 (P[P00508585]); Morovoay, s.d., H. Humbert& H. Perrier de la Bâthie 2350 (P[P00508570, P00508571, P00508572, P00508573, P00508574]); Ambongo et Boïna, 300m, s.d., H. Perrier de la Bâthie 9546 (P[P00508576]), 9551 (P[P00508577]); Prov. Toliara, Forêt de Besomaty, entre le Fiherena et l’Isahaina (Mangoky), 750-800m, s.d., H. Humbert 11234 (P[P00508575]).— s.l., s.d., Baron 5393 (P[P00508586]), 5450 (P[P00508587]). iconography.— Leandri (1942: 257); Figs 24D; 28D, E. eTymology.— The epithet refers to the former Iboina or Boina Region (now Mahajanga province) in northwestern Madagascar. D isTribuTion anD habiTaT .— Endemic to Madagascar (Sava, Sofia, Boeny, Betsiboka, Melaky, Menabe, Atsimo-Andrefana, Haute Matsiatra, Ihorombe, and Androy). Dry deciduous forest, dry spiny thickets, and medium altitude moist evergreen forest (Analavelona). On Mesozoic limestone, basalt, and sandstone. Altitudinal range (50-)200-1000(-1250) m (Fig. 20). preliminary conservaTion assessmenT.— The EOO of Acalypha boinensis is estimated to be 388 446km2 and its AOO 140km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in western dry deciduous forests (see A.diminuta assessment). It has been collected in some protected areas, but it is also known from several unprotected forests subject to various threats (uncontrolled fires for grazing and crops, logging, mining, etc.). Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha boinensis is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required to list it under the Endangered category. maTerial examineD.— 45 collections. Madagascar. Anonymous 5340 (P[P05547039]); Anonymous 5448 (P[P05547072]); Baron, R. 5393 (P[P00508586]), 5450 (P[P00508587]); Bosser, M.J. 13985 (P[P04779823]), 18203 (P[P04779794]); Capuron, R. 24329SF (G, P[P05547235]); d’Alleizette, Ch. 208 (P[P00324564]); Decary, R. 1038 (P[P05543672]), 1045 (P[P00508585]), 8216 (P[P00508588]), 15006 (P[P05547234]), 16385 (P[P05546982]), 17089 (K, MO[MO-3025004], P[P00324527, P05543670], TAN); Gillespie, L. 4037 (MO, US[US01287287]), 10813 (MO), 10817 (MO); Humbert, H. 2350 (P[P00508570, P00508571, P00508572, P00508573, P00508574]), 2715 (P[P05546983, P05546984]), 2997 (G, K, MO, P[P00508549]), 2998 (P[P00508579]), 2998 bis (P[P04779536, P04779537]), 6766 (P[P00324579, P05543684]), 11234 (P[P00508575]), 19373 (P[P05547240]), 19402 (P[P05547241]), 19521 (P[P05547236]), 19693 (P[P05547238]); Keraudren-Aymonin, M. 25935 (P[P00508583, P00508584]); Leandri, J. 401 (P[P00508578]), 3886 (G, P[P05547237]), 3976 (P[P05547239]); Nusbaumer, L. LN 1852 (G[G00090225], P[P05481890]); Perrier de la Bâthie, H. 116 (P[P00508581]), 3676 (P[P00508580]), 9546 (P[P00508576]), 9551 (P[P00508577]), 9823 (P[P00508582]), s.n. (P[P04779845]); Phillipson, P.B. 6220 (K, MO[MO-2965766]); Rakotovao, C. 1052 (MO, P[P04779370], TEF, WAG[WAG0304986]); Ramananjanahary, R. 115 (MO[MO-2965740], P[P04779320], TEF), 266 (MO[MO-3025031], P[P05481881]); Razakamalala, R. 5162 (MO[MO-3025027], P[P01152838], TAN), 6044 (MO[MO-3025025], P[P00887485]); Seyrig, A. 81B (P[P05547034]). references.— Govaerts etal. (2000: 51); Montero Muñoz etal. (2018a: 93). DescripTion Shrubs, deciduous, to 2m tall, monoecious. Branches velutinous and with sparse, simple, erect trichomes to 0.5mm long, glabrescent when mature. Axillary buds ovoid, to 2.5× 2mm, perulate, perules 2, valvate, membranous, brownish, puberulent with simple, appressed trichomes. Stipules to 5mm long, elliptic-lanceolate to linear-lanceolate, pubescent with simple, short, appressed trichomes, denser on midrib; margin ciliate and with some minute glands. Petioles (1- )1.5-4cm long, indumentum similar to that of the young branches. Leaf blades 3-7× (2-)2.5-5(-5.5) cm, ovatelanceolate to subtriangular, membranous; base rounded to subcordate; apex obtuse to acute; margin serrate, teeth acute, reddish, sometimes ending in a minute sessile gland; upper and lower surfaces subvelutinous; venation actinodromous, basal veins 5, secondary veins 4-5 per side. Stipels glandular, 45°E 15°S 20° 25° 300 km 50° Acalypha boinensis fig. 20. — Distribution map of Acalypha boinensis Leandri Madagascar.
425 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) to 0.5mm long, sparsely hairy. Inflorescences spiciform, androgynous, axillary, to 9cm long, mostly female with short male segment; peduncle to 15mm long, it and rachis velutinous. Female segment to 7.5cm long; bracts 4-15, sessile, enlarging in fruit to 5× 7mm, elliptic, pubescent with short, simple trichomes; margin dentate, teeth 20-30, triangular, sometimes ending in a minute, sessile gland, cen - tral tooth not prominent; bracteoles to 2mm long, linearlanceolate, indumentum similar to that on stipules. Male segment persistent, to 2cm long; flowers glomerate; bracts to 2mm diameter, suborbicular, margin irregular, scarious, appressed-pubescent, ciliate. Male flowers: pedicel to 1 mm long, sparsely hairy; buds to 0.4 mm diameter, sparsely hairy. Female flowers 1 per bract, sessile; sepals 3, to 1mm long, triangular, pubescent with simple, short trichomes; ovary c.1 mm diameter, 3-lobed, apparently smooth, surface hispid with simple, short, erect trichomes; styles 3, to 4mm long, distinct, sparsely hairy on rachis, each divided into 8-10 seg - ments. Allomorphic flowers not seen. Capsules to 2.5 mm diameter, papillose, papillae rounded, surface sparsely hairy with simple, short trichomes and hyaline trichomes to 1mm long. Seeds c.1.5 × 1.2 mm, pyriform, foveolate. 8. Acalypha burmanii I.Montero& Cardiel PhytoKeys 108: 93 (Montero Muñoz etal. 2018a). Tragia filiformis Poir., Encyclopédie méthodique. Botanique. Paris 7: 727 (Poiret 1806).— Acalypha acuminata Vahl ex. Baill., Adansonia, Recueil d’observations botaniques 1: 267 (Baillon 1861) nom. illeg., non A.acuminata Benth. (Bentham 1854).— Type: Madagascar?: s.l., s.d., P. Commerson? s.n. (holo-, P-LAM [P00367371]). iconography.— Baillon (1891: pl. 188.); Fig.24E. eTymology.— The epithet honors Dutch botanist Nicolaus Laurens Burman (1743-1793). DisTribuTion anD habiTaT.— Endemic to Madagascar, probably from northeast Madagascar (Fénérive, Toamasina Province). Littoral forest. On sandstone. Altitudinal range unknown (Fig. 21). preliminary conservaTion assessmenT.— Acalypha burmanii is known from two collections, one collected in 1771 by P. Commerson without locality but probably from northeastern Madagascar, and the other collected in Fénérive by d’Alleizette in 1906. It occurs in littoral forests, which are the most threatened vegetation zone in Madagascar (Bollen& Donati 2006; Andriamandimbiarisoa etal. 2015). Although data are too limited to estimate its EOO and AOO, Acalypha burmanii is assigned a preliminary conservation status of Critically Endangered, CR B2ab(ii,iii) (probably EX). m aTerial examineD .— 2 collections. Madagascar. Commerson, P.s.n. (G[G00324451], P[P04779529, P00324459, P00324460, P00799456], P-LAM[P00367371]); d’Alleizette, Ch. s.n. (L[L0242529]). r eferences .— Müller Argoviensis (1865: 40) as A.spiciflora Burm.f.; Müller Argoviensis (1866: 867) as A.spiciflora; Baillon (1891: tab. 188) as A.acuminata; Baillon (1892: 1004) as A.acuminata; Heckel (1903: 198) as A.acuminata; Palacký (1907: 24) as A.acuminata; Pax& Hoffmann (1924: 137) as A.spiciflora; Leandri (1942: 269) as A.spiciflora; Montero Muñoz etal. (2018a: 93). DescripTion Shrubs, probably deciduous, height unknown, monoecious. Branches subglabrous, with sparse, simple, short trichomes. Axillary buds subspherical, to 1× 1.2mm, perulate, perules 2, imbricate, chartaceous, light brown, glabrous. Stipules to 4mm long, linear-lanceolate, margin ciliate and with some sessile glands. Petioles 0.5-1cm long, indumentum similar to that on branches. Leaf blades 2.5-5.5× 1.8-3cm, elliptic to elliptic-lanceolate, membranous; base rounded to acute; apex acuminate, acumen to 12mm long, rounded; margin crenate-serrate, teeth rounded; upper surface glabrous; lower surface subglabrous, with simple, short trichomes on veins; margins ciliate in sinuses; venation actinodromous, basal veins 3, secondary veins 3-4 per side. Stipels glandular, to 0.3mm long, glabrous. Inflorescences spiciform, androgynous, axillary, to 7cm long, mostly male with short female segment; peduncle filiform, to 15mm long, it and rachis subglabrous, with some sparse, simple, short trichomes. Female segment to 2cm long; bracts 1-2, sessile, enlarging in fruit to 10× 11mm, subelliptic, almost glabrous, with some sparse, simple trichomes; mar45°E 15°S 20° 25° 300 km 50° Acalypha burmanii fig. 21 . — Distribution map of Acalypha burmanii I.Montero & Cardiel in Madagascar.
426 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. gin subentire, slightly discolored; bracteoles absent. Male segment persistent, to 4cm long; flowers glomerate; bracts to 0.5mm long, triangular, sparsely hairy. Male flowers: pedicel to 0.7 mm long, sparsely hairy; buds not seen. Female flowers 2 per bract, sessile; sepals 3, to 1mm long, triangular, sparsely hairy with simple, short, arachnoid trichomes; ovary c.1 mm diameter, 3-lobed, papillose, surface puberulent with simple, short trichomes; styles 3, to 3mm long, connate at thickened base, sparsely hairy on rachis, each divided into c.14 segments. Allomorphic flowers not seen. Capsules to 6 mm diameter, papillose, papillae to 1mm long, surface sparsely hairy with simple, short, curved trichomes. Seeds c.3 × 2 mm, pyriform, minutely foveolate. noTes We propose Acalypha burmanii as a new name for Tragia filiformis Poir. (Montero Muñoz etal. 2018a) because combining T.filiformis under Acalypha is blocked by A.filiformis Poir.; nor could we use the illegitimate name A.acuminata Baill. See also comments under the excluded species A.spiciflora. 9. Acalypha cardielii I.Montero& G.A.Levin South African Journal of Botany 146: 636 (Montero Muñoz etal. 2022).— Type: Madagascar. Melaky region (Mahajanga prov.), Tsingy du Bemaraha, 1932-1933, J. Leandri 115 (holo-, P[P00887484]; iso-, P[P05547055]). iconography.— Montero Muñoz etal. (2022); Fig. 25F. e Tymology .— The epithet honors Dr. José María Cardiel, codirector of the first author’s PhD dissertation. He has published extensively about the genus and has contributed to worldwide Acalypha knowledge. DisTribuTion anD habiTaT.— Endemic to Madagascar (Melaky). Dry deciduous forest. On limestone. Altitude c.360m (Fig. 22). preliminary conservaTion assessmenT.— Acalypha cardielii is only known from one collection from the Tsingy de Bemaraha. Its EOO could not be calculated; its AOO is estimated to be 8km2. The Tsingy de Bemaraha lies within a national park and a nature reserve that have been IUCN category II and Ia protected areas since 1927, and a UNESCO World Heritage Site since 1990. The forest of this area has local anthropogenic pressures such as fire associated with the renewal of zebu (cattle) pastures, logging for construction, and deforestation for new agricultural lands, resulting in loss of forest cover during the last decade (Dudley 2008; Goodman etal. 2018). No specimens of this species have been collected for 88 years, so we cannot rule out that this species has become extinct. In conclusion, due to habitat loss, the restricted geographic range, and the absence of recent collections, A.cardielii is assigned a preliminary conservation status of Critically Endangered: CR B2ab(ii,iii) (probably EX). maTerial examineD.— 1 collection. Madagascar: Leandri, J. 115 (P[P05547055, P00887484]). DescripTion Shrubs, probably deciduous, height unknown, monoecious. Branches whitish, pubescent with simple, curved, antrorse trichomes, glabrescent when mature. Axillary buds spherical, to 1mm diameter, perulate, perules 2, imbricated, chartaceous, dark brown, glabrous. Stipules to 1mm long, triangular-lanceolate, apex acute, sparsely hairy with simple, short trichomes, margin with some sessile glands. Petioles canaliculate, 0.5-1.5(-2) cm long, indumentum similar to that on young branches. Leaf blades 1.8-2.6(-3)× 1.1-1.6cm, subrhombic, membranous; base acute; apex obtuse to retuse; margin crenate-serrate, revolute, reddish, teeth rounded; upper surface sparsely hairy with some simple, short trichomes; lower surface glabrous; venation actinodromous, basal veins 3, secondary veins 3-4 per side. Stipels glandular, to 0.2mm long, glabrous. Inflorescences spiciform, androgynous and male, axillary. Androgynous inflorescences to 5.5cm long, mostly male with short female segment; peduncle to 15mm long, it and rachis with indumentum similar to that on petioles. Female segment to 2.5cm long; bracts 2, sessile, enlarging in fruit to 8× 6mm, reniform, glabrous; margin crenate, teeth 6-12, rounded, central tooth not prominent; bracteoles absent. Male segment persistent, to 2.5cm long; flowers glomerate; bracts to 1mm long, elliptic-lanceolate, glabrous. Male inflorescences laxly flowered, to 3.5cm long; subsessile, peduncle to 0.3mm long; bracts like those on 45°E 15°S 20° 25° 300 km 50° Acalypha cardielii fig. 22 . — Distribution map of Acalypha cardielii I.Montero & G.A.Levin in Madagascar.
427 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) androgynous inflorescences. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds to 0.5 mm diameter, glabrous. Female flowers 1 per bract, sessile; sepals 3, to 1mm long, triangular-lanceolate, glabrous, margin with some sessile glands; ovary c.1 mm diameter, 3-lobed, papillose-hispid, papillae acute, ending in simple trichome to 0.5mm long, surface with some subsessile glandular trichomes; styles 3, to 4mm long, distinct, glabrous, each divided into 7-9 segments. Allomorphic flowers sometimes present at inflorescence apex; pedicel filiform, to 1.5cm long, glabrous; sepals 3, similar to those of normal flowers; ovary 1-lobed, to 2× 1.5mm, sparsely hairy, distally fimbriate; style 1, to 4mm long, glabrous. Capsules to 2 mm diameter, papillosehispid, papillae triangular. ending in simple trichome to 1mm long, surface glabrous. Seeds not seen. 10. Acalypha chibomboa Baill. Adansonia, recueil d’observations botaniques 1: 269 (Baillon 1861).— Type: Comoros, Anjouan, 1850, L.H. Boivin s.n. (holo-, P[P00196274]). Acalypha codonocalyx Baill., Adansonia, recueil d’observations botaniques 1: 271 (Baillon 1861).— Type: Comoros. Mohéli, “Île Mohilla”, s.d., M. Richard 286 (lecto-, designated by Montero Muñoz etal. [2018a: 94]: P[P00196282]; isolecto-, P[P00196283]). Acalypha comorensis Pax, Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie. Leipzig 19(1): 95 (Pax 1894).— Type: Comoros. Anjouan, “Johanna”, VIVIII.1875, J. M. Hildebrandt 1662 (holo-, B or WRSL, presumably destroyed; lecto-, designated by Montero Muñoz etal. [2018a: 94]: BREM[BREM0001792]; isolecto-, K[K000186524, K000186525], L[L0241274], P[P00196280], W[W0004243]).— Former syntype: Comoros. Grande Comore, 1886, C. W. Schmidt 192 [n.v.]. iconography.— Baillon (1891: pl. 192); Fig. 24G. eTymology.— The epithet refers to the vernacular name of this plant, known as “chibomboa/choubomboa” in Comoros, according Baillon (1861). DisTribuTion anD habiTaT.— Madagascar (Diana) and Comoros Archipelago (Grande Comoro, Anjouan, Moheli, and Mayotte). In Madagascar: dry deciduous forest on basalt; altitudinal range 120-1020m. In Comoros Archipelago: in valley forest on clay soil: altitudinal range 200-700m (Fig. 23). preliminary conservaTion assessmenT.— Acalypha chibomboa is known from Madagascar and Comoros. In the Comoros, its EOO is estimated to be 8 131km2 and its AOO 44km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. The Comoros archipelago has lost at least 80% of its native vegetation, and is now dominated by cash-crops and subsistence cultivation. Deforestation rate has decreased but is still ongoing, making Comoros the archipelago with the highest annual net loss of forest in WIOR. In addition, alien species have a detrimental effect on the native flora (Safford 2001; Chakravarty etal. 2012; Baret etal. 2013). Ongoing habitat loss will cause continued decline of its EOO and AOO. In the Comoros, A.chibomboa is assessed as Near Threatened (NT) under criterion B, because it meets the values needed for a threatened category for the AOO, but the number of locations is too high to meet the conditions required for a listing under Endangered category. In Madagascar, its EOO is estimated to be 253.391km2 and its AOO 20km2. This species is only known from six locations in dry deciduous forests in the north of the island (see assessment of A.diminuta). This species has been collected in some protected areas, as Ankarana Special Reserve (see assessment of A.ankaranensis), and in the Montagne d’Ambre National Park, where the forests are affected mainly by slash-and-burn agriculture. Currently, Montagne d’Ambre is under additional threat because qat (Catha edulis, Celastraceae) cultivation is increasing (Goodman etal. 2018). Due to ongoing habitat loss that will cause continued decline of its EOO and AOO, A.chibomboa is assessed as Endangered (EN) EN: B1ab(i,iii,iv) + B2ab(ii,iii,iv) in Madagascar. In the whole region, Acalypha chibomboa can be assessed as Near Threatened (NT). It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 23 collections. Comoros Archipelago. Attoumane, M. 15 (MO[MO-2965775]); Barthelat, F. 1436 (P[P00631096]); Boivin, L.H.s.n. (P[P00196274]); Bosser, M.J. 18011 (P[P00196228, P00196229]); Coulon, P. 205 (P[P00184949]); d’Alleizette, Ch. s.n. (L[L0241232]); Hildebrandt, J.M. 1662 (BREM[BREM_0001792], K[K000186524, K000186525], L[L0241274], P[P00196280], W[W0004243]); Humblot, L. 206 (K), 1206 (P[P00196275, P00196276, P00196277]); 1377 (P[P00196278, P00196279]); Jacquemin, H. 837 (P[P00324456]), 837-J (P[P00538326]); Rakotozafy, A. 1152 (P[P00196236]); Richard, M. 225 (P[P00196281]), 286 (P[P00196282, P00196283]); Service Forestier Madagascar 16651-SF (P[P00334973, P00184984]); Waterlot, M. 964 (P[P00196284]). Madagascar. Andrianantoanina, O. 960 (P[P04779784]); BardotVaucoulon, M. 1227 (K, MO[MO-3025011], P[P00455522], TAN); Cours, G. 5485 (P[P00324546, P05510111]); Dorr, L.J. 3622 (BR[BR0000021450280], C, L[L0449357], MO, P[P00508514, P01295515], WAG[WAG0133270]); Malcomber, S. 1239 (CAS, K, MO[MO-2965805]); Razanajatovo, M. G. MHR027 (G[G00170087], K, P[P05516023]). references.— Baillon (1891: pl. 192 as A.codonocalyx); Palacký (1907: 24); Voeltzkow (1917: 447); Pax& Hoffmann (1924: 165); Leandri (1942: 280); Govaerts etal. (2000: 55); Montero Muñoz etal. (2018a: 94). DescripTion Small trees, evergreen, to 5m tall, monoecious. Branches striate, pubescent with simple, short, curved, antrorse trichomes, glabrescent when mature. Axillary buds ovoid, to 1.5× 1mm, perulate, perules 2, valvate, membranous, brownish, sparsely 45°E 15°S 300 km 50° Acalypha chibomboa fig. 23. — Distribution map of Acalypha chibomboa Baill. in Comoros Archipelago and Northern Madagascar.
428 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. AB CD EF GH fig. 24. — Mature female bracts: A, Acalypha andringitrensis Leandri (H. Perrier de la Bâthie 13640); B, A. ankaranensis I.Montero & Cardiel (A. J. M. Leeuwenberg 14374); C, A. bailloniana Müll.Arg. (G. Viscardi 439); D, A. boinensis Leandri (R. Ramananjanahary 115); E, A. burmanii I.Montero & Cardiel (P. Commerson s.n.); F, A. cardielii I.Montero & G.A.Levin (J. Leandri 115); G, A. chibomboa Baill. (F. Barthelat 1436); H, A. claoxyloides Hutch. (F. Friedmann 5633). Illustration by Iris Montero Muñoz. Scale bars: A, 2 mm; B-H, 1 mm.
429 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) hairy with simple, short trichomes. Stipules to 3mm long, linear-lanceolate, ciliate with simple, erect trichomes to 0.5mm long. Petioles very slender, 2.5-9cm long, indumentum similar to that on young branches but less dense, glabrescent. Leaf blades (8-)9-14(-15.5)× 3.5-6cm, elliptic-lanceolate, membranous; base rounded to subcuneate; apex acuminate, acumen to 25mm long, acute; margin serrate, teeth acute; upper surface puberulent with sparse, simple, short trichomes, and with simple, short, appressed trichomes on veins, glabrescent; lower surface subglabrous, with sparse simple, short trichomes and flattened resinous glands; venation actinodromous, basal veins 3 or 5, secondary veins 6-9 per side. Stipels absent. Inflorescences spiciform, usually unisexual, sometimes androgynous, axillary. Androgynous inflorescences laxly flowered, to 11cm long, about equally male and female; peduncle filiform, to 40mm long, indumentum similar to that on petioles. Female segment to 5.5cm long; bracts 2-6, sessile, enlarging in fruit to 5mm diameter, orbicular, sparsely hairy with simple, short, arachnoid trichomes at base, with flattened resinous glands, and ciliate with simple, short trichomes; margin denticulate, teeth 17-19, triangular, ending in sessile gland, central tooth not prominent; bracteoles absent. Male segment deciduous, 6cm long; bracts to 1mm long, ovate, pubescent with simple, short, arachnoid trichomes; flowers glomerate. Male inflorescences laxly flowered, to 12cm long; peduncle to 30mm long, it and rachis with indumentum similar to that on petioles. Female inflorescences laxly flowered, to 5cm long; peduncle filiform, to 20mm long, it and rachis with indumentum similar to that on petioles; bracts 2-5, they and bracteoles like those on androgynous inflorescences. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds to 1mm diameter, glabrous. Female flowers 1 per bract, sessile; sepals 3-4, to 1.3mm long, triangular, subglabrous, ciliate with simple, minute trichomes; ovary c.1 mm diameter, 3-lobed, smooth. surface pubescent with simple, short, arachnoid trichomes and flattened resinous glands; styles 3, to 4mm long, slightly connate at base, sparsely hairy on rachis, each divided into c.6 segments. Allomorphic flowers not seen. Capsules to 4 mm diameter, smooth, surface pubescent with simple, arachnoid trichomes and flattened resinous glands. Seeds c.3 × 2.3 mm, pyriform, minutely foveolate. 11. Acalypha claoxyloides Hutch. Bulletin of Miscellaneous Information, Royal Gardens, Kew 1918: 205 (Hutchinson 1918).— Type: Seychelles. Astove, Cosmoledo and Aldabra, IV.1907, H. P. Thomasset 243 (lecto-, designated by Montero Muñoz etal. [2018a: 94]: K[K000186504]). Acalypha fryeri Hutch., Bulletin of Miscellaneous Information, Royal Gardens, Kew 1918: 206 (Hutchinson 1918).— Type: Seychelles. Aldabra, s.d., J. Fryer 92 (lecto-, designated here, or possibly holotype: K[K000186506]). Acalypha aldabrica Pax& K.Hoffm., Das Pflanzenreich (Engler) 147, 16 (Heft 85): 136 (Pax& Hoffmann 1924).— Type: Seychelles. Aldabra, s.d., W. L. Abbott s.n. (holo-, B?, presumably destroyed; lecto-, designated by Montero Muñoz etal. [2018a: 94]: P[P00887488]; isolecto-, P[P00887489]). iconography.— Fig. 24H. eTymology.— The epithet refers to Claoxylon A.Juss. (Euphorbiaceae), the genus to which the type specimen was first assigned by Baker. D isTribuTion anD habiTaT .— Endemic to Seychelles (Aldabra group). Mixed xeric scrub on coral limestone, at sea level (Fig. 25). preliminary conservaTion assessmenT.— Acalypha claoxyloides is known from the Aldabra Group Islands (Seychelles). Its EOO is estimated to be 5 233km2, close to the threshold of the B1 subcriterion of the Vulnerable category, and its AOO 52km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. The ecosystems in the Aldabra atoll are remarkably intact and it has been a UNESCO World Heritage Site since 1982, but on the other atolls the forests have been seriously degraded by conversion to plantations, mainly coconut, and by guano extraction. Currently the main threat is alien species (Roberts 1988; Safford 2013; Baret etal. 2013). Acalypha claoxyloides seems to be a very common shrub on the atolls of this group of islands, and it meets the EOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category;, it therefore could be assessed as Near Threatened (NT) under criterion B. However, ongoing habitat loss on some atolls and the presence of alien species will likely cause continued decline of its EOO and AOO in the near future, so we assess this species as Endangered under subcriteria B1 and B2. maTerial examineD.— 35 collections. Seychelles. Abbott, W.L.s.n. (P[P00887488, P00887489]); Fitzgerald, D.V. 5989 (BR[BR0000014624568]), 6017 (BR[BR0000014624575], P[P04779371]); Fosberg, F.R. 48741 (MO[MO-2965769]), 48742 (MO[MO-2965771], NY[NY03091181], US[US01287312]), 48771 (M[MO-2965756], NY[NY3091176], [US01287313]), 48900 (MO[MO-2965770], NY[NY3091177], US[US01287308]), 49065 (MO[MO-2965773], NY[NY3091179], US[US01287316]), 49627 (US[US01287309]), 49661 (GH[GH01097234], MO[MO-2965772], NY[NY3091178]), 49662 (MO[MO-2965768], 45 km Seychelles Acalypha claoxyloides fig. 25 . — Distribution map of Acalypha claoxyloides Hutch. in Seychelles (Aldabra group).
430 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. NY3091180, US01287315), 49730 (US01287304), 49804 (US01287307); Frazier, J. 3 (US01287328), 29 (K), 602 (US01287306); Friedmann, F. 5633 (P[P00870926, P00870927]), 5634 (P[P00870925]); Fryer, J.C.F. 18 (K[K000186505]), 92 (K[K000186506]); Gardiner, J.S. 18 (K); Gwynne 1240 (K); Merton 7010 (K); Renvoize, S. A. 748 (BR[BR0000014624629], L[L0241259, L0241260]), 768 (L[L0241258], P[P05481893]), 1001 (BR[BR0000014624605], L[L0241257], P[P05481872]), 1288 (BR[BR0000014624582], L[L0241255, L0241256]), 1289 (BR[BR0000014624599], L[L0241253, L0241254], P[P05481871]), 1314 (K); Rhyne, C. 863 (US[US01287317]); Ridgway, T. 111 (US[US01287305]); Stoddart, D.R. 748 (K), 998 (K), 1231 (K); Thomasset, H.P. 243 (K[K000186504]). references.— Hemsley (1919: 130); Hemsley (1919: 131) as A.fryeri; Pax& Hoffmann (1924: 136) as A.aldabrica; Fosberg (1974: 263); Renvoize (1975: 152); Robertson (1989: 199); Govaerts etal. (2000: 56); Sagun etal. (2006: 124); Montero Muñoz etal. (2018a: 94). DescripTion Small trees, deciduous, to 3m tall, monoecious. Branches reddish, subglabrous, with some simple, arachnoid trichomes, greyish, with thick nodes, and glabrescent when mature. Axillary buds spherical, to 1.5mm diameter, perulate, perules 2, valvate, membranous, light brown, woolly with simple, arachnoid trichomes. Stipules to 1.5mm long, triangular-lanceolate, pubescent with simple, arachnoid trichomes. Petioles 2-3cm long, indumentum similar to that on young branches. Leaf blades (5-)6-11(-14)× (3-)4-8cm, ovate-lanceolate to ellipticlanceolate, chartaceous; base rounded to acute; apex acuminate, acumen to 5mm long, obtuse; margin crenate, slightly revolute, teeth rounded; upper and lower surfaces subglabrous, with some simple, arachnoid trichomes on veins, lower surface also with flattened resinous glands; venation pinnate, secondary veins to 8 per side. Stipels glandular, to 0.5mm long, glabrous. Inflorescences spiciform, androgynous and male, axillary. Androgynous inflorescences to 5.5cm long, mostly male with short female segment; peduncle to 30mm long, it and rachis laxly pubescent with simple, arachnoid trichomes. Female segment to 2cm long; bracts 1-2, sessile, enlarging in fruit to 10-19mm, reniform, subglabrous, appressed-pubescent at base, and with flattened resinous glands, margin denticulate, teeth 19-23, triangular, sometimes ending in sessile gland, central tooth not prominent; bracteoles sessile glands, glabrous. Male segment deciduous, to 3cm long; flowers glomerate, with stalked, flattened resinous glands between pedicels; bracts to 1mm long, ovate-lanceolate, sparsely hairy with simple, arachnoid trichomes. Male inflorescences to 4cm long; subsessile, peduncle to 3mm long, it and rachis appressed-pubescent with simple, short trichomes; flowers glomerate; bracts similar to those on the male segment in androgynous inflorescences. Male flowers: pedicel to 2 mm long, sparsely hairy; buds to 1mm diameter, sparsely hairy. Female flowers 2 per bract, sessile; sepals 3, to 1mm long, ovate-triangular, woolly with simple, arachnoid trichomes; ovary c.1 mm diameter, 3-lobed, smooth, surface woolly with simple, arachnoid trichomes, and with flattened resinous glands; styles 3, to 5mm long, distinct, sparsely hairy on rachis with simple, arachnoid trichomes, each divided into c.10 segments. Allomorphic flowers not seen. Capsules to 3.5 mm diameter, smooth, surface appressed-pubescent with simple, short, trichomes and flattened resinous glands. Seeds c.1.3 × 1 mm, pyriform, foveolate. noTe 1) Hutchinson did not indicate the herbarium where the holotype is located, so the single specimen found is designated as the lectotype; and 2) Acalypha claoxyloides is widespread in the Seychelles archipelago. It is very closely related to A.pubiflora (Klotzsch) Baill., known from southeastern Africa (Botswana, Malawi, Mozambique, South Africa and Zimbabwe), of which it may be a synonym. More study of the African material, as well as the Australian material treated as A.pubiflora var. australica Radcl.-Sm. (Radcliffe-Smith 1990, 1996), is needed to unravel the A.pubiflora Baill. complex. 12. Acalypha crateriana (Coode) I.Montero& Cardiel, comb. nov. Acalypha integrifolia Willd. subsp. marginata (Poir.) Coode var. crateriana Coode, Kew Bulletin 34: 44 (Coode 1979).— Type: Mauritius. Tamarin Falls, 2.III.1975, D. Lorence 1138 (holo-, K[K000431104]; iso-, K[K000431105], MAU[n.v.]). iconography.— Unknown. eTymology.— The epithet probably refers to the Kanaka crater in Mauritius. D isTribuTion anD habiTaT .— Endemic to the Mascarene Islands (Mauritius). Degraded lowland evergreen moist forest. Altitudinal range 700-800m (Fig. 26). preliminary conservaTion assessmenT.— Acalypha crateriana comb. nov. is only known from Kanaka crater and Grand Bassin peak in Mauritius. Its EOO is estimated to be 12.9km2 and its AOO 12km2. Native forest remnants of Mauritius are very restricted, covering less than 5% of the total landmass. This massive deforestation is due mainly to agriculture and urban expansion. Another threat to the Mauritius native flora is the widespread occurrence of alien species. Native plants are seriously affected by invasive plants, reducing their growth rate so regeneration of forests is extremely low (Safford 1997; Florens 2008; Baider& Florens 2013; Norder etal. 2017). No specimens of this species have been collected for 45 years. In recognition of its restricted geographic range (a single known location), the cited threats, and the absence of recent collections, A.crateriana comb. nov. is assigned a preliminary conservation status of Critically Endangered: CR B2ab(ii,iii,iv). maTerial examineD.— 16 collections. Mauritius. Coode, M.J.E. 4694 (K), 4700 (K), 4701 (P[P04779343]), 4704 (K), 4705 (K, MO[MO-2452045]), 4718 (K), 4720 (K), 4721 (P[P04779349]), 4722 (K), 4729 (K), 4732 (K, MO[MO-2452041]), 4737 (K), 4744 (K, MO[MO-2452042]), 4748 (P[P04779342]); Lorence, D. 1138 (K[K000431104, K000431105]), DL1858 (MO[MO-2452048], P[P04779346]). references.— Coode (1982: 74); Montero Muñoz etal. (2018a: 104) as A.marginata. DescripTion Shrubs, leaf persistence unknown, to 2.7m tall, probably dioecious. Branches appressed-pubescent, glabrescent when mature. Axillary buds naked, hispid. Stipules to 5mm long, linear-
431 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) lanceolate, densely pubescent with simple, appressed trichomes, margin papillate. Petioles 0.5-0.8cm long, indumentum similar to that on young branches, glabrescent. Leaf blades 4-7(-10)× 1.5-3cm, elliptic to subobovate, chartaceous; base acute to cuneate; apex rounded to acute; margin crenate to crenate-dentate, slightly revolute, usually reddish or pinkish, teeth rounded; upper surface glabrous; lower surface sparsely hairy with appressed trichomes on veins; venation pinnate, secondary veins 7-8 per side. Stipels absent. Inflorescences spiciform (male) or glomerulate (female), unisexual, axillary. Male inflorescences to 5cm long; peduncle to 5mm long, indumentum similar to that on petioles; flowers glomerate; bracts to 1mm long, triangular, appressed-pubescent. Female inflorescences 2-4mm long; sessile; bracts 1-4, not enlarging in fruit, to 0.5mm long, triangularlanceolate, appressed-pubescent; margin entire; bracteoles absent. Male flowers: pedicel to 0.6 mm long, sparsely hairy; buds to 0.5mm diameter, sparsely hairy. Female flowers 1 per bract, subsessile; sepals 3, to 1mm long, triangular-lanceolate, appressed-pubescent; ovary c.1 mm diameter, 3-lobed, papillose, surface densely hispid; styles 3, to 4mm long, distinct, sparsely hairy on rachis, each divided into 10-11 segments. Allomorphic flowers not seen. Capsules to 3 mm diameter, papillose-hispid, papillae ending in a simple, trichome to 1mm long, surface sparsely hairy with simple, short trichomes, glabrescent. Seeds c.1× 0.7 mm, pyriform, minutely foveolate. noTe Acalypha crateriana comb. nov. was treated by Coode (1979) as variety of A.integrifolia subsp. marginata, treated here as A.marginata. We consider A.crateriana comb. nov. to be a distinct species distinguished mainly by having petioles 0.50.8cm long and leaf blades 4-7(-10)× 1.5-3cm, elliptic to subobovate; base acute to cuneate; apex rounded to acute; margin crenate to crenate-dentate, usually reddish or pinkish, slightly revolute, teeth rounded; vs petioles 0.5-3cm long and leaf blades (4-)6-17-(21)× (2-)2.5-7(-8) cm, usually elliptic to elliptic-lanceolate, sometimes obovate; base rounded to cuneate; apex acute to subacuminate; margin crenate-serrate to serrate, discolorous, teeth rounded to acute, slightly calloseedged in A.marginata. 13. Acalypha decaryana Leandri Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 284 (Leandri 1942).— Type: Madagascar. Prov. Toliara, Ambovombe, 20.VIII.1924, R. Decary 2985 (lecto-, designated here: P[P00508553]; isolecto-,W[W1962-0013399]). iconography.— Leandri (1942: 285); Figs 28F; 35A. eTymology.— The epithet honors French botanist Raymond Decary (1891-1973). DisTribuTion anD habiTaT.— Endemic to Madagascar (Menabe, Atsimo-Andrefana, Ihorombe, Androy, Anosy, and AtsimoAtsinanana). Dry deciduous forest and spiny thickets; sometimes in riparian forest. On Mesozoic limestone, basalt, basement rocks, sandstone, and unconsolidated sand. Altitudinal range (10-) 30-900 (-1000) m (Fig. 27). p reliminary conservaTion assessmenT .— The EOO of Acalypha decaryana is estimated to be 130 409km2 and its AOO 172km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in dry deciduous forest (see assessment of A.diminuta) and spiny thickets, one of the most threatened ecosystems in Madagascar (Aronson etal. 2018). Acalypha decaryana has been collected in some protected areas, but it is also known from several unprotected forests subject to various threats (uncontrolled fires for grazing and crops, logging, mining, etc.). Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha decaryana is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 78 collections. Madagascar. Allorge, L. 2185 (P[P00122670]); Birkinshaw, C.R. 424 (P[P04779214]); Boiteau, P. 2539 (P[P00508569]); Bosser, M.J. 4022 (P[P04779809]), 4073 (P[P04779789]), 10208 (P[P04779811]), 13994 (P[P04779791]), 17213 (P[P00324548]), 17275 (MO[MO-3025010], P[P00324551], TAN), 17417 (P[P00324550, P05543683]), 17840 (P[P00324555, P05543685]); De Block, P. 2383 (BR[BR0000005518647], K, P[P04786280], TAN); Decary, R. 2964 (P[P00508568]), 2985 (P[P00508553], W[W1962-0013399]), 3115 (P[P00508567]), 3247 (L[L0241288], P[P00508566]), 3433 (P[P00508565]), 3728 (P[P00508564, P04786273], S[S13-13801]), 3789 (G, P[P00508563]), 4264 (G, P[P00508562]), 4278 (P[P00508561]), 4343 (P[P00508560]), 8433 (P[P00508559]), 8526 (P[P00508558]), 8878 (P[P00508557], US[US01287334]), 8910 (P[P00508556]), 8956 (P[P00508555]), 8985 (P[P00508554]), 9030 (P[P00508552]), 9079 (B[B100273581], P[P00508551]), 9206 (P[P04779838]), 9864 (P[P00508550]), 10452 bis (P[P04779825]), s.n. (L[L0241289]); Dequaire, J. 24109 (P[P05547013]); Descoings, B. 3108 (P[P05547016]), 3625 (P[P05547012]), 3626 (P[P05547014]); Eboroke, S. 959 (P[P05547078]); Groeninckx, I. 317 (BR[BR0000005513222]); Homolle, A.M. 1706 (P[P05547015]); Humbert, H. 2997 (G, K, MO, P[P00508547, P00508548]), 2997 bis (P[P00508546], WAG[WAG.1930096]), 5405 (B[B100351094], BR[BR0000005045341], GH[GH01097245], 75 km Mauritius La Réunion Acalypha crateriana comb. nov. fig. 26 . — Distribution map of Acalypha crateriana (Coode) I.Montero & Cardiel, comb. nov. in Mascarene Islands.
432 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. MA[MA-01-00820754], NY[NY3091182], P[P00508544, P00508545], S, US[US01287332]), 5640 (GH[GH01097253], P[P00508542, P00508543], US[US01287333]), 11412 (P[P05547248]), 12423 (L[L3795925], MEXU[MEXU1364277], P[P00508537, P00508538, P00508539, P00508540, P00508541]), 12835 (P[P00508536]), 12902 (P[P00508535]), 13098 (B[B100174358], G, P[P00508532, P00508533, P00508534], WAG[WAG1930098]), 13809 (P[P00508530, P00508531]), 14166 (P[P00508529]), 29130 (P[P05547080]), 29182 (P[P05547251]), 29434 (G, P[P05547252], NY[NY2710390]), 29449 (P[P05547253]), 29450 (G, P[P05547254], WAG[WAG.1930092]); Leandri, J. 3564 (P[P05547079]), 3566 (G, P[P05547247], WAG[WAG.1930091]); Lowry II, P.P. 6930 (MA, MO), 6991 (MA, MO, P[P05481880]); McPherson, G. 14916 (P[P00508528]); Messmer, N. NM 184 (G[G00420832], WAG[WAG.1930097]); O’Connor, S. 38 (K), 53 (K); Peltier, J. 2820 (P[P05547017]); Perrier de la Bâthie, H. 9763 (P[P00508527]), 9780 (P[P00508526]); Phillipson, P.B. 1751 (MO, P[P00508525], WAG[WAG.0133186]); Rakotomalaza, P. 211 (MO[MO-2965752], P[P05547994]), 217 (MAUAM, MO, P[P00460980], TAN), 305 (P[P04779767]); Rakotonasolo, F. 6580-RN (P[P05547077]); Randriatsivery, M.F. 351 (MO); Ravelonarivo, D. 4648 (MO[MO-3086130]); Richard, M. 0128 (K); Schlieben, H.J. 8214 (BR[BR0000014625152], G); Service Forestier Madagascar 8486-SF (P[P00508524]). references.— Govaerts etal. (2000: 59); Seebaluck etal. (2015: 150); Montero Muñoz etal. (2018a: 95). DescripTion Shrubs, deciduous, to 3m tall, monoecious. Branches reddish, appressed-pubescent and with rounded resinous glands, glabrescent when mature; brachyblasts present. Axillary buds ovoid, to 1× 0.7mm, perulate, perules 2, imbricate, chartaceous, blackish, sparsely hairy. Stipules to 1.5mm long, triangular-lanceolate, midrib appressed-pubescent with simple trichomes, margin ciliate. Petioles 2-5cm long, pubescent with simple, short trichomes. Leaf blades 1.5-2.5(-4)[-7.5]× 0.4-0.6(-2)[-3.5] cm, elliptic to oblong, sometimes obovate, membranous; base obtuse to acute; apex rounded to obtuse, sometimes acute; margin serrate, teeth subacute; upper and lower surfaces pubescent with simple, short trichomes; lower surface also with rounded resinous glands; venation pinnate, secondary veins 6-7 per side. Stipels absent. Inflorescences spiciform, unisexual, axillary. Male inflorescences densely flowered, to 1.5cm long; subsessile, peduncle to 1mm long, densely pubescent; flowers glomerate; bracts to 0.5mm long, ovate-lanceolate, ciliate with simple, short, trichomes. Female inflorescences laxly flowered, to 3cm long; peduncle filiform, 6-10mm long, it and rachis pubescent with simple, short trichomes; bracts 1-3, sessile, enlarging in fruit to 4× 5.5mm, orbicular-reniform, with two reddish papillae at base, pubescent with simple, short trichomes and rounded resinous glands; margin dentate to crenate-dentate, teeth c.20, rounded to subtriangular, central tooth not prominent; bracteoles to 0.3mm long, triangular, sparsely hairy. Male flowers: pedicel to 1 mm long, sparsely hairy; buds to 0.5mm diameter, sparsely hairy and with some rounded resinous glands. Female flowers 1 per bract, sessile; sepals 3-4, to 1mm long, triangular, appressed-pubescent with simple, short trichomes; ovary c.1 mm diameter, 3-lobed, smooth, surface pubescent with simple, short, appressed trichomes and rounded resinous glands; styles 3, to 4mm long, distinct, sparsely hairy on rachis, with simple trichomes, each divided into 8-10 slender segments. Allomorphic flowers sometimes present at female inflorescence apex; pedicel filiform, to 2mm long, pubescent with simple, curved, antrorse trichomes; sepals 3, to 0.5mm long, triangular, pubescent with simple, short trichomes and rounded resinous glands; ovary 1-lobed, to 1mm diameter, densely pubescent with simple trichomes; style 1, to 4mm long, glabrous. Capsules to 3 mm diameter, smooth, surface pubescent with simple, short, appressed trichomes and rounded resinous glands. Seeds c.1.8 × 1.2 mm, pyriform, foveolate. noTe Leandri designated Decary 2985 as the type, but he did not indicate the herbarium. We located two specimens of this collection. The one at P, where Leandri worked, is the only one that bears the word “type” written by Leandri. We therefore designate this as the lectotype. 14. Acalypha emirnensis Baill. Adansonia, recueil d’observations botaniques 1: 270 (Baillon 1861).— Type: Madagascar. Prov. Antananarivo: Antananarivo “in prov. Emirna, prope Tananarivou”, 1833, W. Bojer s.n. (lecto-, designated here: P[P00536723]; isolecto-, GD-C[GDC005713], P[P00536725]). 45°E 15°S 20° 25° 300 km 50° Acalypha decaryana fig. 27. — Distribution map of Acalypha decaryana Leandri in Madagascar.
433 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) AB CD EF GH fig. 28 . — A, Acalypha diminuta Baill. (photo by L. Gillespie, L. Gillespie 10704); B, C, A. andringitrensis Leandri (photo by P. Phillipson, P. Phillipson 5669); D, E, A. boinensis Leandri (photo by L. Gillespie, L. Gillespie 10813); F, A. decaryana Leandri (photo by P. Lowry II, P. Lowry II 6991); G, H, A. emirnensis Baill. (photo by L. Nusbaumer, P. Ranirison PR994).
440 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in littoral forests on sand. Due to their low elevation and subsequent easy accessibility, littoral forests are under constant pressure and are regarded as the most threatened vegetation zone in Madagascar (Bollen& Donati 2006; Andriamandimbiarisoa etal. 2015). This species has been collected in some protected areas, but it is also known from several unprotected forests subject to constant pressure. We predict that ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha gracilipes is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threat - ened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 29 collections. Madagascar: Baron, R. 1391 (K), 5980 (P[P00513164]), 6420 (P[P00324562]); Commerson, P.s.n. (P[P00513166]), s.n. (P[P04022747]), s.n. (P[P05604465]); Dorr, L.J. 4463 (K, MO, P[P00508455]); Dumetz, N. 764 (MO[MO-2965758], P[P04779867]); Faliniaina, L. 9 (K, MO); Hoffmann, P. 279 (K); Humbert, H. 5835 (P[P00513167, P00513168]); Jussieu, A.s.n. (P[P04804753]); Leandri, J. 4296 (P[P04804743]); McPherson, G. 14325 (MO[MO-2965762]), s.n. (P[P04804702]); Miller, J.S. 10464 (K, MO[MO-2966182], P[P05547263]); Perrier de la Bâthie, H. 9707 (P[P00513169]); Rabenantoandro, J. 1553 (P[P05481907]), 1580 (MO[MO-3025022], P[P05481889]); Rabevohitra, R. 2054 (MO[MO-2965759]); Raharimampionona, J. 38 (P[P05481876]); Rajeriarison, Ch. EUPH-26 (P[P04804488]); Ratovoson, F. 1752 (MAUAM, MO[MO-2966280], P, TAN), 1941 (MAUAM, MO[MO-2966287], TAN); Razaka - malala, R. 6897 (MO[MO-2965737], TAN); Rogers, Z.S. 901 (MO[MO-2965730], P, TAN); Schatz, G.E. 2745 (K, P[P00508422]), 3436 (K, MO[MO-2966191], P[P00508420]); Scott-Elliot, G.F. 2493 (K, P[P00513171]). references.— Müller Argoviensis (1866: 850) as A.commersoniana var. gracilipes; Baillon (1892: 1004); Palacký (1907: 25); Pax& Hoffmann (1924: 106) as A.integrifolia var. gracilipes; Govaerts etal. (2000: 69, 100); Montero Muñoz etal. (2018a: 98). DescripTion Sprawling shrubs, leaf persistence unknown, to 4m tall, monoecious. Branches flexuous, glabrous. Axillary buds spherical, to 0.5mm diameter, perulate, perules 2, valvate, membranous, glabrous, light brown, margin with some sessile glands. Stipules to 1.2mm long, triangular, slightly fleshy, sparsely hairy, margin papillate. Petioles canaliculate, (0.4-)0.7-1.4(-1.7) cm long, glabrous. Leaf blades (4-)5-7(- 7.5)× 1.5-2cm, elliptic-lanceolate, chartaceous; base acute to cuneate; apex acuminate, acumen to 1.3mm long, rounded to subacute, mucronate; margin crenate-serrate to subentire, slightly revolute, sometimes reddish, teeth obtuse; upper and lower surfaces glabrous; venation pinnate, secondary veins 5-7 per side. Stipels absent. Inflorescences spiciform, axillary, androgynous and male. Androgynous inflorescences to 5cm long, mostly male with 1 female bract; sessile; rachis glabrous. Female segment: bract 1, sessile, enlarging in fruit to 3× 4mm, reniform, glabrous; margin dentate, slightly thickened, teeth c.10, rounded, central tooth not prominent; bracteoles absent. Male segment persistent, to 4.5cm long; flowers glomerate; bracts to 0.7mm long, triangularlanceolate, sparsely hairy with some simple, short trichomes at apex. Male inflorescences filiform, to 10cm long; peduncle to 7mm long, glabrous; bracts and bracteoles like those on androgynous inflorescences. Male flowers: pedicel to 0.5 mm long, glabrous; buds to 0.5 mm diameter, glabrous. Female flowers 1 per bract, sessile; sepals 3, to 0.5mm long, triangular, glabrous; ovary c.1 mm diameter, 3-lobed, papillose-hispid, papillae to 0.5mm long, ending in simple, short trichome, surface glabrous; styles 3, to 5mm long, slightly connate at base, rachis thick, glabrous, each divided into c.15 segments. Allomorphic flowers not seen. Capsules to 2 mm diameter (immature), papillose-hispid, papillae to 0.5mm long ending in simple, short trichome, surface glabrous. Seeds too young to describe. noTes 1) Acalypha gracilipes has been usually treated as a variety of A.integrifolia (or its synonym, A.commersoniana Baill. ex Müll. Arg.) or as a variety of A.reticulata (for which the correct name is A.filiformis), both of which are endemics of Mascarene Islands, but A.gracilipes can be clearly distinguished from these species by its denticulate female bracts and glabrous ovary and capsule surfaces, vs entire female bracts and hairy ovary and capsule surfaces in A.filiformis and A.integrifolia; 2) Acalypha gracilipes can be distinguished from both A.urophylla and 45°E 15°S 20° 25° 300 km 50° Acalypha gracilipes fig. 33. — Distribution map of Acalypha gracilipes Baill. in Madagascar.
441 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) A.pervilleana mainly by its glabrous leaves with crenate to sub - dentate margins and obtuse to subacute apices, glabrous female bracts, and papillose-hispid and glabrous ovaries vs pubescent leaves with serrate margins and acuminate (A.urophylla) or usually caudate (A.pervilleana) apex, pubescent female bracts, and hispidulous ovaries with long papillae. In addition, the female bracts of A.pervilleana have a prominent central tooth, which is absent in both A.gracilipes and A.urophylla; and 3) Leandri (1942) included the collection H. Perrier de la Bâthie 9746 as Acalypha reticulata var. cloiselana Leandri, which we treat as a synonym of A.gracilipes; however, the specimen of this collection in P (P00513170) corresponds to A.urophylla. 19. Acalypha hispida Burm.f. Flora Indica 303, pl. 61, f. 1 (Burman 1768).— Ricinocarpus hispidus (Burm.f.) Kuntze, Revisio Generum Plantarum 2: 618 (Kuntze 1891).— Type: Habitat in India. Burman 1768: 302, table 61. iconography.— André (1898: 227); Hooker (1899: t. 7632). e Tymology .— The epithet refers to the long and laciniate styles of the pistillate inflorescence according Sagun etal. (2010), or perhaps to the hispid surface of the ovary. DisTribuTion anD habiTaT.— Cultivated species in Madagascar (Anosy) and Seychelles (Mahé) (Fig. 34). preliminary conservaTion assessmenT.— Acalypha hispida is widely cultivated in warm areas worldwide and not known in the wild; it sometimes escapes and is found along forest edges and roads. We assess A.hispida as Least Concern (LC). maTerial examineD.— 3 collections. Madagascar. Seligson, D. 652 (GH[GH01097235]). Seychelles. Hein, G. 10 (GH[GH01097236]); Jeffrey, C. 599 (K). references.— Bojer (1837: 25); Baillon (1861: 274); Palacký (1907: 25); Robertson (1989: 199); Montero Muñoz etal. (2018a: 99). DescripTion Shrubs or subshrubs, probably evergreen, to 4m tall, possibly dioecious, only female plants known. Branches tomentose and short-hispid, glabrescent when mature. Axillary buds naked, densely pubescent with simple, short trichomes. Stipules to 1cm long, triangular-lanceolate, appressedpubescent. Petioles (2-)4-11cm long, indumentum similar to that on young branches, glabrescent. Leaf blades not variegated, (8-)10-17(-19)× (5.5-)7-10(-13) cm, usually ovate to elliptic-lanceolate, sometimes subrhombic, membranous; base usually rounded, sometimes subcuneate to subcordate; apex acute to subacuminate, acumen to 5mm long, obtuse, mucronate; margin serrate to crenate-serrate, teeth acute or obtuse, sometimes slightly callose-edged; upper and lower surfaces subglabrous, with some sparse, simple trichomes on veins; venation actinodromous, basal veins 5 or 7, secondary veins 7-9 per side. Stipels absent. Inflorescences spiciform, unisexual, axillary. Male inflorescences unknown. Female inflorescences pendulous, densely flowered, to 40cm long; peduncle to 30mm long, pubescent with simple, curved trichomes; bracts numerous, sessile, not enlarging in fruit, to 1mm long, elliptic-lanceolate, sparsely hairy and ciliate; margin entire; bracteoles absent. Male flowers unknown. Female flowers 4-6 per bract, sessile; sepals 3-4, to 1mm long, triangular-lanceolate, ciliate; ovary c.1 mm diameter, 3-lobed, smooth, surface densely hispid; styles 3, to 8mm long, distinct, glabrous, each divided into 6-16 segments. Allomorphic flowers not seen. Capsules and seeds not seen. noTe This shrub, native to Melanesia or Malesia (Sagun etal. 2010), is frequent in gardens throughout the tropics and rarely appears naturalized. As all plants are pistillate, it can only reproduce clonally. We found collections from Madagascar and Seychelles where it is cultivated. It has been reported from Madagascar (Palacký 1907), Mauritius (Bojer 1837; Baillon 1861), La Réunion (Baillon 1861: 274) and the Seychelles (Robertson 1989). 20. Acalypha humbertii Leandri Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 274 (Leandri 1942).— Type: Madagascar. Prov. Toliara, vallées du Mangoky et de l’Isahaina, aux environs de Beroroha, 200m, X.1933, H. Humbert 11289 (lecto-, designated by Montero Muñoz etal. [2018a: 99]: P[P00508400]; isolecto-, P[P00508399]).— Former syntypes: Madagascar. Prov. Toliara, Bassin supérieur de Mandrare du Sud-Est, entre le col de Vavara et la vallée de la Manambolo, 700-1200m, 20-22.XII.1928, H. Humbert 6758 (P[P00508401, P00508402, P00508403, P00508404]). 5°S 45°E 50° 55° 10° 15° 20° 25° Acalypha hispida fig. 34 . — Distribution map of Acalypha hispida Burm.f. in Madagascar and Seychelles.
442 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. AB CD EF G fig. 35. — Mature female bracts: A, Acalypha decaryana Leandri (R. Decary 2985); B, A. emirnensis Baill. (G. McPherson 14590); C, A. filiformis Poir. (P. Commerson s.n.); D, A. fimbriata Schumach. & Thonn. (J. Audru 40466); E, A. gillespieae G.A.Levin & I.Montero (L. Gillespie et al. 10692); F, A. gracilipes Baill. (R. Rabevohitra 2054); G, A. humbertii Leandri (W. Kaudern 222). Illustration by Iris Montero Muñoz. Scale bars: A, C-G, 1 mm; B, 2 mm.
443 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) iconography.— Fig. 35G. eTymology.— The epithet honors French botanist Henri [JeanHenri] Humbert (1887-1967). D isTribuTion anD habiTaT .— Endemic to Madagascar (Menabe, Atsimo-Andrefana, Androy, and Anosy). Dry deciduous forest and dry spiny thickets. On basement rocks and sandstone. Altitudinal range (50-) 200-800 (-1000) m (Fig. 36). preliminary conservaTion assessmenT.— The EOO of Acalypha humbertii is estimated to be 37 244km2 and its AOO 20km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in dry deciduous forest and spiny thickets (see assessment in A.decaryana). The remaining original vegetation is still decreasing. Due to its small AOO, the small number of locations (5), and ongoing habitat loss, Acalypha humbertii is assessed as Endangered: EN B2ab(ii,iii,iv). maTerial examineD.— 11 collections. Madagascar: Bosser, M.J. 10234 (P[P00324559]); Decary, R. 15637 (P[P00324536, P05510104]); Humbert, H. 6758 (P[P00508401, P00508402, P00508403, P00508404]), 11289 (P[P00508399, P00508400]); Jongkind, C.C.H. 3708 (MO, TAN, WAG[WAG.1578689]); Kaudern, W. 222 (S[S17-37631]), 289 (S[S17-37632]); Leandri, J. 3532 (P[P00324507, P05543682]), 3535 (P[P05547075]), 3882 (P[P00324510]), 3900 (P[P00324511, P05543681]). references.— Govaerts etal. (2000: 67); Montero Muñoz etal. (2018a: 99). DescripTion Shrubs, deciduous, to 1.5m tall, highly divaricately branched, monoecious. Branches densely pubescent to subvelutinous, glabrescent when mature. Axillary buds subfusiform, to 4× 2mm, perulate, perules 2, imbricate, chartaceous, dark brown, glabrous. Stipules to 3mm long, linear, densely pubescent. Petioles 0.5-1.5 long, indumentum similar to that on young branches. Leaf blades 2.4-3.7× 1.4-2.6cm, ovate-lanceolate to elliptic-lanceolate, subchartaceous; base rounded to subacute; apex rounded to subacuminate, acumen to 2mm long, rounded; margin crenate-serrate, teeth rounded, callose-edged; upper and lower surfaces velutinous with simple, short trichomes; venation actinodromous, basal veins 3, secondary veins 4-5 per side. Stipels absent. Inflorescences spiciform, androgynous, axillary, to 3cm long, mostly male with 1 female bract at base; peduncle to 5mm long, it and rachis pubescent with simple, short trichomes. Female segment: bract 1, sessile, enlarging in fruit to 6× 8mm, reniform, pubescent with simple, short trichomes; margin subentire to irregularly dentate, teeth c.13, triangular, central tooth not prominent; bracteoles to 0.8mm long, oblong-lanceolate, sparsely hairy. Male segment persistent, to 1.8cm long; flowers glomerate; bracts to 0.5mm long, triangular, sparsely hairy. Male flowers: pedicel to 0.6 mm long, sparsely hairy; buds not seen. Female flowers 1 per bract, sessile; sepals 3, to 0.7mm long, ovate-lanceolate, ciliate; ovary c.1 mm diameter, 3-lobed, echinate, projections linear, to 1.5mm long, surface hispid with simple, rigid trichomes; styles 3, to 3.5mm long, connate at base, sparsely hairy with simple, long, appressed trichomes at rachis, each divided into c.8 segments. Allomorphic flowers not seen. Capsules to 2.5 mm diameter, echinate, projections linear, to 1.5mm long, surface hispid with simple, rigid trichomes. Seeds c.1.5 × 1.2 mm, pyriform, minutely foveolate. 21. Acalypha indica L. Species Plantarum 2: 1003 (Linnaeus 1753).— Type: India. s.l., s.d., Herb. Hermann 3: 2, 434. (lecto-, designated by Radcliffe-Smith [1986: 65]: BM[BM000621782]; isolecto-, BM[BM000621236]). iconography.— Rheede tot Drakestein (1692: t. 81); Anonymous (1694-1718: t. 215); Kirtikar& Basu (1918: t. 874); Fig. 46A. eTymology.— The epithet refers to India, the locality of the type collection. DisTribuTion anD habiTaT.— Acalypha indica is widely distributed in the Paleotropics and introduced in the Americas (Cardiel etal. 2023 in press); it probably is introduced in Madagascar (Diana, Analamanga, and Ihorombe), the Comoros Archipelago (Anjouan, Moheli and Mayotte), the Mascarene Islands (Mauritius and LaRéunion), the Seychelles (Inner and Outer islands), and the Scattered Islands (Glorioso Islands). It is a weedy species associated with cultivated areas, paths, and beaches. Altitudinal range sea level to 450m (Fig. 37). preliminary conservaTion assessmenT.— Acalypha indica is widely distributed in the Paleotropics; it has a large EOO and AOO, exceeding the threshold of any threat category. Therefore, we assess A.indica as Least Concern (LC). 45°E 15°S 20° 25° 300 km 50° Acalypha humbertii fig. 36. — Distribution map of Acalypha humbertii Leandri in Madagascar.
444 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. maTerial examineD.— 94 collections. Comoros Archipelago: Barthelat, F. 113 (K, G, MAO, MO[MO-2966222], P[P00229090]), 170 (G, K, MAO, MO[MO-2966225], P[P00229156]); Boivin, L.H.s.n. (P[P00196285]), s.n. (P[P00196286]), s.n. (P[P00196287]); D’Arcy, W.G. 17553 (MO[MO-2966220]); Floret, J.J. 1196 (P[P00184951]), 1197 (P[P00184952]); Hildebrandt, J.M. 1660 (P[P00196289]); Hoffmann, P. 413 (K, MAO, MO[MO-2966227], P[P00538301], TAN); Labat, J.-N. 3314 (P[P00209769]); Lartigau, C. 19 (P[P00246081]); Pignal, M. 1075 (P[P00157183]); Pobéguin, C.H.O. 14 (P[P00196234, P00196235]); Rakotozafy, A. 1194 (P[P05547071]); Rouhan, G. 33 (P[P00205950]); Waterlot, M. 884 (P[P00196290]). Madagascar. Bosser, M.J. 13231 (P[P04779813]); d’Alleizette, Ch. 1240 M (P[P00508397]); Debray, M.M. 1584-D (P[P04779834, P04779835]); Hildebrandt, J.M. 2915 (BREM[BREM_0001791], G, L[L0241703], P[P00508393, P00508394, P00508395, P00508396]); Humblot, L. 18 (P[P00508392]); Perrier de la Bâthie, H. 9592 (P[P00508391]); Pervillé, A. 729 (P[P00508390]); Rotereau, L.s.n. (P[P05510099]); Viguier, R. 104 (P[P00508389, P05481896]). Mauritius. Anonymous s.n. (G[G00324726]); Ayres, P. 62 (K); Bonnay, M.s.n. (P[P04779496]); Bouton, L. 22 (P[P04779493]); s.n. (G[G00324716]), s.n. (G[G00324720]); Commerson, P. 649 (L[L0241705]), 651 (P[P04779489]); Coode, M.J.E. 4423 (K); Cowper, S.G. 59 (K); Guého, J. 17350 (K); Hering? s.n. (P[P04779502]); Jussieu, A.s.n. (P[P04779495]); Lorence, D. 1339 (MO[MO-2966226]). La Réunion. Anonymous s.n. (P[P04779504]); Boivin, L.H. 1389 (P[P04779494]); Bosser, M.J. 9389 (P[P04779491, P04779492]); Commerson, P.s.n. (P[P04779488]); Desvaux, A.N. 10 (P[P04779503]); du Petit-Thouars, L.M.A.s.n. (P[P04779499]), s.n. (P[P04779500]); Friedmann, F. 1585 (P[P04779501]); Richard, M. 667 (GH, P[P04779498]), s.n. (P[P04779490]), s.n. (P[P04779497]). Scattered Islands. Frazier, J. 105 (US[US01287375]). Seychelles. Alluaud, R.s.n. (GH, P[P04779487]); Anonymous s.n. (P[P04779486]); Blackmore, S.B. 77407 (K); Brunet 035 (K); Cherbonnier s.n. (P[P04779829]); Fosberg, F.R. 48688 (GH[GH01097240], MO[MO-2966229], NY[NY3091187], US[US01287383]), 49844 (MO[MO-2966224]), 52070 (MO[MO-2966230], US[US01287382]); Frazier, J. 30 (K), 513 (US[US01287374]), 802 (US[US01287373]); Friedmann, F. 3545 (P[P00887492]), 4673 (P[P00887491]); Gardiner, J.S. 9 (K), 27 (K), 143 (K); Gwynne 874 (K); Hebert, H. I. 103 (GH[GH01097241]); Horne, J. 312 (K); Jeffrey, C. 709 (L[L0241693], NY, P[P00887490]); Lombat, M. 408 (K); Procter, J. 4045 (K), 4224 (K), 4322 (K), 4347 (K); Renvoize, S. A. 721 (BR[BR0000014627996], L[L0241691], P[P05481874]), 1151 (BR[BR0000014628009], P[P05481873]), 1155 (BR[BR0000014624612], L[L0241251, L0241252]); Robertson, S.A. 2443 (MO[MO-2966228], PRE, US[US01287372], WAG[WAG.1931643]), 2494 (K), 2716 (K), 3044 (K); 3146 (K); Rotereau, L.s.n. (P[P05510098]); Stodd, D.R. 1382 (K), 1459 (K); Stoddart, D.R. 7082 (MO[MO-2966221], NY[NY3091188]); Wood, D. 1452 (K), 1469 (K); Wright, P.A. 16 (K), 25 (K). references.— Bojer (1837: 285); Baillon (1861: 274); Müller Argoviensis (1865: 42; 1866: 868); Baker (1877: 314); Müller Argoviensis (1882: 27); Baillon (1895b: 1197); Cordemoy (1895: 342); Palacký (1907: 25); Voeltzkow (1917: 447); Hemsley (1919: 148); Pax& Hoffmann (1924: 33); Leandri (1935: 43); Leandri (1942: 256); Renvoize (1975: 152); Coode (1982: 69, 78); Robertson (1989: 200); Govaerts etal. (2000: 68); Sagun etal. (2006: 124); Montero Muñoz etal. (2018a: 99). DescripTion Annual herbs, to 0.6(-0.8) m tall, monoecious. Branches tomentose to laxly pubescent with simple, curved trichomes, sometimes also with sparse short, subsessile, glandular trichomes, glabrescent when mature. Axillary buds naked, pubescent with simple, short, appressed trichomes. Stipules to 1mm long, triangular-lanceolate, margin hispidulous-ciliate and with some sessile glands. Petioles slender, (1.5-)2.5-6.5cm long, indumentum similar to that on young branches. Leaf blades (1.5-)2-4(-6)× (1-)1.5-3(-3.5) cm, usually broadly ovate-lanceolate, sometimes spathulate or subrhombic, membranous; base cuneate; apex acute to subacuminate; margin entire toward base, serrate above, teeth subacute, callose-edged; upper surface subglabrous, with simple, short, appressed trichomes on veins; lower surface laxly pubescent to subglabrous; venation actinodromous, basal veins 3 or 5, secondary veins 4-5 per side. Stipels absent. Inflorescences spiciform, androgynous, axillary, to 9cm long, mostly female with short male segment; peduncle to 10mm long, pubescent with simple trichomes, sometimes also with some subsessile glandular trichomes. Female segment laxly flowered, to 8cm long; bracts 7-9, sessile, enlarging in fruit to 8× 12mm, reniform, sparsely hairy with simple, curved trichomes; margin crenate-denticulate, teeth 8-9, broadly triangular, central tooth not prominent; bracteoles absent. Male segment persistent, to 1.5cm long; flowers glomerate; bracts to 0.8mm long, elliptic-lanceolate, sparsely hairy. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds 0.5 mm diameter, sparsely hairy. Female flowers 1 per bract, sessile; sepals 3, to 0.5mm long, triangular-lanceolate, ciliate; ovary c.0.5 mm diameter, 3-lobed, smooth, surface pubescent; styles 3, to 4mm long, distinct, glabrous, each divided into 1-2 segments. Allomorphic flowers sometimes present at inflorescence apex; pedicel filiform, to 5mm long, sparsely hairy; sepals 3, to 0.5mm long, lanceolate, ciliate; ovary 1-lobed, to 5°S 45°E 50° 55° 10° 15° 20° 25° Acalypha indica fig. 37. — Distribution map of Acalypha indica L. in the whole WIOR region.
445 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) 2× 3mm, pubescent, distally fimbriate; style 1, to 1.5mm long, glabrous. Capsules to 2 mm diameter, smooth, surface pubescent. Seeds c.1.5 × 1 mm, pyriform, minutely foveolate. 22. Acalypha integrifolia Willd. Species Plantarum 4 (1): 530 (Willdenow 1805).— Type: Mauritius. s.l., s.d., Anonymous, s.n. (lecto-, designated by Coode [1978: 39]: B[B-W17834-020]). Tragia colorata Poir., Encyclopédie méthodique. Botanique 7: 725 (Poiret 1806).— Acalypha colorata (Poir.) Spreng., Systema vegetabilium 3: 879 (Sprengel 1826).— Acalypha commersoniana var. obtusifolia f. colorata (Poir). Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Acalypha integrifolia var. colorata (Poir.) Pax& K.Hoffm., Das Pflanzenreich (Engler) 147, 16 (Heft 85): 106 (Pax& Hoffmann 1924).— Type: Mauritius. “Cette plante croît dans les indes orientales& à l’Île de France”, s.l. s.d., P. Commerson s.n. (holo-, P-LAM[P00382140]). Acalypha reticulata var. longifolia Müll.Arg., Linnaea 34: 32 (Müller Argoviensis 1865).— Acalypha integrifolia var. longifolia (Müll.Arg.) Coode, Kew Bulletin 34: 41 (Coode 1979).— Type: Mauritius. s.l., s.d., L. Bouton s.n. (holo-, GDC[G00324522]). Acalypha reticulata var. longifolia f. aberrans Müll.Arg., Linnaea 34: 32 (Müller Argoviensis 1865).— Type: Mauritius. “In sylvis Mauritii”, 1833, W. Bojer s.n. (holo-, GDC[G00324521]). Acalypha commersoniana Baill. ex Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 849 (Müller Argoviensis 1866).— Acalypha commersoniana var. brevifolia Baill. ex Müll. Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Type: Mauritius. “Cum var. praecedentibus”, s.d., Anonymous, s.n. (Hb. Willd. fol. 17834 page1) (lecto-, designated by Montero Muñoz etal. [2018a: 100]: B[BW17834-010]).— Former syntypes: Mauritius. s.l., s.d., L. Bouton s.n. GDC[G00324543]. Acalypha commersoniana var. acutifolia Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 849 (Müller Argoviensis 1866).— Acalypha commersoniana var. acutifolia f. purpurea Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 849 (Müller Argoviensis 1866).— Type: Mauritius. s.l., s.d., F. W. Sieber 181 pr. p. (lecto-, designated by Montero Muñoz etal. [2018a: 100]: GDC[G00324550]).— Former syntype: Mauritius. s.l., s.d., L. Bouton s.n. GDC[G00324559]. Acalypha commersoniana var. acutifolia f. purpureo-marginata Müll. Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 849 (Müller Argoviensis 1866).— Type: Mauritius. s.l., s.d., Hb. Boiss. s.n. (holo-, GDC[G00324557]; iso-, K[K000431097]). Acalypha commersoniana var. acutifolia f. concolor Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 849 (Müller Argoviensis 1866).— Acalypha integrifolia var. concolor (Müll.Arg.) Pax& K.Hoffm., Das Pflanzenreich (Engler) 147, 16 (Heft 85): 106 (Pax& Hoffmann 1924).— Type: Mauritius. “Cum form. praecedentibus”, s.d., J. B. G. M. Bory s.n. (lecto-, designated by Montero Muñoz etal. [2018a: 100]: GDC[G00324554]).— Former syntype: Mauritius. s.l., s.d., F. W. Sieber 182 GDC[G00324556]. Acalypha commersoniana var. longifolia Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Type: Mauritius. s.l., s.d., L. Bouton s.n. (lecto-, designated by Montero Muñoz etal. [2018a: 101]: GDC[G00324553]; isolecto-, GDC[G00324551, G00324552], K[K000431101]). Acalypha commersoniana var. parvifolia Baill. ex Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Acalypha integrifolia var. parvifolia (Baill. ex Müll.Arg.) Pax& K.Hoffm., Das Pflanzenreich (Engler) 147, 16 (Heft 85): 106 (Pax& Hoffmann 1924).— Type: Mauritius. “Cum var. praecedentibus”, s.d., F. W. Sieber 369 pr.p. (holo-, GDC[G00324538]). Acalypha commersoniana var. obtusifolia Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Acalypha commersoniana var. obtusifolia f. discolor Müll. Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Type: Mauritius. “Cum praecedentibus”, s.d., F. W. Sieber 181 pr. p. (holo-, GDC[G00324558]). Acalypha commersoniana var. obtusifolia f. unicolor Müll.Arg., Prodromus Systematis Naturalis Regni Vegetabilis 15 (2): 850 (Müller Argoviensis 1866).— Type: Mauritius. s.l., s.d., F. W. Sieber 178 (holo-, GDC[G00324539]; iso-, P[P04779345, P04780015]. Acalypha discolor Bojer, Hortus Mauritianus 286 (Bojer 1837) nom. nud. Tragia macrophylla Wall., A Numerical List of dried specimens of plants in the East India Company’s Museum n. 7796 (Wallich 1847) nom. nud. Tragia lobata Wall., A Numerical List of dried specimens of plants in the East India Company’s Museum n. 7796 (Wallich 1847) nom nud. Acalypha commersoniana Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. Acalypha commersoniana var. concolor Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. Acalypha commersoniana var. discolor Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. Acalypha commersoniana var. parvifolia Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. Caturus sessilis Thouars ex Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. Tragia fruticosa Commers. ex Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. Tragia obtusata Vahl. ex Baill., Adansonia, recueil d’observations botaniques 1: 267 (Baillon 1861) nom. nud. iconography.— Figs 38; 46B; 54A, B. eTymology.— The epithet refers to the entire leaf margins. DisTribuTion anD habiTaT.— Endemic to Mascarene Islands (Mauritius and La Réunion). In Mauritius, in lowland evergreen moist forest, dry forest, and grasslands. In La Réunion in lowland evergreen moist forest. Altitudinal range in Mauritius (150-) 200700m and in La Réunion (150-) 380-1300m (Fig. 39). preliminary conservaTion assessmenT.— Acalypha integrifolia is known from Mauritius and La Réunion. Its EOO is estimated to be 9 004.818km2 and its AOO 80.000km2, which is less than the 500km 2 threshold of the B2 subcriterion of the Endangered category. For the status of forests on Mauritius forests, see the assessment of A.crateriana comb.nov., for those on La Réunion, see the assessment of A.filiformis. Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha integrifolia is assessed as Near Threatened (NT). It meets the EOO and AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category.
446 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. maTerial examineD.— 152 collections. Mauritius: Allan, D.s.n. (K); Anonymous s.n. (B[B-W17834-020]); Anonymous s.n. (B[BW17834-010]); Anonymous s.n. (G[G00324520]); Anonymous s.n. (G[G00324541]); Anonymous s.n. (G[G00324542]); Anonymous s.n. (G[G00324547]); Anonymous s.n. (B[B-W17834-020]); Anonymous s.n. (P[P04779337]); Anonymous s.n. (P[P04779383]); Anonymous G E D A F H B C fig. 38. — Acalypha integrifolia Willd.: A, flowering branch with young leaves; B, flowering branch with mature leaves; C, detail of node, stipules, and petiole base; D, mature female bract; E, ovary and styles; F, calyx of the female flower; G, capsule; H, seed. Based on P. commerson s.n. (A, D-G), W. Bojer s.n. (B), F. R. Fosberg 52683 (C), L. Bernardi 14866-bis (H). Illustration by Laura González Hernández. Scale bars: A, B, 1 cm; C, 2 mm; D-H, 1 mm.
447 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) s.n. (P[P04779384]); Anonymous s.n. (P[P04779399]); Ayres, P.B.s.n. (P[P04779484]); Barlay, C. 2804 (K); Bélanger, M.s.n. (G); Bernardi, L. 14719 (C, G, K, L[L0438912], MO[MO-2965780], P[P04779456]), 14754 (C, G, K, MO[MO-2452044], P[P04779459]), 14866 (C, G, K, P[P04779471]), 14868 (C, K, MO[MO-2452043], P[P04779461]); Blackburn, J.s.n. (P[P04779482]); Boivin, L.H. 1558 (P[P04779408]), s.n. (P[P04779323]) s.n.(P[P04779410]), s.n. (P[P04779411]), s.n. (P[P04779455]) ), s.n. (P[P04779458]); Bojer, W.s.n. (G[G00324521]), s.n. (P[P04779478]), s.n. (P[P04779479]); Bory de Saint-Vincent, J.B.G.s.n. (G[G00324531]), s.n. (G[G00324554]); Bouton, L. 24 (P[P04779437]), s.n. (G[G00324522]), s.n. (G[G00324543]), s.n. (G[G00324545]), s.n. (G[G00324548]), s.n. (G[G00324549]), s.n. (G[G00324551, G00324552, G00324553], K[K000431101]), s.n. (G[G00324555]), s.n. (G[G00324559]); Brown, Ab. s.n. (P[P04779418]); Commerson, P. 655 (L[L0160752], P[P04779325]), 656 (L[L0160754], P[P04779339]), s.n. (P[P00382140]), s.n. (P[P04779327]), s.n. (P[P04779457]), s.n. (P[P04779466]), s.n. (P[P04779467]); Coode, M.J.E. 4836 (K, P[P04779409]), 4843 (K, P[P04779407]), 4845 (P[P04779406]), 4846 (K); Desvaux, A.N. 184 (P[P04779412]); D’Urville, D.s.n. (P[P04779385]), s.n. (P[P04779386]), s.n. (P[P04779387]), s.n. (P[P04779388]), s.n. (P[P04779389]), s.n. (P[P04779390]); Fosberg, F.R. 52683 (US[US01287368]), 52685 (K, MO[MO-2965776], US[US01287369]); Guého, J. 15166 (P[P04779336]); Hb. Boiss. s.n. (G[G00324557], K[K000431097]); Herb. de Tristan s.n. (P[P04786006]); Herb. Parker s.n. (P[P04779419]); Hooker, W.J.s.n. (BR[BR0000014628047]); Hugenie, M? s.n. (G); Jussieu, A.s.n. (P[P04779404]); Lahaie 28 (P[P04779392]); Lorence, D. 133 (MO[MO-2965778]), 134 (MO[MO-2965777]), 1599 (MO[MO-2965779]), 1838 (G, MO[MO-2452047], P[P04779462]), 1960 (K, MO[MO-2452049], P[P04779448]), 4525 (MO[MO-2452050]); Martin 589 (G); Néraud s.n. (G); Perrotet, M.s.n. (P[P04779340]), s.n. (P[P05481894]); Pierre, L. 5338 (P[P04779483]); Richard, M. 178 (P[P04779344]), s.n. (P[P04779405]), s.n. (P[P04779441]); Schleiden, M.J.s.n. (P[P04779481]); Sieber, F.W. 65 (G), 66 (G), 178 (G[G00324539, G00324540], L[L0241390, L0241391, L0241392], P[P04780015, P04779345]), 181 (BR[BR0000014628054], G[G00324550, G00324558], L[L0241755, L0241756, L0241757, L0241758], P[P04779364, P04779396, P04779398]), 182 (G[G00324556], L[L0160809, L0160810], P[P04779391, P04779400, P04779401]), 182 pr. p. (G[G00324546]), 369 (G[G00324538]), 2042 (P[P04779397]); Splitgerber, F.L.s.n. (L[L0241754]); Thonin s.n. (G); Vesco, M.s.n. (P[P04779324]), s.n. (P[P04779326]), s.n. (P[P04779334]), s.n. (P[P04779393]), s.n. (P[P04779394]), s.n. (P[P04779468]), s.n. (P[P04779470]). La Réunion. Anonymous 492 (P[P04779378]); Anonymous s.n. (G[G00324544]); Badré, F. 794 (BR[BR0000014628078], K, P[P04779445, P04779447]); Balfour, I.B.s.n. (K); Bernier, M. M21 (P[P04779416]); Boivin, L.H. 655 (G), 1391 (P[P04779469, P04779414, P04779415]); Bosser, M.J. 21468 (K, P[P04779449, P04779464]), 22576 (P[P04779453]); Bouton, L.s.n. (P[P04779480]); Breón, M.s.n. (P[P04779827]); Cadet, Th. 647 (K), 742 (K, P[P04779452]), 3917 (K, P[P04779460]), 4474 (K); Commerson, P.s.n. (BR[BR0000021450174]), s.n. (P[P04779357]); Coode, M.J.E. 5221 (K); d’Alleizette, Ch. 902 (L[L0241753]); Delessert, A.s.n. (G); Desvaux, A.N. 181 (P[P04779372]), 186 (P[P04779413]), s.n. (P[P04779374]); du Petit-Thouars, L.M.A.s.n. (P[P04779395]), s.n. (P[P04779402]), s.n. (P[P04779403]), s.n. (P[P04779472]); Frappier, M. 2 (GH, P[P04779477]), 12 (P[P04779474]), s.n. (P[P04779475]), s.n. (P[P04779476]); Friedmann, F. 1007 (BR[BR0000014628061], G, P[P04779446, P04779463]), 1026 (P[P04779443]), 1027 (P[P04779444]), 1947 (K, P[P04779442]); Giraudy, H. 18107 (P[P04780018]); Léman, D.S.s.n. (P[P04779341]), s.n. (P[P04779373] pr.p.); Monin, M.s.n. (P[P04779365]); Richard, M. 192 (P[P04779376]), 193 (P[P04779450]), 201 (P[P04779379, P04779381, P04779436, P04779440]), 202 (P[P04779382, P04779439]), 490 (P[P04779465]), 658 (L[L0242527]), s.n. (P[P04779367]), s.n. (P[P04779377]), s.n. (P[P04779380]), s.n. (P[P04779438]), s.n. (P[P04779579] pr.p.), s.n. (P[P05604455] pr.p.); Rivals, P.s.n. (P[P04779451]). references.— Bojer (1837: 286); Baillon (1858: 443) as A.colorata; Müller Argoviensis (1866: 850); Baker (1877: 315) as A.colorata; Cordemoy (1895: 342) as A.colorata; Palacký (1907: 24) as A.commersoniana; Palacký (1907: 25); Pax& Hoffmann (1924: 105); Coode (1982: 69); Robertson (1989: 200); Govaerts etal. (2000: 69); Montero Muñoz etal. (2018a: 100). DescripTion Shrubs, leaf persistence unknown, to 2m tall, monoecious. Branches densely appressed-pubescent, rugose and glabrescent when mature. Axillary buds ovoid, to 1.2× 0.7mm, perulate, perules 2, valvate, membranous, brownish, appressed-pubescent. Stipules to 8mm long, triangular-lanceolate, with reddish midrib and two greenish wings, subglabrous, with some simple, minute, appressed trichomes on midrib. Petioles thick, 0.5-2.2cm long, indumentum similar to that on young branches, glabrescent. Leaf blades (5-)9-15.5× (1.5-)3.5-6.5cm, usually elliptic-lanceolate to oblong-lanceolate, sometimes slightly panduriform, chartaceous to coriaceous; base truncate to subcordate; apex rounded, mucronate; margin entire to denticulate, revolute, sometimes reddish or pinkish, teeth, if present, minute, subacute; upper and lower surfaces glabrous; venation prominent on lower surface, sometimes reddish, pinnate, secondary veins 15-18 per side, straight, parallel. Stipels absent. Inflorescences spiciform, male, and solitary female bracts, axillary. Male inflorescences to 9cm long; subsessile, peduncle to 2mm long, indumentum similar to that on petioles; flowers glomerate; bracts to 0.7mm long, triangular-rhombic, appressed-pubescent. Solitary female bracts sessile, enlarging in fruit to 6× 6.5mm, suborbicular, becoming coriaceous, glabrous; margin entire; bracteoles absent. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds to 75 km Mauritius La Réunion Acalypha integrifolia fig. 39 . — Distribution map of Acalypha integrifolia Willd. in the Mascarene Islands.
448 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. 0.5mm diameter, sparsely hairy. Female flowers 2-3 per bract, sessile; sepals 3, to 1.5mm long, ovate-lanceolate, pubescent with simple, appressed trichomes, especially at margin; ovary c.1 mm diameter, 3-lobed, papillose-hispid, papillae to 1mm long, ending in a simple trichome to 0.5mm long, with appressed trichomes, surface appressed-pubescent; styles 3, to 5mm long, distinct, sparsely hairy with simple, appressed trichomes on rachis, each divided into c.12 segments. Allomorphic flowers sometimes present, axillary; pedicel filiform, to 19mm long, glabrous; sepals to 2mm long, ovate-lanceolate, subglabrous, with some simple appressed trichomes, margin with sessile glands; ovary not seen. Capsules to 4 mm diameter, papillose-hispid, papillae triangular, to 1mm long, with simple, appressed trichomes, surface appressed-pubescent. Seeds c.2 mm diameter, subglobose, foveolate. noTes 1) Coode (1979, 1982) accepted three subspecies and six varieties within Acalypha integrifolia, but we find the varieties he placed within subsp. integrifolia overlap too much to accept them as distinct taxa. See notes for A.marginata for our treatment of what Coode treated within A.integrifolia subsp. marginata and subsp. panduriformis; 2) Boivin mixes two species under collection Boivin 1391 (A. filiformis [P04779576, P05604483] and A. integrifolia [P04779414, P04779415, P04779417, P04779329, P04779469]); and 3) the following specimens P00324571, P00324572, P00324573 and P00324574 are labeled as being from Madagascar (without a specific locality), but the labels are not original and A. filiformis is endemic to the Mascarene Islands. The location given for these specimens almost certainly is incorrect. 23. Acalypha isaloensis I.Montero& Cardiel Systematic Botany 45 (1): 122 (Montero Muñoz etal. 2020a).— Type: Madagascar. Ihorombe (Fianarantsoa prov.), Massif de l’Isalo, à l’est de Ranohira, Andohakandrareza, au SE de la Piscine, 810m, 22°33’S, 45°21’E, 14.II.1990, J. N. Labat 2119 (holo-, P[P00513133]; iso-, ILLS, K).— Paratypes: Madagascar. Ihorombe (Fianarantsoa prov.), Isalo, canyon des Singes, 746m, 22°29’12.239”S, 45°22’43.381”E, 15.IV.1967, H. Jacquemin H278J (P[P04804757, P04804758]); Isalo, canyon des Rats, pied des falaises le long du ruisseau, 899m, 22°28’47.759”S, 45°22’39.601”E, III.1960, M.Keraudren-Aymonin 402 (P[P04804755]); Ranohira, Isalo-Gebirge, 900m, 22°33’20.307”S, 45°25’09.167”E, 22.XII.1959, H.-J. Schlieben 8236 (B, BR[BR0000021450358], G, K). iconography.— Montero Muñoz etal. (2020a); Fig. 46C. eTymology.— The epithet refers to type locality, the Isalo massif (Madagascar), to which this species appears to be endemic. DisTribuTion anD habiTaT.— Endemic to Madagascar (Ihorombe). Dry deciduous forest. On sandstone. Altitudinal range 746-900m (Fig. 40). preliminary conservaTion assessmenT.— The EOO and AOO of A.isaloensis are estimated to be 8km 2 . This species appears to be a narrow endemic known only from the Isalo massif. All collections are from the Isalo National Park, which has been a category II protected area (Dudley 2008) since 1962. This protected area is threatened by burning to improve grazing, which is a cultural heritage of the Bara people, as well as logging and the impact of introduced species (Kull 2004; Swierkosz 2007; Andreone etal. 2013). In recognition of its restricted geographic range (a single known location) and the cited threats, A.isaloensis is assigned a preliminary conservation status of Critically Endangered: CR B2ab(i,iii,iv). maTerial examineD.— 3 collections. Madagascar: Jacquemin, H. H278J (P[P04804757, P04804758]); Keraudren-Aymonin, M. 402 (P[P04804755]); Labat, J.-N. 2119 (K, P[P00513133]). DescripTion Shrubs or subshrubs, leaf persistence unknown, to 0.8m tall, monoecious. Branches puberulent with sparse simple, short, trichomes, glabrescent when mature. Axillary buds ovoid, to 1× 0.7mm, perulate, perules 2, valvate, membranous, brownish, densely pubescent with simple, short, appressed trichomes. Stipules to 1.2(-2) mm long on young branches, sometimes reduced to 0.5mm long on older branches, triangular-lanceolate, apex acute, with appressed trichomes, margin with sessile glands. Petioles (2-)3-4.5cm long, indumentum similar to that on young branches. Leaf blades (5-)6-10(-11.5)× 2-3.5(-4) cm, elliptic to oblong-lanceolate, membranous; base rounded to subcuneate; apex subacuminate, acumen to 10mm long, rounded, mucronate; margin sparsely serrulate, teeth minute, obtuse, callose-edged; upper surface laxly pubescent with simple, short trichomes; lower surface indumentum similar to that on upper surface but denser; axils of the secondary veins with pocket-shaped domatia; venation prominent on upper surface, pinnate, secondary veins 5-10 per side. Stipels absent. Inflorescences spiciform, male, and solitary female bracts, axillary. Male inflorescences to 8.5cm long, filiform; peduncle to 7mm long, rachis clearly visible, both with indumentum similar to that on young branches; flowers glomerate; bracts to 0.3mm long, ovate, sparsely hairy with simple, short trichomes, and with some sessile glands at base. Solitary female bracts sessile, enlarging in fruit to 3.5× 5mm, subreniform, appressed puberulent, with sparse resinous drops, margin crenate to subentire; sterile bracts sometimes present, solitary, similar to the female bracts but longer, to 1× 1.2cm, suborbicular; bracteoles absent. Male flowers subsessile; buds to 0.5mm diameter, puberulent. Female flowers 1 per bract, sessile; sepals 3, to 0.3mm long, triangular, puberulent; ovary c.1mm diameter, 3-lobed, papillose, papillae acute, ending in hyaline trichome to 0.5mm long, surface hispidulous; styles to 8mm long, connate at base, sparsely hairy, each divided into 5-6 slender segments. Allomorphic flowers not seen. Capsules to 3mm diameter, papillose-hispid, papillae acute, ending in hyaline trichome to 0.8mm long, surface puberulent. Seeds too young to describe. 24. Acalypha lamiana (Leandri) I.Montero& Cardiel PhytoKeys 108: 101 (Montero Muñoz etal. 2018a).— Acalypha reticulata var. urophylla f. lamiana Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 263 (Leandri 1942).— Type: Madagascar. Prov. Mahajanga, Réserve de Marohogo, 28.XII.1938, H. J. Lam& A. D. J. Meeuse 6127 (lecto-, designated by Montero Muñoz etal. [2018a: 102]: P[P05604417]; isolecto-, L[L0242109]).— Former syntypes: Madagascar. Prov. Antananarivo, Tsarasaotra, II.1898, H. Perrier de la Bâthie 457 (P [P05604408, P05604409, P05604410, P05604413]); Prov.Fianarantsoa, Ankiri-
449 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) hitra près du mont Tsiltondroina, III.1902, H. Perrier de la Bâthie 9817 (P[P05604403]), 9817bis (P[P05604404, P05604405]); Prov.Mahajanga, NW of Ankazobe, Vallée de l’Ikopa, 14.III.1930, R. Decary 7535 (P[P05604421]); Massif de l’Ankarafantsika, 11.I.1938, R. Decary 12876 (P[P05604420]); Bekodoka, 17.IX.1930, R. Decary 8109 (P[P05604422]); Région d’Antsalova, 1932-1933, J. Leandri 998 (P[P05604411]); Tsingy du Bemaraha, 3-6.X.1932, J. Leandri 176 (P[P05604415]); Dokolahy, II-IV.1933, J. Leandri 602 (P[P05604414]); Prov. Toliara, Soahazo Forest, 100m, 22.X.1932, J. Leandri 414 (P[P05604416]). iconography.— Leandri (1942: 263); Figs 41; 46D. eTymology.— The epithet honors Dutch botanist Herman Johannes Lam (1892-1977). DisTribuTion anD habiTaT.— Endemic to Madagascar (Boeny, Melaky, Bongolava, Menabe, Atsimo-Andrefana, and Anosy). Dry deciduous forest, secondary grasslands, and pastures (savanna). Frequently on Mesozoic limestone. Altitudinal range (25-) 100-800 (-1200) m (Fig. 42). p reliminary conservaTion assessmenT .— The EOO of Acalypha lamiana is estimated to be 376 831km 2 and its AOO 120km 2 , which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in the western dry deciduous forests (see comments in A.diminuta). It has been collected in some protected areas, but it is also known from several unprotected forests subject to various threats (uncontrolled fires for grazing and crops, logging, mining, etc.). Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha lamiana is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 50 collections. Madagascar. Afzelius, K. 81 (P[P00508497]); Baron, R. 6808 (K); Boivin, L.H.s.n. (K[K000186503]); Bolliger, R.F. RFB 083 (G[G00340310]); Capuron, R. 1935 (P[P04804747]), 1953 (P[P04779856, P04779858, P04779859]); Decary, R. 8109 (P[P05604422]), 8161 (K, P[P00324533]), 12876 (P[P05604420]), 15198 (P[P00508466]), 15725 (G, MA, P[P05547224]), 17069 (P[P05510110, P00324526]); Gillespie, L. 4141 (K, MO, P[P00513154], US[US01287449]), 10691 (MO), 10701 (CAN), 10702 (CAN), 10703 (CAN), 10832 (CAN); Harmelin 10247 (P[P04804764]); Humbert, H. 5102 (P[P04779863]), 12638 (P[P00508441, P00508442]), 12644 (P[P04804526, P04804527]), 19738 (P[P00508432]), 19990 (P[P00508431]), 20005 (P[P04779862]); Lam, H.J. 6127 (L[L0242109], P[P05604417]), bis 27 (P[P05604418]); Leandri, J. 176 (P[P05604415]), 414 (P[P05604416], US[US01287453]), 602 (P[P05604414]), 998 (P[P05604411]), 2060 (P[P04804746]), 2652 (P[P05543667, P00324505]), 2711 (G, K, P[P04804745]), 2856 (G, K, MO[MO-2965757], P[P04779857]), 2921 (P[P04779855]), 3018 (P[P04804744]), 4252 (P[P04804750]), s.n. (P[P05604419]); Nicoll, M.F. 377 (K, MO, P[P05604412]); Nusbaumer, L. LN 875 (K), LN 3084 (BR[BR0000015215604V], G[G00376211], MO[MO-2965725]); Perrier de la Bâthie, H. 457 (P[P05604408, P05604409, P05604410, P05604413]), 9817 (P[P05604403]), 9817 bis (P[P05604404, P05604405]); Rakotomalaza, P.J. 280 (P[P05516018]); Rakotovao, C. 4638 (P[P04804767]), 6237 (G, MO[MO-2965731], TAN); Ravelonarivo, D. 4528 (MO); Thunberg s.n. (S[S17-37572]). references.— Montero Muñoz etal. (2018a: 101). DescripTion Shrubs, probably deciduous, to 3m tall, monoecious. Branches laxly pubescent with curved, antrorse trichomes, glabrous when mature. Axillary buds ovoid, to 1× 0.5mm, perulate, perules 2, valvate, membranous, blackish, glabrous. Stipules to 6mm long, triangular, subglabrous, with some hyaline trichomes to 1mm long, margin papillate. Petioles filiform, 1-2.5(-4) cm long, indumentum similar to that on young branches. Leaf blades 5-7× 2-2.5(-3.5) cm, ovate-lanceolate to elliptic-lanceolate, membranous; base rounded to obtuse, rarely subcuneate; apex acuminate, acumen to 10mm long, acute; margin serrulate, reddish, teeth minute, sometimes with simple trichome at apex; upper surface subglabrous, with some simple, appressed trichomes at base of midrib; lower surface subglabrous, with some sparse, simple trichomes; margins sparsely ciliate; venation actinodromous, basal veins 3, secondary veins 4-5 per side. Stipels glandular, to 0.5mm diameter, glabrous. Inflorescences spiciform, androgynous, and solitary female bracts, axillary. Androgynous inflorescences to 4cm long, mostly male with short female segment; sessile, rachis filiform, subglabrous, with some curved trichomes at base. Female segment: bracts 1-(4), sessile, enlarging in fruit to 7× 6mm, subrounded, glabrous; margin slightly crenatedentate, revolute, reddish, teeth rounded, central tooth some45°E 15°S 20° 25° 300 km 50° Acalypha isaloensis fig. 40 . — Distribution map of Acalypha isaloensis I.Montero & Cardiel in Madagascar.
456 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. 29. Acalypha leptomyura Baill. Histoire physique, naturelle et politique de Madagascar Atlas, 2, Fasc. 27: t. 191 (Baillon 1891).— Type: Madagascar. Baillon 1891: pl. 191, holotype. iconography.— Baillon (1891: t. 191); Figs 54C, D; 57A. eTymology.— The epithet is taken from the Greek ‘leptos’, fine or delicate, ‘myos’, mouse, and ‘oura’, tail. It possibly refers to the thin rachis of the inflorescences. DisTribuTion anD habiTaT.— Endemic to Madagascar (Diana, Sofia, Betsiboka, Alaotra-Mangoro, Analamanga, Bongolava, Melaky, Menabe, Atsimo-Andrefana, Androy, Anosy, Atsimo-Atsinanana, and Ihorombe). Lowland evergreen moist forest, dry deciduous forest, and sclerophyllous woodland. On sandstone, unconsolidated sand, and Mesozoic limestone. Altitudinal range (15-) 100-1000 (-1400) m (Fig. 48). p reliminary conservaTion assessmenT .— The EOO of Acalypha leptomyura is estimated to be 385 705km2 and its AOO 120km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows in a variety of forests: dry deciduous forests, (see assessment of A.diminuta), lowland evergreen moist forests (see assessment of A.emirnensis), and sclerophyllous woodlands, which mostly have been converted to secondary grasslands by human activities such as fires for agriculture or cattle grazing (Kull 2003; Gautier etal. 2018). This species has been collected in some protected areas, but it is also known from several unprotected forests subject to various threats. Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha leptomyura is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. m aTerial examineD .— 42 collections. Madagascar. Andriamihajarivo, T. 1108 (MO[MO-3025023], P[P01104326], TAN); Baron, R. 4576 (P[P04779799]), 4685 (P[P05547007, P05547009]); Bosser, M.J. 9078 (P[P04779786]), 9089 (P[P04779785]), 10256 (P[P00324462]); Capuron, R. 2020 (P[P05547053, P05547054]); d’Alleizette, Ch. s.n. (L[L0241867]); Decary, R. 1128 (P[P05547011]), 7312 (C, P[P00513075]), 7535 (G, P[P05604421], S), 15818 (P[P05510105, P00324523]), 15823 (P[P05547221]), 17043 (P[P05547217]), 18845 (P[P05547220]); Dequaire, J. 27125 (P[P05547219]); Grevé, H. 109 (K, P[P00513081, P00513082, P00513083]); Humbert, H. 3533 (G, K, P[P00513076, P00513077, P00513078]), 4089 (P[P00513079, P00513080]), 7089 (P[P05547223]), 13829 (P[P00513074]), 18819 (P[P00513067, P00513068]), 19671 (K, P[P05547222]); Labat, J.-N. 2320 (P[P023912], WAG[WAG.1578522]); Leandri, J. 150 (P[P05547050]), 154 (P[P05547051]); 167 (P[P00324570]), 600 (P[P00513065, P00513066]), 930 (P[P05547052]), 2121 (G, P[P05547225]); Morat, P. 4714 (P[P05547008, P05547010]), 4736 (P[P05546999, P05547000, P05547001, P05547002, P05547003]); Nicoll, M.F. 162 (K, MO, P[P00513064]); Perrier de la Bâthie, H. 9812 (P[P00513069, P00513070, P00513071, P00513072]); Rakotonao 11308 (P[P05547005]); Randrianarivony, T. 749 (P[P01084145]); Randrianasolo, A. 6866 (P[P05547006]); Rasamison 12296 (P[P05547049]), 12314 (P[P05547218]); Service Forestier Madagascar 54 (G, P[P00513073]), 166-SF (P[P00513163]); Waterlot, M. 624 (P[P05547004]). references.— Baillon (1892: 1004); Koehne (1892: 131); Palacký (1907: 25); Pax& Hoffmann (1924: 112); Leandri (1942: 271); Govaerts etal. (2000: 72); Montero Muñoz etal. (2018a: 103). DescripTion Shrubs, deciduous, to 1.5m tall, divaricately branched, monoecious. Branches pubescent with simple, curved, antrorse trichomes, glabrescent when mature. Axillary buds spherical, to 0.5mm diameter, perulate, perules 2, imbricate, chartaceous, brownish, ciliate. Stipules to 1.5mm long, linear-lanceolate, glabrous except margin with some sessile glands and apex with some hyaline trichomes to 1mm long. Petioles filiform, 0.3-0.7(-1.5) cm long, indumentum similar to that on young branches, base also with hyaline trichomes to 1mm long. Leaf blades (1.5-)2.5-3.5(-4.5)×1-2(-2.5) cm, ovate-lanceolate to subtriangular, membranous; base obtuse to truncate; apex subacuminate, acumen to 8mm long, rounded to subacute; margin crenate to serrate, teeth rounded to subacute, sometimes with gland at apex; upper surface laxly pubescent with sparse, simple, long, erect trichomes, and with simple, short, curved trichomes on veins; lower surface indumentum similar to that on upper surface but denser; venation actinodromous, basal veins 3, secondary veins 4-5 per side. Stipels slightly fleshy, linear-lanceolate, to 0.5mm long, glabrous. Inflorescences spiciform, androgynous, and solitary female bracts, axillary. Androgynous inflorescences to 4cm long, mostly male with 1 female bract; sessile; rachis filiform, pubescent. Female seg - ment: bract 1, sessile, enlarging in fruit to 5mm diameter, 45°E 15°S 20° 25° 300 km 50° Acalypha lepidopagensis fig. 47 . — Distribution map of Acalypha lepidopagensis Leandri in Madagascar.
457 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) suborbicular, subglabrous, with some sparse, simple, erect trichomes; margin entire to crenate, teeth 5-7, rounded, central tooth sometimes prominent; sterile female bracts sometimes present, similar to regular female bracts, to 1mm diameter; bracteoles absent. Male segment persistent, to 3.5cm long; flowers glomerate; bracts to 0.3mm long, triangular-lanceolate, sparsely hairy with sparse trichomes, margin with some sessile glands. Solitary female bracts sessile, similar to those of androgynous inflorescence. Male flowers: pedicel to 0.3 mm long, sparsely hairy; buds to 0.5 mm diameter, subglabrous, papillose. Female flowers 1 per bract, sessile; sepals 3, to 0.5mm long, triangular, ciliate; ovary c.0.6 mm diameter, 3-lobed, papillose-hispid, papillae minute, rounded, ending in simple trichome to 1mm long, surface pubescent with simple, short trichomes; styles 3, to 3mm long, distinct, subglabrous, with some long, hyaline trichomes, each divided into c.10 segments. Allomorphic flowers not seen. Capsules to 2 mm diameter, papillose-hispid, papillae rounded, ending in hyaline trichome to 1mm long., surface pubescent with simple, short trichomes. Seeds c.1.5× 1.2mm, pyriform, minutely foveolate. 30. Acalypha levinii I.Montero& Cardiel South African Journal of Botany 146: 638 (Montero Muñoz etal. 2022).— Type: Madagascar. Haute Matsiatra region (Fianarantsoa prov.), Parc d’Ánja Community Reserve, E of RN7, c.9.5km of Ambalavao, 990m, 21°51’7.5”S, 46°50’43.1”E, 29.XI.2012, L. J. Gillespie, G. A. Levin& J. Razanatsoa 19809 (holo-, CAN).— Paratypes: Madagascar. Haute Matsiatra region (Fianarantsoa prov.), Haute Matsiatra, Parc d’Ánja Community Reserve, E of RN7, c.9.5km of Ambalavao, 990m, 21°51’6.1”S, 46°50’43.7”E, 29.XI.2012, L. J. Gillespie, G. A. Levin& J. Razanatsoa 19808 (CAN); Atsimo-Andrefana region (Toliara prov.), Beza Mahafaly Reserve, near Betioky, parcelle 1, Along the Sakamena river bank, 140m, 23°38’60”S, 44°32’60”E, 2.XI.1987, P.B. Phillipson 2503 (K, MO, US, WAG); Plateau calcaire au N de la basse vallée du Fiherenana, 200-300m, I.1947, H. Humbert 19866 (P[P00324585]). iconography.— Montero Muñoz etal. (2022); Fig. 57B. eTymology.— The epithet honors Dr. Geoffrey A. Levin, codirector of the first author’s PhD dissertation. He has published many articles about Acalypha and contributed to the knowledge of this genus worldwide. DisTribuTion anD habiTaT.— Endemic to Madagascar (AtsimoAndrefana and Haute Matsiatra). Degraded sclerophyllous woodland. On basement rocks and sandstone. Altitudinal range 140-990m (Fig. 49). preliminary conservaTion assessmenT.— Acalypha levinii is known from four collections from different localities. Its EOO is estimated to be 12 578km 2 and its AOO 16km 2 , which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows in sclerophyllous woodlands in the highlands (see assessment of A.leptomyura). This species was collected in degraded forests and in both protected (community reserve and national reserve) and unprotected areas. Even in the reserves, the habitat of this species is subject to various threats (uncontrolled fires for grazing and crops), so ongoing habitat loss and degradation will cause continued decline of its EOO and AOO. Acalypha levinii is assessed as Endangered (EN): EN B2ab(ii,iii,iv). m aTerial examineD .— 4 collections. Madagascar: Gillespie, L. 10808 (CAN), 10809 (CAN); Humbert, H. 19866 (P[P00324585]); Phillipson, P.B. 2503 (K, MO, US[US01287331], WAG[WAG.1578523]). DescripTion Shrubs, deciduous, to 2m tall, divaricately branched, monoecious. Branches red-tinged, densely pubescent with simple, curved, antrorse trichomes, glabrescent when mature. Axillary buds spherical, to 1mm diameter, perulate, perules 2, overlapping (superposed), membranous, brownish, sparsely hairy. Stipules to 4mm long, lanceolate, apex acute, pubescent with simple, appressed trichomes. Petioles 1.5-3(-4) cm long, indumentum similar to that on young branches. Leaf blades (3.5-)4-6.5× 2-3.3cm, ovate-lanceolate to elliptic-lanceolate, membranous; base rounded to subcordate; apex slightly acuminate, acumen 10mm long, rounded to subacute, mucronate; margin serrate to crenate, teeth subrounded, slightly reddish, callose-edged; upper surface laxly pubescent with simple, erect trichomes and subsessile reddish glandular trichomes; lower surface indumentum similar to that on upper surface but denser, axils of secondary veins with hair-tuft domatia; venation actino - dromous, basal veins 3, secondary veins 5-6 per side. Stipels 45°E 15°S 20° 25° 300 km 50° Acalypha leptomyura fig. 48. — Distribution map of Acalypha leptomyura Baill. in Madagascar.
458 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. filiform, to 0.3mm long, glabrous. Inflorescences spiciform, androgynous, and solitary female bracts, mainly axillary, some androgynous inflorescences terminal. Androgynous inflorescences to 4cm long, mostly male with short female segment; subsessile, peduncle to 0.3mm long; rachis indumentum similar to that on petiole. Female segment to 2.5cm long; bracts 1-4, sessile to slightly petiolate, enlarging in fruit to 10× 12mm, oblate to suborbicular, indumentum similar to that on leaves and ciliate, glabrescent; margin dentate, teeth c.15, rounded, callose-edged, slightly reddish, central tooth not prominent; bracteoles absent. Male segment persistent, to 2cm long; flowers glomerate; bracts to 0.5mm long, lanceolate, sparsely hairy. Solitary female bracts sessile, similar to those of androgynous inflorescences. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds to 0.8 mm diameter, sparsely hairy, papillose. Female flowers 1 per bract, sessile; sepals 3, to 0.5mm long, triangular, sparsely hairy; ovary c.1 mm diameter, 3-lobed, apparently smooth, surface hispid with simple, erect trichomes to 1mm long; styles 3, to 4mm long, slightly connate at base, sparsely hairy, each divided into c.12 segments. Allomorphic flowers sometimes present at inflorescence apex; pedicel filiform, to 12mm long, sparsely hairy; sepals 3, similar to those of normal flowers; ovary 1-lobed, to 1mm diameter, hispid with simple, erect trichomes to 1mm long; style 1, to 6mm long, sparsely hairy. Capsules to 2.5 mm diameter, papillose-hispid, papillae rounded, surface densely pubescent with simple trichomes. Seeds c.1.8× 1.5mm, subglobose, foveolate. 31. Acalypha linearifolia Leandri Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 275 (Leandri 1942).— Type: Madagascar. Prov. Toliara, Ambovombe, Kotoala, 21.I.1931, R. Decary 8423 (lecto-, designated by Montero Muñoz etal. [2018a: 103]: P[P00513090]; isolecto-, S[S07-14664], TAN[TAN000510], US[US01014148]).— Former syntype: Madagascar. Prov. Toliara, delta de la Linta, 17-24.VIII.1928, H. Humbert& C. F. Swingle 5385 (P[P00513086, P00513087, P00513088, P00513089], US[US00096361]). iconography.— Leandri (1942: 276); Fig. 57C. eTymology.— The specific epithet refers to the linear leaf blades characteristic of this species. DisTribuTion anD habiTaT.— Endemic to Madagascar (AtsimoAndrefana, Androy, and Atsinanana [one known collection]). Littoral forest and littoral thicket. On unconsolidated sand and Tertiary limestone. Altitudinal range sea level to 10m (Fig. 50). p reliminary conservaTion assessmenT .— Acalypha linearifolia is known from five collections. Its EOO is estimated to be 84 834km 2 and its AOO 16km2. The large EOO is due to one collection presumably made in Toamasina by G. Cours. The other collections are from southern Madagascar. This species grows mainly in littoral forests (see assessment of A.gracilipes). There are no collections made in protected areas. In addition, no specimens of this species have been collected for 70 years, so we cannot rule out that this species has become extinct. Due to habitat loss and the absence of recent collections, A.linearifolia is assigned a preliminary conservation status of Endangered: EN B2ab(ii,iii,iv); although it meets the values needed for a threatened category based on the B2 subcriterion, this species could probably be assessed as extinct (EX). m aTerial examineD .— 4 collections. Madagascar: Cours, G. 4637 (P[P00324458, P00513085]), 4638 (G, P[P00513084]); Decary, R. 8423 (P[P00513090], S[S07-14664], TAN[TAN000510], US[US01014148]); Humbert, H. 5385 (P[P00513086, P00513087, P00513088, P00513089], US[US00096361]). references.— Govaerts etal. (2000: 72); Montero Muñoz etal. (2018a: 103). DescripTion Shrubs, probably evergreen, to 1.5m tall, monoecious. Branches densely pubescent with stellate trichomes, glabrescent when mature. Axillary buds ovoid, to 1× 0.7mm, perulate, perules 2, valvate, membranous, brownish, pubescent with stellate trichomes; Stipules to 1mm long, lanceolate, pubescent with stellate trichomes, margin with some sessile glands. Petioles to 0.2cm long, indumentum similar to that on young branches. Leaf blades ericoid, 3.5-4.7× 0.1-0.2cm, linear, coriaceous; base acute; apex obtuse to rounded; margin entire, revolute; upper and lower surfaces densely pubescent with stellate trichomes; venation prominent on lower surface, pinnate, secondary veins to 15 per side. Stipels absent. Inflorescences spiciform, male, and solitary female bracts, 45°E 15°S 20° 25° 300 km 50° Acalypha levinii fig. 49 . — Distribution map of Acalypha levinii I.Montero & Cardiel in Madagascar.
459 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) axillary. Male inflorescences to 1.4cm long, with glossy exudate; subsessile, peduncle to 2mm long, pubescent with stellate trichomes; flowers glomerate; bracts orbicular, to 0.5mm diameter, pubescent with stellate trichomes. Solitary female bracts subsessile, peduncle to 2mm long, pubescent with stellate trichomes; bract short-petiolate, enlarging in fruit to 4× 5mm, suborbicular, densely pubescent with stellate trichomes; margin entire to denticulate, central tooth not prominent; bracteoles absent. Male flowers: pedicel to 0.5 mm long, glabrous; buds to 0.5mm diameter, pubescent with stellate trichomes. Female flowers 1 per bract, sessile; sepals 3, to 1mm long, ovate-lanceolate, pubescent with stellate trichomes; ovary c.1 mm diameter, 3-lobed, papillose, surface pubescent with stellate trichomes; styles not seen. Allomorphic flowers not seen. Capsules to 3 mm diameter, papillose, surface pubescent with stellate trichomes. Seeds c.2 × 1.8 mm, subglobose, minutely foveolate. 32. Acalypha magistri I.Montero& Cardiel South African Journal of Botany 146: 640 (Montero Muñoz etal. 2022).— Type: Madagascar. Sava region (Antsiranana prov.), au nord d’Andapa, dans la Réserve naturelle intégrale de Marojejy, aux environs du sommet d’Ambatosoratra, 1583m, 14°32’S, 49°42’E, 17-24.VI.1994, D. Ravelonarivo 273 (holo-, MO; iso-, ILLS, MAUAM, P, TAN). iconography.— Montero Muñoz etal. (2022); Fig. 57D. eTymology.— The epithet honors teachers at the Alonso de Ercilla High School (Ocaña, Toledo, Spain), where the first author studied. The epithet magistri is a nominative plural noun. DisTribuTion anD habiTaT.— Endemic to Madagascar (Sava). Medium altitude evergreen moist forest. On basement rocks, 1583m altitude (Fig. 51). preliminary conservaTion assessmenT.— Acalypha magistri is known only from one collection from Marojejy. Its EOO could not be calculated; its AOO is estimated to be 8km 2 . Marojejy has been a protected area since 1952, and a National Park since 1998 (Category II; Dudley 2008). The main threats that affect the Marojejy forests are illegal logging of precious wood, uncontrolled fires, and mining for topaz and beryl, mainly at the boundaries of the National Park. However, at the high elevations where this species occurs (it was collected near the top of Ambatosoratra peak), habitat disturbance from ecotourism likely is the only threat (Patel 2007; Almeda& Ranarivelo 2019). In view of its limited AOO, Acalypha magistri is assigned a preliminary conservation status of Vulnerable: VU D2. maTerial examineD.— 1 collection. Madagascar: Ravelonarivo, D. 273 (MAUAM, MO). DescripTion Small trees, leaf persistence unknown, to 6m tall, monoecious. Branches pubescent with simple, short, erect trichomes, glabrescent when mature. Axillary buds ovoid, to 1.5× 0.7mm, perulate, perules 2, imbricate, chartaceous, dark brown, glabrous. Stipules to 2mm long, triangular-lanceolate, apex acute, base with some sparse, simple trichomes, margin ciliate with simple, short trichomes. Petioles 1.2-3.5cm long, indumentum similar to that on young branches. Leaf blades 7.5-12× 2-5.5cm, elliptic-lanceolate to ovate-lanceolate, subchartaceous; base rounded; apex acuminate, acumen to 13mm long, acute; margin serrate, teeth acute; upper surface subglabrous, with simple, short, erect, simple trichomes, especially on veins; lower surface indumentum similar to that on upper surface but denser; margin ciliate; venation pinnate, secondary veins 6-8 per side. Stipels absent. Inflorescences spiciform, androgynous, axillary, to 9.5cm long, mostly male with a female segment; peduncle to 22mm long, it and rachis pubescent with simple, curved, antrorse trichomes. Female segment to 3.7cm long; bracts 4, sessile, enlarging in fruit to 5mm diameter, orbicular, pubescent with simple, erect trichomes to 0.5mm long; margin subentire to crenate, central tooth not prominent; bracteoles absent. Male segment persistent, to 3.8cm long; flowers glomerate; bracts to 0.7mm long, triangular, sparsely hairy. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds not seen. Female flowers 1 per bract, sessile; sepals 3, to 1.5mm long, triangular-lanceolate, sparsely hairy; ovary not seen; styles 3, to 3mm long, distinct, sparsely hairy. Allomorphic flowers not seen. Capsules to 5 mm diameter, echinate, projections conical, to 2mm long, surface pubescent with simple, short, erect trichomes. Seeds c.4× 3mm, pyriform, minutely foveolate. 45°E 15°S 20° 25° 300 km 50° Acalypha linearifolia fig. 50. — Distribution map of Acalypha linearifolia Leandri in Madagascar.
460 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. 33. Acalypha marginata (Poir.) Spreng. Systema vegetabilium 3: 879 (Sprengel 1826).— Tragia marginata Poir., Encyclopédie méthodique. Botanique 7: 725 (Poiret 1806).— Acalypha integrifolia subsp. marginata (Poir.) Coode, Kew Bulletin 34: 42 (Coode 1979).— Type: Mauritius. “Les Indes Orientales”, s.d. Anonymous (probably P. Commerson) s.n. (holo-, P-LAM[P00382145]). Acalypha integrifolia subsp. panduriformis Coode, Kew Bulletin 34: 42 (Coode 1979).— Type: La Réunion. Cliff between St Philippe and St Joseph near Basse vallée, c.100m, 26.II.1975, M. J. E. Coode& T. H. Cadet 4968 (holo-, K[K000431108]; iso-, K[K000431107]). Acalypha integrifolia subsp. marginata var. saltuum Coode, Kew Bulletin 34: 43 (Coode 1979).— Type: Mauritius. Macabé, 650m, 15.II.1975, M. J. E. Coode etal. 4874 (holo-, K[K000431106]; iso-, P[P04779351], MAU[n.v.]). iconography.— Fig. 54E. eTymology.— The epithet probably refers to the discolorous margin of the leaf blades. DisTribuTion anD habiTaT.— Endemic to Mascarene Islands (Mauritius and La Réunion). Lowland evergreen moist forest and secondary forest. Altitudinal range in Mauritius 420-650m, in La Réunion 100-1200m (Fig. 52). preliminary conservaTion assessmenT.— Acalypha marginata is known from Mauritius and La Réunion. Its EOO is estimated to be 8 916km2 and its AOO 44km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. For the status of forests on Mauritius, see the assessment of A.crateriana comb.nov.; for those on La Réunion, see the assessment of A.filiformis. Ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha marginata is assessed as Near Threatened (NT) under criterion B. It meets the EOO and AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 38 collections. Mauritius. Anonymous 490 (P[P04779328]); Anonymous s.n. (G[G00324533]); Anonymous s.n. (G[G00324534]); Anonymous s.n. (G[G00324537]); Anonymous s.n. (P[P04779363] pr.p.); Ayres, P.B.s.n. (P[P04779841]); Bouton, L. 23 (P[P04779354]), s.n. (G[G00324535]); Commerson, P.s.n. (P[P00382145]), s.n. (P[P04779358]), s.n. (P[P04779360]); Coode, M.J.E. 4370 (K, P[P04779362]), 4778 (K, MO[MO-2452040]), 4867 (K, P[P04779348]), 4868 (K, P[P04779350]), 4869 (K), 4870 (K), 4874 (K[K000431106], P[P04779351]), 4876 (K); Lorence, D. 1954 (K, MO[MO-2452046], P[P04779347]). La Réunion. Billiet, F. 585 (BR[BR0000014628085], K, WAG[WAG.1578518]); Boivin, L.H.s.n. (P[P04779331]), s.n. (P[P04779352]); Bosser, M.J. 21043 (P[P04779330]); Brown, Ab. s.n. (P[P04779375]); Cadet, Th. 4925 (K); Commerson, P.s.n. (P[P04779356]), s.n. (P[P04779357] pr.p.), s.n. (P[P04779359]); Coode, M.J.E. 4967 (K), 4968 (K[K000431107, K000431108]), 5006 (K); d’Alleizette, Ch. s.n. (L[L0241920]); Friedmann, F. 847 (P[P04779333]), 919 (K, P[P04779335, P04779353]); Léman, D.S.s.n. (P[P04779332]), s.n. (P[P04779355]); Richard, M.s.n. (P[P04779361]). references.— Bojer (1837: 286); Baillon (1858: 443); Baillon (1861: 267); Baker (1877: 315); Cordemoy (1895: 343); Palacký (1907: 25); Pax& Hoffmann (1924: 106); Coode (1982: 73) as A.integrifolia subsp. panduriformis, A.integrifolia subsp. marginata; Coode (1982: 74) as A.integrifolia var. saltuum; Govaerts etal. (2000: 69) as A.integrifolia subsp. marginata; Seebaluck etal. (2015: 152); Montero Muñoz etal. (2018a: 104). DescripTion Shrubs, leaf persistence unknown, to 3m tall, monoecious. Branches velutinous, glabrescent when mature. Axillary buds naked, hispid. Stipules to 10mm long, linear-lanceolate, densely pubescent with simple, appressed trichomes, margin papillate. Petioles thick, 0.5-3cm long, indumentum similar to that on young branches, glabrescent. Leaf blades (4-)6-17-(21)× (2- )2.5-7(-8) cm, usually elliptic to elliptic-lanceolate, sometimes obovate, chartaceous; base rounded to cuneate; apex acute to subacuminate; margin crenate-serrate to serrate, discolorous, teeth rounded to acute, slightly callose-edged; upper and lower surfaces pubescent with simple, short, appressed trichomes, glabrescent; venation pinnate, secondary veins 13-14 per side. Stipels absent. Inflorescences spiciform (male) or glomerulate (female), unisexual, axillary. Male inflorescences to 11cm long; peduncle to 2mm long, pubescent with simple, short, erect trichomes; flowers glomerate; bracts to 1mm long, triangular, densely appressed-pubescent. Female inflorescences 0.2-0.4cm long; sessile;; bracts 3-4, sessile, not enlarging in fruit, to 0.5mm long, triangular, appressed-pubescent; margin entire; bracteoles absent. Male flowers: pedicel to 0.6 mm long, sparsely hairy; buds 45°E 15°S 20° 25° 300 km 50° Acalypha magistri fig. 51. — Distribution map of Acalypha magistri I.Montero & Cardiel in Madagascar.
461 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) to 0.8mm diameter, sparsely hairy. Female flowers 1 per bract, sessile; sepals 3, to 1.2mm long, elliptic, appressed-pubescent; ovary c.1 mm diameter, 3-lobed, papillose-hispid, papillae acute, ending in a simple trichome to 1mm long, surface pubescent with simple, short trichomes; styles 3, to 2.5mm long, distinct, rachis thick, sparsely hairy with simple, short, appressed trichomes, each divided into c.10 segments. Allomorphic flowers not seen. Capsules to 3.5 mm diameter, papillose, papillae to 0.5mm long, ending in a simple trichome to 1mm long, surface sparsely hairy with simple, erect trichomes to 0.5mm long. Seeds c.3× 2.2 mm, pyriform, minutely foveolate. noTe Acalypha marginata was treated by Coode (1979) as A.integrifolia subsp. marginata. We consider A.marginata to be a distinct species differentiated mainly by its variegated leaf blades and minute female bracts that are not accrescent in fruit, vs uniformly colored leaf blades and conspicuous female bracts that are accrescent in fruit in A.integrifolia. We include as synonyms A.integrifolia subsp. panduriformis and Acalypha integrifolia subsp. marginata var. saltuum because the morphology of these taxa overlap too much with those of A.marginata to accept them as distinct. 34. Acalypha mayottensis I.Montero& Cardiel PhytoKeys 140: 65 (Montero Muñoz etal. 2020b).— Type: Mayotte. Mamoudzou commune, îlot M’bouzi, 12°48’50”S, 45°14’08”E, 10-50m, 22.XI.2000, J.-N. Labat, F. Barthelat, C.M. Hladik& A.B. Sifary 3268 (holo-, G[G00034240]; iso-, K, MAO, MO[MO2965774], P[P00209719]).— Paratypes: Mayotte. Mamoudzou commune, îlot M’Bouzi, 12°48’57”S, 45°14’06”E, 90m, 22.XI.2000, J.-N. Labat, F. Barthelat, C.M. Hladik& A.B. Sifary 3272 (G[G00034255], K, MAO, MO[MO-2966248], P[P00209724, P00209725]); îlot M’Bouzi, 12°48’39”S, 45°14’06”E, 26.XII.2002, F. Barthelat, A.de Vanssay& G. Rembert 1112 (MAO, P[P00339165]).— Precise location unknown, probably from M’Bouzi islet, 01.I.2010, G. Viscardi 310 (HKM, P[P02439826]). iconography.— Montero Muñoz etal. (2020b); Figs 54F; 57E. e Tymology .— The epithet refers to Mayotte island, to which the small Mbouzi islet belongs, and to which this species appears to be endemic. DisTribuTion anD habiTaT.— Endemic to Comoros Archipelago (Mayotte). Dry forest and secondary forest. Altitudinal range 1090m (Fig. 53). p reliminary conservaTion assessmenT .— Acalypha mayottensis is only known from Mbouzi islet. Its EOO and AOO are estimated to be 8km 2 . Mbouzi islet was declared a “Réserve Naturelle Nationale” in 2007, a category IV protected area (Dudley 2008). In the 1990s the islet had lost 70% of its original forests to agriculture. Mbouzi now conserves 10% of its natural and secondary forest. Currently, the most serious threat are invasive species, both animals, such as Eulemur fulvus, and plants, such as Antigonon leptopus, Lantana strigocamara, Leucaena leucocephala, Litsea glutinosa, Spathodea campanulata and Furcraea foetida (Boullet& Traclet 2018; Quintard etal. 2019). Due to its restricted geographic range and the threats from alien species, A.mayottensis is assigned a preliminary conservation status of Critically Endangered: CR B2ab(ii,iii). maTerial examineD.— 4 collections. Mayotte. Barthelat, F. 1112 (MAO, P[P00339165]); Labat, J.-N. 3268 (G[G00034240], K, MAO, MO[MO-2965774], P[P00209719]), 3272 (G[G00034255], K, MAO, MO[MO-2966248], P[P00209724, P00209725]); Viscardi, G. 310 (P[P02439826]). DescripTion Shrubs, deciduous, to 5m tall, monoecious. Branches laxly pubescent with simple, erect trichomes to 1mm long, glabrous when mature. Axillary buds ovoid, to 3× 2.3mm, perulate, perules 2, imbricate, chartaceous, blackish, glabrous. Stipules to 6mm long, linear to triangular-lanceolate, becoming filiform when mature, margins scarious, sparsely hairy, glabrescent, and with some glands. Petioles slender, (2-)3-5(-6) cm long, pubescent with simple, curved, antrorse trichomes. Leaf blades 5-10× 3-6cm, ovate-lanceolate to elliptic-lanceolate, membranous; base rounded to subcordate; apex subacuminate to acuminate, acumen to 15mm long, rounded; margin crenulate-serrulate to subentire, slightly revolute, teeth rounded; upper surface pubescent with simple, thin, erect trichomes, glabrescent; lower surface indumentum similar to that on upper surface but denser; axils of secondary veins with minute, sparsely hairy, pocket-shaped domatia, sometimes with hair-tuft domatia; margins ciliate in sinuses; venation actinodromous, basal veins 3, secondary veins 4-6 per side. Stipels absent. Inflorescences spiciform androgynous, axillary, to 6cm long, mostly male with short female segment; peduncle thick, to 15mm long, laxly pubescent with trichomes similar to those on young branches, glabrescent. Female segment to 2.5cm long; bracts 1-2, sessile, enlarging in fruit to 19× 21mm, subreniform, veins and margin sparsely hairy with simple, erect trichomes to 1.5mm long, glabrescent; margin crenate to subentire, sometimes dentate in young bracts, central tooth 75 km Mauritius La Réunion Acalypha marginata fig. 52. — Distribution map of Acalypha marginata (Poir.) Spreng. in the Mascarene Islands.
462 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. not prominent; bracteoles absent. Male segment persistent, to 4cm long; flowers glomerate; bracts to 0.5mm long, triangular, sparsely hairy. Male flowers: pedicel to 1mm long, glabrous; buds to 0.7mm diameter, sparsely hairy with simple, arachnoid trichomes. Female flowers 1 per bract, sessile; sepals 3[4], to 0.7mm long, ovate-triangular, sparsely hairy with simple, arachnoid trichomes; ovary c.1mm diameter, 3-lobed, echinate, projections and surface hispid; styles 3, to 3mm long, distinct, rachis thick, appressed-pubescent, each divided into 8-10 segments. Allomorphic flowers not seen. Capsules to 4mm diameter, erinaceous projections conical, subacute, to 1mm long, they and surface hispid with simple, erect trichomes to 1mm long. Seeds 2.5× 2mm, pyriform, minutely foveolate. 35. Acalypha medibracteata Radcl.-Sm.& Govaerts Kew Bulletin 52 (2): 477 (Radcliffe-Smith& Govaerts 1997).— Acalypha gagnepainii Leandri Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 274 (Leandri 1942) nom. illeg., non A.gagnepainii Merr. (Merrill 1938).— Type: Madagascar. Prov. Toliara, Massif du Vohitsiombe (Fort-Dauphin), 31.VII.1926, R. Decary 4664 (lecto-, designated by Montero Muñoz etal. [2018a: 104]: P[P00508417]; isolecto-, S[S07-14667]).— Former syntypes: Madagascar. Prov. Toliara, Vallée du Mandrare, s.d., R. Decary 2620 (P[P00508418]); Vallée de l’Ikonda, au N d’Ambovombe, R. Decary 8913 (P[P00887487]); Imangory, s.d., R. Decary 8948 (P[P00508416]); Beteny (limite Nord-Est de l’Androy), 22.XI.1931, R. Decary 9355 (P[P00508415], G[G00034242], GB[GB0047682]). Acalypha medibracteata var. calcicola (Leandri) Radcl.-Sm.& Govaerts, Kew Bulletin 52 (2): 477 (Radcliffe-Smith& Govaerts 1997).— Acalypha gagnepainii var. calcicola Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 275 (Leandri 1942).— Type: Madagascar. Prov. Toliara, vallée moyenne du Mandrare près d’Anadabolava, X.1933, H. Humbert 12422 (lecto-, designated by Montero Muñoz etal. [2018a: 104]: P[P00508409]; isolecto-, P[P00508405, P00508406, P00508407, P00508408]).— Former syntypes: Madagascar. Prov. Toliara, Fort-Dauphin, J. Cloisel 18 (P[P00508410, P00508411]); Basse vallée du Fiherenana, s.d., H. Humbert 11573 (P[P00508412, P00508413]). iconography.— Leandri (1942: 269); Figs 54 G, H; 57F. eTymology.— The epithet refers to the position of the female bracts near the middle of the bisexual inflorescences (RadcliffeSmith& Govaerts 1997). D isTribuTion anD habiTaT .— Endemic to Madagascar (Menabe, Atsimo-Andrefana, Androy, and Anosy). Dry deciduous forest, dry spiny thickets, and sometimes in riparian forest. On basement rocks, Tertiary limestone, sandstone, and unconsolidated sand. Altitudinal range 100-800m (Fig. 55). p reliminary conservaTion assessmenT .— The EOO of Acalypha medibracteata is estimated to be 85 039km2 and its AOO 92km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows mainly in dry deciduous forests and spiny thickets (see assessment of A.decaryana). The remaining original vegetation is still decreasing (Moat& Smith 2007, Gautier etal. 2018). A.medibracteata has been collected in some protected areas, but it is also known from several unprotected forests subject to various threats (uncontrolled fires for grazing and crops, logging, mining, etc.). We predict that ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha medibracteata is assessed as Near Threatened (NT) because its AOO meets the threshold needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. m aTerial examineD .— 41 collections. Madagascar. Afzelius, K.s.n. (P[P05547249]); Baron, R. 646 (K, P[P04779782]); Bernardi, L. 11576 (BR[BR0000021450181], G, P[P04779802]); Bosser, M.J. 3700 (P[P04779804]), 3726 (P[P04779807]), 3756 (P[P04779806]), 13871 (P[P04779812, P04779815]), 13950 (P[P04779817, P04779818]); Cloisel, J. 18 (P[P00508410, P00508411]); Decary, R. 2620 (P[P00508418]), 4664 (P[P00508417], S[S07-14667]), 8913 (L[L0241358], P[P00887487]), 8948 (NY, P[P00508416]), 9355 (G[G00034242], GB[GB-0047682], P[P00508415]), 15956 (P[P05547255]), 15963 (P[P05547041]), 15970 (P[P05547232]), 16029 (P[P00324525, P05510102]), 16103 (P[P05547231]), 16123 (P[P05547233]), s.n. (L[L0241301]); Gillespie, L. 10828 (MO); Goudot, J. 192 (G[G00420838]); Homolle, A.M. 1503 (P[P05547178]), 1504 (P[P05547226]); Humbert, H. 11573 (P[P00508412, P00508413]), 12422 (P[P00508405, P00508406, P00508407, P00508408, P00508409]), 19885 (P[P05547275, P05547276]), 29015 (P[P05547048]), 29467 (P[P04779860]); Lam, H.J. 5455 (L[L0241359], P[P00508414], WAG[WAG.1578538]); O’Connor, S. 22 (K); Phillipson, P.B. 1750 (P[P04779820]), 2368 (MO, P[P04779820], WAG[WAG.1578536]), 2447 (MO, P[P04779819], WAG[WAG.1578537]), 2701 (BR[BR0000016205871], P[P05547045]); Randrianasolo, A. 152 (P[P05547042]); Ravelonanahary 2770-RN (P[P05547046]), 4762-RN (P[P05547043, P05547044]); Ravelonarivo, D. 4643 (MO[MO-3029388]); Sauther, M. 33 (P[P04779822]). references.— Govaerts etal. (2000: 75, 100); Montero Muñoz etal. (2018a: 105). 75 km Acalypha mayottensis Comoros Mayotte N fig. 53 . — Distribution map of Acalypha mayottensis I.Montero & Cardiel in Comoros Archipelago.
463 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) AB CD EF GH fig. 54. — A, B, Acalypha integrifolia Willd. (photo by J. M. Sarrailh); C, D, A. leptomyura Baill. (photo by T. Randrianarivony, T. Randrianarivony 749); E, A. marginata (Poir.) Spreng. (photo by F. Duban); F, A. mayottensis I.Montero & Cardiel (photo by J.-N. Labat, J.-N. Labat 3268); G, H, A. medibracteata Radcl.-Sm. & Govaerts (photo by L. Gillespie, L. Gillespie 10828).
464 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. DescripTion Shrubs, deciduous, to 2m tall, monoecious. Branches cinereous, nodose, pubescent with curved, antrorse trichomes, glabrescent when mature. Axillary buds ovoid, to 2× 1.5mm, perulate, perules 2, imbricate, chartaceous, light brown, woolly with simple, arachnoid trichomes. Stipules to 1.5mm long, lanceolate, woolly with simple, arachnoid trichomes, base with some sessile glands. Petioles 0.5-1(-3.5) cm long, indumentum similar to that on young branches. Leaf blades 3.5-5(-7)× (0.5-)1-1.5(-2.5) cm, usually linear-lanceolate to oblong-lanceolate, sometimes ovate-lanceolate, membranous; base rounded; apex acute to subacuminate, acumen to 6mm long, acute, mucronate; margin crenulate-serrulate, teeth with gland at apex; upper and lower surfaces laxly pubescent with simple, erect trichomes, and with simple, curved trichomes on veins, glabrescent; venation pinnate, secondary veins to 7-8 per side. Stipels glandular, to 0.2mm diameter, glabrous. Inflorescences spiciform, androgynous, axillary, to 7.5cm long; peduncle to 20mm long, pubescent with simple, arachnoid trichomes. Female segment to 3.5cm long; bracts 1-2(-3), sessile, enlarging in fruit to 12× 10mm, reniform, pubescent with short, simple trichomes and flattened, resinous glands, glabrescent; margin denticulate to subentire, central tooth not prominent; bracteoles absent. Male segment deciduous, to 2cm long; flowers glomerate; bracts to 1.2mm diameter, orbicular, woolly with simple, arachnoid trichomes, glabrescent; flowers glomerate, with stalked, flattened, resinous glands between pedicels. Male flowers: pedicel to 0.6 mm long, sparsely hairy; buds to 0.6mm diameter, sparsely hairy. Female flowers 2 per bract, sessile; sepals 3, to 0.5mm long, triangular, sparsely hairy; ovary c.1 mm diameter, 3-lobed, smooth, surface densely pubescent with simple, erect trichomes and flattened resinous glands; styles 3, to 5mm long, distinct, sparsely hairy, each divided into c.8 segments. Allomorphic flowers not seen. Capsules to 3 mm diameter, smooth, laxly pubescent with simple, short, curved trichomes and flattened resinous glands. Seeds c.1.9 × 1.5 mm, pyriform, minutely foveolate. 36. Acalypha menavody (Leandri) I.Montero& Cardiel PhytoKeys 108: 105 (Montero Muñoz etal. 2018a).— Acalypha spiciflora var. menavody Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 270 (Leandri 1942).— Type: Madagascar. Prov. Antsiranana, collines et plateaux calcaires de l’Analamera, I.1938, H. Humbert 19149 (lecto-, designated by Montero Muñoz etal. [2018a: 105]: P[P00536737]; isolecto-, P[P00536736, P00536738]). iconography.— Leandri (1942: 269); Fig. 57G. eTymology.— The epithet refers to the vernacular name of this plant, known as Menavody [cul rouge] (Leandri 1942). D isTribuTion anD habiTaT .— Endemic to Madagascar (Diana and Sava). Dry deciduous forest. On Mesozoic limestone. Altitudinal range 40-315m (Fig. 56). P relimiNary coNservatioN assessmeNt .— The EOO of Acalypha menavody is estimated to be 2 994km2 and its AOO 28km2. This species has been collected from several protected areas: Montagne des Français, which has been a protected area since 2006 (Category V; Dudley 2008), but most parts of the forest have been degraded by agriculture, grazing, and charcoal extraction (Sabel etal. 2009; De Block 2018); Ankarana Special Reserve (see assessment of A.ankaranensis); and Loky-Manambato, which has been a protected area since 2005 (Category V; Dudley, 2008), but the forest of which has been extensively degraded and continues to be threatened by subsistence farming, fires to clear land for grazing, logging, and gold extraction. The most serious threat in this last site is the illegal cutting of precious woods (Rambeloson 1999; Vargas etal. 2002; Rakotondravony 2009; Goodman etal. 2018). This species has also been collected in unprotected areas. Habitat loss and continued degradation of forests in these areas lead us to assess A.menavody as Endangered: EN B1ab(i,iii,iv) + B2ab(ii,iii,iv). maTerial examineD.— 8 collections. Madagascar: Bardot-Vaucoulon, M. 1016 (K, MO, P[P00309589, P00309590], TAN); Be, J. 325 (K, MO); d’Alleizette, Ch. s.n. (L[L0241933]); De Block, P. 1123 (BR[BR0000009728301, BR0000009728400], G, K, MO, P[P04779776], TAN); Humbert, H. 19149 (P[P00324461, P00536736, P00536737, P00536738]); Nusbaumer, L. LN 2178 (G[G00074391]); Ranaivojaona, R. 1659 (CNARP, MO[MO-2965747], P, TAN); Razafimandimbison, S. 1771 (S[S17-48170]). references.— Leandri (1942: 270); Montero Muñoz etal. (2018a: 105). 45°E 15°S 20° 25° 300 km 50° Acalypha medibracteata fig. 55. — Distribution map of Acalypha medibracteata Radcl.-Sm. & Govaerts in Madagascar.
465 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) DescripTion Shrubs, probably deciduous, to 3m tall, monoecious. Branches glabrous (youngest with some hyaline trichomes at base). Axillary buds ovoid, to 3× 2mm, perulate, perules 2, imbricate, chartaceous, blackish, glabrous; Stipules to 8mm long, linearlanceolate, ciliate with simple, arachnoid trichomes. Petioles slender, canaliculate, 1-2.5(-3) cm long, subglabrous, with some simple, arachnoid trichomes. Leaf blades 6-8(-9.5)× 4-6(-7.5) cm, broadly ovate-lanceolate, thin-membranous; base usually cordate, sometimes rounded to acute; apex acuminate, acumen to 10mm long, rounded, mucronate; margin denticulate, teeth rounded to obtuse, slightly callose-edged; upper and lower surfaces subglabrous, with some simple, appressed trichomes on veins; venation actinodromous, basal veins 5, secondary veins 5-6 per side. Stipels glandular, to 0.2mm long, sparsely hairy with some arachnoid trichomes. Inflorescences spiciform, androgynous, axillary, filiform, laxly flowered, to 10cm long, mostly male with short female segment; peduncle to 45mm long, glabrous. Female segment to 3.8cm long; bracts (1-)2-3, sessile, enlarging in fruit to 13mm diameter, suborbicular, glabrous; margin entire; bracteoles absent. Male segment persistent, to 5cm long; flowers glomerate; bracts to 0.3mm diameter, orbicular, sparsely hairy with some simple, arachnoid trichomes. Male flowers: pedicel to 0.5 mm long, glabrous; buds not seen. Female flowers 1 per bract, sessile; sepals 3, to 0.5mm long, triangular, sparsely hairy with simple, arachnoid trichomes; ovary c.1 mm diameter, 3-lobed, papil - lose, papillae acute, to 1mm long, ending in gland, surface glabrous; styles 3, to 4mm long, connate at base, rachis thick, glabrous, each divided into c.12 segments. Allomorphic flowers not seen. Capsules to 4 mm diameter, papillose, papillae to 2mm long, surface subglabrous, with some, sparse, simple trichomes. Seeds c.3 × 2 mm, pyriform, minutely foveolate. 37. Acalypha nusbaumeri I.Montero& Cardiel PhytoKeys 140: 69 (Montero Muñoz etal. 2020b).— Type: Madagascar. Reg. Sava [Prov. Antsiranana], sous-préfecture de Vohemar, commune rurale de Daraina, forêt de Bekaraoka, partie nord, 13°06’S, 49°42’E, 177m, 13.II.2004, L. Nusbaumer& P. Ranirison LN1169 (holo-, G[G00028080]; iso-, K, MO, P[P01152829, P05547228]). iconography.— Montero Muñoz etal. (2020b); Figs 57H; 69A. eTymology.— The epithet honors Louis Nusbaumer, researcher and curator at the Conservatoire et Jardin botaniques de la Ville de Genève, Switzerland. Nusbaumer works on the systematics, phylogeny, biogeography, and conservation of Malagasy plants and is also the collector, with Patrick Ranirison, of the type specimen of this species. DisTribuTion anD habiTaT.— Endemic to Madagascar (Sava). Mainly in dry deciduous forest. On basement rocks. Altitudinal range 200-351m (Fig. 58). P relimiNary coNservatioN assessmeNt .— Acalypha nusbaumeri is only known from one collection from Loky-Manambato site (see assessment of A.menavody). Its EOO could not be calculated; its AOO is estimated to be 8km2. Acalypha nusbaumeri is assigned a preliminary conservation status of Critically Endangered: CR B2ab(ii,iii). maTerial examineD.— 1 collection. Madagascar: Nusbaumer, L. LN 1169 (G[G00028080], K, MO, P[P05547228, P01152829]). DescripTion Shrubs, probably deciduous, to 0.8m tall, divaricately branched, monoecious. Branches blackish, pubescent with simple, curved, antrorse trichomes, glabrous when mature. Axillary buds spherical, to 1mm diameter, perulate, perules 2, imbricate, membranous, brownish, external perule dentate, sparsely hairy. Stipules to 2mm long, triangular-lanceolate, midrib prominent, sparsely hairy with simple, short, erect trichomes. Petioles slender, 2-3.5(-4.5) cm long, indumentum similar to that on young branches. Leaf blades 5-8.5× 2.5-5cm, ovatelanceolate to elliptic-lanceolate, thin-membranous; base obtuse to rounded; apex subacuminate, acumen to 0.5mm long, rounded, mucronate; margin crenate-serrate, teeth rounded; upper and lower surfaces subglabrous, with some simple, short, curved, antrorse trichomes on veins; axils of secondary veins with sparsely hairy, pocket-shaped domatia; venation actinodromous, basal veins 3 or 5, secondary veins 3-4 per side. Stipels absent. Inflorescences spiciform, androgynous, axillary, to 45°E 15°S 20° 25° 300 km 50° Acalypha menavody fig. 56. — Distribution map of Acalypha menavody (Leandri) I.Montero & Cardiel in Madagascar.
472 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. DisTribuTion anD habiTaT.— Endemic to Madagascar (AlaotraMangoro, Analamanga, Itasy, Vakinankaratra, Amoron’i Mania, and Haute Matsiatra). Sclerophyllous woodlands and secondary forest. On basement rocks. Altitudinal range (800-) 1500-2000m (Fig. 63). p reliminary conservaTion assessmenT .— The EOO of Acalypha radula is estimated to be 33 908km2 and its AOO 60km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows in the sclerophyllous woodlands in the highlands (see assessment of A.leptomyura). This species has been collected in degraded forests and in unprotected areas, and in many localities. Its habitat is subject to various threats (uncontrolled fires for grazing and crops, charcoal production), so we predict that ongoing habitat loss and degradation will cause continued decline of its EOO and AOO. Acalypha radula is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 28 collections. Madagascar. Anonymous 9628 (P[P00513109]); Anonymous s.n. (P[P00513097]); Baron, R. 272 (P[P00513096]), 1818 (K[K000186509], P[P00513119, P00513120]), 4068 (K[K000384473]); Benoist, M.R. 244 (P[P00513098]); Bojer, W.s.n. (P[P00724168]); Bosser, M.J. 15133 (P[P04779790, P04779793]), 19481 (P[P04779797]); Catat, L.D.M. 413 (P[P00513099]); d’Alleizette, Ch. s.n. (L[L0242131]); Decary, R. 13008 (P[P00513100]); Gillespie, L. 10757 (MO), 10758 (MO); Hildebrandt, J.M. 3865 (BREM[BREM_0001787], G[G00034245], P[P00513101, P00513102, P00513103]), 3900 (BREM[BREM_0001783], G[G00074184, G00190630], JE[JE00000288, JE00000289], K[K000186507], M[M0110604], P[P00513121, P00513122]); Humbert, H. 4575 (P[P00513107]), 29970 (G, P[P05547266]); Jard. Bot. Tananarive 2528 (P[P00513106]), 4537 (P[P00513104, P00513105]); Le Myre de Villers, M.s.n. (P[P00513108]); Perrier de la Bâthie, H. 9672 (P[P00513110, P00513111]), 9862 (P[P00513116]), 13357 (P[P00513114, P00513115]); Razafimandimbison, S. 1359 (S[S16-50647]); Service Forestier Madagascar 45-SF (P[P00513112]), s.n. (P[P00513113]); Viguier, R. 1836 (P[P00513118]). references.— Baillon (1861: 268) as A.salviifolia; Müller Argoviensis (1866: 889) as A.salviifolia; Baillon (1891: 193); Baillon (1892: 1004) as A.salviifolia; Baillon (1895a: 1180) as A.hilde - brandtii; Palacký (1907: 25) as A.hildebrandtii; Palacký (1907: 26); Palacký (1907: 26) as A.salviifolia; Pax& Hoffmann (1924: 102) as A.salviifolia; Pax& Hoffmann (1924: 156) as A.hildebrandtii; Leandri (1942: 278); Leandri (1952); Jenkins (1987: 347); Jenkins (1990: 408, 433); Govaerts etal. (2000: 67) as A.hildebrandtii; Govaerts etal. (2000: 85, 86, 105) as A.salviifolia; Schatz (2001: 142); Sagun etal. (2006: 124); Seebaluck etal. (2015: 149, 153); Montero Muñoz etal. (2018a: 108). DescripTion Shrubs, evergreen, to 2.5m tall, monoecious or dioecious. Branches densely pubescent with simple, short, more or less appressed trichomes and hyaline trichomes to 0.8mm long, glabrescent when mature. Axillary buds ovoid, to 2× 1.5mm, perulate, perules 2, imbricate, chartaceous, blackish, pubescent with simple, short trichomes. Stipules to 8mm long, triangular-lanceolate, midrib thick, margin scarious, appressed-pubescent, especially on midrib. Petioles 0.5-2.2(- 3) cm long, indumentum similar to that on young branches and with scattered short, glandular trichomes. Leaf blades conspicuously bullate, 6-10× (1.5-)2-3(-3.5) cm, narrowly triangular-lanceolate to linear-lanceolate, subchartaceous; base rounded to truncate, sometimes subcordate; apex rounded to acute; margin serrate, teeth subacute, irregular; upper surface pubescent with simple, short, erect, trichomes on top of blisters and on main veins and scattered, short, glandular trichomes, denser on veins near base, and sometimes stellate or fasciculate trichomes; lower surface pubescent with simple, thin trichomes, denser on veins, and sometimes stellate or fasciculate trichomes; venation actinodromous, basal veins 5, secondary veins 6-10 per side. Stipels absent. Inflorescences spiciform, unisexual, axillary. Male inflorescences densely flowered, to 5cm long; peduncle to 8mm long, indumentum similar to that on young branches; flowers glomerate; bracts to 0.5mm long, narrowly spatulate, sparsely hairy and ciliate with simple trichomes, and with short, glandular trichomes. Female inflorescences densely flowered, to 6cm long; peduncle to 8mm long, indumentum similar to that on young branches; bracts 4-9, sessile, enlarging in fruit to 12× 14mm, subtriangular-reniform, pubescent with simple, thin, erect, trichomes and thick, glandular trichomes to 1mm long, ciliate; margin subentire to shallowly dentate; teeth c.39, triangular, with glandular trichomes, central tooth not prominent; 45°E 15°S 20° 25° 300 km 50° Acalypha rabesahalana fig. 62. — Distribution map of Acalypha rabesahalana I.Montero & Cardiel in Madagascar.
473 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) bracteoles to 1.5mm long, triangular, pubescent with simple, short trichomes and ciliate. Male flowers: pedicel to 0.8 mm long, sparsely hairy; buds to 1mm diameter, sparsely hairy. Female flowers 1 per bract, sessile; sepals 3, to 0.8mm long, oblong-lanceolate, sparsely hairy and ciliate; ovary c.1 mm diameter, 3-lobed, smooth, surface hispid with simple, short, erect trichomes, and with glandular trichomes; styles 3, to 5mm long, slightly connate at base, sparsely hairy on rachis, each divided into 9-10 segments. Allomorphic flowers sometimes present at female inflorescence apex, occasionally at male inflorescence apex; pedicel filiform, to 20mm long, sparsely hairy; ovary 1-lobed, to 3mm diameter, puberulent with simple, short trichomes, distally fimbriate; style 1, to 1mm long, glabrous. Capsules to 5 mm diameter, smooth, surface sparsely hairy with simple, short, erect trichomes and glandular trichomes to 1.3mm long. Seeds c.3 × 2 mm, pyriform, minutely foveolate. noTes 1) Baker designated R. Baron 1818 as the type specimen of Acalypha radula but did not indicate the herbarium. We found three specimens of this collection, designating the best preserved of them as the lectotype; and 2)Baillon designated J. M. Hildebrandt 3900 as the type of Acalypha hildebrandtii but did not indicate the herbarium. We found nine specimens of this collection, designating the best preserved of them as the lectotype. 43. Acalypha richardiana Baill. Adansonia, recueil d’observations botaniques 1: 268 (Baillon 1861).— Type: Comoros. Mohéli, “ÎleMohilla”, s.d., M. Richard 287 (lecto-, designated by Montero Muñoz etal. [2018a: 108]: P[P04779566]; isolecto-, P[P04779562, P04779564, P04779565]).— Former syntypes: Madagascar. s.l., s.d., M. Richard 544 (P[P04779563]).— Mayotte. s.l., s.d., L. H. Boivin 3373 (GDC[G00324505], P[P00196299, P00196300], W[WRchb. 1889-0166704]). Acalypha ovalifolia Baill., Adansonia, recueil d’observations botaniques 1: 269 (Baillon 1861).— Acalypha reticulata var. ovalifolia (Baill.) Müll.Arg., Linnaea 34: 32 (Müller Argoviensis 1865).— Acalypha filiformis var. ovalifolia (Baill.) Govaerts, World Checklist and Bibliography of Euphorbiaceae (and Pandaceae) 62 (Govaerts etal. 2000).— Type: Mayotte. XI.1848, L. H. Boivin 3372 (holo-, P[P00196298]). iconography.— Fig. 65G. eTymology.— The epithet honors French botanist Jean Michel Claude Richard (1787-1868). DisTribuTion anD habiTaT.— Endemic to the Comoros Archipelago (Anjouan, Moheli and Mayotte). Dry forest and secondary forest. Altitudinal range 20-680m (Fig. 64). p reliminary conservaTion assessmenT .— Acalypha richardiana is only known from the Comoros. Its EOO is estimated to be 4 395km2 and its AOO 36km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category (see assessment of A.chibomboa from Comoros). We predict that ongoing habitat loss and the impact of alien species in this archipelago will cause continued decline of its EOO and AOO, so A.richardiana is assessed as Endangered (EN): EN B1ab(i,iii,iv) + B2ab(ii,iii,iv). maTerial examineD.— 15 collections. Comoros Archipelago. Barthelat, F. 680 (P[P00273174, P00273175]), 1163 (P[P00437836]); Boivin, L.H. 3372 (P[P00196298]), 3373 (G[G00034250, G00324505], P[P00196299, P00196300, P02712256, P02712257], W[WRchb. 1889-0166704]), s.n. (P[P00196291]); d’Alleizette, Ch. s.n. (L[L0242103]); Hildebrandt, J.M. 1661 (BREM[BREM_0001789], L[L0242104], P[P00635340]); Labat, J.-N. 2908 (P[P00078905]); Pascal, O. 954 (P[P00144572]); Richard, M. 224 (P[P04779560, P04779561]), 287 (P[P04779562, P04779564, P04779565, P04779566]), 544 (P[P04779563]), s.n. (P[P00196297]); Rouhan, G. 845 (P[P00684849]); Tinguy, H. 1001 (P[P00144571]). references.— Müller Argoviensis (1866: 852 as A.reticulata var. ovalifolia, 855); Baillon (1892: 1004); Palacký (1907: 25 as A.ovalifolia, 26); Voeltzkow (1917: 447); Pax& Hoffmann (1924: 127); Leandri (1942: 272); Govaerts etal. (2000: 62, 85, 105); Montero Muñoz etal. (2018a: 109). DescripTion Shrubs, leaf persistence unknown, to 2m tall, monoecious. Branches densely pubescent with simple, short, curved trichomes, glabrous when mature. Axillary buds spherical, to 1mm diameter, perulate, perules 2, valvate, membranous, brownish, pubescent with simple, short trichomes. Stipules to 6mm long, linear, sparsely hairy. Petioles 2-6(-9) cm 45°E 15°S 20° 25° 300 km 50° Acalypha radula fig. 63. — Distribution map of Acalypha radula Baker in Madagascar.
474 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. long, indumentum similar to that on young branches. Leaf blades 7-13.5× 2-7.5cm, ovate-lanceolate, membranous; base rounded to truncate; apex acute to acuminate, acumen to 15mm long, rounded, mucronate; margin crenate to crenate-dentate, slightly revolute, teeth rounded; upper surface laxly pubescent with simple, short, trichomes, denser on veins, glabrescent; lower surface subvelutinous, glabrescent; venation actinodromous, basal veins 3 or 5, secondary veins 5-8 per side. Stipels glandular, to 0.2mm long, sparsely hairy. Inflorescences spiciform, androgynous, and sometimes solitary female bracts, axillary. Androgynous inflorescences to 6cm long, mostly male with 1 female bract; peduncle to 2.5mm long, it and rachis sparsely hairy. Female segment: bract 1, sessile, enlarging in fruit to 7× 6mm, suborbicular, pubescent; margin usually subentire, sometimes slightly crenate, with some sessile glands; bracteoles absent. Male segment persistent, filiform, laxly flowered, to 5.5cm long; flowers glomerate; bracts to 0.8mm long, triangular, sparsely hairy and ciliate. Solitary female bracts sessile, similar to those of androgynous inflorescences. Male flowers: pedicel to 0.5 mm long, sparsely hairy; buds to 0.5 mm diameter, sparsely hairy. Female flowers 1-2 per bract, sessile; sepals 3, to 0.5mm long, triangular, glabrous, margin with some sessile glands; ovary c.1 mm diameter, 3-lobed, apparently smooth, surface densely hispid with hyaline trichomes to 1mm long; styles 3, to 4mm long, distinct, sparsely hairy, each divided into 10-12 segments. Allomorphic flowers sometimes present, axillary; pedicel filiform, to 19mm long, pubescent with simple, short, curved trichomes; sepals 3, to 0.5mm long, triangular, ciliate and with some sessile glands; ovary 1-lobed, to 1mm diameter, echinate, pubescent with simple, flattened trichomes; style 1, to 3mm long, sparsely hairy. Capsules to 2 mm diameter, papillose-hispid, papillae acute, to 0.5mm long, surface pubescent with simple, short trichomes. Seeds c.1.5mm diameter, subglobose, minutely foveolate. noTe The specimens indicated as isolectotypes do not have Richard’s original labels. They instead have labels with Baillon’s handwriting giving the location as “Madagascar”. We believe that the correct location is the one indicated on the selected lectotype, “Île Mohilla” (currently named Mohéli), in the Comoros Archipelago. A search of specimens at P showed that Richard’s collections numbered 284-286, 288, 290, 291 and 293 are also from Mohéli (289has no locality and 292 is not listed). Richard 544has an apparently original label showing the locality only as “Madagascar.” Collections at P with nearby numbers are labelled as being from either “Nord de Madagascar” or “Île Nos-bé” (now Nossi-bé), so this collection could be from northern Madagascar, although we have seen no other specimens from outside the Comoros Archipelago. 44. Acalypha rottleroides Baill. Adansonia, recueil d’observations botaniques 1: 270 (Baillon 1861).— Type: Madagascar. Prov. Antsiranana: Nossibé, 1837, M. Richard 215 (holo-, P[P00536728]). Acalypha polynema Baill., Histoire physique, naturelle et politique de Madagascar Atlas, 2, fasc. 27: t. 187 (Baillon 1891).— Type: Madagascar. Baillon 1891: pl. 187, holotype. Acalypha juliflora Pax, Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie. Leipzig 19: 95 (Pax 1894).— Type: Madagascar. Prov. Antsiranana: Nossibé, “Urwald von Loko-bé”, XII.1879, J. M. Hildebrandt 3279 (lecto-, designated here: W[W1889-0089773]; isolecto-, JE[JE00004291], K[K000186527], P[P00536729, P00536730, P00536731]). iconography.— Baillon (1891: t. 185, t. 187); Figs 65H; 69E. eTymology.— The epithet likely refers to a resemblance to the genus Rottlera Roxb. (now treated as a synonym of Mallotus Lour.; Euphorbiaceae). D isTribuTion anD habiTaT .— Endemic to Madagascar (Diana and Sava). Degraded lowland evergreen moist forest and dry deciduous forest. On Mesozoic limestone and on basement rocks. Altitudinal range 200-370m (Fig. 66). preliminary conservaTion assessmenT.— The EOO of Acalypha rottleroides is estimated to be 5 643km2, which is near to the threshold of the Endangered category, and its AOO 32km2. It is known from only four locations. Acalypha rottleroides grows mainly in dry deciduous forest, but in Nosy Bé this species grows in lowland evergreen moist forest. The collections found in Nosy Bé were made in 1879 and this island has been largely deforested for conversion into coffee plantations, rice fields, ylang-ylang (Cananga odorata Hook.) orchards, and sugar cane fields (Andreone etal. 2013). Currently, the area where these specimens were collected is the National Park of Lokobé (Category II; 75 km Acalypha richardiana Comoros Mayotte N fig. 64 . — Distribution map of Acalypha richardiana Baill. in Comoros Archipelago.
475 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) AB CD EF GH fig. 65. — Mature female bracts: A, Acalypha perrieri Leandri (R. Decary 15792); B, A. pervilleana Baill. (J. J. Floret 682); C, A. pervilleana (J. M. Hildebrandt 3356); D, A. poiretii Spreng. (L. Boivin s.n.); E, A. rabesahalana I.Montero & Cardiel (H. Humbert 32480); F, A. radula Baker (Service Forestier Madagascar 45-SF); G, A. richardiana Baill. (M. Richard 224); H, A. rottleroides Baill. (L. Nusbaumer LN2629). Illustration by Iris Montero Muñoz. Scale bars: A, D, E, G, H, 1 mm; B, C, F, 2 mm.
476 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. Dudley 2008). The remaining collections of A.rottleroides were made in Ankarana Special Reserve, Loky-Manambato, and in the unprotected forests of Ambilanivy and Rangoty (Ampasindava). These areas are threatened by mining (sapphire and gold), illegal cutting of precious woods, and fires to improve grazing and agriculture (Kull 2000; Rakotondravony 2009; Goodman etal. 2018). Continued habitat loss leads us to assess A.rottleroides as Endangered: EN B2ab(ii,iii,iv). specimen examineD.— 8 collections. Madagascar. Cours, G. 5626 (P[P00324530]); Hildebrandt, J.M. 3279 (G, JE[JE00004291], K[K000186527], L[L0241761], P[P00536729, P00536730, P00536731], W[W1889-0089773]); Humbert, H. 19065 (P[P00324502]); Nusbaumer, L. LN 1933 (G[G00090298], P[P05547268]), LN 2629 (G[G00180763], P[P04786259]); Ranirison, P. PR 1085 (G[G00090513], P[P05547267]); Richard, M. 181 (P[P04779559]), 215 (P[P00536728]). references.— Müller Argoviensis (1866: 854); Baillon (1891: 182); Baillon (1895b: 1197); Baillon (1895b: 1197) as A.polynema; Palacký (1907: 25) as A.juliflora; Palacký (1907: 26) as A.polynema; Palacký (1907: 26); Nitschke (1923: 280) as A.juliflora; Pax& Hoffmann (1924: 127); Leandri (1942: 277); Govaerts etal. (2000: 70, 105) as A.juliflora; Govaerts etal. (2000: 86); Montero Muñoz etal. (2018a: 109). DescripTion Shrubs, deciduous, to 2m tall, monoecious. Branches whitish, hispid with simple, hyaline trichomes to 2mm long, and with glandular trichomes to 1.5mm long, glabrescent when mature but glandular trichomes persistent. Axillary buds spherical, to 1mm diameter, perulate, perules 4, overlapping (superposed), membranous, scarious, margin irregular, glabrous. Stipules to 7mm long, ovate-lanceolate to oblong-lanceolate; margin scarious, glabrous except margin ciliate with some short, glandular trichomes. Petioles (3-)3.5-7.5(-8) cm long, indumentum similar to that on young branches, hyaline trichomes deciduous. Leaf blades 7-13× 5-7.5cm, ovate-lanceolate, thin-membranous; base cordate; apex acute to acuminate, acumen to 10mm long; margin crenate to denticulate, teeth rounded to acute; upper and lower surfaces subglabrous, with some thin, glandular trichomes to 1.5mm long; margin ciliate with simple, short trichomes, and with glandular trichomes to 1mm long, venation actinodromous, basal veins 5, secondary veins 4-6 per side. Stipels absent. Inflorescences spiciform, androgynous, axillary, to 9cm long, mostly female with short male segment (sometimes appearing female due to male segment falling); peduncle to 10mm long, with glandular trichomes to 1mm long. Female segment laxly flowered, to 8cm long; bracts 6-9, sessile, enlarging in fruit to 6× 9mm, reniform, with glandular trichomes to 1.5mm long; margin subentire to crenate, teeth triangular, central tooth not prominent; bracteoles absent. Male segment deciduous, to 1cm long; flowers glomerate; bracts to 1.5mm diameter, suborbicular, scarious, glabrous; margin irregular, papillose. Male flowers: pedicel to 0.5 mm long, glabrous; buds to 0.5 mm diameter, glabrous. Female flowers 1 per bract, sessile; sepals 3, to 1.2mm long, triangularlanceolate, reddish, margin irregular, with sparse glandular trichomes; ovary c.1 mm diameter, 3-lobed, smooth, surface with some glandular trichomes to 0.5mm long; styles 3, to 6mm long, slightly connate at base, glabrous, each divided into 9-10 segments. Allomorphic flowers not seen. Capsules to 2.5 mm diameter, smooth, surface with sparse glandular trichomes. Seeds c.1.5 × 1.2 mm, pyriform, minutely foveolate. noTe Baillon designated J. M. Hildebrandt 3279 as the type of Acalypha juliflora but did not indicate the herbarium. We found six specimens of this collection, designating the best preserved of them as the lectotype. 45. Acalypha spachiana Baill. Adansonia, recueil d’observations botaniques 1: 272 (Baillon 1861).— Acalypha spachiana var. latifolia Baill., Adansonia, recueil d’observations botaniques 1: 272 (Baillon 1861).— Type: Madagascar. Prov. Antsiranana, Baies de Rigny et de Diego-Suarès, 1848, L. H. Boivin 2654 (lecto-, designated by Montero Muñoz etal. [2018a: 109]: P[P00536733]; isolecto-, G[G00034251], GDC[G00324359], P[P00536734]). 45°E 15°S 20° 25° 300 km 50° Acalypha rottleroides fig. 66. — Distribution map of Acalypha rottleroides Baill. in Madagascar.
477 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) Acalypha spachiana var. acutifolia Baill., Adansonia, recueil d’observations botaniques 1: 272 (Baillon 1861).— Type: Madagascar. Prov. Antananarivo, Antananarivo, 12.II.1840, J. P. Goudot s.n. (holo-, G[G00383582]). Acalypha spachiana var. minor Baill., Adansonia, recueil d’observations botaniques 1: 272 (Baillon 1861).— Type: Madagascar. s.l., s.d., W. Bojer s.n. (holo-, P[P00536735]; iso-, TUB[TUB002081, TUB002082]). Acalypha buchenavii Müll.Arg., Abhandlungen herausgegeben vom Naturwissenschaftlichen Vereins zu Bremen. Bremen 7: 27 (Müller Argoviensis 1882).— Type: Madagascar. Prov. Antananarivo, Antananarivo, 18.XII.1877, D.C. Rutenberg s.n. (lecto- [probably holotype], designated by Montero Muñoz etal. [2018a: 110]: BRNU[BRNU347926]). Acalypha squarrosa Pax, Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie. Leipzig 19: 97 (Pax 1894).— Type: Madagascar. Prov. Antananarivo, Antsirabe, “Sírabé”, VIII.1880, J. M. Hildebrandt 3560 (holo-, B or WRSL, presumably destroyed; lecto-, designated by Montero Muñoz etal. [2018a: 110]: JE[JE00004308]; isolecto-, JE[JE00004309], K[K000186510], P[P00536732]). Tragia saxatilis Bojer ex Pax& K.Hoffm., Das Pflanzenreich (Engler) 147, 16 (Heft 85): 33 (Pax& Hoffmann 1924), nom. nud. iconography.— Leandri (1942: 257, 276); Figs 69F; 73A. e Tymology .— The epithet honors French botanist Édouard Spach (1801-1879). DisTribuTion anD habiTaT.— Endemic to Madagascar (Diana, Sava, Sofia, Alaotra-Mangoro, Analamanga, Vakinankaratra, Haute Matsiatra, and Anosy). Lowland evergreen moist forest, moist semideciduous forest, dry deciduous forest, riparian forest, and secondary forest. On Mesozoic limestone, sandstone, and basement rocks. Altitudinal range (30-) 100-1600 (-1900) m (Fig. 67). p reliminary conservaTion assessmenT .— The EOO of Acalypha spachiana is estimated to be 298 644km2 and its AOO 172km2, which is less than the 500km2 threshold of the B2 subcriterion of the Endangered category. This species grows in evergreen moist forests (see assessment of A.emirnensis) and dry deciduous forests (see assessment of A.diminuta). Acalypha spachiana has been collected in some protected areas, but it is also known from several unprotected forests subject to various threats. We predict ongoing habitat loss will cause continued decline of its EOO and AOO. Acalypha spachiana is assessed as Near Threatened (NT) under criterion B. It meets the AOO values needed for a threatened category, but the number of locations is too high to meet the conditions required for a listing under the Endangered category. maTerial examineD.— 85 collections. Madagascar. Anonymous 1240 (P[P05547038]); Anonymous 1274 (P[P05547030]); Anonymous 3891 (P[P04779550]); Anonymous 3911 (P[P04779547]); Anonymous 3927 (P[P04779548]); Anonymous 5455 (P[P05547040]); Anonymous 5731 (P[P05547028]); Anonymous s.n. (P[P04779552]); Anonymous s.n. (P[P04779555]); Baron, R. 574 (P[P04779833]), 4067 (K[K000384479]), 4426 (P[P04779542, P04779543]), 5103 (K[K000384478], P[P04779801]); Be, J. 168 (P[P05481878]); Benoist, M.R. 177 (P[P04779544]), 196 (P[P04779800]); Blackburn, J. 106 (46?) (K[K000384482]); Boivin, L.H. 2654 (G[G00034251, G00324359], P[P00536733, P00536734]); Bojer, W.s.n. (P[P00536735], TUB[TUB002081, TUB002082]); Bosser, M.J. 235 (P[P04779810]), 2792 (P[P04779805]), 9895 (P[P05547256]); Campenon, R.P.s.n. (P[P04779796]); Cours, G. 399 (P[P04779541]), 428 (P[P04779540]), 478 (P[P05547261]), 1820 (P[P00324542]), 1968 (P[P05547183]); Croat, T.B. 28526 (P[P04779836]); d’Alleizette, Ch. s.n. (L[L0242170]); Decary, R. 553 (P[P04779837]), 5972 (P[P04779538]), 6032 (P[P04779839]), 6055 (P[P04779534]), 6197 (P[P05547185]); Foury, C.s.n. (P[P05481886]); Gautier, L. LG 4904 (G[G369044/1], P[P05547021]), LG 5076 (G[G00170072], P[P05543673]), LG 5214 (G[G00170081], P[P05543675]); Gillespie, L. 10856 (MO); Goudot, J.s.n. (G[G00383582]); Herb. d’Alleizette 324 (P[P04779828]); Hildebrandt, J.M. 3560 (JE[JE00004308, JE00004309], K[K000186510], P[P00536732]); Homolle, A.M. 179 (P[P05547029]), 1968 (P[P04779866]); Humbert, H. 2180 (P[P05481895]), 11698 (P[P04779532, P04779535]), 18593 (P[P04779553]), 18954 (P[P04779556, P04779557, P04779558]), 19741 (P[P05547258]), 19791 (P[P05547257]), 32613 (P[P05547058]), 32721 (G, P[P05547273]); Jacquemin, H.s.n. (P[P04779554]); Jard. Bot. Tananarive 5739 (P[P00324569]); Jongkind, C.C.H. 3309 (BR[BR0000005555499], K, TAN, WAG[WAG.1578524, WAG.1578525]); Le Myre de Villers, M.s.n. (P[P04779798]), s.n. (P[P04779830]), s.n. (P[P05547180]); Leandri, J. 3482 (P[P05547262]), 3582 (P[P05547037]), 3890 (P[P05547036]); Maître 83 (P[P05547024]); Manjato, N. 67 (P[P05510106]); Morat, P. 1431 (P[P05547023, P05547025, P05547026, P05547027]); Nusbaumer, L. LN 1817 (G[G00090193], P[P05547260]), LN 1843 (G[G00090216], P[P05547259]); Parker, W.G.s.n. (K[K000384483]); Peltier, J. 1645 (P[P05547032]); Perrier de la Bâthie, H. 9599 (P[P00324503]), 9859 (P[P05547182]); Rakotondranony 5 (K[K000384476]); Ramandimbimanana, S.D. SDR 150 (G[G00376942]), 328 (G[G00377119]); Razafindrakoto 12569-RN (P[P04779533]); Razafitsalama, L. J. 776 (G[G00382426], P[P04779545]); Rotereau, L.s.n. (P[P05510095]); Rutenberg, C.s.n. (G[G00190632]), s.n. (BRNU[BRNU347926]); Schlieben, H.J. 8156 (B, G[G00420848]); Scott-Elliot, G.F. 2042 (P[P04779530, P04779531]); Seyrig, A. 52 (P[P05547033]); Waterlot, M.s.n. (P[P05547031]). references.— Müller Argoviensis (1866: 827); Müller Argoviensis (1882: 27) as A.buchenavii; Baillon (1892: 1003); Baillon (1895b: 1199) as A.buchenavii; Palacký (1907: 26); Pax& Hoffmann (1924: 33); Pax& Hoffmann (1924: 33) as A.squarrosa; Leandri (1942: 255, 257); Jenkins (1987: 347); Jenkins (1990: 408, 433); Govaerts etal. (2000: 89, 96); Govaerts etal. (2000: 89) as A.squarrosa; Sagun etal. (2006: 124); Seebaluck etal. (2015: 153); Montero Muñoz etal. (2018a: 109). DescripTion Shrubs, deciduous, to 3m tall, monoecious. Branches pubescent with simple, curved, retrorse trichomes, glabrescent when mature. Axillary buds ovoid, to 3× 2mm, perulate, perules 4, overlapping (superposed), membranous, brownish, glabrous. Stipules to 5mm long, linear-lanceolate, midrib prominent, sparsely hairy with simple, appressed trichomes on midrib. Petioles (0.5-)1.5-2(-4) cm long, indumentum similar to that on young branches. Leaf blades 2-6(-13)× 1.3-4(-7) cm, usually ovate-lanceolate, sometimes elliptic-lanceolate to obovate-lanceolate, membranous; base rounded to subcordate; apex acute to acuminate, acumen to 10mm long, rounded to subacute, mucronate; margin serrate, teeth rounded to acute, callose-edged; upper and lower surfaces sparsely hairy with simple, short, erect trichomes, and with simple, curved trichomes on veins; venation actinodromous, basal veins 5, secondary veins 4-5 per side. Stipels glandular, to 0.5mm long, sparsely hairy. Inflorescences spiciform, androgynous, terminal, to 6cm long, mostly female with short male segment (sometimes appearing female due to male segment falling); peduncle to 10mm long, indumentum similar to that on young
478 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. branches. Male segment deciduous, to 1.5cm long; flowers glomerate; bracts to 1mm long, triangular, ciliate. Female segment to 4.5cm long; bracts c.18, sessile, enlarging in fruit to 6× 9mm, reniform, pubescent with simple, short trichomes and glandular trichomes to 1mm long; margin dentate, teeth 11-14, usually triangular, sometimes rounded, central tooth not prominent; bracteoles absent. Male flowers: pedicel to 1 mm long, sparsely hairy; buds to 1 mm diameter, sparsely hairy. Female flowers 1 per bract, sessile; sepals 3, to 0.5mm long, triangular, sparsely hairy; ovary c.1 mm diameter, 3-lobed, muricate, surface hispid with simple trichomes to 1.5mm long; styles 3, to 3mm long, distinct, sparsely hairy on rachis, each divided into 8-10 segments. Allomorphic flowers sometimes present, axillary; pedicel filiform, to 3mm long, puberulent with simple, short, curved, antrorse trichomes; sepals 3, to 0.5mm long, lanceolate, sparsely hairy; ovary 2-lobed, to 1mm diameter, papillose-hispid; styles 2, to 2mm long, glabrous. Capsules to 2.3 mm diameter, papillose, papillae rounded, minute, surface pubescent with appressed, hyaline trichomes. Seeds c.1.8 × 1.2 mm, pyriform, foveolate. noTe The holotype of Acalypha buchenavii should be at BREM, but Rutenberg’s specimens arrived at BRNU herbarium after World War II. Originally in the Überseemuseum in Bremen (BREM), they were transferred to northern Moravia (Czechia was at that time a Protectorate of Nazi Germany) to save them from potential destruction by bombardment. After the collapse of the Nazi regime, these collections were confiscated as “German property” and sent to BRNU. 46. Acalypha tremula I.Montero& Cardiel Systematic Botany 45 (1): 127 (Montero Muñoz etal. 2020a).— Type: Madagascar. Sava [Antsiranana], Province de Diego-Suarez, Vohemar, commune rurale de Daraina, forêt d’Ankaramy, 181m, 13°17.1’S, 49°40.91’E, 26.II.2004, L. Nusbaumer& P. Ranirison 1227 (holo-, G[G00006790], iso-, K, P[P04786267, P05547264], TAN).— Paratypes: Madagascar. Sava region (Antsiranana prov.), Fivondronana, Diego II, commune Mahavanona, Montagne des Français, forêt d’Ampitiliantsambo, à 3 heures de marche à pied, à l’Est d’Andranomanitra, 205m, 12°23’13”S, 49°23’04”E, 14.I.2005, R. Randrianaivo etal. 1160 (BR[BR0000005242887], CNARP, MO [MO2965736], P, TAN); Ambilobe, Mahamasina, Réserve Spéciale d’Ankarana, chemin d’Ambohimalaza, sous-bois sur basalte, 110m, 12°58’1.20”S, 49°6’5.11”E, 08.II.2003, M. Bardot-Vaucoulon 1400 (P[P00455711], K, MO [MO-2966207], TAN); Vohemar, commune rurale de Daraina, forêt de Solaniampilana-Maroadabo, 422m, 13°06.50’S, 49°33.97’E, 03.II.2006, L. Nusbaumer& P. Ranirison LN 2181 (G[G00074394]). iconography.— Montero Muñoz etal. (2020a); Figs 69G; 73B. eTymology.— The epithet refers to the resemblance of the long petiolate leaves of this species to those of the Eurosiberian tree Populus tremula L. (Salicaceae). DisTribuTion anD habiTaT.— Endemic to Madagascar (Diana and Sava). Dry deciduous forest. On clay soil on basement rocks. Altitudinal range 110-420m (Fig. 68). p reliminary conservaTion assessmenT .— The EOO of Acalypha tremula is estimated to be 2 609km2 and its AOO 16km2. It has been collected from three protected areas: Montagne des Français (see assessment of A.gillespieae), Ankarana Special Reserve (see assessment of A.ankaranensis), and Loky-Manambato (see assessment of A.menavody). The small number of locations, together with continuing habitat loss and degradation in these areas, lead us to assess A.tremula as Endangered: EN B1ab(i,iii) + B2ab(ii,iii). maTerial examineD.— 4 collections. Madagascar: Bardot-Vaucoulon, M. 1400 (K, MO[MO-2966207], P[P00455711], TAN); Nusbaumer, L. LN 1227 (G[G00006790], K, P[P05547264, P04786267]), LN 2181 (G[G00074394]); Randrianaivo, R. 1160 (BR[BR0000005242887], CNARP, MO[MO-2965736], TAN). DescripTion Shrubs, probably deciduous, to 4m tall, divaricately branched, monoecious. Branches slightly angular, dark brown, rugose, pubescent with simple, antrorsely curved trichomes, glabrescent when mature. Axillary buds, spherical, to 1mm diameter, perulate, perules 2, imbricate, chartaceous, scarious, glabrous. Stipules to 3mm long, triangular-lanceolate, apex acute, midrib prominent, scarious toward margins, glabrous. Petioles slender, (3-)4-6cm long, glabrous. Leaf blades (4-)5-7.5× (3.5-)4-6.5cm, broadly 45°E 15°S 20° 25° 300 km 50° Acalypha spachiana fig. 67. — Distribution map of Acalypha spachiana Baill. in Madagascar.
479 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) ovate to subtriangular, subchartaceous; base rounded to truncate; apex acuminate, acumen 0.5-1mm long, acute; margin crenate to serrate, teeth rounded to subacute; upper surface subglabrous, with some sparse, simple, erect trichomes; lower surface subglabrous, with some simple trichomes on veins; axils of secondary veins with hair-tuft domatia; margin ciliate in sinuses (except on older leaves); venation actinodromous, basal veins 3, secondary veins 3-5 per side. Stipels absent. Inflorescences spiciform, androgynous and male, axillary, sometimes bearing erinaceous galls. Androgynous inflorescences to 7cm long, mostly male with 1 female bract; sessile. Female segment: bract 1, sessile, enlarging in fruit to 3× 4mm, reniform, glabrous; margin subentire, with minute blackish papillae; bracteoles absent. Male segment persistent, to 6.5cm long; flowers glomerate; bracts to 0.5mm long, ovate-lanceolate, sparsely hairy with some simple trichomes at apex. Male inflorescences to 7cm long, filiform; sessile; rachis thin, glabrous; bracts like those on androgynous inflorescences. Male flowers: pedicel to 0.5mm long, glabrous; buds to 0.5mm diameter, glabrous. Female flowers 1 per bract, sessile; sepals 3, to 1mm long, triangular-lanceolate, ciliate; ovary c.1.5mm diameter, 3-lobed, echinate with linear projections to 2mm, hispidulous and ending in simple trichome(s) or gland, surface hispidulous; styles 3, to 5mm long, distinct or slightly connate at base, sparsely hairy with simple, short trichomes, each divided into 5-7 slender segments. Allomorphic flowers not seen. Capsules to 3mm diameter, erinaceous, projections linear, to 3.5mm long, acute, hispidulous, ending in trichome(s) or gland, surface sparsely hispidulous. Seeds c.1.3× 1mm, pyriform, smooth. 47. Acalypha urophylla Boivin ex Baill. Adansonia, recueil d’observations botaniques 1: 273 (Baillon 1861).— Acalypha reticulata var. urophylla (Boivin ex Baill.) Müll.Arg. Linnaea 34: 32 (Müller Argoviensis 1865).— Acalypha filiformis var. urophylla (Boivin ex Baill.) Govaerts, World Checklist and Bibliography of Euphorbiaceae (and Pandaceae) 63 (Govaerts etal. 2000).— Type: Madagascar. Prov. Antsiranana, Nossibé, plateau de Hell-Ville, VI.1847, L. H. Boivin 2178 (lecto-, designated by Montero Muñoz etal. [2018a: 110]: P[P00536752]; isolecto-, G[G00034246], GDC[G00324519], P[P00536751, P00536753]).— Former syntypes: Madagascar. s.l., s.d., L. M. A. Du Petit-Thouars s.n. (P[P00536748]); Prov. Antsiranana, Nossibé, XII.1840, A. Pervillé 364 (P[P00536749]); M. Richard 385 (P[P00536750]). Acalypha goudotiana Baill., Adansonia, recueil d’observations botaniques 1: 268 (Baillon 1861).— Acalypha reticulata var. goudotiana (Baill.) Müll.Arg., Linnaea 34: 32 (Müller Argoviensis 1865).— Acalypha filiformis var. goudotiana (Baill.) Govaerts, World Checklist and Bibliography of Euphorbiaceae (and Pandaceae) 62 (Govaerts etal. 2000).— Type: Madagascar. s.l., 1830, J. P. Goudot s.n. (holo-, G; iso-, P[P00536727] fragment). Acalypha meiodonta Baill., Bulletin mensuel de la Société linnéenne de Paris 2: 1197 (Baillon 1895b).— Acalypha reticulata var. meiodonta (Baill.) Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 267 (Leandri 1942).— Type: Madagascar. “Centr. Madag.”, XII.1883, R. Baron 2826 (lecto-, designated by Montero Muñoz etal. [2018a: 106]: P[P05604378]; isolecto-, K[K000186508]).— Former syntype: Madagascar. “Centr. Madag.”, s.d., R. Baron 6581 (K[n.v.]), syn. nov. Acalypha reticulata var. urophylla f. meeusei Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 264 (Leandri 1942).— Type: Madagascar. Prov. Toamasina, Analamazaotra forest, 1912, H. Perrier de la Bâthie 9741 (lecto-, designated by Montero Muñoz etal. [2018a: 111]: P05604377).— Former syntypes: Madagascar. Prov. Fianarantsoa, base Est du Pic d’Ivohibe, 19.IX.1926, R. Decary 5352 (P[P00224690]); Prov. Toamasina, Analamazaotra, 1000m, 10.XI.1938, H. J. Lam& A. D. J. Meeuse 5290 (L[L0242106]); Moramanga, 900m, 11.XI.1938, H. J. Lam& A. D. J. Meeuse 5363 (L[L0242105]); Soanierana, Andasibé, 9.XII.1938, H. J. Lam& A. D. J. Meeuse 5814 (WAG[WAG0133229], L[L0242110]). Acalypha reticulata var. urophylla f. longa Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 265 (Leandri 1942).— Type: Madagascar. Prov. Toliara, bassin de la Manampanihy, Col de Fitana, 700m, 15.X.1928, H. Humbert 6015 (lecto-, designated by Montero Muñoz etal. [2018a: 111]: P[P05604383]).— Former syntypes: Madagascar. Prov. Toliara, Col d’Ivolo (District de Fort-Dauphin), 500m, 5.IX.1932, R. Decary 10558 (P[P05604384]); Fort-Dauphin, col de Tsitongabarika, 600m, 9.IX.1932, R. Decary 10595 (P[P05604399]). Acalypha reticulata var. urophylla f. vohitrae Leandri, Notulae Sys - tematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 265 (Leandri 1942).— Type: Madagascar. Prov. Toamasina, Andevorante, rive droite de la Vohitra près de Lohariandava, 45°E 15°S 20° 25° 300 km 50° Acalypha tremula fig. 68 . — Distribution map of Acalypha tremula I.Montero & Cardiel in Madagascar.
480 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. AB CD EF GH fig. 69. — A, Acalypha nusbaumeri I.Montero & Cardiel (photo by L. Nusbaumer, L. Nusbaumer LN1169); B, A. pervilleana Baill. (photo by L. Gillespie, L. Gillespie 10649); C, A. perrieri Leandri (photo by L. Gillespie, L. Gillespie 10835); D, A. radula Baker (photo by L. Gillespie, L. Gillespie 10758); E, A. rottleroides Baill. (photo by L. Nusbaumer, L. Nusbaumer 1933); F, A. spachiana Baill. (photo by L. Gillespie, L. Gillespie 10856); G, A. tremula I.Montero & Cardiel (photo by R. Randrianaivo, R. Randrianaivo 1160); H, A. urophylla Boivin ex Baill. (photo by L. Nusbaumer, L. Nusbaumer LN1663).
481 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) 200-250m, 10.X.1912, R. Viguier& H. Humbert 661 (lecto-, designated by Montero Muñoz etal. [2018a: 111]: P[P05604437]; isolecto-, P[P05604440, P05604441]). Acalypha reticulata var. urophylla f. glabrescens Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 266 (Leandri 1942).— Type: Madagascar. Prov. Antsiranana, Massif du Tsaratanana, 2200m, H. Perrier de la Bâthie 16180 (lecto-, designated by Montero Muñoz etal. [2018a: 111]: P[P05604386]).— Former syntypes: Madagascar. Prov. Antsiranana, Sambirano, XI-XII.1937, H. Humbert 18659 (P[P05604397, P05604400]); Massif du Tsaratanana, 1600m, H. Perrier de la Bâthie 15371 (P[P05604385]); IX.1912, H. Perrier de la Bâthie 18614 (P[P05604382]); Prov. Fianarantsoa, Ranohira, Isalo, 30.VII.1928, H. Humbert 5014 (P[P05604401, P05604402]); Prov. Mahajanga, Beritsoka, H. Perrier de la Bâthie 413 (P[P05604390, P05604392]); Beritsoka, XII.1897, H. Perrier de la Bâthie 422 (P[P05604387, P05604388, P05604391]); Beritsoka, H. Perrier de la Bâthie 9822 (P[P05604389]); without exact location: s.l., R. Baron 5987 (P[P05604407]); s.l., L. Humblot 335 (P[P05604393, P05604394, P05604395, P05604396]). Acalypha reticulata var. meiodonta f. andronea Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 268 (Leandri 1942).— Type: Madagascar. Prov. Mahajanga, Bemarivo (Boïna), XII.1906, H. Perrier de la Bâthie 9561 (lecto-, designated by Montero Muñoz etal. [2018a: 106]: P[P00513143]; isolecto-, P[P00513144]).— Former syntypes: Madagascar. Prov. Mahajanga, Manongarivo (Ambongo), X.1904, H. Perrier de la Bâthie 1677 (P[P00513146, P00513147]); bord du massif du Manongarivo, versant du Sambirano, 1909, H. Perrier de la Bâthie 9934 (P[P00513149]); haut Bemarivo (Andranofosy), Boïna, I.1907, H. Perrier de la Bâthie 9635 (P[P00513145]); Massif du Manongarivo, Sambirano, IV.1909, H. Perrier de la Bâthie 9928 (P[P00513142]); Massif du Manongarivo, versant du Sambirano, IX.1909, H. Perrier de la Bâthie 9939 (P[P00513150]), syn.nov. Acalypha reticulata var. urophylla f. typique Leandri, Notulae Systematicae. Herbier du Muséum de Paris. Phanérogramie. Paris 10: 262 (Leandri 1942) nom. inval. iconography.— Baillon (1891: t. 189); Figs 69H; 73C-F. eTymology.— The epithet is taken from the Greek words ‘uro’, tail, and ‘phyllus’, leaf. It probably refers to the acuminate apex of the leaf blades. DisTribuTion anD habiTaT.— Endemic to Madagascar (Diana, Sava, Sofia, Analanjirofo, Boeny, Betsiboka, Alaotra-Mangoro, Melaky, Bongolava, Analamanga, Vakinankaratra, Amoron’i Mania, Haute Matsiatra, Vatovavy-Fitovinany, Atsimo-Atsinanana, Ihorombe, Menabe, Atsimo-Andrefana, Androy and Anosy). Medium altitude moist evergreen forest, lowland altitude moist evergreen forest, moist semi-deciduous forest, sclerophyllous woodland, dry deciduous forest, and riparian forest. On basement rocks, sandstone, and unconsolidated sand. Altitudinal range 100-2000 (-2280) m (Fig. 70). p reliminary conservaTion assessmenT .— The EOO of Acalypha urophylla is estimated to be 552 176km2 and its AOO 812km2, which is less than the 2 000km2 threshold of the B2 subcriterion of the Vulnerable category. However, A.urophylla has a large EOO, a wide elevational range, and occurs in many different ecosystems, so it is likely that its AOO actually exceeds the threshold of any threat category. Therefore, we assess A.urophylla as Least Concern (LC). maTerial examineD.— 361 collections. Madagascar. Académie Malgache s.n. (P[P00508499]); Adamainty, J.F. 3 (G, MAUAM, MO); Afzelius, K.s.n. (P[P00508498]); Almeda, F. 7914 (CAS[CAS-0064833], G, MO[MO-2965739]), 8100 (CAS[CAS-0064830]); Ammann, M.Y. MYA 026 (G[G00180173]), MYA 106 (G, K, P[P04786263]), MYA 322 (G, P[P04786264]), MYA 410 (G[G00180542]); Andriamihajarivo, T. 1979 (MO[MO-3037278], P[P01200027]); Andrianjafy, N.M. 337 (K, MO[MO-2966179], P[P05604406]), 494 (P[P05604423]), 668 (P[P05604373]), 1466 (MO[MO-2965754], TAN); Anonymous 1677 (P[P05604376]); Anonymous 169820 (P[P04804779]); Anonymous 2727 (P[P04804528]); Anonymous 3774 (P[P04804536]); Anonymous 5756 (P[P04804537]); Anonymous 9891 (P[P04804512]); Anonymous s.n. (P[P04804507]); Anonymous s.n. (P[P04804534]); Antilahimena, P. 307 (P[P04779788]), 3220 (G[G00420842], MO[MO-2965863], TAN), 4195 (K, MO[MO-2965842], P[P05572501]), 4271 (MO[MO-3025014], P[P05481539]), 4905 (G, MO[MO-2966260], P[P05481544]), 4990 (G, MO[MO-3025017], P[P05481457]), 5260 (MO[MO-3025013], P[P05481541]), 5378 (MO[MO-3025015], P[P01152830], TAN), 8161 (MAUAM, MO[MO-2965746], P[P01152832], TAN); Baron, R. 2309 (P[P04779832]), 2826 (P[P05604378]), 2874 (P[P04804725]), 3048 (K), 3086 (K), 3107 (P[P04804722]), 3701 (K), 3786 (K), 3874 (K), 4066 (K), 4403 (P[P04779783]), 4619 (P[P04779780]), 4793 (P[P04804728]), 4848 (K), 4991 (K), 4992 (K), 4993 (K), 4994 (K), 5560 (P[P04804726]), 5581 (P[P05604375]), 5671 (P[P04804729]), 5823 (P[P04804730]), 5987 (P[P05604407]), s.n. (P[P04804727]); Benoist, M.R. 1135 (P[P04804720]), 1484 (G, K, P[P04804719]), 1512 (MA[MA-01-00730257], P[P04804724]), 1519 (P[P04804723]), s.n. (P[P04779781]); Bernard, R. 1295 (G, P[P04786265]); Bernardi, L. 11139 (G[G00034253], L[L0242525], P[P04779803]), 11825 (G[G60429]), s.n. (P[P04779775]); Birkinshaw, C.R. 521 (P[P00513131]), 759 (BR[BR0000009679566]), 872 (G, ILL, MAUAM, MO[MO-2967237], P[P00513132], TAN), 1464 (P[P05481883]); Boivin, L.H. 52 (P[P00513160]), 2178 (G[G00324519, G00034246], P[P04804721, P00536751, P00536752, P00536753]), s.n. (P[P00324574]), s.n. (P[P04804508]), s.n. (P[P04804511]), s.n. (K[K000186501]), s.n. (K[K000186502]); Bosser, M.J. 1201 (P[P04779808]), 2191 (P[P04804716]), 2374 (P[P00508489]), 2690 (P[P00508490]), 5952 (P[P04804715]), 6985 (P[P04804718]), 8508 (P[P00508491, P00508492]), 16709 (P[P04804717]), 17510 (P[P00508493, P00508494]), 18256 (P[P00508495]); Buerki, S. 83 (G, MO[MO-2966262], P[P05604433], TAN); Burivalova, Z. ZB 021 (G[G00304003]); Casimir, Z. 597 (P[P00872737]); Catat, L.D.M. 459 (P[P04804712]); Ceplitis, A. AA957 (UPS[UPS V-095281]); Cloisel, J. 136 (P[P00513165]); Cours, G. 810 (P[P04804711]), 928 (P[P04804714]), 1007 (G, P[P04804710]), 1606 (B[B100138967], G, MA[MA-01-00730255], P[P04804713]), 2074 (G, P[P00508486]), 2078 (B[B100120929], MA[MA-01-00730258], P[P00508488]), 3240 (B, MA[MA-01-00730259], P[P00508487], WAG[WAG.1930071]); Croat, T.B. 29479 (P[P00508483]), 29487 (P[P00508485]), 30146 (P[P00508481]), 30697 (P[P00508482]), 32280 (P[P00508484]), 32284 (WAG[WAG.1578691]); Cynthia, F. 177 (P[P05510094]); d’Alleizette, Ch. 846 (P[P00513161]), s.n. (L[L0242102]), s.n. (L[L0242179]); Decary, R. 90 (C), 1306 (P[P00513159]), 1317 (P[P00513158]), 1334 (P[P00513157]), 1524 f (G[G00420845]), 1966 (P[P04804515]), 5069 (B[B100273583], P[P00508480]), 5352 (P[P00224690]), 5431 (P[P04804498]), 5711 (C, G, P[P00508479]), 6969 (P[P00508478]), 7057 (K, P[P00508477], US[US01287452]), 7113 (P[P00508476, P04786260], S[S13-13803]), 7232 (P[P00508475]), 7418 (P[P00508474]), 7567 (P[P00508473]), 7956 (P[P00508472]), 8134 (B[B100120931], P[P00508471]), 9432 (P[P04804538]), 10558 (P[P05604384]), 10595 (P[P05604399]), 13772 (P[P00508470]), 14692 (K, P[P00508469]), 14865 (P[P00508468]), 15186 (K, P[P00508467]), 15247 (P[P00508465], WAG[WAG.1930072]), 15596 (P[P04779778]), 17457 (G, K, P[P00508464]), 17508 (B[B100346702], BR[BR0000005095049], P[P00513156]), 17730 (BR[BR0000009311879], G, P[P00508463]), 17821 (BR[BR0000009311978], P[P00508462]), 17829 (P[P00508461]), 18246 (G, K, P[P00508460]), 18406
488 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. Acalypha venosa Poir. Encyclopédie méthodique. Botanique 6: 204 (Poiret 1804).— Type: Commerson s.n. remarKs The type specimen of Acalypha venosa (Commerson s.n., from Madagascar) corresponds to Leptonema venosum (Poir.) A.Juss. (Phyllanthaceae), as pointed out by Steudel (1841: 31). Acknowledgements This research received support from the SYNTHESYS Project (http://www.synthesys.info/; FR-TAF 6307, DE-TAF 3319, and SE-TAF 5590) financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program, the European Molecular Biology Organization (https://www. embo.org; STF-8222), and the United States National Science Foundation (DEB-0128872). We kindly thank the curators and staff of all the herbaria mentioned for facilitating the study of their specimens. 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493 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) appenDix 1. — List of the species and specimens of Acalypha included in the micromorphological studies. Taxon Country Specimens (Herbarium) A. andringitrensis Leandri Madagascar J. Perrier de la Bâthie 9671 (P) A. ankaranensis I.Montero & Cardiel Madagascar A. Leeuwenberg 14374 (G) A. baretiae I.Montero & Cardiel Madagascar R. Razakamalala 6064 (MO) A. chibomboa Baill. Comoros F. Barthelat 1436 (P) A. claoxyloides Hutch. Seychelles F.R. Fosberg 49627 (MO) A. crateriana (Coode) I.Montero & Cardiel, comb. nov. Mauritius D. Lorence 1858 (K) A. decaryana Leandri Madagascar P.J. Rakotomalaza 211 (P) A. diminuta Baill. Madagascar H. Grevé 223 (P), L.P.G. Nusbaumer LN928 (G), H. Ralimanana 239 (G) A. emirnensis Baill. Madagascar H. Humbert 6514 (GH), S. Malcomber 2118 (P), R. Rakoto 15 (G), L. Gautier LG 3889 (G), C. Rakotovao 2677 (P) A. filiformis Poir. Mauritius M.J.E. Coode 5062 (P) La Réunion Th. Cadet 899 (P) A. fimbriata Schumach. & Thonn. Ghana C.C.H. Jongkind 2277 (P) A. hispida Burm. f. Colombia H. García-Barriga 6477 (COL) Ecuador C.H. Dodson 11285 (SEL) United States J.R. Abbott 10611 (SEL) A. indica L. French Guiana R.M. Harley 24801 (U) Russian Federation V. Vasak s.n. (W) A. lamiana (Leandri) I.Montero & Cardiel Madagascar H.J. Lam 6127 (P) A. leandrii I.Montero & Cardiel Madagascar J. P. Leandri 2996 (P) A. leptomyura Baill. Madagascar P. Morat 4736 (P) A. linearifolia Leandri Madagascar G. Cours 4638 (P) A. marginata (Poir.) Spreng. La Réunion P. Commerson s.n. (G) A. mayottensis I.Montero & Cardiel Mayotte J.-N. Labat 3268 (G), J.-N. Labat 3272 (G), G. Viscardi 310 (P) A. menavody (Leandri) I.Montero & Cardiel Madagascar R. Ranaivojaona 1659 (MO) A. nusbaumeri I.Montero & Cardiel Madagascar L. Nusbaumer LN 1169 (P) A. perrieri Leandri Madagascar L. Gautier LG 5615 (G) A. perrieri aff. Leandri Madagascar P. De Block 1082 (G) A. pervilleana Baill. Madagascar L. Razafitsalama 1063 (G), A. Mocquerys 159 (G) A. poiretii Spreng. Argentina C.A. O’Donell 890 (GH) A. rabesahalana I.Montero & Cardiel Madagascar H. Humbert 32480 (G) A. radula Baker Madagascar Service Forestier Madagascar 45-SF (P), Service Forestier Madagascar s.n. (P), J.M. Hildebrandt 3865 (BREM) A. richardiana Baill. Comoros J.M. Hildebrandt 1661 (BREM) A. rottleroides Baill. Madagascar P. Ranirison PR 1085 (G) A. indet. 1 Madagascar A. Mocquerys 405 (G) A. indet. 2 Madagascar F. Ratovoson 1347 (G) A. tremula I.Montero & Cardiel Madagascar L. Nusbaumer LN 1227 (G) A. urophylla Boivin ex Baill. Madagascar S. Malcomber 1954 (G), J.M. Hildebrandt 2914 (BREM) A. vulneraria Baill. Madagascar F. Randriatafika 812 (MO) A. wilkesiana Müll.Arg. Peru J. Schunke 1941 (COL) United States T. Walters 2694 (SEL)
494 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. appenDix 2. — List of Acalypha names cited in the Western Indian Ocean Region (WIOR). Accepted names in bold letters. Published names Accepted names A. acuminata Vahl ex Baill. nom. illeg. A. burmanii I.Montero & Cardiel A. aldabrica Pax & K.Hoffm. A. claoxyloides Hutch. A. amentacea Roxb. subsp. wilkesiana (Müll.Arg.) Fosberg A. wilkesiana Müll.Arg. A. andringitrensis Leandri A. ankaranensis I.Montero & Cardiel A. arborea Comm. in Poir. nom. nud. A. filiformis Poir. A. aspretorum Leandri nom. nud. A. medibracteata Radcl.-Sm. & Govaerts A. bailloniana Müll.Arg. A. bakeriana Baill. A. emirnensis Baill. A. baretiae I.Montero & Cardiel A. baronii Baker A. emirnensis Baill. A. boinensis Leandri A. berryi I.Montero & Cardiel, sp. nov. A. buchenavii Müll.Arg. A. spachiana Baill. A. burmanii I.Montero & Cardiel A. cardielii I.Montero & G.A.Levin A. chibomboa Baill. A. claoxyloides Hutch. A. cloiselana M. Denis in Leandri nom. nud. A. gracilipes Baill. A. codonocalyx Baill. A. chibomboa Baill. A. colorata (Poir.) Spreng. A. integrifolia Willd. A. commersoniana Baill. nom. nud. A. integrifolia Willd. A. commersoniana Baill. ex Müll.Arg. A. integrifolia Willd. A. commersoniana var. acutifolia Müll.Arg. A. integrifolia Willd. A. commersoniana var. brevifolia Baill. ex Müll.Arg. A. integrifolia Willd. A. commersoniana var. concolor Baill. nom. nud. A. integrifolia Willd. A. commersoniana var. discolor Baill. nom. nud. A. integrifolia Willd. A. commersoniana var. gracilipes (Baill.) Müll.Arg. A. gracilipes Baill. A. commersoniana var. longifolia Müll.Arg. A. integrifolia Willd. A. commersoniana var. obtusifolia Müll.Arg. A. integrifolia Willd. A. commersoniana var. parvifolia Baill. nom. nud. A. integrifolia Willd. A. commersoniana var. parvifolia Baill. ex Müll.Arg. A. integrifolia Willd. A. commersoniana f. colorata (Poir.) Müll.Arg. A. integrifolia Willd. A. commersoniana f. concolor Müll.Arg. A. integrifolia Willd. A. commersoniana f. discolor Müll.Arg. A. integrifolia Willd. A. commersoniana f. purpurea Müll.Arg. A. integrifolia Willd. A. commersoniana f. purpureo-marginata Müll.Arg. A. integrifolia Willd. A. commersoniana f. unicolor Müll.Arg. A. integrifolia Willd. A. commersonii Baill. nom. nud. A. gracilipes Baill. A. comorensis Pax A. chibomboa Baill. A. crateriana (Coode) I.Montero & Cardiel, comb. nov. A. crenata var. glandulosa Müll.Arg. A. lanceolata var. glandulosa (Müll.Arg.) Radcl.-Sm. A. decaryana Leandri A. diminuta Baill. A. discolor Bojer nom. nud. A. integrifolia Willd. A. emirnensis Baill. A. emirnensis var. bara Leandri A. emirnensis Baill. A. emirnensis var. jabohaziana Leandri A. emirnensis Baill. A. fasciculata Müll.Arg., syn. nov. A. emirnensis Baill. A. fasciculata var. humbertiana Leandri, syn. nov. A. emirnensis Baill. A. fasciculata var. lyallii (Baker) Leandri, syn. nov. A. emirnensis Baill. A. filiformis Poir. A. filiformis var. arborea Poir. A. filiformis Poir. A. filiformis var. goudotiana (Baill.) Govaerts A. urophylla Boivin ex Baill. A. filiformis var. ovalifolia (Baill.) Govaerts A. ovalifolia Baill. A. filiformis var. pervilleana (Baill.) Govaerts A. pervilleana Baill. A. filiformis var. urophylla (Boivin ex Baill.) Govaerts A. urophylla Boivin ex Baill. A. filiformis var. urophylloides (Pax & K.Hoffm.) Govaerts A. pervilleana Baill. A. fimbriata Schumach. & Thonn. A. fryeri Hutch. A. claoxyloides Hutch. A. gagnepainii Leandri nom. illeg.A. medibracteata Radcl.-Sm. & Govaerts A. gagnepainii var. calcicola Leandri A. medibracteata Radcl.-Sm. & Govaerts A. gillespieae G.A.Levin & I.Montero A. goudotiana Baill. A. urophylla Boivin ex Baill. A. gracilipes Baill. A. hildebrandtii Baill. A. radula Baker A. hispida Burm.f. A. hologyna Baker A. emirnensis Baill. A. humbertii Leandri
495 Monograph of Acalypha L. (Euphorbiaceae) of the Western Indian Ocean Region ADANSONIA, sér. 3 • 2023 • 45 (26) Published names Accepted names A. humblotiana Baill. A. pervilleana Baill. A. indica L. A. indica var. bailloniana (Müll.Arg.) Hutch. A. bailloniana Müll.Arg. A. integrifolia Willd. A. integrifolia subsp. marginata (Poir.) Coode A. marginata (Poir.) Spreng. A. integrifolia subsp. panduriformis Coode A. marginata (Poir.) Spreng. A. integrifolia var. colorata (Poir.) Pax & K.Hoffm. A. integrifolia Willd. A. integrifolia var. concolor (Müll.Arg.) Pax & K.Hoffm. A. integrifolia Willd. A. integrifolia var. crateriana Coode A. crateriana (Coode) I.Montero & Cardiel, comb. nov. A. integrifolia var. gracilipes (Baill.) Pax & K.Hoffm. A. gracilipes Baill. A. integrifolia var. longifolia (Müll.Arg.) Coode A. integrifolia Willd. A. integrifolia var. parvifolia (Baill. ex Müll.Arg.) Pax & K.Hoffm. A. integrifolia Willd. A. integrifolia var. saltuum Coode A. marginata (Poir.) Spreng. A. isaloensis I.Montero & Cardiel A. juliflora Pax A. rottleroides Baill. A. lamiana (Leandri) I.Montero & Cardiel A. lanceolata var. glandulosa (Müll.Arg.) Radcl.-Sm. A. lantanaefolia Bojer nom. nud. A. filiformis Poir. A. leandrii I.Montero & Cardiel A. leonii Baill. A. leonii var. perrierana Leandri A. leonii Baill. A. lepidopagensis Leandri A. leptomyura Baill. A. levinii I.Montero & Cardiel A. linearifolia Leandri A. lyallii Baker A. emirnensis Baill. A. madagascariensis Pax & K.Hoffm., syn. nov. A. emirnensis Baill. A. madreporica Baill. A. emirnensis Baill. A. magistri I.Montero & Cardiel A. marginata (Poir.) Spreng. A. mayottensis I.Montero & Cardiel A. medibracteata Radcl.-Sm. & Govaerts A. medibracteata var. calcicola (Leandri) Radcl.-Sm. & Govaerts A. medibracteata Radcl.-Sm. & Govaerts A. meiodonta Baill., syn. nov. A. pervilleana Baill. A. menabeana Leandri nom. nud. A. medibracteata Radcl.-Sm. & Govaerts A. menavody (Leandri) I.Montero & Cardiel A. neptunica Müll.Arg. A. nusbaumeri I.Montero & Cardiel A. ovalifolia Baill. A. richardiana Baill. A. paxii Aug.DC., syn. nov. A. pervilleana Baill. A. perrieri Leandri A. pervilleana Baill. A. poiretii Spreng. A. polynema Baill. A. rottleroides Baill. A. rabesahalana I.Montero & Cardiel A. radula Baker A. reticulata (Poir.) Müll.Arg. A. filiformis Poir. A. reticulata var. arborea (Poir.) Müll.Arg. A. filiformis Poir. A. reticulata var. cloiselana Leandri A. gracilipes Baill. A. reticulata var. goudotiana (Baill.) Müll.Arg. A. urophylla Boivin ex Baill. A. reticulata var. longifolia Müll.Arg. A. integrifolia Willd. A. reticulata var. meiodonta (Baill.) Leandri, syn. nov. A. pervilleana Baill. A. reticulata var. ovalifolia (Baill.) Müll.Arg. A. richardiana Baill. A. reticulata var. pervilleana (Baill.) Müll.Arg. A. pervilleana Baill. A. reticulata var. urophylla (Boivin ex Baill.) Müll.Arg. A. urophylla Boivin ex Baill. A. reticulata var. urophylloides Pax & K.Hoffm. A. pervilleana Baill. A. reticulata f. aberrans Müll.Arg. A. integrifolia Willd. A. reticulata f. andronea Leandri A. pervilleana Baill. A. reticulata f. glabrescens Leandri A. urophylla Boivin ex Baill. A. reticulata f. humblotiana (Baill.) Leandri A. pervilleana Baill. A. reticulata f. lamiana Leandri A. lamiana (Leandri) I.Montero & Cardiel A. reticulata f. longa Leandri A. urophylla Boivin ex Baill. A. reticulata f. meeusei Leandri A. urophylla Boivin ex Baill. A. reticulata f. typique Leandri nom. inval. A. urophylla Boivin ex Baill. A. reticulata f. vohitrae Leandri A. urophylla Boivin ex Baill. A. richardiana Baill. A. rottleroides Baill. A. salviifolia Baill. nom. nud. A. radula Baker Appendix 2. — Continuation.
496 ADANSONIA, sér. 3 • 2023 • 45 (26) Montero-Muñoz I. et al. Published names Accepted names A. spachiana Baill. A. spachiana var. acutifolia Baill. A. spachiana Baill. A. spachiana var. latifolia Baill. A. spachiana Baill. A. spachiana var. minor Baill. A. spachiana Baill. A. spiciflora Burm.f. Claoxylon spiciflorum (Burm.f.) A.Juss. A. spiciflora var. menavody Leandri A. menavody (Leandri) I.Montero & Cardiel A. squarrosa Pax A. spachiana Baill. A. tremula I.Montero & Cardiel A. urophylla Boivin ex Baill. A. urophylla Pax nom. illeg.A. pervilleana Baill. A. venosa Poir. Leptonema venosum (Poir.) A.Juss. A. vulneraria Baill. A. wilkesiana Müll.Arg. Caturus sessilis Thouars ex Baill. nom. nud. A. integrifolia Willd. Ricinocarpus baillonianus (Müll.Arg) Kuntze A. bailloniana Müll.Arg. Ricinocarpus hispidus (Brum.f.) Kuntze A. hispida Brum.f. Ricinocarpus poiretii (Spreng.) Kuntze A. poiretii Spreng. Ricinocarpus wilkesianus (Müll.Arg.) Fosberg A. wilkesiana Müll.Arg. Tragia colorata Poir. A. integrifolia Willd. Tragia filiformis Poir. A. burmanii I.Montero & Cardiel Tragia fruticosa Commers. in Baill. nom. nud. A. integrifolia Willd. Tragia lobata Wall. nom. nud. A. integrifolia Willd. Tragia macrophylla Wall. nom. nud. A. integrifolia Willd. Tragia marginata Poir. A. integrifolia Willd. Tragia obtusata Vahl. in Baill. nom. nud. A. integrifolia Willd. Tragia reticulata Poir. A. filiformis Poir. Tragia salviifolia Bojer in Baill. nom. nud. A. radula Baker Tragia saxatilis Bojer in Pax & K.Hoffm. nom. nud. A. spachiana Baill. Appendix 2. — Continuation.