Acalypha L.

Genus Acalypha L. Species Plantarum: 1003 (Linnaeus 1753); Genera Plantarum, ed. 5: 436 (Linnaeus 1754). — Type: Acalypha virginica L. (lectotype designated bySmall 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 (Linnaeus 1767). — 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. 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 claus - seni 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 et al. (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 et al. 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 et al. 2018b, 2020a, 2020 b, 2022, and another one in this paper). 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 0.5 cm 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 5 mm long, usually more than 10 mm long), with more than 2 bracts, usually many, rarely 1 ............................................................................................................ 9 5. Leaf blades linear, c. 0.2 cm wide .................................................................................. A. linearifolia Leandri — Leaf blades not linear, 1-7(-8) cm wide ....................................................................................................... 6 6. Petioles (2-) 3-4.5 cm 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

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