A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar Wendy L. Applequist Abstract APPLEQUIST, W.L. (2018). A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar. Candollea 73: 27-48. In English, English and French abstracts. DOI: http://dx.doi.org/10.15553/c2018v731a4 Homalium sect. Odontolobus Warb. (Salicaceae) is endemic to Madagascar and formerly included three species. In this revision, nine species are recognized, three of them newly described (Homalium densispicatum Appleq., Homalium masoalense Appleq., Homalium ovatifolium Appleq.) and three transferred from Homalium sect. Blackwellia Benth. Homalium densispicatum has highly reduced flowers like those of Homalium moniliforme H. Perrier, but the leaves are larger and the inflorescences thicker, with flowers in elongated many-flowered glomerules. Homalium masoalense and Homalium ovatifolium are northern species that have frequently ovate leaves and long styles and filaments; the flowers of Homalium masoalense are larger, with longer sepals, while Homalium ovatifolium has often canescent inflorescences. Two new subspecies are described: Homalium moniliforme subsp. littorale Appleq. and Homalium planiflorum subsp. roseiflorum Appleq. A key for identification of the species of Homalium sect. Odontolobus is provided. Résumé APPLEQUIST, W.L. (2018). Révision de Homalium sect. Odontolobus (Salicaceae) endémique de Madagascar. Candollea 73: 27-48. En anglais, résumés anglais et français. DOI: http://dx.doi.org/10.15553/c2018v731a4 Homalium sect. Odontolobus Warb. (Salicaceae) est endémique de Madagascar et comprenait trois espèces. Dans cette révision, neuf espèces sont reconnues, dont trois sont nouvellement décrites (Homalium densispicatum Appleq., Homalium masoalense Appleq., Homalium ovatifolium Appleq.) et trois sont transférées de Homalium sect. Blackwellia Benth. Homalium densispicatum possède des fleurs très réduites comme Homalium moniliforme H. Perrier, mais ses feuilles sont plus grandes et ses inflorescences plus épaisses, avec des glomérules allongés à fleurs nombreuses. Homalium masoalense et Homalium ovatifolium sont des espèces du nord avec des feuilles généralement ovales et de styles et filets allongés. Les fleurs de Homalium masoalense sont plus grandes, avec des sépales plus longs, tandis que Homalium ovatifolium possède des inflorescences souvent canescentes. Deux sous-espèces nouvelles sont décrites: Homalium moniliforme subsp. littorale Appleq. et Homalium planiflorum subsp. roseiflorum Appleq. Une clé de détermination des espèces de Homalium sect. Odonotolobus est proposée. Keywords SALICACEAE – Homalium – Homalium sect. Odontolobus – Madagascar – New species – Taxonomy Address of the author: Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A. E-mail: [email protected] Submitted on October 30, 2017. Accepted on March 5, 2018. First published online on May 3, 2018. ISSN : 0373-2967 – Online ISSN : 2235-3658 – Candollea 73(1) : 27-48 (2018) © CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE 2018 28 – Homalium sect. Odontolobus (Salicaceae) in Madagascar Candollea 73, 2018 Introduction Material and methods Homalium Jacq. is a pantropical woody genus traditionally Herbarium specimens at P and MO (herbarium acronyms placed in Flacourtiaceae; along with a large portion of that according to Index herbariorum, 2018) were examined, as formerly recognized family, its proper affinities are now known were then-undistributed duplicates for exchange available at to be with the taxa placed in an expanded Salicaceae (Chase those herbaria and images available through JSTOR Global et al., 2002). The genus includes over 150 currently recognized Plants website [http://plants.jstor.org] of types held by other species (see Applequist, 2013). Characters these species have institutions. Most recent collections have duplicates at TAN or in common include hermaphroditic flowers with two perianth TEF, including all those derived from the Missouri Botanical whorls, semi-inferior ovaries, oppositipetalous stamens, single Garden’s Madagascar program. As these were not seen they are conspicuous large glands at the base of each sepal, and very not listed among material examined; however, the locations of small fruits with few maturing seeds; in other floral characters, many such duplicates are available through Tropicos (2018). they encompass an unusual degree of diversity. Species were defined according to a taxonomic species Madagascar is perhaps the primary center of diversity of concept (e.g., Grant, 1981), the most widely used and usually Homalium. Warburg (1894) published a complex classifica- most practical approach in plant taxonomy, with three mor- tion of the genus that recognized two subgenera and nine phological features differing among groups of populations sections. With some emendations and additions such as the considered sufficient to warrant recognition at the species level. publication of Homalium sect. Rhodonisa (Tul.) Sleumer, that Flowers undergo little expansion in fruit, but are usually cadu- classification has been mostly maintained since. The subgenera, cous at or before fruit maturity. Because fruits with mature defined by whether stamens were only one per petal (Homalium seeds are rarely seen, only flowering characters are used in subg. Blackwellia (Benth.) Warb.) or fasciculate (Homalium species delimitation. In descriptions of taxa, character states subg. Homalium), were implicitly known by early authors to or size ranges given in parentheses are uncommon. be artificial and were apparently accepted and maintained for Known distributions and habitat are described for each convenience. The present author (Applequist, 2016) has pro- species or subspecies. To save space, locality data are not posed to emend this classification to recognize ten sections and provided for all specimens seen of common taxa, and those omit recognition of subgenera, since the relationships among that are provided are edited for brevity, especially when full some sections are unclear. Of those ten sections, six are found label data are available from Tropicos (2018). “Fkt.” is used in Madagascar and five are endemic there. throughout as an abbreviation for “fokontany”. A complete One of these endemic Malagasy sections is Homalium sect. index of specimens seen is provided as an appendix. Maps Odontolobus Warb., which traditionally is characterized by a of georeferenced specimens may also be generated within reduction of the sepals to tiny deltoid teeth; the flowers are the Madagascar Catalogue (2018), which is continually usually sessile and quite small, in two species much reduced updated with new determinations and specimens. and borne in clusters, and the filaments and styles are unusu- A preliminary, unofficial assessment of conservation status ally short. Warburg (1894) initially included two species, using the categories and criteria of IUCN (2012) is provided H. parkeri Baker and H. lucidum Scott-Elliot, in this section. for each taxon recognized. In instances when the extent of Perrier de la Bâthie (1940) did not recognize Homalium occurrence (EOO) and area of occupancy (AOO) might affect sect. Odontolobus; he included these two species within the assessed status, GeoCAT (Bachman & Moat, 2012) was Homalium sect. Blackwellia Benth. and described another used to estimate those values. Geographic coordinates were species in that section, H. moniliforme H. Perrier, that was later taken from label data or from the Missouri Botanical Garden’s transferred to Homalium sect. Odontolobus by Sleumer (1973), gazetteer (Tropicos, 2018). author of the most recent treatment of the Malagasy species of Homalium. Recent study of existing herbarium material has Taxonomic treatment indicated that several additional, unrecognized taxa are assign- able to this section. Moreover, three species traditionally placed Homalium sect. Odontolobus Warb. in Engl. & Prantl, Nat. in Homalium sect. Blackwellia were more similar to members Pflanzenfam. III(6a): 35. 1894. of Homalium sect. Odontolobus in overall morphology, though Typus (designated by Sleumer, 1973: 306): Homalium lacking the extreme sepal reduction. A modernized treatment parkeri Baker. of Homalium sect. Odontolobus was thus needed and is herein presented. Trees (occasionally shrubby in H. parkeri). Stipules axillary, free. Inflorescences spicate to racemose (partly paniculate with racemoid branches in H. planiflorum subsp. roseiflorum) with most flowers borne in small clusters or glomerules (singly in H. brachystylis); bracts small, often broad, usually persistent; Candollea 73, 2018 Homalium sect. Odontolobus (Salicaceae) in Madagascar - 29 Key to the species of Homalium bracteoles caducous or persistent (rapidly caducous or pos- sect. Odontolobus sibly absent in H. masoalense), minute to broad and larger than bracts, usually thick-textured, in 2 species fleshy, approaching 1. Sepals reduced to small teeth, barely visible from above subterete at base. Leaves alternate (sometimes opposite or ........................................................................................2 subopposite), glabrous (occasionally glabrate with few tri- 1a. Sepals from nearly as long as petals to less than half as long, chomes on midrib or sparsely pubescent when young). Flowers sometimes quite small but readily visible from above ......5 sessile or short-pedicellate, pedicels not articulated; perianth 2. Inflorescences canescent; most flowers borne in elongated, 5-8-merous. Sepals