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A Revision of the Genus Actephila (Phyllanthaceae) in the Malesian Region

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A Revision of the Genus Actephila (Phyllanthaceae) in the Malesian Region Blumea 62, 2017: 7–25 ISSN (Online) 2212-1676 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE https://doi.org/10.3767/000651917X694985 A revision of the genus Actephila (Phyllanthaceae) in the Malesian region M. Heijkoop1, P.C. van Welzen1,2 Key words Abstract The genus Actephila (Phyllanthaceae) is distributed from India in the north-west to Thailand and via Malesia to the Melanesian Islands and Australia in the south-east. Eleven species are recognized of which three Actephila are newly described, A. discoidea with long leaf apices and a fleshy nectar discs, A. emarginata with an emarginate Asia leaf base, and A. stipularis with large stipules. Further, A. javanica is synonymized with A. excelsa; and a former Malesia synonym or separate variety, A. excelsa var. acuminata, is recognized as distinct species, but had to be synonymized Phyllanthaceae under A. subsessilis. Actephila puberula, not present in Malesia, but often synonymized with A. excelsa, is kept revision separate because of the hairy ovary and different base of the leaf blades. taxonomy Published on 31 January 2017 INTRODUCTION century Pax & Hoffmann (1922, 1931) included Actephila in the subfamily Pyllanthoideae subtribe Amanoinae. The placement Actephila Blume consists of (sub)shrubs to small trees. Its changed again when Webster (1994, 2014) included the genus distribution ranges from India in the north-west to Thailand and in subfamily Phyllanthoideae tribe Wielandieae, a classification via Malesia to the Melanesian Islands and Australia in the south- followed by Radcliffe-Smith (2001). More recently, it has been east. Actephila is classified in family Phyllanthaceae (formerly proposed to recognize the subfamily Phyllanthoideae again at part of Euphorbiaceae s.l.), tribe Poranthereae (Forster 2005, the family level as the Phyllanthaceae (Chase et al. 2002, APG Hoffmann et al. 2006, Vorontsova & Hoffmann 2008). 2003). Molecular analysis of genera placed in Phyllanthaceae The genus was first described by Blume in 1826, but he did not revealed that Actephila is closely related with the genera An- mention any species; he only referred to a collecting locality in drachne L. and Leptopus Decne., and less so to Meineckia Java and additional collection data. Later Hasskarl (1844) in- Baill., Poranthera Rudge, and Zimmermannia Pax (Wurdack et cluded the genus in Savia Willd. and described Savia actephila al. 2004, Samuel et al. 2005). In the most recent classification Hassk. Actephila was reinstated by Miquel (1859) and he re- by Vorontsova & Hoffmann (2008) Actephila was placed in the named S. actephila to Actephila javanica Miq. This is the only tribe Poranthereae. species from the same location (Nusa Kambangan Island) Unfortunately most revisions are outdated. The last complete that was mentioned by Blume in his genus description. Miquel revision of Actephila was by Pax & Hoffmann (1922). The only (1859) also described A. bantamensis Miq. from W Java, here recent revisions of the genus were for Thailand (Chayamarit considered to be a synonym of A. javanica. 2005) and the Australian region, where the number of species The species are common in the rain forest understorey with increased from five to 14 (Forster 2005). We had similar results mostly small trees (up to 5 m) and a few species as higher trees with the revision of the Malesian species, new species are (up to 15 m). The genus is characterized by the presence of described and several names are synonymized. stiff, caducous, somewhat deltoid or triangular stipules at the The present revision of the Malesian species of Actephila shows base of the petioles; very small, straight, simple, caducous hairs that the variation within certain species is very high, especially on all parts; penninerved leaves; axillary, usually single, small in A. excelsa (including A. javanica var. javanica) and A. lindleyi unisexual flowers with 0–6 petals much smaller than the 5 or 6 (Steud.) Airy Shaw. Because of the high variability and small sepals; a circular disc surrounding the stamens or ovary, and differences between species, species delimitation is difficult, the woody fruits, with often a knobbly surface, which are septi- which explains why in the past many specimens were incorrectly cidally or loculicidally dehiscent and supported by a persistent identified or new species were not recognized. For example, and sometimes enlarging calyx. Typical for Phyllanthaceae is Actephila excelsa (Dalzell) Müll.Arg. and A. javanica represent the presence of two ovules per locule and solitary or fasciculate a complex species within Malesia. They are variously treated axillary flowers (partly after Radcliffe-Smith 2001, Vorontsova as separate species (e.g., Chayamarit 2005), as synonyms & Hoffmann 2008). (this treatment) or as separate varieties of a single species The taxonomic placement of the genus Actephila has changed (e.g., Airy Shaw 1972). several times. An initial classification of Euphorbiaceae s.l. by In this revision eleven species are recognized, of which six were Müller (1866) placed the genus Actephila in the tribe Phyl- already known as species and one as a variety. Actephila col- lantheae subtribe Savieae (Forster 2005). During the 20th linsae Hunter is excluded as it only occurs in Thailand, outside the Malesian region. Actephila mooreana Baill. (‘mooriana’) was 1 Naturalis Biodiversity Center, Botany, P.O. Box 9517, 2300 RA Leiden, The formerly synonymized with A. lindleyi, but is distinct. It occurs Netherlands; corresponding author e-mail: [email protected]. 2 Institute of Biology Leiden, Leiden University, P.O. Box 9505, 2300 RA exclusively in Australia, where A. lindleyi is in fact absent (a Leiden, The Netherlands. description of A. mooreana can be found under A. lindleyi in © 2017 Naturalis Biodiversity Center You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 8 Blumea – Volume 62 / 1, 2017 the revision by Forster 2005). Actephila puberula Kurz, usually The base of the leaf blade varies from attenuate (e.g., A. lindleyi) considered as a synonym or variety of A. excelsa, is regarded as to acute or somewhat obtuse (e.g., A. excelsa); A. emarginata a distinct species of the Andaman islands as the leaf blade base is unique with an emarginate leaf base (Fig. 4). The apex var- is rounder to even slightly emarginate and the ovary is densely ies from obtuse to acute (e.g., A. trichogyna), to acuminate to hairy instead of glabrous. Actephila excelsa var. acuminata Airy cuspidate (e.g., A. excelsa). Shaw appeared to be distinct from the rest of A. excelsa, but conspecific with the Vietnamese A. subsessilis Gagnep., the Inflorescences and flowers older and thus accepted name. Recently, A. alanbakeri Welzen Inflorescences are usually absent and the flowers generally & Ent (Van der Ent et al. 2016) was described and here three grow solitary or paired in leaf axils, sometimes in larger fas- species are newly recognized: A. discoidea Heijkoop & Welzen, cicles. Repeated flowering from the same axils (leafless after A. emarginata Heijkoop & Welzen, and A. stipularis Heijkoop the first season) results in thickened brachyblasts on the stem & Welzen. This revision was solely based on morphological or thick branches from which short racemes grow with new features. generations of flowers (e.g., A. lindleyi; Fig. 5). The aim of this study is to revise the Malesian species in the All species appear to be dioecious, because only either stami- genus Actephila, which will be part of the ongoing revision of nate or pistillate specimens were found on the same specimen. the Euphorbiaceae for the international Flora Malesiana Project All flowers are 5-merous with five sepals, petals and stamens (published work already visible via www.nationaalherbarium. when staminate. The sepals are often somewhat ovate to more nl/euphorbs). elliptic or obovate. Petals are very small, much smaller than the sepals; they are obovate and distinctly narrowed towards MORPHOLOGICALLY important CHARACTERS the base. The stamens are either free (e.g., A. excelsa) or connate at base Characters used for species recognition are indumentum, (e.g., A. lindleyi). Stamens are 2-thecate with the thecae dor- stipule shape and size, petiole length, size of the leaf blade, sally connected to the connective. The staminate flowers have shape of the leaf blade base and apex, pedicel length and a circular, somewhat lobed disc surrounding the stamens, which shape, flower structures, thickness of the nectar disc, structure is either thin (e.g., A. excelsa) or fleshy (e.g., A. trichogyna). In of the fruit wall surface, size and shape of the columella, and between the stamens a 3-branched pistillode is present except size of the calyx. in A. trichogyna, which shows a bundle of hairs instead. The pistillate flowers usually have longer pedicels than the Indumentum staminate flowers. A 3-locular ovary is present, with two ovules All species are subglabrous and possess small, sericeous or per locule. Three sessile stigmas are found directly on top of puberulous simple hairs only, which can be seen with high the ovary, each is apically bifid. The circular disc resembles magnification only. These hairs occur on all young plant parts that of the staminate flowers in shape. and are often early caducuous, which makes the plants seem- ingly glabrous with the exception of A.
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