Search

Deparia chinensis (Ching) X.S.Guo & C.Du in Chen (2016: 18). ≡ Dryoathyrium chinense Ching in Jin (1988: 36). Lectotype (designated by Hsu and Kuo in Kuo et al. 2018):— CHINA. Chongqing [Szechuan]: Chengkou [Chen-kou Hsien], 2100 m, 17 October 1958, T. L.Tai 106702 (PE barcode 00049936 [photo!], isolectotypes HNWP barcode 0219360 [photo!], IBK barcode 00034139 [photo!], IBSC barcode 0676556 [photo!], NAS barcode 00082856 [photo!], PE barcode 00049937 [photo!], SZ barcodes 00149268 [photo!] & 00149269 [photo!], WUK barcode 0130652 [photo!]). – Dryoathyrium chinense Ching (1974: 214), not validly published, with two gatherings indicated as types, contrary to Art. 40.2. – Dryoathyrium chinense Ching in Hsieh (1985: 153), not validly published, with the identity of the type not explicitly indicated, contrary to Art. 40 Note 2. – Deparia chinensis (Ching) Z.R.Wang in He et al. (2013: 422), new combination not validly published, with incorrect reference to basionym, contrary to Art. 41.5. – Deparia chinensis (Ching) T.C.Hsu & L.Y.Kuo in Kuo et al. (2018: 651), isonym., Published as part of Zhang, Jian-Hang, Du, Cheng, Applequist, Wendy L. & Liao, Shuai, 2023, An overview on the valid publication and typification of Dryoathyrium chinense and Deparia chinensis (Athyriaceae, Aspleniineae, Polypodiales), pp. 67-69 in Phytotaxa 579 (1) on page 68, DOI: 10.11646/phytotaxa.579.1.8, http://zenodo.org/record/7532153, {"references":["Chen, Y. - S. (2016) The Checklist of Shaanxi Vascular Plants. Higher Education Press, Beijing, 525 pp.","Jin, S. - Y. (1988) Bibliography of new taxa (Cormophytes) and type specimens in the herbaria of China (I). Bulletin of Botanical Research, Harbin, Supplement: 1 - 117.","Kuo, L. - Y., Ebihara, A., Hsu, T. - C., Rouhan, G., Huang, Y. - M., Wang, C. - N., Chiou, W. - L. & Kato, M. (2018) Infrageneric Revision of the Fern Genus Deparia (Athyriaceae, Aspleniineae, Polypodiales). Systematic Botany 43: 645 - 655. https: // doi. org / 10.1600 / 036364418 X 697364","Ching, R. - C. (1974) Dryoathyrium Ching. In: Editorial Committee of Flora Tsinlingensis (Eds.) Flora Tsinlingensis 2. Science Press, Beijing, pp. 100 - 101 & 213 - 215.","Hsieh, Y. - T. (1985) The taxonomy of Dryoathyrium Ching in China. Bulletin of Botanical Research, Harbin 5 (3): 147 - 157. Available from: https: // bbr. nefu. edu. cn / CN / abstract / abstract 1781. shtml (accessed 12 January 2022)","He, Z. - R., Wang, Z. - R. & Kato, M. (2013) Deparia Hooker & Greville. In: Wu, Z. - Y., Raven, P. H. & Hong, D. - Y. (Eds.) Flora of China 2 - 3. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis, pp. 418 - 442. Available from: http: // www. efloras. org / florataxon. aspx? flora _ id = 2 & taxon _ id = 109640 (accessed 12 January 2022)"]}

Homalium sect. Rhodonisa (Tul.) Sleumer in Bull. Jard. Bot. Natl. Belg. 43: 300. 1973. ��� Nisa [unranked] Rhodonisa Tul., as ��� Rhodonisae ��� in Ann. Sci. Nat., Bot. s��r. 4, 8: 70. 1857. Typus (designated by SLEUMER, 1973: 300): Nisa sanguinea Boivin ex Tul. (��� Homalium sanguineum (Boivin ex Tul.) Baill.). Stipules axillary, free, usually rapidly caducous. Inflorescences axillary, spiciform to racemose or paniculate; bracts broad (sometimes rather short), bracts and bracteoles persistent. Flowers short-pedicellate, with pedicels articulated usually just below the receptacle, to subsessile or sessile; perianth 4���5(���6)-merous; sepals oblong, sometimes broadly or narrowly, to somewhat obovate, ovate, or elliptical (seldom to oblanceolate or lanceolate), slightly accrescent and usually curving over the fruit; calyx tube tubular with a narrowed base, often visibly ribbed or grooved, to funnelform, in fruit tubular to turbinate or ellipsoid; petals obovate to oblanceolate with rounded (to obtuse) apex, much longer than sepals, ascending (spreading), at least slightly accrescent (where known); sepals and petals ciliate, usually shortly, or not; sepal glands of moderate size, broadly elliptical to rounded (somewhat oblong or trapezoid), glabrous; stamens 1 per petal, inserted between glands; anthers varying from broadly oblong-elliptical with oblong-elliptical locules, the slits of dehiscence nearly parallel, to small, transversely elliptical with subglobose locules, widely separated lateral slits towards apex (sometimes apparently or nearly basifixed with connective not prominent); upper surface of ovary sometimes quite narrow, nearly flat to shallowly conical in flower, in fruit convex to conical (or little expanded); styles (3���)4���5, usually fused (or connivent) for at least half of length (sometimes only shortly). Locule of fruit narrowly cylindrical or ellipsoid (to narrowly obovoid), pubescent throughout; seeds 2���4 per fruit, small (possibly no mature seeds seen). Notes. ��� Adequate molecular data to elucidate relationships among the very diverse, probably paraphyletic lineages of Homalium do not exist. The relationships of species of sect. Rhodonisa would be of particular interest. The genus as now circumscribed has an apparent division between Homalium sect. Homalium and similar sections (e.g., in Madagascar, Eumyriantheia and Nisa) and sect. Blackwellia Benth. and related sections (primarily Polyanthera). Species that are typical of the former group have relatively few (4���7) broad sepals and petals; relatively large, broadly elliptical anthers; large sepal glands; and broadly funnelform calyx cups in flower. Sepals and petals are usually different in shape and may differ in size or accrescence. Typical species of sects. Blackwellia and Polyanthera have often more numerous, narrow sepals and petals, which usually are similar in shape; often small broad anthers with subglobose locules (though broadly elliptical anthers are also seen); and often narrowly funnelform to tubular, ridged calyx cups. The upper surface of the ovary is often prominently conical. Sect. Rhodonisa includes species that have relatively few, broad petals and sepals, like sects. Eumyriantheia and Nisa, but often elongated, narrow, ridged calyx cups, and sometimes also small ��� Blackwellia -like��� anthers with subglobose locules. These species are not easily classed as belonging to either group of sections, which is one reason for this author���s preference not to recognize subgenera (APPLEQUIST, 2016a). SLEUMER (1973) recognized two red-flowered species, H. sanguineum (Boivin ex Tul.) Baill. and H. rubriflorum Sleumer, and one white-flowered species, H. albiflorum, within sect. Rhodonisa. Red flowers are rather unusual in Homalium, though not unknown from other sections, and the two red-flowered species are also distinguished from the H. albiflorum complex in having long-branched paniculate inflorescences and flowers borne only 1 or 1���2 per node and bract (rather than in clusters of 2 or more). The taxonomy of the red-flowered species is not problematic, and no undescribed similar species were identified. However, the populations with clustered whitish (occasionally to yellowish or greenish) flowers, which SLEUMER (1973) subsumed within H. albiflorum, appear actually to represent no fewer than ten distinct taxa. These seem to fall roughly into two clusters. Homalium albiflorum s.str. (Fig. 1) and three new species, H. analavelonae Appleq., H. ihosyense Appleq., and H. phillipsonii Appleq., have usually five petals (varying to four or six); the outer surfaces of the sepals are sparsely pubescent or pilose, and the petals are at least sparsely pubescent (to pilose or glabrescent) with ciliate margins. Two previously described segregate species, H. leucophloeum (Tul.) Baill. and H. baillonii Scott-Elliot, and four new species, H. megaphyllum Appleq., H. pseudoracemosum Appleq., H. rakotovaoi Appleq., and H. vohitsiandrianense Appleq., have usually four petals (seldom to five in two species), glabrous or glabrate sepals, and glabrous petals without ciliate margins (except that in H. pseudoracemosum sepals are sometimes sparsely pubescent and petal margins are sometimes ciliate). These groups differ in ecological preferences, with the H. albiflorum group preferring arid to subhumid western and northern habitats and the H. leucophloeum group mostly in central and eastern (or extreme northern) forests that are humid or at least less arid (Fig. 2). An outlier is H. pseudoracemosum, a western species which is morphologically more similar to H. albiflorum than others in this group, and which may represent a transitional form between the two groups. Key to the species ofHomalium sect. Rhodonisa 1. Inflorescences usually panicles with well-developed branches (reduced to racemes); flowers borne singly or 1���2 per node; petals red............................................... 2 1a. Inflorescences racemes, spikes, or short-branched racemiform panicles; flowers mostly borne in clusters of at least 2���3 per node (very rarely, most borne singly); petals whitish (to yellow or pale green)................................. 3 2. Leaves elliptical to broadly (narrowly) elliptical, (4.5���)5���16 �� 2.1���5.5(���8) cm with petiole 4���8(���10) mm; flowers 1���2 per node, pedicellate; petals 2.4���4 mm, sparsely appressed-pubescent on abaxial surface mostly towards base; Alaotra-Mangoro, mid-elevation forests........................................................................... 10. H. rubriflorum 2a. Leaves broadly elliptical (elliptical, somewhat obovate), (5.6 ���)7.7 ��� 11.7 �� (3.8 ���) 4.2 ��� 8.2 cm with petiole (9 ���) 12 ��� 22 mm; flowers 1 per node, sessile; petals 4.4 ��� 6.3 mm, glabrous except for sometimes ciliate margins; Analanjirofo, low-elevation forests............................................................................. 11. H. sanguineum 3. Young twigs pilose; leaves pilose, especially on veins; secondary veins strongly prominent, sometimes with small domatia in vein axils............................ 7. H. phillipsonii 3a. Young twigs glabrous (rarely pubescent); leaves glabrous (rarely sparsely pubescent); secondary veins moderately to little prominent, almost always without (very rarely with) domatia....................................................................... 4 4. Inflorescences at least partly short-branched racemiform panicles........................................................................ 5 4a. Inflorescences racemose to spicate............................... 6 5. Leaves usually mostly lanceolate to ovate, to narrowly elliptical or elliptical; inflorescences consistently racemiform panicles; rachis glabrous or glabrate (branches may be sparsely pubescent); petals 4(5), 3���5.5 mm, glabrous except margins sometimes sparsely short-ciliate............................................................... 8. H. pseudoracemosum 5a. Leaves elliptical to broadly elliptical (ovate, perhaps very rarely lanceolate or narrowly elliptical); inflorescences usually mostly racemose; portions of rachis moderately to densely pubescent; petals 5(6), 2.6���8 mm, sparsely to moderately short-pubescent on both surfaces (rarely glabrescent)............................................... 1. H. albiflorum 6. Free portions of sepals sparsely pubescent or pilose; petals pubescent (or pilose), usually sparsely (rarely glabrescent), the margins ciliate....................................................... 7 6a. Free portions of sepals glabrous or glabrate except for sometimes ciliate margins; petals glabrous, the margins not ciliate (in H. leucophloeum rarely sparsely ciliate)... 9 7. Peduncles (0.6���) 2.5���6 cm; rachis at least partly pubescent; anthers transversely (to broadly) elliptical; widespread................................................................. 1. H. albiflorum 7a. Peduncles H. analavelonae 8a. Leaves elliptical (to somewhat obovate or broadly elliptical), (4.7���)6.7���14 �� (3���) 4���6.3 cm, thick-textured; racemes (3���) 4.5���11 cm with rachis moderately pilose to glabrate; pedicels 0.5���4.5 mm; sepals narrowly oblong, 1.8���3 mm, sparsely pilose outside, ciliate; petals 3.5���12 mm, strongly accrescent, sparsely pubescent to sparsely pilose; near Ihosy....................................................... 4. H. ihosyense 9. Leaves ovate to broadly ovate (to lanceolate, elliptical), with margins irregularly and shallowly crenulate to crenate or crenate-denticulate (somewhat revolute); pedicels (1���)2.5���6(���8.5) mm............................................ 10 9a. Leaves elliptical, broadly elliptical or oblong-elliptical (to narrowly elliptical, ovate) with margins subentire to shallowly repand or undulate; pedicels short, 0.5���1 mm, or flowers sessile............................................................ 11 10. Leaves ovate (lanceolate to elliptical), (5.5���)6.7 ��� 15.7 �� (2.8 ���) 3.5 ��� 6.5 cm with acuminate (acute, obtuse, or emarginate) apex; racemes (3 ���) 5 ���16 cm; sepal glands 0.4���0.6 �� 0.3���0.4 mm; northern Madagascar............................................................................. 9. H. rakotovaoi 10a. Leaves of type ovate to broadly ovate (aberrantly broadly elliptical), (4 ���)5 ��� 7.8 �� (2 ���) 2.6 ��� 4.8 cm with acute (rounded, emarginate) apex; racemes (1.7���) 3���4 cm; sepal glands 1.2���1.3 �� 0.8���0.9 mm; southeastern Madagascar (Vohitsiandriana).................. 12. H. vohitsiandrianense 11. Leaves elliptical to oblong-elliptical (narrowly elliptical, ovate), 13���35 �� (5���) 5.7���11.5 cm, usually thin-textured in proportion to their size (to moderately thick); inflorescences often partly cauliflorous on twigs below leaves; northern Madagascar...................... 6. H. megaphyllum 11a. Leaves elliptical or broadly elliptical to broadly obovate (obovate), (4.3���)5.2��� 12(���15) �� 2.1��� 8.5 cm, moderately thick-textured; inflorescences usually all borne in leaf axils, on distal portions of twigs; southern or central Madagascar............................................................... 12 12. Tree to 12(���20) m or shrub; leaves broadly elliptical to elliptical, (5.3���)6.2��� 12(��� 15) �� (2.8���) 3.5��� 8.5 cm, with petiole (10���)14���28(���33) mm; flowers short-pedicellate or sessile; southeastern Madagascar, littoral or low-elevation forests...................................................... 3. H. baillonii 12a. Tree to 30 m; leaves elliptical (ovate, narrowly elliptical), (4.3���)5.2���10(���12.4) �� 2.1���4.5(���5.8) cm, with petiole (6���)9���14(���27) mm; flowers normally sessile (in rare individuals short-pedicellate); central to south-central Madagascar, mid-elevation forests............. 5. H. leucophloeum, Published as part of Applequist, Wendy L., 2020, A revision of Homalium sect. Rhodonisa (Salicaceae) endemic to Madagascar, pp. 245-268 in Candollea 75 (2) on pages 247-249, DOI: 10.15553/c2020v752a8, http://zenodo.org/record/5724885, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","APPLEQUIST, W. L. (2016 a). A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae). Candollea 71: 231 - 256."]}

9. Homalium retusum (Blume) Wassel & Appleq., comb. nov. ��� Nisa retusa Blume, Mus. Bot. Lugd. -Bat. 2: 28. 1856. Lectotypus (designated here): MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Ile Ste. Marie, VIII���IX, fl., Bernier 173 (L [L0010974]; isolecto-: K [K000231491, K000231492] images seen, L [L0010975, L0010976] images seen, P [P00418083, P00418084, P00418085]!, TAN [TAN000593] image seen). = Homalium scleroxylon (Tul.) Baill. in Bull. Mens. Soc. Linn. Paris 1: 575. 1886. ��� Nisa scleroxylon Tul. in Ann. Sci. Nat., Bot. s��r. 4, 8: 69���70. 1857. Lectotypus (first step designated by SLEUMER, 1973: 299; second step designated here): MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Ile Ste. Marie, VIII ��� IX, fl., Bernier 173 (P [P00418085]!; isolecto-: K [K000231491, K000231492] images seen, L [L0010974, L0010975, L0010976] images seen, P [P00418083, P00418084]!, TAN [TAN000593] image seen). Syntypus: Galdich. [Gaudichaud] 296 [in phytotheca Jalbertiana] (not located). Tree 6���7 m tall, 10 cm dbh; twigs gray to medium brown, glabrous, leafing internodes 1.6���3.5(���4.5)mm diam.; stipules 2��� 6 mm, sparsely ciliate. Leaves obovate to broadly elliptic, (5���)7���16 �� 3.5���9 cm (unusually small leaves sometimes subtending inflorescences); base cuneate to convex; apex rounded to retuse or cuspidate; margin entire, rarely slightly crenulate, glands small; secondary veins (5���)6 ���9(��� 14) per side; both surfaces glabrous, drying light to medium brown on adaxial surface and darker on abaxial surface; petiole glabrous, (10���) 15���30 mm. Inflorescences paniculate, axillary, 8 ���12 cm; rachis short-pubescent; bracts and bracteoles 1���3 mm, usually ciliate, otherwise glabrous to sparsely pubescent; flowers mostly paired with 2 per bract. Flowers 5���6-merous, white or yellow; sepals obovate at anthesis becoming oblanceolate, 4���8(��� 9) mm, adaxial and abaxial surfaces glabrous; sepal glands glabrous; petals oblong-ovate, 1.5���3 mm, adaxial surface usually pubescent apically, basally glabrous, abaxial surface glabrous except often pubescent at extreme apex, margin usually ciliate; filaments glabrous, 1.2���1.9 mm; anthers (0.3���) 0.35���0.45 mm; styles (0.7���)1.2���1.7(���2) mm. Distribution, ecology and conservation status. ��� Homalium retusum is found near the eastern coast in the northern part of Madagascar at low elevations. Few specimens are known, suggesting that this is an uncommon to rare species. Only five clearly distinct subpopulations have been recorded (Ile Ste.-Marie in Analanjirofo, Foulpointe and Ambila-Lemaitso in Atsinanana, and Anjana and Andempona in SAVA). The Extent of Occurrence (EOO) is calculated as c. 9847 km �� and the Area of Occupancy (AOO) as 28 km ��. Much of the habitat is subject to ongoing anthropogenic damage, especially in view of its relative accessibility. Therefore, a preliminary assessment of its conservation status is Endangered [EN B1ab(iii)+2ab(iii)]. Vernacular names and use. ��� ���Hazombato��� (Service Forestier 2587); ���Longotra��� (Service Forestier 9646); ���Marangu�� Touditch��� (Bernier 173, Chapelier s.n.); ���Voapak��� (Humblot 63); ���Zana��� (Service Forestier 4918); ���Zanakanivato��� (Service Forestier 2597). Wood is good for construction (Bernier 173). Notes. ��� Homalium retusum has previously been lumped into H. nudiflorum s.l. (SLEUMER, 1973), which as herein circumscribed, differs in having smaller, narrower, shorterpetioled leaves; its inflorescences are usually racemose, not over 9 cm, glabrous or sparsely pubescent, with flowers borne singly. Homalium retusum most closely resembles H. pachycladum: both have relatively large leaves, paniculate inflorescences, and flowers with glabrous sepals and near-glabrous petals. Homalium retusum is known only from low-altitude and littoral forests, while H. pachycladum occurs farther inland and at much higher elevations. Homalium pachycladum has elliptic to obovate (or rarely ovate) leaves with shorter petioles (6���15 mm) and flowers all borne singly; its twigs seem usually to be thick, giving a robust appearance. Another inland species with large leaves, H. confertum, is distinguished by having usually elliptic to oblong-elliptic leaves with a serrate to crenate margin, racemose inflorescences with flowers borne singly, and petals densely pubescent on most of both surfaces. The history of the names applying to this species resembles that of Homalium planiflorum (Boivin ex Tul.) Baill. (Blackwellia planiflora Boivin ex Tul.) versus Blackwellia gracilis Blume (see APPLEQUIST, 2017, 2018b), but without disruptive nomenclatural consequences. Blume published Nisa retusa based on material of Bernier 173 a year before Tulasne published N. scleroxylon based mostly on the same collection. Blume specified that his name was based only on material sent from P to L. That material was described in the protologue (BLUME, 1856 ���1857: 28) as ���Nomine Asteropeia et Mourangia Touditsch ex Herb. Mus. Paris. mecum benevole communicata.��� Three sheets at L bore labels initially marked ��� Asteropeia [or Asteropeja] multiflora ��� then ��� Mourangia [or Morangia] touditch ��� (a vernacular name, though not identified as such on those sheets); Blume later added ��� Nisa retusa ��� in the top margins. Much more recent pink typed labels identify these sheets as being type material of N. scleroxylon, ���leg. Bernier 173 ���. One year later, Tulasne published N. scleroxylon, citing ���Bernerii herb., n. 173; Galdich. herb., n. 296, in phytotheca Jalbertiana��� (TULASNE, 1857: 70). These are Latinizations for Bernier and Gaudichaud-Beaupr�� respectively. Although Tulasne presumably used only those duplicates of Bernier 173 to be found in France, he did not say so. The protologue (TULASNE, 1857: 70) further states: ��� Marangue-Touditch (1), auctore Bernerio, vernaculare audit������, explaining the labeling seen, and apparently misinterpreted, by Blume. Three duplicates of Bernier 173 exist at P; two were labeled as having the vernacular name ��� Marangu�� touditch ��� and one had initially been labeled ��� Asteropeia ���. Though the sheets sent to L were not initially labeled as Bernier 173, they can be confidently assigned to that collection. SLEUMER (1973: 299) referred to Bernier 173 (L) as the ��� holotype ��� of Nisa retusa and to Bernier 173 (P) as the ��� holotype ��� of Nisa scleroxylon. For N. retusa, the original material was already restricted to the three sheets at L, which are syntypes, and Sleumer made no choice between them. The sheet that appears most complete is here chosen as a lectotype. For N. scleroxylon, Sleumer���s statement constitutes a first-step lectotypification (Art. 9.17 of ICN; TURLAND et al., 2018). This action means that N. retusa and N. scleroxylon are not nomenclatural synonyms, since their types are different specimens, even though they are from the same gathering. The best of the sheets at P is here chosen as a second-step lectotypification. Additional material examined. ��� MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: Foulpointe?, s.d., fl., Chapelier [Herb. Boivin] s.n. (P [2 sheets]); Foulpointe, Morarano, Andohanakoho, 17��41'52"S 49��27'28"E, 34 m, 9.III.2005, fl., Lehavana et al. 304 (MO, P); Foulpointe, for��t d���Analalava, 17��42'00"S 49��26'00"E, 35 m, 19.V.2004, fl., Randrianarivelo et al. 2 (MO, P); Sahapanpana, Foulpointe, 20.VI.1950, fl., Service Forestier 2587 (P); Ambila-Lemaitso, 1.II.1952, fr., Service Forestier 4918 (P); Ambila-Lemaitso, 27.IV.1954, fl., Service Forestier 9646 (P). Reg. SAVA [Prov. Antsiranana]: W de Nosiarina, sommet d���Anjana, 14��12'23"S 50��04'26"E, 190 m, 29.VIII.2007, fr., Andriamihajarivo et al. 1341 (MO); Antalaha, Andempona, 13.VI.1950, fl., Service Forestier 2597 (MO, P). Sine loco: Madag. bor., Andahoul, s.d., fr., Humblot 63 (P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 15-16, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","APPLEQUIST, W. L. (2017). Proposal to conserve the name Blakwellia (' Blackwellia') paniculata (Homalium paniculatum) (Salicaceae). Taxon 66: 206 - 207.","APPLEQUIST, W. L. (2018 b). A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae). Candollea 73: 221 - 244.","BLUME, C. L. (1856 - 1857). Museum Botanicum Lugduno-Batavum. Vol. 2. E. J. Brill, Lugduni-Batavorum.","TULASNE, L. - R. (1857). Florae Madagascariensis fragmenta. Ann. Sci. Nat., Bot. ser. 4, 8: 44 - 163.","TURLAND, N. J., J. H. WIERSEMA, F. R. BARRIE, W. GREUTER, D. L. HAWKSWORTH, P. S. HERENDEEN, S. KNAPP, W. - H. KUSBER, D. - Z. LI, K. MARHOLD, T. W. MAY, J. MCNEILL, A. M. MONRO, J. PRADO, M. J. PRICE & G. F. SMITH (2018). International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Veg. 159."]}

8. Homalium pachycladum Wassel & Appleq., sp. nov. (Fig. 5). Holotypus: MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: fiv. Maroantsetra, comm. Antsirabesahatany, fkt. Anjiahely, 15��23'06"S 49��26'58"E, 809 m, 20.XII.2002, fl., fr., Antilahimena, Pascal & Sassidy 1540 (MO [MO- 6144913]!; iso-, P!, TAN). Homalium pachycladum Wassel & Appleq. differs from H. retusum (Blume) Wassel & Appleq. in having thicker twigs, shorter petioles, glabrous (to sparsely pubescent) inflorescences with flowers borne singly, and larger petals. Shrub or tree to 10 m tall, 30 cm dbh; twigs medium brown or pale gray, glabrous, leafing internodes (1.7 ���)2 ��� 4(��� 5.5) mm diam.; stipules fused, caducous. Leaves broadly elliptic to elliptic or obovate (somewhat ovate), (3 ���)5.8 ��� 14.5 �� (1.8���) 3.5���7 cm; base rounded to cuneate; apex rounded to obtuse, apiculate, or emarginate; margin entire to subentire, with small, inconspicuous glands; secondary veins 8���11 per side; both surfaces glabrous, drying yellowish brown to greenish on adaxial surface, darker to pale brown on abaxial surface; petiole glabrous, 6���15 mm. Inflorescences paniculate, axillary, 4.5���11 cm; rachis glabrous to sparsely short-pubescent; bracts and bracteoles glabrous, (1���) 1.8���3.5 mm; flowers borne singly. Flowers 5���6(��� 7)-merous; sepals yellow to green, oblanceolate or oblong-oblanceolate to broadly spatulate in flower, 5.1���7.8 mm in flower, 7���10 mm in fruit, adaxial and abaxial surfaces glabrous; sepal glands glabrous; petals yellow to yellow-green, broadly ovate-elliptic to broadly ovate, 2.6���3.3 mm in flower, 3���5.6 mm in fruit, both surfaces sparsely to moderately pubescent only at apex and margins minutely ciliate; filaments glabrous, 2���3 mm; anthers 0.5���0.7 mm; styles ca. 2.5���4 mm. Distribution, ecology and conservation status. ��� Homalium pachycladum is endemic to mid-elevation humid forest west of Maroantsetra. The Area of Occupancy is 12 km ��; the Extent of Occurrence (EOO) would be calculated as c. 6 km ��, but following the IUCN (2017) guidelines for application of Red List Criteria, the EOO must be at least equal to the AOO and is therefore considered to be 12 km ��. All three collections were made inside the eastern edge of the recently delineated protected area of Makira. A preliminary assessment of this species��� conservation status is therefore ���Least Concern��� [LC]. Notes. ��� Homalium pachycladum often has a particularly robust appearance, especially during fruiting, with thick twigs and sturdy inflorescences. Though twig diameter is too variable within species to serve as a primary taxonomic character, it may be of use in recognizing this species. It most closely resembles H. retusum, which has obovate to broadly elliptic leaves with a long petiole, (10 ���) 15 ��� 30 mm, a cuneate to convex base, and more marginal glands; its flowers are smaller and mostly borne in pairs. That species favors low altitudes. Another species of moderate elevations with large, sometimes broad leaves, H. confertum, usually has a toothed leaf margin with conspicuous, elongated glands. The inflorescences of that species are racemose and its petals are densely pubescent. Paratypi. ��� MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Maroantsetra, Marovovonana, Ankirindro forest, 15��17'35"S 49��23'59"E, 664 m, 15.I.2003, fr., Antilahimena 1734 (MO, P); Maroantsetra, Abinanitelo, fkt. Ambalamahogo, Maimborondro summit, 15��19'03"S 49��24'06"E, 800 m, 8.IX.2004, fl., Antilahimena 2861 (G, MO, P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 14-15, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["IUCN (2017). Guidelines for Using the IUCN Red List Categories and Criteria: Version 13. Prepared by the IUCN Standards and Petitions Subcommittee. IUCN Species Survival Commission, Gland & Cambridge."]}

Homalium sect. Nisa (Noronha ex Thouars) Baill. ex Warb. in Engl. & Prantl, Nat. Pflanzenfam. 98���99: 36. 1893. ��� Nisa Noronha ex Thouars, Gen. Nov. Madagasc.: 24. 1806. ��� Homalium [unranked as ��] Nisa (Noronha ex Thouars) Baill. in Bull. Mens. Soc. Linn. Paris 1: 575. 1886. Typus (designated by SLEUMER, 1973: 295): Homalium nudiflorum (DC.) Baill. (��� Nisa nudiflora DC.). Stipules opposite the petiole base and fused (in H. intercedens Sleumer axillary, free), usually caducous. Leaves with venation more or less brochidodromous. Inflorescences spicate; bract and bracteoles large, usually broad, thicktextured, persistent. Flowers sessile (rarely subsessile); perianth 5���6(���8)-merous. Sepals obovate to oblanceolate (to nearly ligulate), oblanceolate-oblong or narrowly elliptic (in H. louvelianum H. Perrier oblong-ovate to ovate-deltoid), spreading, accrescent (only slightly so in H. louvelianum); calyx tube short, broadly funnelform to cup-shaped, or in fruit hemispherical. Petals ovate to oblong, usually smaller than sepals, modestly accrescent and curving over fruit (in H. louvelianum similar to sepals in size and spreading); sepals and petals not ciliate, or ciliolate in conjunction with overall surface pubescence. Sepal glands large, elliptic to elongated oblong-elliptic or roughly trapezoidal. Stamens 1 per petal, inserted between glands; anthers dorsifixed, broadly oblongelliptic with oblong-elliptic locules and a large connective, the slits of dehiscence nearly parallel. Upper surface of ovary often nearly flat in flower, in fruit becoming hemispherical or broadly conical; styles 3���4, fused basally (rarely fused for most of length). Locule of fruit subglobose or vertically compressed, internally glabrous (short-pubescent); seeds sometimes 1 per fruit, subglobose, largely filling the locule (or several small immature seeds, possibly most never maturing). Key to the species of Homalium sect. Nisa 1. Sepals about equal to petals at anthesis, broadly oblongovate (to ovate-deltoid), in fruit slightly accrescent, H. louvelianum 1a. Sepals longer than petals at anthesis, oblanceolate to narrowly oblong-oblanceolate, spatulate, narrowly elliptic, or nearly ligulate (in flower obovate to elliptic), in fruit conspicuously accrescent,> 3.5 mm and much longer than petals; petals usually curving inward as fruit matures.... 2 2. Sepal glands densely pubescent on upper (inward-facing) surface............................................... 10. H. stelliferum 2a. Sepal glands glabrous or glabrate throughout.............. 3 3. Leaves lanceolate; stipules free............ 4. H. intercedens 3a. Leaves elliptic, oblong-elliptic, obovate or oblanceolate (rarely to ovate, very rarely a few lanceolate); stipules fused, opposite petioles................................................ 4 4. Calyx densely pubescent to sericeous at least on cup, basal portion of abaxial sepal surface.................................... 5 4a. Calyx glabrous or sparsely pubescent, often more densely short-pubescent on cup, with trichomes confined to medial or marginal areas of one or both sepal surfaces............. 7 5. Leaves elliptic to narrowly elliptic, obovate, or oblanceolate, (2.5���)3���6(���9) �� 1.3���2.5(���3) cm, without marginal glands; bracts and bracteoles often ciliate, pubescent only along midrib or glabrous; adaxial surface of sepals glabrous; petals 1.5���2.5 mm, densely pubescent abaxially (to sparsely at margins), densely to sparsely pubescent adaxially; filaments 0.9���1.1 mm, anthers 0.2���0.3 mm................................................................................ 11. H. tenue 5a. Leaves elliptic to broadly or narrowly elliptic or obovate, (3.3���)5���9.5 �� (1.8���)2���4.5(���5.4) cm, usually with inconspicuous marginal glands; bracts and bracteoles ciliate and usually pubescent throughout (mostly along midrib); adaxial surface of sepals short-pubescent at least basally; petals 2���5 mm, velutinous on both surfaces; filaments (1.2���) 1.9���2.8 mm, anthers 0.4���0.65 mm..................... 6 6. Flowers mostly borne in pairs or closely clustered; bracts and bracteoles 3.5���5 mm; calyx cup long-sericeous, both sepal surfaces densely pubescent; southern Madagascar............................................................. 2. H. ciliolatum 6a. Flowers usually borne singly; bracts and bracteoles 1.5��� 2.5(��� 3) mm; calyx cup densely pubescent, sepals densely pubescent towards base of both surfaces, becoming sparsely pubescent (to glabrous) near apical margins; northeastern Madagascar............... 1. H. antilahimenae 7. Petioles (10���) 15���30 mm; inflorescences paniculate, pubescent; flowers mostly paired...................... 9. H. retusum 7a. Petioles 3���15 mm; inflorescences racemose or paniculate, glabrous or seldom sparsely or patchily pubescent; flowers borne singly................................................................. 8 8. Leaves oblanceolate, (2.7���)3.4 ��� 6.8 �� (0.9 ���) 1.1 ���2 cm, with revolute margins; inflorescences racemose, (0.5���)1���2.5(���3.3) cm; filaments 0.5���0.8 mm, anthers ca. 0.2 mm.............................................. 6. H. mandenense 8a. Leaves elliptic to broadly or narrowly elliptic, oblong-elliptic, or obovate (to oblanceolate in H. nudiflorum), 4.9���16 �� 1.5���9 cm, with entire (sometimes partly, not strongly revolute), crenate, or serrate to dentate margins; inflorescences racemose or paniculate, (2���) 3��� 12 cm; filaments 1.6���3.1(���3.7) mm, anthers (0.3���)0.5���0.7(���0.8) mm..... 9 9. Leaves (5 ���)6 ��� 16 �� (1.8 ���)3 ��� 6.2(��� 7) cm with petiole 3���7(���10) mm; leaf margins usually prominently toothed (subentire) with elongated glands in tooth apices; both surfaces of petals densely (to moderately) pubescent especially on apical half; humid forests to drier central forests, at elevations of 445���1860 m................. 3. H. confertum 9a. Leaves 4.9��� 14.5 �� 1.5���7 cm with petiole (3���)4���13(���15) mm; leaf margins entire to repand (crenulate, partly somewhat revolute), with glands small and round or absent; petals usually moderately (to densely) pubescent only at apex of one or both surfaces (rarely on entire apical portion), otherwise sparsely pubescent with often ciliate margins, or glabrous; humid (to littoral) forests, at elevations of 12���1220 m............................................ 10 10. Leafing internodes (1.7 ���)2��� 4(���5,5) mm diam.; leaves 8.5��� 14.5 �� 5 ���7 cm; inflorescences paniculate; petals in fruit 3���5.6 mm................................. 8. H. pachycladum 10a. Leafing internodes (0.9���)1.1���2.4(���3.2) mm diam.; leaves 4.9���9(���10.5) �� 1.5���4.2(���5.4) cm; inflorescences racemose (paniculate); petals 1.5���4 mm.............. 7. H. nudiflorum, Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 3-4, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328."]}

5. Homalium louvelianum H. Perrier in M��m. Mus. Natl. Hist. Nat. 13: 292. 1940. Lectotypus (designated by SLEUMER, 1973: 296): MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: Tampina, s.d., fl. & fr., Louvel 87 (P [P04704068]!). Syntypi: MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: Ambila au S de Tamatave, 4.V.1928, fl., Decary 6381 (P [P04704069]!, PRE [PRE0602227 -0] image seen, S [S10-10168] image seen, US [US00603578] image seen); for��ts c��ti��res [Tampina?], s.d., fl., Louvel 109 (P [P04704067]!). Reg. Vatovavy-Fitovinany [Prov. F ianarantsoa]: Mananjary, zone c��ti��re, III���IV.1909, fl., Geay 7425 (P [P04704065]!); ibid. loco, fl., Geay 7426 (P [P04704066]!); ibid. loco, fl., Geay 8069 (P [P04704064]!). Tree to 16 m tall, 40 cm dbh; bark orange or yellowish, exfoliating in plaques; twigs reddish brown when young, becoming grayish brown, glaucous to glabrous (minutely papillate), leafing internodes 1���2.5(���4.2) mm diam.; stipules fused, caducous. Leaves narrowly elliptical to oblanceolate or elliptical (obovate), (3.5���)5.5���12(���14) �� (1.5���)2���3.9(���4.5) cm; base cuneate to narrowly convex (slightly attenuate); apex acute to rounded, sometimes partly rounded-cuspidate, short-acuminate, or retuse; margin subentire, usually somewhat revolute at least at base, with glands in inconspicuous concavities or projections; secondary veins (6���)8���12(���14) per side, sometimes with very similar intersecondary veins; both surfaces glabrous, drying brown to dark (or greenish) brown on adaxial surface, usually somewhat paler brown on abaxial surface; petiole glabrous, (2 ���)3��� 11(��� 15) mm. Inflorescences racemose, axillary, 0.7���3.7(���5.6) cm; rachis glabrous; bracts and bracteoles glabrous except for sometimes ciliate margins, 0.5 ��� 1.5 mm; flowers borne singly (but usually clustered close together) or paired. Flowers 5��� 6-merous (aberrantly 4-merous), pale yellow to pale green or green sometimes with pinkish margins, or abaxial surface of calyx reddish; sepals broadly oblong-ovate (to ovate-deltoid), (1 ���) 1.2 ��� 1.8 mm in flower, (1���)1.5���2.6(��� 3.5) mm in fruit, adaxial and abaxial surfaces glabrous (calyx cup seldom sparsely pubescent); sepal glands yellow, glabrous; petals oblong-elliptic (broadly oblongovate), not curving inwards in fruit, 1.3 ���1.8 mm in flower, (1 ���)1.5���2(���2.5) mm in fruit, abaxial surface densely shortpubescent (to glabrous at margins toward base), adaxial surface densely to moderately short-pubescent towards apex, ciliate on most of margin; filaments glabrous, (0.8���)0.9���1.2(���1.3) mm; anthers 0.3���0.4(���0.45) mm; styles (0.4���)0.5���0.5(���1) mm. Vernacular names and use. ��� ���Fotsiakara��� (Ludovic & Ratiana 666, Ludovic 703, 735, Service Forestier 15375); ���Fotsiakarjay fotsy��� (Ludovic 391), ���Gavoala��� (Service Forestier 1562, 4702, 5690, 9531, 16869, 19188, 19502 [Betsimaraka dialect]); ���Goaviala��� (Service Forestier 4916); ���Goviala��� (Louvel 87, 109); ���Hafeala��� (Service Forestier 2936); ���Hazombato��� (Birkinshaw et al. 355, Rabevohitra et al. 4985, Randrianaivo et al. 1875); ���Hazombatovavy��� (Service Forestier 14837); ���Langipasina��� (Service Forestier 9514); ���Ramirisa��� (Service Forestier 2851, 3361); ���Ramirisy��� (Service Forestier 6975 [Tanosy dialect]); ���Rotra��� (Cours 2952); ���Zahandambo��� (Service Forestier 2110). Wood is used for construction (Ludovic & Ratiana 666, Ludovic 703, Service Forestier 19188, 19502). Distribution, ecology and conservation status. ��� Homalium louvelianum is widely distributed along most of the eastern coast of Madagascar. It is found in littoral forests, or less often low-altitude humid forests near the coast, on sand and ferralitic soil. Although littoral forests are highly fragmented and threatened, H. louvelianum is still frequently collected over a wide range. Its conservation status is assessed as ���Least Concern��� [LC]. Notes. ��� Homalium louvelianum displays less perianth accrescence than any other species of Homalium sect. Nisa. It has the smallest flowers within the section, and is unique in having sepals similar in size to the petals and broadest towards the base. The flowers often appear crowded on short racemes, though occasionally the rachis is relatively elongated and flowers more widely spaced. In these characters and the unusually short styles and filaments, the species bears a strong resemblance to those species of Homalium sect. Odontolobus Warb. s.l. (APPLEQUIST, 2018a) that display less extreme floral reduction. While no molecular data adequate to resolve relationships within Malagasy Homalium are available, this species suggests an intermediate or transitional form between those two sections which may provide a clue as to their relationships. Razakamalala et al. 3796 is identified as a hybrid between H. louvelianum and H. tenue. The leaves are consistent with H. louvelianum, narrowly elliptical with acute to acuminate apices. The sepals are only slightly longer than the petals, but resemble those of most species of Homalium sect. Nisa in shape and texture and are slightly reflexed, while the petals are densely pubescent and slightly inward-curving. Homalium tenue is found at this locality and its leaf shape and dense petal indument make it the presumptive second parent. Three specimens are from farther north than any population of typical Homalium louvelianum and occur at unusually high altitudes. Birkinshaw et al. 2015 has large, unusually shaped leaves and pubescent inflorescences and sepals. Two others from the forest of Ampondrabe have small leaves; their floral morphology is largely consistent with H. louvelianum. If typical H. louvelianum occurred in these regions, these specimens would be presumed to be hybrids, but since they are far from the known range of H. louvelianum, it is possible that they represent distinct species for which material is insufficient to allow recognition. Further investigation of these populations would be desirable. Additional material examined. ��� MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: 1 km S Mandrisy along road, 16��29'02"S 49��50'45"E, 10 m, 9.VII.1996, fr., Birkinshaw et al. 355 (MO, P); Ile Sainte-Marie, for��t d���Ambohidena, 16��51'11"S 49��57'10"E, 10 m, 13. V.2003, fr., McPherson et al. 18894 (MO, P); Soanierana-Ivongo, Antanambao-Ambodimanga, 16��46'19"S 49��43'52"E, 18.V.2003, fr., Rabenantoandro et al. 1481 (MO, P); fkt. Ambohidena, for��t d���Ambohidena, 16��51'11"S 49��57'18"E, 18.II.2004, fl., Rabevohitra et al. 4985 (MO, P); fkt. Manompana, for��t de Pointalare [Pointe �� Larr��e], 16��47'18"S 49��41'08"E, 24 m, 14. VI.2011, fr., Randrianaivo et al. 1875 (P); Tampolo, 17��17'S 49��25'E, 0���5 m, 5���15.V.1997, fl., Ravololonanahary et al. 82 (MO, P); fkt. Ambohidena, for��t d���Ambohidena, 16��50'25"S 49��57'09"E, 10 m, 19.II.2004, fl., Razakamalala et al. 899 (MO, P); Mahambo, au S de F��n��rive, 30.VIII.1957, fr., Service Forestier 18143 (P [2 sheets]); S de la Mandrisy, S d���Antanambe, Mananara, 13.XI.1964, fr., Service Forestier 23771 (P [2 sheets]). Reg. Anosy [Prov. Toliara]: Mandena, 24��57'S 47��00'E, 0���10 m, 19.IV.1989, fl., Gereau et al. 3388 (MO, P); Ste. Luce, 24��47'S 47��10'E, 20 m, 15. I.1990, fl., McPherson et al. 14796 (MO, P); Ft-Dauphin, fkt. Tsiharoa, for��t d���Etazo (S 5), 24��49'36"S 47��06'44"E, 20 m, 8.VIII.2012, Rakotonirina et al. 870 (MO); Mandena, 24��57'S 47��00'E, 0 m, 22.VI.1996, fr., Razafimandimbison 211 (G, K, MO, P); Mandena, Ft-Dauphin, 7.III.1951, fl., Service Forestier 2851 (P); Mandena, 11 km au N de Ft. Dauphin, IV���V.1953, fl., Service Forestier 6975 (P). Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: fkt. Baboaka, 1.5 km SW d���Analamena, 23��09'37"S 47��44'41"E, 15 m, 6.VI.2003, fr., Ludovic 391 (MO); Vohimasy, Agnalazaha Ampoakafo, 23��10'58"S, 47��42'27"E, 12 m, 13.VIII.2006, fr., Razafitsalama 961 (MO); Mahabo Mananivo, for��t de Manombo, 23��10'12"S 47��42'50"E, 21 m, fl., Razakamalala & Rabehevitra 701 (MO [2 sheets], P); Manombo RS, parcelle I, for��t d���Anaviavy, 22��59'16"S 47��42'26"E, 65 m, 14.IX.2005, fl., Razakamalala et al. 2127 (P); Analazaka, Efotsy, 30.VIII.1955, fl., Service Forestier 15375 (P); Vohimasy, Agnalazaha Ampoakafo, 23��10'58"S 47��42'27"E, 12 m, 13.VIII.2006, fr., Razafitsalama 961 (MO). Reg. Atsinanana [Prov. Toamasina]: Ambila-Lemaitso, 18��49'28"S 49��09'19"E, 10 m, 25.IX.2000, fl., Andrianjafy et al. 108 (MO, P); Vohibola, N of Lac Ampitabe, 18��33'34"S 49��15'01"E, 5 m, 12.II.2003, fr., Lowry et al. 6074 (MO, P); fkt. Andranonkoditra, for��t de Vohibola, 18��35'32"S 49��14'02"E, 5 m, 11.II.2003, fr., Rabenantoandro et al. 1259 (MO, P); Akanin���ny Nofy, for��t de Vohibola, 18��33'46"S 49��15'06"E, 3.VIII.2003, fr., Razakamalala & Rabehevitra 649 (MO, P); Ambila-Lemaitso, 25.X.1950, fr., Service Forestier 1562 (P); Ambila-Lemaitso, 14.IV.1952, fr., Service Forestier 4702 (MO, P); ibid. loco, 25.IX.1952, fr., Service Forestier 5690 (P); ibid. loco, 16.III.1954, fr., Service Forestier 9531 (P); Tampina, JB 20, 12.IX.1958, fr., Service Forestier 19188 (P); Ambila-Lemaitso, Jardin Botanique no. 2, 22.V.1959, fl., Service Forestier 19502 (P). Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa]: fkt. Marohita, for��t d���Ambahisosotra, 21��29'12"S 48��16'20"E, 13 m, fr., Ranaivojaona et al. 541 (MO, P); fkt. Ambalavontaka, for��t d���Antaimby, 20��23'26"S 48��32'43"E, 10 m, 20.IV.2004, fr., Ranaivojaona et al. 651 (MO, P); Pangalane, sud[?] Mananjary, 24.2.1954, fl., Service Forestier 9514 (P); for��t de Manampano, Mananjary, 17.VI.1955, fr., Service Forestier 14837 (P). Sine loco: 3���5 m, 10.X.1946, fr., Cours 2952 (MO, P). Hybrid specimen. ��� MADAGASCAR. Reg. Anosy [Prov. Toliara]: Iaboko, Antsotso Avaratra, 24��34'16"S 47��12'06"E, 271 m, 8.XII.2007, fr., Razakamalala et al. 3796 (P). Specimens incertae sedis. ��� MADAGASCAR. Reg. SAVA [Prov. Antsiranana]: Tsihomanaomby access from Antanambaonisokitra, 14��05'52"S 50��00'26"E, 331 m, 23.IV.2014, fl., Birkinshaw et al. 2015 (MO); Daraina, for��t d���Ampondrabe, 12��57'14"S, 49��42'13"E, 517 m, 9.IV.2004, fl., Ranirison 612 (P); Tsaratanana, Ambarilao, for��t d���Ampondrabe (Antsanjoana), 12��57'29"S 49��41'32"E, 600 m, 9.XI.2005, fr., Ratovoson et al. 1094 (MO, P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 9-10, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","APPLEQUIST, W. L. (2018 a). A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar. Candollea 73: 27 - 48."]}

7. Homalium nudiflorum (DC.) Baill. (Fig. 4). ��� Nisa nudiflora DC., Prodr. 2: 55. 1825. Lectotypus (designated here): MADAGASCAR: sine loco, s.d., fr., Petit-Thouars s.n. (P [P00418087]!; isolecto-: P [P00418086]! pro parte). Tree to 20 m tall, 50 cm dbh; bark gray, yellowish inside, sometimes platanoid; twigs light to medium gray or brown, glabrous, leafing internodes (0.9���)1.1���2.4(���3.2) mm diam.; stipules fused, caducous. Leaves obovate to elliptic, broadly or narrowly elliptic, or oblanceolate, 4���9(���10.5) �� 1.5���4.2(���5.4) cm; base convex to cuneate, rounded, or attenuate; apex rounded to slightly acuminate or apiculate, rarely acute with a rounded tip or retuse; margin entire (seldom shallowly repand, rarely slightly crenate-denticulate or partly and not strongly revolute), glands if present small, round, usually inconspicuous, rarely prominent in young leaves; secondary veins (5���)6���9(���11) per side; adaxial surface glabrous (glaucous), abaxial surface glabrous or seldom weakly glaucous, both surfaces drying light to medium brown or green to olive, sometimes the adaxial surface brown, young leaves sometimes dark brown; petiole glabrous, (3���)4���13(���16) mm. Inflorescences racemose or occasionally paniculate (especially in Alaotra-Mangoro region), axillary (rarely paired), (3���)4���9(���11) cm; rachis glabrous to moderately (partially densely) minutely pubescent); bracts and bracteoles 1.5���3.5(���4) mm, glabrous or occasionally ciliate; flowers borne singly. Flowers 5 ��� 6(7)-merous, pale yellow, pale green to yellowish green, or white; sepals oblanceolate to narrowly oblong-oblanceolate or spatulate (in early flower obovate), 3���6 mm in flower, 5.6���9.5(���10) mm in fruit, adaxial surface glabrous, abaxial surface glabrous or glabrate (rarely with margin ciliate or calyx cup pubescent); sepal glands orange to bright yellow, glabrous; petals ovate (broadly ovate at anthesis) to elliptic, 1.5���4 mm, both surfaces glabrous or short-pubescent near apex (seldom moderately or rarely densely pubescent over much of surface), sometimes also on midrib abaxially, margins sometimes ciliate; filaments glabrous, 1.6���3(���3.7) mm; anthers (0.3���)0.5���0.7(���0.8) mm; styles (0.8���)1.5���3.4(���4) mm. Distribution, ecology and conservation status. ��� Homalium nudiflorum as circumscribed herein is a very widespread, though scattered species in eastern forests. It usually occurs in mid-elevation humid forests up to 1220 m, but is occasionally found near sea level. Reported substrates are lateritic soils, including ferruginous crust and cuirasse, volcanic soil, and gneiss or granite. It occurs in several protected areas, including Galoko, Manombo, Mantadia, Midongy du Sud, Montagne d���Ambre, and Zahamena. Its conservation status is estimated as ���Least Concern��� [LC]. However, it should be noted that regional variants of the species may be genetically distinct and at greater risk. Vernacular names and uses. ��� ���Antevaratra ��� (Randriamanarivo et al. 10); ���Fotsiakara��� (Service Forestier 10150); ���Hazofotsy �� [petits] feuilles��� (Service Forestier 3794); ���Hazomalany��� (Andriamahefarivo et al. 90); ���Hazombato��� (Service Forestier 11013, 13239); ���Hazondroka��� (Service Forestier 16192 [Antaisaka dialect]); ���Hazoporofoka��� (Service Forestier 2231); ���Lamotiala��� (Service Forestier 10435); ���Lapivahatra��� (Razanatsima et al. 1364); ���Menavahatra��� (Antilahimena et al. 3005); ���Remirisa��� (Antilahimena et al. 4826). Wood is used for construction of houses (Randriamanarivo et al. 10, Service Forestier 16192) and for heating and charcoal production (Service Forestier 16192). Notes. ��� Homalium nudiflorum as defined by SLEUMER (1973) was much too broadly circumscribed, including in a single species with one variety all of sect. Nisa that was not obviously excluded by having greatly different perianth morphology (H. louvelianum), stipule morphology (H. intermedium), or sepal gland and foliar indument (H. stelliferum). In this treatment, eight species are recognized within what was formerly the H. nudiflorum complex. Homalium nudiflorum as herein defined is much less heterogeneous, but remains a widespread and morphologically variable taxon. Variation in morphology is probably geographically correlated, but since most parts of the range are represented by few individuals, it is impossible to determine how consistently so. Most conspicuously, most collections from the far north (DIANA and SAVA) have broadly elliptical leaves (providing the upper extreme of the size range) with usually rounded bases, a glaucous upper surface, and often shallowly repand rather than entire margins. Additionally, these collections have moderate (to dense) indument on the apical half of the petals, while material from other parts of the country usually has trichomes only at the extreme petal apices. These populations might merit recognition as a species. However, most of their distinctive characters may be found individually in specimens from other regions (e.g., there are southern collections with large, broad-based, broadly elliptical leaves or with heavy petal indument) and there are both typical northern collections and potential intermediates. Because hybridization between H. nudiflorum and H. confertum is suspected to occur primarily in the north (see below), it is also possible that some characters of these specimens are attributable to gene flow or incomplete separation from H. confertum. SLEUMER (1973: 299) stated that Petit Thouars s.n. ���P & P-Juss 14 411��� was the ��� holotype ��� of Homalium nudiflorum, and that no duplicate was found at G-DC, where material possessed by De Candolle might have been expected to be located. Two sheets of Petit Thouars s.n. were located at P, both of which were labeled ��� holotype ��� by Sleumer; these should be considered syntypes. Both are of poor quality, but the sheet barcoded [P00418086] includes a fragment packet containing flowers of H. nudiflorum along with mounted material that appears to be H. louvelianum. Therefore, [P00418087] is chosen as lectotype. The type collection has narrowly elliptic to obovate or narrowly obovate leaves (some fairly large, though not broad for their size), a glabrous rachis, bracts (except for cilia) and calyx, and sparsely pubescent petals. Without locality data, it is not possible to identify this as pertaining to any specific geographic variant of H. nudiflorum, which favors the maintenance of a relatively broad species concept for pragmatic reasons, while excluding several segregate taxa that are clearly not conspecific with the type. Several specimens (see below) are observed that have the fairly small leaves and most other features of Homalium nudiflorum s.str. combined with features of H. confertum, in particular dense indument on most, or sometimes all, of both petal surfaces. A few have variable leaf morphology with some leaves being crenulate or dentate-crenulate, with some elongated glands. A few other specimens with atypical morphology are herein treated as H. nudiflorum. All of the putative intermediates come from regions where both H. nudiflorum and H. confertum are found. Hybridization is observed in other sections of Homalium (APPLEQUIST 2016a, 2018b). Hybridization or gene flow between these species, which are probably closely related, is suggested to be responsible for the features of these specimens. One or two specimens, e.g., Service Forestier 23750 with very small anthers and short filaments and styles, might be suspected of other hybrid ancestry, but their morphology is otherwise consistent with H. nudiflorum. Service Forestier 18334 has leaves similar to those of one form of H. nudiflorum (except that the adaxial surface is glaucous) but relatively gigantic flowers, with sepals reaching 20 mm in fruit. No other material of this section has flowers of similar size. As three characters to distinguish it from H. nudiflorum, as herein circumscribed, are not seen, it is tentatively placed in that species. However, further collections of this population would be highly desirable. Baron 6251 is a specimen in poor condition with a few small flowers and a few elliptical to ovate leaves, one of them very large. Though leaf shape is quite variable in this species complex, ovate leaves are normally not seen. This may represent a hybrid with a larger-leaved species, possibly of another section, or a distinct species. Baron 6270 bears a fragment with short inflorescences and very small (2.2���3.3 �� 1.5���2.2 cm), closely spaced, broadly elliptical leaves with entire margins, which are not glaucous. As the inflorescence is well past anthesis, the small size of the leaves and internodes is not due to immaturity. The specimen might be an aberrant individual of H. nudiflorum or an otherwise unknown related taxon. No locality data are available for Baron���s collections, so it is not possible to attempt to relocate the collected populations, and the available material is not adequate to recognize new species. Additional material examined. ��� MADAGASCAR. Reg. Alaotra-Mangoro [Prov. Toamasina]: Menalamba, Ambatovy forest, 18��48'29"S 48��18'50"E, 1060 m, 16.XII.2004, fl., Antilahimena et al. 3005 (MO, P); Ampanatovana, Ambatovy forest, 18��51'35"S 48��19'12"E, 1116 m, 6.XII.2008, fl., Antilahimena et al. 6959 (G, K, MO); Ambatondrazaka, Cours 1095 (P);] near Andasibe, forest of Mantadia, 18��55'S 48��25'E, 950���1150 m, 7.XI.1994, fl., McPherson & van der Werff 16535 (MO); Ampitambe, for��t d���Analamay, 18��48'39"S 48��20'35"W, 1077 m, 3.X.2008, fr., Rakotondrajaona 609 (MO); for��t de Berano, Ampitambe, Ambatovy, 18��51'21"S 48��20'04"E, 1048 m, 12.I.2005, fr., Ranaivojaona et al. 1043 (MO, P). Reg. Analanjirofo [Prov. Toamasina]: Maroantsetra, Anjahana, Ambanizana, 15��37'30"S 49��58'35"E, 190 m, 20.IX.2002, fl., Antilahimena & Daiky 1423 (MO, P); Marovovonana, Andampy forest, 15��19'03"S 49��24'06"E, 800 m, 3.IX.2004, fl., Antilahimena 2719 (G, MO); Parc National de Zahamena, 17��33'49"S 48��54'01"E, 794 m, fl., Randriamanarivo et al. 10 (MO); Antanambe, au S de Mananara, 10���14. XI.1964, fl., Service Forestier 23750 (P [2 sheets]). Reg. Anosy [Prov.Toliara]: fkt. Antsontso, Ivohibe forest, 24��34'13"S 47��12'01"E, 286 m, 23.V.2006, fr., Antilahimena et al. 4826 (P); for��t d���Analalava, 24��13'S 47 ��21'E, 40 m, 29.X.1989, fr., McPherson 14289 (MO, P); Antsotso, Vohimena, for��t d���Ivohibe, 24��33'45"S 47��12'19"E, 421 m, 28.I.2015, fr., Razanatsima et al. 1364 (G, MO); for��t de Vohibe-Manantenina, 14.VIII.1951, fl., Service Forestier 3794 (P). Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: Farafang��na, for��t class��e de Manombo, 23��04'00"S 47��40'33"E, 42 m, 27.IX.2002, fl., Rabevohitra et al. 4128 (MO, P); Farafangana, Ankarana, village le plus proche Manombo, 23��00'39"S 47��44'08"E, 12 m, 16.IX.2005, fl., Rakotonirina et al. 300 (MO, P); Midongy du Sud, Anaviavy, Kelilahy, 16.IV.1956, fr., Service Forestier 16192 (P). Reg. DIANA [Prov. Antsiranana]: Ampasindava, for��t d���Andranomatavy, 13��40'04"S 47��58'53"E, 410 m, 1.XII.2009, fr, Madiomanana et al. 282 (P); for��t s��che d��grad��e d���Andavakoera, �� 3 km �� l���Est de Marivorahona, 13��05'56"S 49��08'33"E, 54 m, 2.IX.2005, fl., fr., Randrianaivo et al. 1221 (MO, P); Marivorahona, W du mont Andavakoera, 13��05'14"S 49��10'29"E, 100 m, 4.II.2005, fl., Ratovoson et al. 881 (MO); 9 km SE d���Anketrabe, for��t de Kalabenono, chaine de Galoko, 13��38'30"S 48��40'53"E, 976 m, 23.XI.2006, fr., Razafitsalama & Torze 1105 (MO, P); Ambilobe, Andampy, 14.V.1954, fl., Service Forestier 10435 (P); Ambilobe, comm. rurale Beramanja, Cha��ne Galoka [Galoko], Mont Galoka [Galoko], fkt. Anketrabe-Belinta, revel�� 2, 13��35'12"S 48��43'37"E, 22.II.2005, fr., Wohlhauser et al. 767 (MO). Reg. Haute Matsiatra [Prov. Fianarantsoa]: for��t d���Andrambovato �� l���E de Fianarantsoa, 24.I.1955, fr., Service Forestier 11593 (P). Reg. Ihorombe [Prov. Fianarantsoa]: Farafangana, Ankarenambe, 24.II.1950, fr., Service Forestier 2231 (P). Reg. SAVA [Prov. Antsiranana]: fkt. d���Ankijabe, for��t de Binara, camp I (13��15��� S, 049��37��� E), 1.6 km au SW du camp, 900 m, 9.XI.2001, fl., Gautier & Ravelonarivo 4142 (MO, P); Montagne d���Ambre near Grand Cascade, 12��31��� S, 049��10��� E, 900 m, 22.I.1994, fr., Leeuwenberg et al. 14292 (P); Montagne d���Ambre, 12��32'S 49��09'E, 1000���1100 m, 20.XI.1989, fr., McPherson 14484 (MO, P); Daraina, for��t d���Antsahabe, 13��13'S 49��33'E, 896 m, 21.XI.2004, fl., Nusbaumer & Ranirison 1261 (MO, P). Reg. Sofia [Prov. Mahajanga]: Mandritsara, commune de Makira, 15��31'S 49��05'E, 1169 m, 26.IV.2007, fl., Lehavana et al. 481 (MO, P). Selected hybrid or intermediate specimens. ��� MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: fkt. Ambatolampy, for��t d���Ankerana, 18��23'34"S 48��48'37"E, 952 m, 23.I.2012, fr., Ravelonarivo et al. 4149 (MO). Reg. SAVA [Prov. Antsiranana]: Montagne d���Ambre, 12��32'35"S 49��08'46"E, 1300 m, fl., 6.XII.2007, Ramandimbimanana et al. 9 (MO, P); J.B. 19 Roussettes, Diego-Suarez, 11.XI.1954, fl., Service Forestier 11013 (P); Montagne d���Ambre, 12��30'30"S 49��10'09"E, 1000 m, fr., 1.VI.2008, Trigui et al. 485 (MO). Specimens incertae sedis. ��� MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Baie d���Antongil, massif de Farankaraina entre Navana et Andranofotsy, 0���150 m, 19.IX.1957, fr., Service Forestier 18334 (P). Sine loco: s.d., fr., Baron 6251 (P); s.d., fr., Baron 6270 (P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 11-14, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","APPLEQUIST, W. L. (2016 a). A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae). Candollea 71: 231 - 256.","APPLEQUIST, W. L. (2018 b). A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae). Candollea 73: 221 - 244."]}

2. Homalium ciliolatum (Sleumer) Wassel & Appleq., comb. & stat. nov. ��� Homalium nudiflorum var. ciliolatum Sleumer in Bull. Jard. Bot. Belg. 43: 299. 1973. Lectotypus (designated here): MADAGASCAR. Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa]: Ifanadiana, for��t d���Ambohimiera [Tsaratanana], 20.VIII.1954, fl., fr., Service Forestier 14566 (P [P00375093]!; isolecto-: L [L0010977] image seen, P [P04734282]!). Tree to 25 m tall, 1 m dbh; bark gray-green; twigs gray to light brown, sometimes minutely pubescent on new growth, otherwise glabrous, leafing internodes (0.8���)1.2���2.4(���3.2) mm diam.; stipules fused, caducous. Leaves elliptic to broadly elliptic or obovate, (3.6���)4.5���9.5(���11.6) �� (1.6���) 2.3���4.9 cm; base convex to broadly cuneate; apex rounded to cuspidate or obtuse (retuse, short-acuminate); margin entire, glands inconspicuous; secondary veins (5���)7���10 per side, sometimes with intersecondary veins almost equally large; both surfaces glabrous, drying pale to medium or olive-brown; petiole glabrous, 4���10 mm. Inflorescences mostly pseudoterminal, confined to twig apices, racemose to paniculate, 3.5���11.5(���20) cm; rachis pubescent, especially towards apex; bracts and bracteoles 3.5���5(���7) mm long, short-pubescent and ciliate; flowers mostly borne in pairs or gathered in widely-spaced clusters. Flowers 6���7(���8)-merous, white; sepals elliptic when young to narrowly elliptic or oblong-elliptic, becoming oblanceolate, 4.8���9 mm in flower, (7���) 8.5���10 mm in early fruit, entire adaxial and abaxial surfaces of sepals short-pubescent, base of calyx cup sericeous; sepal glands glabrous; petals lanceolate to narrowly ovate or narrowly oblong-ovate, 3���4 mm in flower, 4���5.5 mm in early fruit, both surfaces velutinous; filaments glabrous (rarely sparsely pilose), (1.5���) 2.2���2.8 mm; anthers 0.5���0.6 mm; styles (2���) 2.4���3 mm. Vernacular names and use. ��� ���Fotsiakara��� (Service Forestier 21504); ���Fotsiankara��� (Service Forestier 13984, 21504 [Antsakag��n��ral dialect]); ���Fotsivony��� (Service Forestier 6390); ���Hazofotsy �� grandes feuilles��� (Service Forestier 3796); ���Hazombato��� (Service Forestier 14719); ���Kamirisa��� (Antilahimena et al. 4863); ���Ramanerika��� (Service Forestier 14566). Wood is used for construction and firewood (Service Forestier 21504). Distribution, ecology and conservation status. ��� Homalium ciliolatum is confined to the southeastern regions of AtsimoAtsinanana, Vatovavy-Fitovinany, and Anosy. It is usually found near the coast at low elevations, but the type was probably from a mid-elevation forest. Reported substrates include basalt and rocky soil; habitats include forest and savoka (disturbed grassland). As many as ten subpopulations have been collected in the past (older locality data are imprecise). However, it has only been collected twice in the past 54 years, though much of its habitat is relatively accessible, so it is evidently rare. From all historical localities, the Extent of Occurrence is calculated as c. 15,980 km �� and the Area of Occupancy as 40 km ��. Most historical populations were from land that has been subject to serious anthropogenic damage in the intervening decades and is still not protected. Therefore a preliminary assessment of its conservation status is ���Vulnerable��� [VU B1ab(iii)+ 2ab(iii)], but this is likely to underestimate the degree of threat it faces. Notes. ��� Homalium ciliolatum was described by SLEUMER (1973) as a variety of H. nudiflorum, from which it is most obviously separated by its densely pubescent flowers. The petals are velutinous on both surfaces; the basal part of the calyx externally is densely sericeous, and the remainder of the sepals are short-pubescent and ciliate. Also, the upper portions of the inflorescence rachis and the bracts and bracteoles are pubescent, and the bracts and bracteoles short-ciliate. The petals appear often to remain curved over the ovary before fruit develops. It is likely that no fully mature perianths have been seen and the final size is larger than the range reported here. The distribution of H. ciliolatum is more southerly than that of H. nudiflorum and it is confined to lower altitudes. Two other species herein excluded from the H. nudiflorum complex, the newly described H. antilahimenae Wassel & Appleq. and H. tenue Wassel & Appleq., also have calyces much more pubescent than those of H. nudiflorum. Homalium ciliolatum is distinguished from these by the distinctive long-sericeous indument of the calyx cup, which is not seen in any other species, and by having most flowers borne in pairs. Service Forestier 14566 at P was stated (SLEUMER, 1973: 299) to be the ��� holotype ��� of H. nudiflorum var. ciliolatum. There are in fact two duplicates of this collection at P, which are therefore syntypes. The duplicate designated herein as lectotype is in better condition and has better original label data. Additional materials examined. ��� MADAGASCAR. Reg. Anosy [Prov. Toliara]: Iabakoho, Antsontso, Ivohibe forest, 24��34'10"S 47��12'37"E, 41 m, 24.V.2006, fr., Antilahimena et al. 4863 (MO, P); for��t d���Androkabe, FortDauphin, 15.VIII.1951, fr., Service Forestier 3796 (P). Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: Vangaindrano, Vohipaho, 23��31'40"S 47��29'46"E, 73 m, 17.IX.2009, fl., Andriamihajarivo et al. 1701 (MO); Farafangana, Ihorombe, Mahatsinjoriaka, 23.X.1952, fr., Service Forestier 6390 (P); Farafangana, Amporofo, 26.VII.1955, fr., Service Forestier 13984 (P); Farafangana, Bekaraoka, 8.X.1963, fl., Service Forestier 21504 (MO, P); S de Farafangana, rte de Manombo, aux P.K. 20��� 21, 14���17.X. 1964, fr., Service Forestier 23613 (P). Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa]: Mananjary, Ambohinihaonana, 2.VI.1954, fl., Service Forestier 14476 (P); Mananjary, Vohilaba, Ambodononoka, 20.VI.1954, fl., Service Forestier 14719 (P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 5-6, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328."]}

10a. Homalium stelliferum H. Perrier subsp. stelliferum Tree to 15 m tall, 80 cm dbh; twigs with leafing internodes 1���2.7 mm diam. Leaves narrowly oblong-elliptic to oblongoblanceolate or oblanceolate (rarely obovate), 3.6 ��� 11.5 �� 1.3���4.5(���5.5) cm; apex cuspidate to rounded, obtuse or acute (short-acuminate); margin subentire to inconspicuously toothed (dentate), with glands usually present in marginal convexities; drying brown (to grayish green) on adaxial surface, lighter brown (to green) on abaxial surface; petiole pilose, short-pubescent or glabrous, (2���)3���5(���10) mm. Inflorescences: bracts and bracteoles usually with ciliate margin, densely to moderately pubescent (pilose, glabrate), 2.3��� 3.5(��� 4) mm. Flowers 5 ��� 6-merous; sepals narrowly elliptic to narrowly oblong-elliptic or oblanceolate, (4���) 6���8.4 mm after anthesis, 9.5���11 mm in fruit, glabrous or with dense appressed pubescence on calyx cup, dense to sparse pubescence in medial portion of one or both sepal surfaces. Vernacular names and use. ��� ���Fotsiakara��� (Service Forestier 12394); ���Tanatampotsy��� (Service Forestier 5798); ���Tanatanapotsy��� (Service Forestier 12631); ���Tsianihomposa��� (R��serves Naturelles s.n.); ���Tsitongamposa��� (R��serves Naturelles s.n.); ���Vavarony��� ou ���Voavarony��� (Service Forestier 13709). Wood is used for construction of houses (Service Forestier 19809). Distribution, ecology and conservation status. ��� Homalium stelliferum subsp. stelliferum has been widely distributed in the past but is seldom collected; only two recent collections are known. It is usually found near the eastern coast in lowaltitude forests, extending to coastal or mid-elevation forests. Nine widely separated locations have been collected. The historical habitat is mostly unprotected, but includes the protected area of Zahamena. The Extent of Occurrence (EOO) is calculated as c. 51,200 km �� but the Area of Occupancy as 36 km ��, which is heavily dependent upon historical subpopulations. Low-elevation eastern humid forest has been subject to considerable anthropogenic damage from wood cutting and forest clearing for agriculture, and this continues to occur in unprotected areas. The preliminary conservation status of this subspecies is assessed as ���Vulnerable��� [VU B2ab(iii)], which may underestimate the level of threat it faces. Additional material examined. ��� MADAGASCAR. Reg. Alaotra-Mangoro [Prov. Toamasina]: Ambatondrazaka [Zahamena], au bord de la riv. Ranomainty, 17��45'12"S 48��44'07"E, 5.III.2001, fr., Ratovoson et al. 460 (G, MO, P, K, TEF not seen). Reg. Analanjirofo [Prov. Toamasina]: Ambatomilona, Mananara, 15.XII.1951, fr., Service Forestier 5798 (P). Reg. Anosy [Prov. Toliara]: col Sakatelo (Ambazaha), canton Mahatalaky, distr. Ft-Dauphin, 12.XI.1949, fl., R��serves Naturelles s.n. (P). Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: Boma, distr. de Farafangana, 13.IX.1954, fr., Service Forestier 12394 (P). Reg. Atsinanana [Prov.Toamasina]: Vatomandry, comm. Ambalabe, 19��09'43"S 48��36'05"E, 423 m, 26.XI.2004, fr., Randrianasolo et al. 953 (MO, P); Andriantantely, Brickaville, 22.X.1950, fl., Service Forestier 12631 (P). Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa]: Iaranina[?], Ampasinambo, Nosy-Varika, 24.XI.1954, fl., Service Forestier 13709 (MO, P); Antsampanandalana, Ambahaka, Ankarimbelo, Fort-Carnot, 23.X.1960, fl., fr., Service Forestier 19809 (P [2 sheets]); cr��te de Tsangatsanga, �� l���E de Kianjavato (entre Ifanadiana et Anosivola), 550 m, 6.XII.1964, fl., Service Forestier 23918 (MO)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on page 18, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766

6. Homalium mandenense Wassel & Appleq., sp. nov. (Fig. 3). Holotypus: MADAGASCAR. Reg. Anosy [Prov. Toliara]: pr��fecture de Taolagnaro, for��t de Mandena, piste au N de l���intersection des piste-Camp, 24��57'S 47��00'E, 0���10 m, 8.III.1989, fr., Dumetz 537 (MO [MO-3662611]!; iso-: P [P04734305]!, TAN). Homalium mandenense Wassel & Appleq. differs f rom Homalium nudiflorum (DC.) Baill. in having smaller, consistently oblanceolate leaves with revolute margins, shorter racemes, and flowers smaller at maturity. Tree or shrub to 7 m tall, 6 cm dbh; twigs medium brown to grayish, usually glaucous when young, peeling (glabrous), leafing internodes 1.0��� 2.5(��� 3.2) mm diam.; stipules fused, caducous. Leaves oblanceolate, (2.7���)3.4���6.8 �� (0.9���) 1.1���2 cm; base cuneate; apex rounded to acute or cuspidate; margin more or less revolute, glands inconspicuous or absent; secondary veins 6��� 7(��� 8) per side; both surfaces glabrous, drying brown (seldom to green) on adaxial surface, brown or greenish on abaxial surface; petiole glabrous, (3���)6���12(���14) mm. Inflorescences racemose, axillary, (0.5���)1��� 2.5(��� 3.3) cm; rachis glabrous; bracts and bracteoles glabrous except for ciliate margins, 0.9��� 1.6 mm; flowers borne singly. Flowers 6 ��� 7(��� 8)-merous, white to yellowish; sepals elliptical (to obovate) in early flower, becoming oblanceolate to narrowly oblong-oblanceolate, 2.8���4 mm in flower, 3.5���4.7 mm in fruit, adaxial and abaxial surfaces glabrous or with sparse pubescence only at base of calyx cup; sepal glands yellow, glabrous; petals elliptic, to oblong at maturity, 1.4���2.4 mm, moderately (densely) pubescent on apical portion of both surfaces, sometimes extending to most of adaxial surface, and margins ciliate; filaments glabrous, 0.5��� 0.8 mm; anthers 0.2 mm; styles ca. 0.5(���1?) mm. Vernacular names and use. ��� ���Z��ra��� (Ludovic 1649); ���Zoramena��� (Ratovoson 1719). Wo o d i s u s e d f o r c o n s t r u c t i o n (L u d o v i c 1 6 4 9, Ratovoson 1719). Distribution, ecology and conservation status. ��� Homalium mandenense is or has been endemic to four littoral forests in a very small portion of the southeastern region of Anosy, near Toliara. The potential habitat has been declining in area and quality, but Mandena is now a protected area. The Extent of Occurrence is calculated as 72 km �� and the Area of Occupancy as 20 km ��. While some subpopulations are in protected areas, others are not, and their habitat is threatened by mining and wood harvest for firewood and charcoal. Therefore, the preliminary assessment of this species��� conservation status is ���Endangered��� [ENB1ab(iii)+2ab(iii)]. Notes. ��� Homalium mandenense is distinguishable from H. nudiflorum by its small leaves, racemes, and flowers. The calyx appears to increase in size less than in most species of the section, so that at maturity the flowers are markedly smaller than those of H. nudiflorum. Though the twigs are often strongly glaucous, the leaves are not; they are coriaceous in texture, with unusual strongly revolute margins. This species may also be confused with H. tenue, another smallleaved species newly described from southeastern Madagascar. Homalium tenue has frequently elliptical to obovate leaves with entire margins; its inflorescences are larger, and the outer surface of the calyx is densely pubescent. Both species have unusually short filaments and styles and small anthers, which in H. mandenense are also broader than usual. This suggests the possibility that they share a close relationship with H. louvelianum. Paratypi. ��� MADAGASCAR. Reg. Anosy [Prov.Toliara]: fkt. Ebakika, for��t d���Ampagnoronavoa, 24��43'46"S 47��09'26"E, 34 m, 6.IX.2012, fr., Ludovic 1649 (MO); Sainte Luce, pr��s du segment forestier S8, 24��46'19"S 47��08'55"E, 17 m, 20.X.2012, fl., Ramananjanahary et al. 826 (MO); Sainte Luce S7, 2.5 km au S d���Ambandrika for��t, 24��48'22"S 47��08'17"E, 15 m, 22.XI.2011, fl., Ratovoson 1719 (MO); sables dunaires anciens �� Mandromondromotra, au N de FortDauphin, 9.XII.1968, fl., Service Forestier 28645 (MO, P [2 sheets])., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 10-11, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766

4. Homalium intercedens Sleumer in Bull. Jard. Bot. Belg. 43: 300. 1973. Holotypus: MADAGASCAR. Reg. DIANA [Prov. Antsiranana]: plateau calcaire de l���Ankarana, �� l���E d���Ambondromifehy, 8.III.1951, fr., Service Forestier 3047 (P [P00375092]!; iso-: P [P04677610, P04677611]!). Tree to 25 m tall, 30 cm dbh; bark gray-white, exfoliating, platanoid; twigs dark brown, glabrous, leafing internodes ca. 0.7���1.3 mm diam.; stipules free, caducous. Leaves lanceolate, 3.4���6.7 �� 1.2���2.4 cm; base convex; apex acuminate; margin serrulate with round to elongated glands in tooth apices; secondary veins 10���12(���14) per side; both surfaces glabrous, drying dark brown with paler veins on adaxial surface, lighter to greenish on abaxial surface; petiole glabrous, 7���16 mm. Inflorescences racemose, axillary, 2.1���2.8 cm; rachis glabrous; bracts and bracteoles glabrous, ca. 1.5���2.5 mm; flowers borne singly. Flowers (5���)6-merous, yellowish-green; sepals narrowly oblanceolate to nearly ligulate, 6���8 mm in fruit, adaxial surface with appressed pubescence on calyx cup and sparsely pubescent on sepal bases, adaxial surface glabrous; sepal glands glabrous; petals deltoid-ovate, 2.5���3 mm in fruit, abaxial surface short-pubescent, adaxial surface glabrous; filaments glabrous, ca. 2 mm; anthers not seen; styles ca. 1.5���1.8 mm. Distribution, ecology and conservation status. ��� Homalium intercedens is known only from Ankarana in the DIANA region in the extreme north of Madagascar. It is reported on limestone and at a relatively low altitude. It is known from only one subpopulation (or perhaps two subpopulations close together) and rarely collected. Ankarana has been visited by many collectors over the years. TROPICOS (2020) has databased over 2100 historical and recent specimens from the area so the small number of collections is likely to indicate that the taxon is either narrowly endemic or uncommon within its range. However, the population is in a protected area that is known for inaccessible terrain, so it is at little risk of loss due to human activity. Therefore, its conservation status is tentatively assessed as ���Least Concern��� [LC]. Notes. ��� Homalium intercedens is easily distinguished from other species of sect. Nisa by its free stipules (caducous, but leaving two stipule scars rather than a single fused scar) and small lanceolate leaves with serrulate margins. Additional material examined. ��� MADAGASCAR. Reg. DIANA [Prov. Antsiranana]: Ambilobe, Mahamasina, RS d���Ankarana, chemin du canyon forestier, 12��55'22"S 049��06'27"E, 130 m, 16.I.2003, fr., Bardot-Vaucoulon et al. 1207 (P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 8-9, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["TROPICOS (2020). Missouri Botanical Garden, Saint Louis. [http: // www. tropicos. org]"]}

1. Homalium antilahimenae Wassel & Appleq., sp. nov. (Fig. 2). Holotypus: MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Fiv. Maroantsetra, comm. Ambinanitelo, fkt. Marovovonana, 15��19'41"S 49��31'29"E, 199 m, 29.VIII.2004, fl., Antilahimena 2621 (MO [MO-6755052]!; iso-: G!, K!, P!, TAN, USMS). Homalium antilahimenae Wassel & Appleq. differs from H. ciliolatum (Sleumer) Wassel & Appleq. in having shorter inflorescences with most flowers borne singly, smaller bracts and bracteoles, and less densely pubescent calyces with indument shorter on cup and sparser towards apical margins of sepals. Tree 10 to 15 m tall, 60 cm dbh; twigs dark to reddish brown when young, becoming gray to medium brown, glabrous, leafing internodes (0.9���)1.3���2.4(���4) mm diam.; stipules fused, caducous. Leaves elliptic to broadly or narrowly elliptic or obovate, (3.3 ���)5��� 8(��� 10.7) �� (1.8���)2��� 4.5(���5.4) cm; base cuneate to convex, often short-attenuate at extreme base; apex rounded to cuspidate (acute to slightly acuminate, retuse); margin entire, glands small or absent; secondary veins 7���11 per side; both surfaces glabrous, drying medium-brown on adaxial surface, darker on abaxial surface; petiole glabrous, 4���14 mm. Inflorescences paniculate to racemose, axillary, 2.5���8(���11.8) cm; rachis pubescent, often densely on apical portion; bracts and bracteoles 1���2.5(���3) mm, densely to sparsely short-pubescent throughout or mostly along midrib, often ciliate; flowers usually borne singly, rarely opposite or subopposite. Flowers 6���7(���8)-merous, pale yellow, yellow-green, or whitish; sepals oblanceolate to narrowly oblong-oblanceolate, narrowly elliptic, or in flower obovate to elliptic, 3���6 mm in flower, 5���8 mm in fruit, both surfaces densely to moderately pubescent at base, always densely so on outer surface of calyx cup, short-pubescent on basal portion or along midrib, more sparsely pubescent (to glabrous) towards apical margins; sepal glands glabrous; petals ovate (to broadly ovate when young), 2���3.5 mm in flower, to 5 mm in fruit, both surfaces velutinous; filaments glabrous, (1.2���) 1.9���2.4 mm; anthers 0.4���0.6 mm; styles 1���2 mm. Etymology. ��� Homalium antilahimenae is named for botanist Patrice Antilahimena, one of the most experienced living field botanists in Madagascar, who has made over 9500 collections, including the type, and additionally made valuable contributions to ecological and floristic studies (e.g., ANDREWS et al., 1998; PHILLIPSON et al., 2010). Vernacular names and use. ��� ���Jaborahoditra��� (Rakotonirina et al. 180); ���Mafaikoditra��� (Antilahimena et al. 1213); ���Maroampototsa��� (Antilahimena 2621); ���Tanampotsy��� (Service Forestier 15944). Wood is used to construct houses (Rakotonirina et al. 180). Distribution, ecology and conservation status. ��� Homalium antilahimenae is native to the northeastern regions of SAVA and Analanjirofo, where it usually occurs in low- to mid-elevation humid forests (with one specimen labeled at a range of 740���1200 m). It appears to be uncommon. As few as six distinct locations may be known; the Extent of Occurrence (EOO) is calculated as c. 11,190 km �� (which includes a significant area of open water) and the Area of Occupancy (AOO) as 32 km ��. Its habitat includes the protected area of Masoala, but other portions of the known habitat are more subject to ongoing anthropogenic degradation. A preliminary assessment of its conservation status is therefore ���Vulnerable��� [VU B1ab(iii)+2ab(iii)]. Notes. ��� Homalium antilahimenae is part of a group of three species, including H. ciliolatum (Sleumer) Wassel & Appleq. and H. tenue Wassel & Appleq., that are separated from H. nudiflorum s.str. by more pubescent flowers and inflorescences. Homalium ciliolatum has often longer inflorescences, 3.5���11.5(���20) cm, larger bracts and bracteoles, 3.5���5(���7) mm long, and sepal indument that covers the entirety of both surfaces, with a markedly longer sericeous indument on the calyx cup. Homalium tenue is distinguished by smaller sepals, 3.5���5(���6) mm in fruit with a glabrous adaxial surface, less pubescent petals, usually smaller leaves, and slender-graceful twigs and rachises. Paratypi. ��� MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Maroantsetra, Anjahana, Amodivangy, 15��25'47"S 49��50'07"E, 200 m, 19.VII.2002, fl., Antilahimena et al.1213 (G, MO); Maroantsetra, Ambanizana, NP Masoala, 15��37'48"S 49��59'03"E, 473 m, IX.2002, fl., Antilahimena 2911 (MO); Anefitrakely private forest, 15��50'04"S 49��40'05"E, 128 m, 28.VII.2007, fl., Antilahimena 5763 (G, K, MO); Ambodimangavolo, Mahasoa, Ambatoharanana, 17��33'57"S 48��53'45"E, 740���1200 m, 25.X.2000, fl., Ratovoson et al. 327 (G, MO); Farankaraina, Maroantsetra, 24.VII.1956, fl., Service Forestier 15944 (P); env. de la Baie d���Antongil, for��t de Fahampanambo, dans la basse vall��e de l���Antanambalana, 25���26.IX.1957, fl., Service Forestier 18278 (P [2 sheets]). Reg. SAVA [Prov. Antsiranana]: Sambava, Marogaona, Ambodivapaza, 14��08'20"S 49��55'27"E, 378 m, 27.VIII.2013, Rakotonirina et al. 180 (MO)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 4-5, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["ANDREWS, J., P. ANTILAHIMENA & C. R. BIRKINSHAW (1998). Use of a day resting box by a wild sportive lemur, Lepilemur dorsalis, on Nosy Be, north-western Madagscar. Folia Primatol. 69 (Suppl. 1): 18 - 21.","PHILLIPSON, P. B., P. P. LOWRY II, L. ANDRIAMAHEFARIVO, P. ANTILAHIMENA & C. BIRKINSHAW (2010). Floristic inventory of the Ambatovy-Analamay mine site and comparison to other sites in Madagascar. Malagasy Nat. 3: 44 - 76."]}

11. Homalium tenue Wassel & Appleq., sp. nov. (Fig. 7). Holotypus: MADAGASCAR. Reg. Anosy [Prov. Toliara]: Comm. Iabokoho, fkt. Antsontso, Ivohibe forest, 24��34'10"S 47��12'37"E, 41 m, 24.V.2006, fl., Antilahimena et al. 4858 (MO [MO-6396271]!; iso-: P [P06161693]!, TAN, TEF). Homalium tenue Wassel & Appleq. differs f rom H. antilahimenae Wassel & Appleq. in having usually narrower leaves with shorter petioles, racemose (to racemiform) inflorescences with rachis not densely pubescent and bracts not pubescent on most of surface, and smaller flowers with sepals adaxially glabrous. Tree to 16 m tall, 8 cm dbh; twigs gray to light brown, glabrous, leafing internodes 0.8���1.6(���2.3) mm diam.; stipules fused, caducous. Leaves elliptic to narrowly elliptic, obovate, or oblanceolate (very rarely a few lanceolate), (2.5���)3���6(���10) �� 1.3���2.5(���3) cm; base cuneate to slightly attenuate; apex cuspidate to short-acuminate, acute, or rounded; margin entire, glands absent or very few; secondary veins 7���10(���12) per side, intersecondary veins sometimes similar; both surfaces glabrous, drying brown, adaxial surface usually paler; petiole glabrous, 3���7(���9) mm. Inflorescences racemose (racemiform panicles), axillary, (2���)3���7(���8) cm; rachis minutely pubescent (to glabrous), markedly thin; bracts and bracteoles pubescent on midrib, ciliate at margins or glabrous, 1���3 mm; flowers borne singly (or partly paired, the second flower probably being borne on a highly reduced branch). Flowers 5���7-merous, yellowish green, greenish white, or white; sepals in fruit narrowly oblong to oblanceolateoblong, 3.5���5(���6) mm, abaxial surface densely pubescent on calyx cup and sepal bases with indument sometimes extending up the medial portion, elsewhere glabrous, adaxial surface glabrous; sepal glands glabrous; petals oblong-ovate to elliptic or lanceolate, 1.5���2.5 mm, abaxial surface densely pubescent (to moderately pubescent, sparsely pubescent near margins), adaxial surface densely to sparsely pubescent; filaments glabrous, 0.9���1.1 mm; anthers 0.2���0.3 mm; styles 0.5���1 mm. Vernacular names and use. ��� ���Fotsiakaraminty��� (Ludovic 1074); ���Fostiakora mena��� (Ludovic 1061); ���Hazofotsy��� (Service Forestier 9742); ���Hazomby��� (Ludovic 1074); ���Zora��� (Service Forestier 9742). Wood is used for construction (Ludovic 1074). Distribution, ecology and conservation status. ��� Homalium tenue is found in low-elevation forests in the southeastern regions of Anosy and Atsimo-Atsinanana; it is reported on sand and laterite. Although the localities in Anosy are variously described, all come from a single small region, so that not more than five distinct subpopulations are known. The Extent of Occurrence and Area of Occupancy are calculated including an unusual collection that may be of hybrid origin (Service Forestier 23622, discussed below) on the grounds that if it is a hybrid, the parental species is likely also to be present in the forest in question. The Extent of Occurrence (EOO) is calculated as c. 908 km �� and the Area of Occupancy as 28 km ��. Habitat is largely unprotected and because low-elevation forest is easily accessed, it is very vulnerable to anthropogenic damage from wood harvest and forest clearing. A preliminary assessment of its conservation status is therefore ���Endangered��� [EN B1ab(iii)+2ab(iii)]. Notes. ��� Homalium tenue overlaps in distribution with H. nudiflorum, which has sometimes similar leaves. Homalium tenue differs in the denser indument on the petals and calyx cup, usually extending to the basal medial portion of the sepals, and its flowers are smaller; its slender twigs and inflorescences give it a more delicate appearance than H. nudiflorum, though it is reportedly a sometimes large tree. The leaves are usually quite small (the inclusion of Service Forestier 23622, which in most other features is consistent with the species, significantly increases the leaf size range). Collections from AtsimoAtsinanana have more prominently acuminate and often larger leaves than those from Toliara. Two other species, H. antilahimenae and H. ciliolatum, have dense pubescence on both petals and portions of the calyx; these two and H. tenue are suspected to be a natural group. Those two species have at least some indument on both sepal surfaces, with longer indument on the calyx cup, and their flowers are larger at maturity; they lack the gracile appearance of H. tenue. Homalium ciliolatum, which overlaps in geographic range, has usually larger and broader leaves, flowers usually borne in pairs, and large pubescent bracts and bracteoles; the calyx cup is long-sericeous and the sepals pubescent throughout on both surfaces. Homalium antilahimenae is native to northern Madagascar and has often larger leaves with some marginal glands, often densely pubescent rachises, and bracts and bracteoles often pubescent throughout. Hybridization between Homalium tenue and H. louvelianum has been observed (see under the latter species above). Paratypi. ��� MADAGASCAR. Reg. Anosy [Prov. Toliara]: Bemangidy Forest, 65 km N of Ft. Dauphin, 24��34'05"S 47��12'38"E, 100 m, 10.II.2006, fr., Lowry et al. 6718 (MO); Fort-Dauphin, Iabakoho, 24��35'33"S 47��12'52"E, 22.V.2006, fr., Rajoharison et al. 166 (MO); Anosy, Taolagnaro, Iaboko, 24��34'16"S 47��12'05"E, 271 m, 1.IV.2008, fr., Razakamalala et al. 4065 (MO); for��t Andrazato, Ft. Dauphin, 5.IV.1954, fr., Service Forestier 9742 (P); for��t de Bemangidy, au N de Mahatalaky (Ft. Dauphin), 100 m, II.1995, fr., Service Forestier 11797 (MO, P). Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: Vangaindrano, Matanga, 23��30'57"S 47��29'33"E, 96 m, 18.IV.2008, fr., Ludovic 1061 (MO); ibid. loco, fr., Ludovic 1074 (MO); Mananivo, 40 km SW Farafangana, for��t Iabomary, 23��03'34"S 47��40'23"E, 0���50 m, 6.II.2001, buds, Rabenantoandro et al. 430 (G, K, MO, P); S de Farafangana rte de Manombo, aux P.K. 20��� 21, 14���17.X. 1964, fl., fr., Service Forestier 23622 (P [2 sheets])., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 20-22, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766

9. Homalium retusum (Blume) Wassel & Appleq., comb. nov. ≡ Nisa retusa Blume, Mus. Bot. Lugd. -Bat. 2: 28. 1856. Lectotypus (designated here): MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Ile Ste. Marie, VIII–IX, fl., Bernier 173 (L [L0010974]; isolecto-: K [K000231491, K000231492] images seen, L [L0010975, L0010976] images seen, P [P00418083, P00418084, P00418085]!, TAN [TAN000593] image seen). = Homalium scleroxylon (Tul.) Baill. in Bull. Mens. Soc. Linn. Paris 1: 575. 1886. ≡ Nisa scleroxylon Tul. in Ann. Sci. Nat., Bot. sér. 4, 8: 69–70. 1857. Lectotypus (first step designated by SLEUMER, 1973: 299; second step designated here): MADAGASCAR. Reg. Analanjirofo [Prov. Toamasina]: Ile Ste. Marie, VIII – IX, fl., Bernier 173 (P [P00418085]!; isolecto-: K [K000231491, K000231492] images seen, L [L0010974, L0010975, L0010976] images seen, P [P00418083, P00418084]!, TAN [TAN000593] image seen). Syntypus: Galdich. [Gaudichaud] 296 [in phytotheca Jalbertiana] (not located). Tree 6–7 m tall, 10 cm dbh; twigs gray to medium brown, glabrous, leafing internodes 1.6–3.5(–4.5)mm diam.; stipules 2– 6 mm, sparsely ciliate. Leaves obovate to broadly elliptic, (5–)7–16 × 3.5–9 cm (unusually small leaves sometimes subtending inflorescences); base cuneate to convex; apex rounded to retuse or cuspidate; margin entire, rarely slightly crenulate, glands small; secondary veins (5–)6 –9(– 14) per side; both surfaces glabrous, drying light to medium brown on adaxial surface and darker on abaxial surface; petiole glabrous, (10–) 15–30 mm. Inflorescences paniculate, axillary, 8 –12 cm; rachis short-pubescent; bracts and bracteoles 1–3 mm, usually ciliate, otherwise glabrous to sparsely pubescent; flowers mostly paired with 2 per bract. Flowers 5–6-merous, white or yellow; sepals obovate at anthesis becoming oblanceolate, 4–8(– 9) mm, adaxial and abaxial surfaces glabrous; sepal glands glabrous; petals oblong-ovate, 1.5–3 mm, adaxial surface usually pubescent apically, basally glabrous, abaxial surface glabrous except often pubescent at extreme apex, margin usually ciliate; filaments glabrous, 1.2–1.9 mm; anthers (0.3–) 0.35–0.45 mm; styles (0.7–)1.2–1.7(–2) mm. Distribution, ecology and conservation status. – Homalium retusum is found near the eastern coast in the northern part of Madagascar at low elevations. Few specimens are known, suggesting that this is an uncommon to rare species. Only five clearly distinct subpopulations have been recorded (Ile Ste.-Marie in Analanjirofo, Foulpointe and Ambila-Lemaitso in Atsinanana, and Anjana and Andempona in SAVA). The Extent of Occurrence (EOO) is calculated as c. 9847 km ² and the Area of Occupancy (AOO) as 28 km ². Much of the habitat is subject to ongoing anthropogenic damage, especially in view of its relative accessibility. Therefore, a preliminary assessment of its conservation status is Endangered [EN B1ab(iii)+2ab(iii)]. Vernacular names and use. – “Hazombato” (Service Forestier 2587); “Longotra” (Service Forestier 9646); “Marangué Touditch” (Bernier 173, Chapelier s.n.); “Voapak” (Humblot 63); “Zana” (Service Forestier 4918); “Zanakanivato” (Service Forestier 2597). Wood is good for construction (Bernier 173). Notes. – Homalium retusum has previously been lumped into H. nudiflorum s.l. (SLEUMER, 1973), which as herein circumscribed, differs in having smaller, narrower, shorterpetioled leaves; its inflorescences are usually racemose, not over 9 cm, glabrous or sparsely pubescent, with flowers borne singly. Homalium retusum most closely resembles H. pachycladum: both have relatively large leaves, paniculate inflorescences, and flowers with glabrous sepals and near-glabrous petals. Homalium retusum is known only from low-altitude and littoral forests, while H. pachycladum occurs farther inland and at much higher elevations. Homalium pachycladum has elliptic to obovate (or rarely ovate) leaves with shorter petioles (6–15 mm) and flowers all borne singly; its twigs seem usually to be thick, giving a robust appearance. Another inland species with large leaves, H. confertum, is distinguished by having usually elliptic to oblong-elliptic leaves with a serrate to crenate margin, racemose inflorescences with flowers borne singly, and petals densely pubescent on most of both surfaces. The history of the names applying to this species resembles that of Homalium planiflorum (Boivin ex Tul.) Baill. (Blackwellia planiflora Boivin ex Tul.) versus Blackwellia gracilis Blume (see APPLEQUIST, 2017, 2018b), but without disruptive nomenclatural consequences. Blume published Nisa retusa based on material of Bernier 173 a year before Tulasne published N. scleroxylon based mostly on the same collection. Blume specified that his name was based only on material sent from P to L. That material was described in the protologue (BLUME, 1856 –1857: 28) as “Nomine Asteropeia et Mourangia Touditsch ex Herb. Mus. Paris. mecum benevole communicata.” Three sheets at L bore labels initially marked “ Asteropeia [or Asteropeja] multiflora ” then “ Mourangia [or Morangia] touditch ” (a vernacular name, though not identified as such on those sheets); Blume later added “ Nisa retusa ” in the top margins. Much more recent pink typed labels identify these sheets as being type material of N. scleroxylon, “leg. Bernier 173 ”. One year later, Tulasne published N. scleroxylon, citing “Bernerii herb., n. 173; Galdich. herb., n. 296, in phytotheca Jalbertiana” (TULASNE, 1857: 70). These are Latinizations for Bernier and Gaudichaud-Beaupré respectively. Although Tulasne presumably used only those duplicates of Bernier 173 to be found in France, he did not say so. The protologue (TULASNE, 1857: 70) further states: “ Marangue-Touditch (1), auctore Bernerio, vernaculare audit…”, explaining the labeling seen, and apparently misinterpreted, by Blume. Three duplicates of Bernier 173 exist at P; two were labeled as having the vernacular name “ Maranguá touditch ” and one had initially been labeled “ Asteropeia ”. Though the sheets sent to L were not initially labeled as Bernier 173, they can be confidently assigned to that collection. SLEUMER (1973: 299) referred to Bernier 173 (L) as the “ holotype ” of Nisa retusa and to Bernier 173 (P) as the “ holotype ” of Nisa scleroxylon. For N. retusa, the original material was already restricted to the three sheets at L, which are syntypes, and Sleumer made no choice between them. The sheet that appears most complete is here chosen as a lectotype. For N. scleroxylon, Sleumer’s statement constitutes a first-step lectotypification (Art. 9.17 of ICN; TURLAND et al., 2018). This action means that N. retusa and N. scleroxylon are not nomenclatural synonyms, since their types are different specimens, even though they are from the same gathering. The best of the sheets at P is here chosen as a second-step lectotypification. Additional material examined. – MADAGASCAR. Reg. Atsinanana [Prov. Toamasina]: Foulpointe?, s.d., fl., Chapelier [Herb. Boivin] s.n. (P [2 sheets]); Foulpointe, Morarano, Andohanakoho, 17°41'52"S 49°27'28"E, 34 m, 9.III.2005, fl., Lehavana et al. 304 (MO, P); Foulpointe, forêt d’Analalava, 17°42'00"S 49°26'00"E, 35 m, 19.V.2004, fl., Randrianarivelo et al. 2 (MO, P); Sahapanpana, Foulpointe, 20.VI.1950, fl., Service Forestier 2587 (P); Ambila-Lemaitso, 1.II.1952, fr., Service Forestier 4918 (P); Ambila-Lemaitso, 27.IV.1954, fl., Service Forestier 9646 (P). Reg. SAVA [Prov. Antsiranana]: W de Nosiarina, sommet d’Anjana, 14°12'23"S 50°04'26"E, 190 m, 29.VIII.2007, fr., Andriamihajarivo et al. 1341 (MO); Antalaha, Andempona, 13.VI.1950, fl., Service Forestier 2597 (MO, P). Sine loco: Madag. bor., Andahoul, s.d., fr., Humblot 63 (P).

3. Homalium confertum Baker in J. Linn. Soc., Bot. 21: 341. 1884 (Fig. 1). Lectotypus (designated by SLEUMER, 1973: 298): MADAGASCAR: ��� Central Madagascar ���, s.d., fr., Baron 3185 (K [K000231489] image seen; isolecto-: BM, L [L0010973] image seen, P [P00418088]!). Syntypus: MADAGASCAR: ���Central Madagascar���, s.d., fr., Baron 3256 (K [K000231490] image seen). Tree to 22 m tall, 60 cm dbh; bark exfoliating in plaques; twigs gray to medium brown, glabrous, leafing internodes (1.2���)1.7���4(���5)mm diam.; stipules fused, sometimes persistent. Leaves elliptic to oblong-elliptic, narrowly or broadly elliptic, or obovate, (5���)6��� 16 �� (1.8���)3��� 6.2(���7) cm; base convex to cuneate; apex cuspidate, obtuse, acute, or rounded (shortacuminate); margin serrate to dentate or slightly crenate, rarely nearly entire, glands in tooth apices elongated and relatively conspicuous; secondary veins 8���14(���16) per side, sometimes with similar intersecondary veins; both surfaces glabrous (aberrantly bearing patches of short trichomes, possibly pathogenic), or abaxial surface moderately glaucous, drying medium to dark brown on both surfaces (rarely light green); petiole 3���7(���10) mm, glabrous. Inflorescences racemose, axillary (often paired), (2���)4���8.5(��� 11) cm; rachis glabrous (sparsely shortpubescent or more densely so in irregular patches); bracts and bracteoles (1.5���) 2���4 mm, glabrous except margins of bracts usually ciliate; flowers borne singly. Flowers 5���6-merous, white to green or yellow-green; sepals 3���7 mm in flower, 8���12.5 mm in fruit, oblanceolate to spatulate or narrowly oblanceolateoblong, adaxial surface usually glabrous (midrib sometimes sparsely short-pubescent), abaxial surface usually glabrous (sparsely short-pubescent at base); sepal glands yellow, glabrous; petals ovate, 2���4 mm, both surfaces densely (to moderately) pubescent especially on apical half; filaments glabrous (sparsely pubescent on the basal portion), 1.7���3.1 mm; anthers (0.45���) 0.5���0.6 mm; styles 1.5���2.5 mm. Vernacular names and use. ��� ���Hazombato��� (Razafindramora 1, Ser vice Forestier 1776, 25277); ���Hazombatofotsy ��� (Razanatsima et al. 1136, Service Forestier 7362, 16801, 25277); ���Lalondalitra��� (Service Forestier 15406); ���Longotra-fotsy��� (Service Forestier 1249); ���Marankoditra��� (R��serves Naturelles 6043, Service Forestier 25277); ���Tsianihiposa��� (Service Forestier 10148). Wood is used to make unspecified products (Service Forestier 25277). Distribution, ecology and conservation status. ��� Homalium confertum is widespread in eastern and central regions of Madagascar. It usually occurs in humid forests at mid- to high elevations (range 445���1860 m), but also extends into relatively dry western forests (Ankarafantsika). Reported substrates include gneiss. It is sometimes common and locally dominant (Randrianaivo et al. 526). It has been collected in several protected areas, including Ambohitantely, Anjanaharibe-Sud, Ankarafantsika, Montagne d���Ambre, and Zahamena. Its conservation status is estimated as ���Least Concern��� [LC]. Notes. ��� Homalium confertum was lumped into H. nudiflorum by SLEUMER (1973). It differs from H. nudiflorum s.s. in its larger, often strongly toothed leaves with many marginal glands and its more densely pubescent petals; the final perianth size may also be larger. The two overlap in range, but H. confertum is more tolerant of dry climate and high altitude, with much of its habitat on the central plateau. It is similar to H. pachycladum and H. retusum, which also overlap in range, in maximum leaf size, but H. pachycladum and retusum are distinguished by entire leaf margins, fewer leaf glands, and petals glabrous on the abaxial surface. Several possible hybrids between H. confertum and H. nudiflorum are seen in two regions where both are known, and gene flow is suspected to be responsible for some of the range of variation in H. nudiflorum; these are discussed under the latter species. Hybridization is seen in most sections of Homalium, even rarely among species of different sections (APPLEQUIST, 2018b), so the ability of these species to hybridize does not suggest that they should not be recognized as distinct, given their significant morphological and ecological differences. Additional material examined. ��� MADAGASCAR. Reg. Alaotra-Mangoro [Prov. Toamasina]: PN Zahamena, 17��38'25"S 48��38'34"E, 989 m, 30.III.2000, buds, Ratovoson et al. 184 (MO, P); Manaka, Ambatondrazaka, 20.II [?].1953, fl., Reserves Naturelles 6043 (MO); P. K. 39 route d���Anosibe, canton and souspr��fecture Anosibe, 4.V.1965, fr., Service Forestier 25277 (P). Reg. Amoron���i Mania [Prov. Fianarantsoa]: commune d���Ambalamanakana, 20��44'01"S 47��11'42"E, 1860 m, 27.III.1996, fl., Rakotomalaza et al. 681A (MO, P). Reg. Analamanga [Prov. Antananarivo]: Tampoketsa-Ankazobe, RS Ambohitantely, 18��11'52.5"S 47��17'3"E, 1620 m, 7.III.2004, fl., Almeda et al. 8636 (MO); Anjozorobe, commune d���Ambongamarina, Andreba, 18��16'55"S 47��54'24"E, 1400 m, 11.III.2000, fl., Randrianaivo et al. 526 (MO); Ambohitantely RS, 18��11'46"S 47��17'24"E, 1620 m, 12.III.1997, Ravololonanahary et al. 69 (MO). Reg. Analanjirofo [Prov. Toamasina]: Beanana, 700 m, 22.II.1954, fl., Service Forestier 9058 (MO, P [2 sheets]). Reg. Anosy [Prov. Toliara]: Antsotso Avaratra, for��t Tsitongambarika, 24��34'S 47��12'E, 2.IV.2008, fr., Razakamalala et al. 4148 (MO, P). Reg. Atsinanana [Prov. Toamasina]: Ambodilendemy, Ankerana, 18��25'49"S 48��47'14"E, 890 m, 24.III.2011, fl., Antilahimena 7778 (MO); Ambodilendemy, vers le sommet d���Ankerana, 18��25'49"S 48��47'14"E, 900m, 14.III.2011, fl., Ravelonarivo & Felix 3663 (MO); Vatomandry, Ambalabe, 19��09'27"S 48��34'57"E, 596 m, 11.III.2013, fl., Razanatsima et al. 1136 (BR, G, K, MO, USMS). Reg. Boeny [Prov. Mahajanga]: R��serve Naturelle Int��grale d���Ankarafantsika, 16��18'00"S 46��49'00"E, 22.IV.1994, fl., Rajemisa 24 (MO). Reg. Bongolava [Prov. Antananarivo]: Tsiroanomandidy, Ambaravaranala, 18��25'S 45��40'E, 1190���1200 m, 25.VI.1997, fl., Rakotomalaza et al. 1353 (G, MO). Reg. DIANA [Prov. Antsiranana]: Montagne d���Ambre (partie N.O.), 1000���1800 m, 23.XII.1967, fr., Bernardi 12003 (MO, P); contreforts occidentaux du Massif de Marojejy (NE) pr��s du col de Doanyanala (limite des bassins de la Lokoho et de l���Andraronga), 800���1200 m, 25.I���25. II.1949, fl., Humbert 23040 (P [3 sheets]); montagnes au N de Mangindrano (haute Maevarano) jusqu���aux sommets d���Ambohimirahavavy (partage des eaux Mahavavy-Androranga), 1500 m, 19.I���12.II.1951, fr., Humbert & Capuron 24888 (P); flanc S du M. Tsaratanana, 1700 m, XII.1912, fl., Perrier de la B��thie 6705 (P). Reg. Haute Matsiatra [Prov. Fianarantsoa]: Ampamaherana, Fianarantsoa, 13.XII.1949, fl., Service Forestier 1249 (P). Reg. SAVA [Prov. Antsiranana]: Daraina, for��t d���Antsahabe, 13��13'03"S 49��32'46"E, 780 m, 2.XII.2004, fr., Gautier 4806 (MO, P); Ampasindava, for��t de Bongomihirahavavy, 13��46'11"S 48��04'58"E, 445 m, 4.XII.2013, fr., Gautier et al. 6092 (G, P); massif de l���Anjanaharibe (pentes et sommet N) a l���W d���Andapa (haute Andramonta, bassin de la Lokoho: NE), 1600���1800 m, 10.XII.1950 ��� 3.I.1951, fl., Humbert et al. 24704 (P); Doany, Anjialavahely, 14��17'41"S 49��32'50"E, 949 m, 15.III.2006, fl., Rakotovao et al. 2986 (MO, P); Anjialavabe, Ankiakabe, 14��10'47"S 49��22'01"E, 1567 m, 23.II.2007, fl., Ravelonarivo et al. 2238 (MO, P); Antalaha, Bealampona, village de Befingotra, 14��45'11"S 049��28'49"E, 762 m, 21.IV.1997, fl., Razafindramora 1 (MO). Reg. Sofia [Prov. Mahajanga]: Marsoandakana, 14��55'39"S 49��25'52"E, 1176 m, 23.II.2008, fl., Bernard et al. 876 (MO); NW du campement Bemafo, 14��13'17"S 49��03'55"E, 1830 m, 26.X.2005, fl., Buerki et al. 106 (MO, P). Reg. Vakinankaratra [Prov. Antananarivo]: Angodona, Tsinjoarivo, 6.VI.1950, fl., Service Forestier 1766 (P [2 sheets]). Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa]: Ampasinambo, Nosy-Varika, 21.IV.1954, fl., fr., Service Forestier 10148 (MO, P)., Published as part of Wassel, Anna C. & Applequist, Wendy L., 2020, A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar, pp. 1-23 in Candollea 75 (1) on pages 6-8, DOI: 10.15553/c2020v751a1, http://zenodo.org/record/5724766, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","APPLEQUIST, W. L. (2018 b). A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae). Candollea 73: 221 - 244."]}

7. Homalium fortunatii Appleq., spec. nova (Fig. 2). Holotypus: MADAGASCAR. Prov. Toamasina: Vatomandry, Ambalabe, along trail to Ambatobe, a deviation from Ambalabe-Sahanionaka trail, 19°18'17"S 48°37'22"E, 302 m, 4.II.2011, fl. & fr., Randrianasolo et al. 1358 (MO-6703445!; isotype: G!, P!; TAN). Homalium fortunatii Appleq. differs from H. thuarsianum (Tul.) Baill. in having minutely pubescent young twigs; elliptical to narrowly elliptical (to broadly elliptical or obovate) leaves, (3.4–)7.5–13 × (1.4–) 3.5–5.3 cm, with base convex (to rounded) and apex cuspidate with rounded tip (to rounded); and sepals and petals slightly shorter than, to slightly longer than, the calyx tube. Tree to 12–15 m; young twigs minutely pubescent becoming glabrous. Leaves elliptical to narrowly elliptical (broadly elliptical to obovate in abnormally small leaves), (3.4–)7.5–13 × (1.4 –) 3.5 – 5.3 cm; petiole 2 – 6.5 mm; base convex (to rounded), often attenuate at extreme base; apex cuspidate with rounded tip (rounded); margins shallowly repand, with glands small, often slightly protruding; adaxial surface drying dark brown, sometimes gray-tinged, abaxial surface brown. Inflorescences paniculate, lateral, sometimes cauliflorous on small twigs, 5–11.5 cm, pubescent; flowers pedicellate with pedicels 0.7– 2 mm; bracts lanceolate to ovate, 1.8–3.5 mm (Flowers 7–9-merous; calyx tube funnelform with rounded base, erect-pubescent; sepals ligulate (to narrowly lanceolate) with acute (to rounded) apex, 2.1–2.5 mm; petals ligulate with acute (to rounded-acute) apex, 1.8–2.7 mm; sepals and petals post-flowering pinkish white or reddish, slightly shorter than calyx tube to slightly longer, ascending, ciliate with long soft hairs, abaxial surface sparsely pubescent, adaxial surface sparsely villous; sepal glands pubescent; filaments 0.6–0.7 mm, basally villous; ovary villous; styles 4–5, 0.6–0.8 mm, basally villous. Etymology. – Homalium fortunatii is named for botanist Fortunat Rakotoarivony; he was among the collectors of the type specimen and is a strong advocate of conservation and botanical research in the region of its collection. Vernacular names and use. – “ Hazombatofotsy ” (Randrianasolo et al. 1358); “Vazanaomby” (Randrianasolo et al. 1358). Wood of H. fortunatii is used for construction (Randrianasolo et al. 1358). Distribution, ecology and conservation status. – Homalium fortunatii is known from two low-elevation humid forests in Toamasina. Because of its very small known Area of Occupancy (only 8 km ²) its conservation status may be estimated as “Vulnerable” [VU D2]. Notes. – Homalium fortunatii is among the H. thuarsianum (Tul.) Baill. group of species with paniculate inflorescences and pubescent sepal glands. Homalium thuarsianum has glabrous young twigs, usually oblanceolate to obovate (to oblongoblanceolate or elliptical leaves) with cuneate to convex bases and variable (often rounded, but rarely rounded-cuspidate) apices, and sepals and petals that are consistently longer than the calyx tube when fully developed. Material of H. fortunatii has been misidentified as H. erianthum, which likewise has relatively large leaves and paniculate inflorescences. Homalium erianthum has glabrous sepal glands and much longer stamens, and is confined to Anstiranana. Paratypus. – MADAGASCAR. Prov. Toamasina: Masoala Peninsula, trail just above Antalavia River on S bank, 1–2 km from coast, 15°45'S 50°02'E, 50–100 m, 19.II.1988, fl. & fr., Schatz et al. 1928 (MO, P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on page 232, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

1. Homalium acuminatifolium Appleq., spec. nova (Fig. 1). Holotypus: MADAGASCAR. Prov. Toliara: forêt au S d’Eminiminy, Parcelle I dans la RNI d’Andohahela, 24°39'00"S 46°48'30"E, 0 – 500 m, 10 – 26.XI.1993, fl., Randriamampionona 682 (MO-6087923!; isotype: MO-6087899!, P [P04735190]!, TAN). Homalium acuminatifolium Appleq. differs from H. axillare (Lam.) Benth. in having lanceolate (to narrowly lanceolate or ovate), acuminate leaves. Tree to 15 m, 30 cm dbh; young twigs glabrous. Leaves lanceolate to narrowly lanceolate (ovate), 4–7.3 × (1.1–)1.3 –2.1(–3) cm; petiole 3– 8 mm; base convex (rounded); apex long-acuminate; margins shallowly serrate, with glands small, usually protruding at apex of tooth; adaxial surface drying grayish to blackish, greenish or darker brown, abaxial surface pale brown to brown. Inflorescences racemose, lateral, (2 –) 4 – 10 cm, short-pubescent; flowers pedicellate with pedicels (0.3–)0.5 – 1.4(– 2.3) mm; bracts broadly elliptical to broadly ovate, 0.4– 1 mm. Flowers 9-merous; calyx tube short with a rounded base, sparsely pubescent; sepals ligulate with acute apex, 1.3–2.3 mm; petals ligulate with acute apex, 1.4– 2.3 mm; sepals and petals yellowish-white, probably somewhat accrescent, usually much longer than calyx tube, often spreading and often becoming somewhat reflexed after flowering, densely ciliate with long trichomes, both surfaces sparsely pubescent or adaxial surface sparsely hispid; sepal glands glabrous; filaments 1.7–2.6 mm, sparsely pubescent; ovary densely hispid to pilose; styles 4–5, 1.3–2 mm, basally hispid to pilose. Vernacular names. – “Hazofotsy” (Réserves Naturelles 2098); “Zôra” (Randriamampionona 682, Réserves Naturelles 2098, 6733). Distribution, ecology and conservation status. – Homalium acuminatifolium is native to low- to mid-elevation humid forests in the southeastern part of the province of Toliara; it sometimes occurs along waterways and on laterite and granite. GeoCAT estimates the Extent of Occurrence at c. 791 km ² and the Area of Occupancy at 28 km ²; there are seven distinct locations, though four of these are within a few km of one another. The species does occur in the protected area of Andohahela, but its habitat outside Andohahela is threatened with continuing degradation. Therefore a conservation status of “Vulnerable” [VU B1ab(iii)+B2ab(iii)] would seem to be warranted. Note. – Sleumer determined available specimens of this taxon as H. axillare (Lam.) Benth. “ forma foliis ovato-lanceol.” The flowers resemble those of H. axillare but the leaves are distinctly different in shape, being usually lanceolate (to narrowly lanceolate or ovate) with a long-acuminate apex. Leaf margins are shallowly serrate and the small marginal glands usually protrude from tooth apices. The racemes are sometimes very numerous and crowded and the petals and sepals are more commonly spreading at anthesis, though these characters are variable. Paratypi. – MADAGASCAR. Prov. Toliara: à la base et en avant de la forêt de Manantantely près de Fort-Dauphin, 50–300 m, 1.III.1947, fl., Humbert 20373 (P); Farafara Vatanibe, W du village de Malamba, au bord de la Belomotse, 24°51'02"S 47°00'20"E, 135 m, 14–18.XI.2009, post-fl., Rakotovao et al. 4517 (MO); SW de la forêt d’Ankiboriabo, Marovato, parcelle I d’Andohahela RNI, 24°36'S 46°50'E, 250–750 m, 14.XI.1993, fl. & fr., Randriamampionona 659 (BM, BR, CANB, CAS, G, K, L, M, MO, P, PRE, S, US, USMS); Magatisaka, 24°35'07"S 47°02'15"E, 669 m, 21.XI.2009, fl., Razakamalala 4774 (MO); Ifarantsa, Fahibe, 11.XII.1950, fl. & fr., Réserves Naturelles 2098 (P); FortDauphin, Naniliha [Enaniliha], 15.IX.1954, Réserves Naturelles 6733 (P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on page 225, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

Homalium microphyllum O. Hoffm., Sert. Pl. Madagasc.: 18. 1881. This species was described from a single collection (Hildebrandt 3329 from Vavatobé [probably = Ambavatoby, Antsiranana]), which has many duplicates; no similar material has been collected since. Though the collected material seems to be well into flower, the sepals and petals are small and often spreading, apparently scarcely accrescent, and the ovary is not strongly conical. In these characters, it resembles the species transferred to Homalium sect. Odontolobus . However, it also has characters more consistent with Homalium sect. Blackwellia sensu stricto; it appears that there is usually only one bract per flower, the calyx tube is cylindrical and well-developed, the upper surface of the ovary is not particularly broad, and the styles are much longer than in most species of Homalium sect. Odontolobus . It is suggested that the collected individual was an intersectional hybrid that is not known to persist and does not merit taxonomic recognition., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on page 242, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

2a. Homalium axillare subsp. axillare Tree to 16 m, 26 cm dbh, or large shrub. Leaves elliptical to broadly elliptical (seldom to ovate or obovate), (2.5–)3.6–7.5 (–9.3) × 1.5–3.8(–4.3) cm; petiole 4–8 mm; margins crenateserrate (crenulate, subentire), with glands usually elongated in tooth apices; adaxial surface drying darker, brown (greenish), abaxial surface brown (greenish). Inflorescences (2.6–) 4–14.5 cm. Filaments (1.3–) 2.1–3.5 mm. Distribution, ecology and conservation status. – Homalium axillare subsp. axillare is found in mid-elevation to moderately high-elevation humid forests in Toamasina and surrounding provinces; it is once reported on crust. It is relatively common and is found in protected areas including Analamazaotra, Zahamena, and Ranomafana; therefore a preliminary assessment of its conservation status is “Least Concern” [LC]. Vernacular names. – “Hazombato” (Service Forestier 1047, 1391, 1603, 6628); Iasiravina (Service Forestier 14394); “Marakoditra” (Service Forestier 7551); “Maranikoditra” (Razanatsoa & Marcellin 234); “Marankoditra” (Louvel 63, Perrier de la Bâthie 6794, Razanatsoa et al. 45, Service de Colonisation 42, Service Forestier 1603, 4841, Thouvenot 42); “Maroankoditra” (Antilahimena et al. 3217, Antilahimena & Félix 3158, Rakotonandrasana et al. 608). Selected material examined. – MADAGASCAR. Prov. Antananarivo: 10 km SE de Tsinjoarivo, forêt de Mahatsinjo, 19°40'86"S 47°46'12"E, 1550 m, 13.I.1999, fl., Messmer & Andriatsiferana 679 (MO). Prov. Fianarantsoa: Ambatofinandrahana, 20°29'58"S 46°51'37"E, 1411 m, 2.XI.2004, fl., Andriamihajarivo et al. 409 (MO); Ranomafana National Park, Vohiparara, 21°14'25"S 47°23'38"E, 1100–1200 m, 2.XI.1997, fl., Malcomber et al. 2856 (BR, G, K, L, MO); Ihorombe, Mahatsijoriaka, 9.XI.1951, fl., Service Forestier 4841 (P); Fort-Carnot, Ambodivonana, 23.X.1954, fl., Service Forestier 14394 (P). Prov. Mahajanga: Matsoandakana, 15°08'25"S 49°21'04"E, 1037 m, 9.II.2008, fl., Bernard et al. 725 (MO). Prov. Toamasina: Ambatovy forest, Berano village, 18°48'29"S 48°18'50"E, 1060 m, 25.I.2005, fr., Antilahimena et al. 3217 (G, K, MO, P); Andasibe, Menalamba, Ambatovy, Dynatec camp, 18°51'06"S 48°18'39"E, 1119 m, 18.XII.2005, fl., Antilahimena & Edmond 4452 (MO, P); Fkt. Andasibe, Analamazaotra, 18°55'22"S 48°25'32"E, 1001 m, 21.XII.2013, fr., Antilahimena et al. 8838 (MO, P); Analamazaotra, X.1925, fr., Louvel 63 (P); Ambatovy, Andranovery, 18°05'57"S 48°18'19"E, 1040 m, 27.I.2008, fl., Phillipson & Antilahimena 6069B (MO, P); RNI Zahamena, à côté du RNI, 17°45'45"S 48°42'10"E, 953 m, 29.XI.2001, fl., Rakotonandrasana et al. 608 (MO, P); Analamazaotra, Lac vert, 18°56'20"S 48°25'15"E, 966 m, 12.XII.2013, fl., Rasoazanany et al. 537 (MO, P); Fkt. Ampitambe, 18°49'12"S 48°20'7"E, 1178 m, 2.II.2005, fl., Razafindraibe & Antilahimena 15 (MO, P); Ambatovy, Torotorofotsy forest, 18°52'05"S 48°21'27"E, 961 m, 13.I.2005, fr., Razanatsoa et al. 20 (MO, P); Tsinjoarivo, Ambatolampy, 11.XII.1952, fl., Service Forestier 6628 (P); Ambotrajanga, Perinet, 17.XII.1952, fr., Service Forestier 7551 (MO, P); près d’Ankarahara, P.K. 102 de la route Tananarive-Moramanga, rive gauche du Mangoro, 21.XI.1963, fr., Service Forestier 22924 (P); Andapabe, Ilalanga, 28.I.1967, fr., Service Forestier 26282 (P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 226-228, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

15. Homalium viguieri H. Perrier in Mém. Mus. Natl. Hist. Nat. 13: 287. 1940. Lectotypus (designated by SLEUMER, 1973: 253): MADAGASCAR. Prov. Toamasina: Prov. d’Andovoranto, district de Moramanga, berges du Mangoro près d’Ankarefo, 800 m, 10.XI.1912, fr., Viguier & Humbert 1188 (P [P04679649]!). Syntypi: MADAGASCAR: sine loco, s.d., fr., Humblot 609 (P [P04679644]!; K [K000231503] image seen, S [S10-10215] image seen, US [US00603579] image seen). Prov . Toamasina: Ambatovola, Vohitra, 700 m, I.1928, fl. & fr., Perrier de la Bâthie 18374 (P [P04679643]!). Shrub to 4 m or small tree, sometimes lianescent; young twigs minutely pubescent. Leaves broadly elliptical to elliptical (obovate to broadly obovate, rarely to broadly ovate or suborbicular), 2–5.6 × 1.1–3.8 cm; petiole 1.5–3(–4) mm; base convex to rounded or cuneate; apex rounded to acute or obtuse (short-acuminate); margins serrulate, with glands usually elongated, forming teeth apices, rarely small and imbedded in margin; adaxial surface drying dark grey to blackish, abaxial surface brown. Inflorescences racemose, lateral, 1.3–6.1(–7.5) cm, pubescent; flowers normally sessile (short-pedicellate); bracts linear to lanceolate or spatulate, 0.8–1.2(–1.5) mm, often persistent. Flowers 6(7)-merous; calyx tube broadly funnelform with rounded base, sparsely pubescent to glabrate; sepals lanceolate-oblong to oblong or obovate with acute (cuspidate, rounded) apex, (0.7–) 1.1–1.9 mm; petals oblong-elliptical to obovate, oblong or spatulate with acute (cuspidate, rounded) apex, (1–) 1.3– 2.4 mm; sepals and petals white to greenish (purplish) white, little accrescent, about as long as calyx tube, soon spreading, usually reflexed after flowering, ciliate with widely spaced hairs especially in fruit, both surfaces sparsely pubescent to glabrate; sepal glands short-pubescent; filaments 1.7–3 mm, glabrous; ovary sparsely to moderately pubescent, sometimes becoming glabrate; styles (3)4, 1.7–3 mm, glabrate with few trichomes near base. Vernacular names. – “Fanazava” (Réserves Naturelles 6891); “Hazondromifehy” (Réserves Naturelles 8704); “Lalona fotsy” (Réserves Naturelles 2807); “Maroankoditra” (Service Forestier 5718); “Marokoditra” (Service Forestier 3292); “Voafotsy” (Cours 289). Distribution, ecology and conservation status. – Homalium viguieri is native to mid-elevation humid forests to 1100 m; it does not occur in the extreme north or southeast. This species appears to be uncommon, as the number of collections is small compared to some other species with equally wide distributions and the number of recent collections particularly small. The Extent of Occurrence is estimated as c. 66,400 km ² and the Area of Occupancy as 56 km ². It was collected several times in or around the protected area of Zahamena (= Perinet, Ambatondrazaka) but most of the habitat is unprotected, has suffered damage since the existing collections were made, and is at threat of further damage. Because over 10 distinct locations have been documented, this species would not meet the criteria for a conservation status of “Vulnerable”, but a status of “Near Threatened” [NT] might be appropriate. Notes. – Homalium viguieri has pubescent sepal glands and racemose inflorescences; it is most similar to H. micranthum and the small-leaved H. perrieri. It is distinguished from similar species by its broad sepals and petals, which are spreading to reflexed shortly after anthesis, and usually sessile flowers. The inside of the ovary locule may be glabrous or sparsely pubescent. In these characters it seems to be the species of Homalium sect. Blackwellia that is most similar to the species of Homalium sect. Odontolobus . One of the syntypes of H. viguieri enumerated above, Perrier de la Bâthie 18374, is herein doubtfully placed in H. micranthum. “ Homalium viguieri var. cuneiforme H. Perrier ”, based on Cours 289 (PERRIER DE LA BÂTHIE, 1946), was not validly published because of the absence of a required Latin diagnosis. It was later sunk into H. viguieri (SLEUMER, 1973). It does not appear to represent a distinct infraspecific taxon. Additional material examined. – MADAGASCAR. Prov. Antananarivo: La Mandraka, VIII.1906, fl., Alleizette 1018 (P); Betsitra, près Analabe (N Imerina), X.1906, fr., Alleizette 1276 (P). Prov. Fianarantsoa: Vangaindrano, vallée de l’Itomampy, 700 m, VI.1914, fr., Perrier de la Bâthie 12660 (P); Manakara, s.d., fr., Service Forestier 2749 (P); P.K. 92, route Farafangana à Vohipena, 3.IV.1951, fr., Service Forestier 3681 (P); Andriodriotra, Antsindra, Ambohimanga Sud, 20.VII.1952, fr., Service Forestier 6381 (P). Prov. Toamasina: Andapanomby, Makira forest, Ampandisanana river, 15°21'04"S 49°06'53"E, 1007 m, 26.IV.2007, fl. & fr., Birkinshaw et al. 1666 (G, MO); Manakambahiny, 1100 m, III.1938, fr., Cours 289 (P); Mahanoro, riv. Andohasira, Ampasimpotsy, 19°53'54"S 48°47'27"E, 5 m, 22.VI.2010, fr., Rakotonirina 535 (MO, P); R.N. III, Manaka Est, Ambatondrazaka, 8.III.1951, fr., Réserves Naturelles 2807 (P); Menalamba, Perinet, 21.III.1951, fl., Réserves Naturelles 3292 (P); ManakaEst, Ambatondrazaka, 6.II.1954, fr., Réserves Naturelles 6891 (P); ibid loco, 30.III.1954, fr., Réserves Naturelles 6896 (P); ibid loco, 12.III.1956, fl., Réserves Naturelles 8704 (P [2 sheets]); Antsahatsaka, Beravina, Perinet, 7.IX.1952, fr., Service Forestier 5718 (P); berges du Mangoro au N d’Ambodimanga, 21.I.1968, fl., Service Forestier 28127 (P [2 sheets])., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on page 242, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328.","PERRIER DE LA BATHIE, H. (1946). Flacourtiacees. Fl. Madagascar Comores 140."]}

9. Homalium martialii Appleq., spec. nova (Fig. 4). Holotypus: MADAGASCAR. Prov. Antsiranana: Vohemar, Antsirabe Nord, Andravinambo, forêt d’Antsolatra, 14°07'27"S 49°58'10"E, 385 m, 31.I.2014, fl. & fr., Martial et al. 431 (MO-6634726!; isotype: P!, TAN!). Homalium martialii Appleq. differs from H. thuarsianum (Tul.) Baill. in having usually elliptical to narrowly elliptical (to obovate) leaves, with often longer petioles and with margins irregularly crenate or shallowly repand (crenate-serrate), and sepals and petals much shorter than calyx tube (seldom only slightly shorter), often somewhat lanceolate. Tree to 10 m, 13 cm dbh; young twigs glabrous. Leaves elliptical (to narrowly elliptical or somewhat obovate), (3–)3.5–7.6(–9) × (1.7–)2–3.3(–3.7) cm; petiole 3–12(–15) mm; base convex or basally attenuate; apex cuspidate with rounded tip to short-acuminate or rounded; margins irregularly crenate or shallowly repand (crenate-serrate), with glands small, elliptical, usually sunk in margins (slightly prominent at tooth apices); adaxial surface drying grayish or dull brown, abaxial surface medium brown. Inflorescences few-branched panicles (racemes with some flowers clustered), lateral, 1–3 per node, (4–)5.5–10(–12) cm, short-pubescent; flowers pedicellate with pedicels (0.5–)1.5–2.7(–3.6) mm; bracts lanceolate, 0.8–1.8(–3?), rapidly caducous. Flowers (7–)8–9-merous; calyx tube funnelform with rounded base, short-pubescent; sepals narrowly lanceolate (ligulate) with acute apex, 1.4 –3 mm; petals narrowly lanceolate (ligulate) with acute apex, 1.3–2.6 mm; sepals and petals greenish-white to grayish or reddish (probably only post-flowering), moderately accrescent, much shorter (to slightly shorter) than calyx tube, ascending (suberect, rarely spreading), ciliate with long soft hairs, abaxial surface short-pubescent, adaxial surface moderately to sparsely pilose; sepal glands moderately long-pubescent; filaments 0.6–1 mm, basally villous; ovary villous to pilose; styles 4–5(–6), 0.6–0.8 mm, basally villous. Etymology. – Homalium martialii is named in honor of Martial, a parataxonomist working in the Makirovana region of northeastern Madagascar, who was the lead collector of three of the four known collections of the species. The species is reported to be visited by bees (Martial et al. 126, 478). Vernacular names and use. – “Jabaorahoditra” (Martial et al. 431). Wood of H. martialii is used for construction (Martial et al. 126, 478). Distribution, ecology and conservation status. – Homalium martialii is native to low-altitude humid forests in a small portion of eastern Antsiranana fairly near the coast, once at 502 m, on sandy soil. The Area of Occupancy is estimated at 16 km ² and the Extent of Occurrence at about 28.1 km ². Three of these collections are within the Makirovana-Tsihomanaomby forest, which for the moment has some protection through a community-based conservation agreement facilitated by the Missouri Botanical Garden, and the fourth is near it. Because all populations could be affected by a single event such as a cyclone or large fire, a conservation status of “Vulnerable” [VU D2] is warranted. Notes. – Homalium martialii belongs to the group of three species, including H. thuarsianum (Tul.) Baill., that share usually well-developed paniculate inflorescences and pubescent sepal glands, as well as usually pubescent inflorescences and calyx. Homalium thuarsianum has mostly oblanceolate to obovate leaves, with petioles to 5(–6) mm, and with margins entire or subentire (to revolute or slightly repand). As in many Malagasy species of sect. Blackwellia, its petals and sepals at maturity are longer than the calyx tube and consistently ligulate. Homalium fortunatii, native to Toamasina, is distinguished by its large leaves. An unusual feature of H. martialii is that the sepals and petals are usually much shorter than the calyx tube and often narrowly lanceolate, which gives the flowers a distinctive appearance as compared to species with longer, ligulate sepals and petals. Homalium martialii also has an unusually large number of styles (as many as 6, which is not seen in any other Malagasy species). Paratypi. – MADAGASCAR. Prov. Antsiranana: Forêt classée entre le village d’Andrangana et la riv. d’Anjombalava, massif de Tsihomanaomby, W d’Ambavala, 14°06'03"S 50°02'52"E, 150 m, 7.V.2000, fr., Andrianjafy et al. 62 (MO); Sambava, Marogaona, Ambodimanganangaiky, forêt de Makirovana, en haut versant, 14°10'42"S 49°56'03"E, 502 m, 17.IV.2013, fl., Martial et al.126 (G, K, MO); Sambava, Anjangoveratra, Ambavala, forêt de Tsihomanaomby, Atsahakasaka, sur crête, 14°06'55"S 50°03'03"E, 110 m, 11.II.2014, fl., Martial et al. 478 (MO)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 235-236, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

3. Homalium brachyrhachis Sleumer in Bull. Jard. Bot. Natl. Belg. 43: 261. 1973. Holotypus: MADAGASCAR. Prov. Toliara: crêtes et barres calcaires au lieu dit Ankiranja, à 30 – 35 km de Manja [21°25'S 44°20'E] sur la route de Bevoay [21°50'S 43°52'E], 3–4.XII.1969, fl., Service Forestier 28953 (P [P04705554]!; isotype: L [L0010886] image seen, TEF [TEF000288] image seen). Tree to 6 – 7 m or large shrub; bark platanoid; young twigs glabrous. Leaves elliptical to ovate, (2.7–)3.9 – 6.2 × (2–) 2.5–3.5 cm; petiole 11–16 mm; base rounded (convex), the extreme base attenuate; apex short-acuminate to acute (obtuse); margins prominently crenate-serrulate, with glands small, round, inside teeth apices; adaxial surface drying olive, abaxial surface pale green. Inflorescences racemose, lateral, 1.5 – 3.8 cm, glabrous; flowers pedicellate with pedicels 2– 3 mm; bracts not seen. Flowers 7 –8-merous; calyx tube funnelform with rounded base, minutely pubescent; sepals ligulate with acute apex, 1.5–2.5 mm; petals ligulate with acute apex, 1.8–2.5 mm; sepals and petals about equal to calyx tube or slightly longer, ascending (partly spreading to somewhat reflexed), short-ciliate, abaxial surface minutely pubescent with narrow dark stripes, adaxial surface sparsely pubescent; sepal glands pubescent; filaments 1–1.4 mm, glabrous; ovary pubescent; styles 3–4, 0.8–1.4 mm, basally pubescent. Distribution, ecology and conservation status. – Homalium brachyrhachis is the only species of this section known from dry western forest on limestone. It is known from only two contemporaneous collections in northwestern Toliara province, on the road from Manja to Bevoay, representing a single population. No further material has been collected for almost 50 years. The natural vegetation in this region is mostly extirpated and the remnants suffer ongoing threats and anthropogenic degradation; some land in the region is protected, but the original locality is not. This species, if it still exists, may therefore be at imminent risk of extinction. An appropriate preliminary assessment of conservation status is therefore “Critically Endangered” [CR B1ab(iii)+B2ab(iii)]. Additional material examined. – MADAGASCAR. Prov. Toliara: crêtes et barres calcaires au lieu dit Ankiranja, à 30–35 km de Manja sur la route de Bevoay, 3–4.XII.1969, fl., Service Forestier 28936 (P [4 sheets])., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 229-230, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

11. Homalium myrtifolium Sleumer in Bull. Jard. Bot. Natl. Belg. 43: 258. 1973. Holotypus: MADAGASCAR. Prov. Toamasina: Vavatenina, Fénérive-Est, 20.I.1957, fl., Réserves Naturelles 8879 (P [P00375094]!; isotype: P [P04679966]!; L image seen, TAN image seen). Tree or shrub to 9 m; young twigs minutely pubescent when very young, soon becoming glabrous. Leaves obovate, 0.9–2.5(–3) × 0.4–1.1(–1.5) cm; petiole (0–) 0.5–2.5 mm; base cuneate to convex; apex rounded; margins few-toothed near apex to subentire, with glands small, rounded, sunk in margins at tooth apices; adaxial surface drying pale brown, abaxial surface greenish; secondary veins strongly ascending, at a narrow angle to midrib. Inflorescences racemose, lateral, 1.4–5 cm, short erectpubescent; flowers pedicellate with pedicels 0.5–2 mm (subsessile); bracts linear to lanceolate, 0.5–1 mm. Flowers 6-merous; calyx tube funnelform to subcylindrical with rounded base, pubescent; sepals ligulate to narrowly oblanceolate with acute apex, 1.4–2.6 mm; petals spatulate or ligulate to narrowly lanceolate with acute apex, 1.7–3 mm; sepals and petals greenish to yellowish white to partly pink or red (perhaps only postanthesis), modestly accrescent, usually much longer than calyx tube (especially in fruit), ascending or partly spreading, seldom becoming reflexed, ciliate, adaxial surface sparsely pubescent, abaxial surface sometimes bearing few long hairs; sepal glands long-pubescent, often rather sparsely; filaments 1.6–2.5 mm, sparsely pilose; ovary pilose to hispid (becoming glabrate); styles 3(4), 1.3–1.7 mm, basally pilose. Vernacular name. – “Hazombato” (Réserves Naturelles 8879). Distribution, ecology and conservation status. – Homalium myrtifolium is native to low- to mid-altitude humid forests in Toamasina and Toliara. Only three distinct locations have been documented. All collections from Toliara come from the area of Tsitongambarika, which is protected, and the species has been collected once near, though not within, Zahamena National Park. The third population, from an unprotected area, has not been recollected since 1957. Though the Area of Occupancy is estimated at only 16 km ², the Extent of Occurrence is estimated at c. 12,600 km ². However, much of the intervening habitat is unsuitable due to ongoing anthropogenic damage. Although the fact that the only apparently thriving population is protected argues against a conservation status of Endangered, the species should be considered “Near Threatened” [NT] based on its limited range and the uncertain status of one of the formerly existing populations. Notes. – Homalium myrtifolium is easily recognized by its small, pale-drying obovate leaves with entire or minutely toothed margins and few, ascending secondary veins. It is the only species that has secondary veins ascending at a narrow angle to the midrib; it is among the group with racemose inflorescences and pubescent sepal glands. Most collections come from a single small region in southeastern Toliara; two collections come from Toamasina, well to the north. The morphology of those collections is not significantly different from that of the southern collections. Additional material examined. – MADAGASCAR. Prov. Toamasina: A l’W du village d’Ambatoaranana, NE de la RNI de Zahamena, 17°33'30"S 48°53'20"E, 900 m, 24.I.1994, fr., Randrianjanaka & Be Maxime 52 (MO, P). Prov. Toliara: Fkt. Antsotso, forêt d’Ivohibe, 24°34'15"S 47°12'36"E, 44 m, 29.I.2015, fl., Randriatsivery et al. 580 (G, MO); Antsotso Avaratra, 24°33'54"S 47°11'55"E, 432 m, 11.XII.2007, fl., Razakamalala et al. 3882 (MO, P [2 sheets]); Antsotso Avaratra, forêt Tsitongambarika, 24°34'16"S 47°12'05"E, 271 m, 1.IV.2008, fl., Razakamalala et al. 4108 (MO, P); Fkt. Antsotso, forêt d’Ivohibe, 24°33'51"S 47°12'20"E, 395 m, 28.I.2015, fl., Razanatsima et al. 1385 (G, MO); forêt de Bemangidy, au N de Mahatalaky, Fort-Dauphin, 1–2.II.1963, fl., Service Forestier 22339 (P); Bemangidy Forest, 24°34'15"S 47°12'36"E, 150 m, 13.IV.2010, Thulin & Razafindraibe 11855 (MO)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 238-239, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

4. Homalium cauliflorum H. Perrier in Mém. Mus. Natl. Hist. Nat. 13: 290. 1940. Lectotypus (designated by SLEUMER, 1973: 262): MADAGASCAR. Prov. Antsiranana: Bemarivo, bords de torrents, 100 m, IX.1912, fl., Perrier de la Bâthie 2199 (P [P04734103]!; isolectotype: K [K000231495] image seen, L [L0010897] image seen). Syntypus: MADAGASCAR. Prov. Antsiranana: Ambahatra, E du massif de Manongarivo, IX.1909, fl., Perrier de la Bâthie 2325 (P [P04734102]!). Shrub or tree to 12 m, sometimes with multiple trunks, 12 cm dbh; bark blackish; young twigs glabrous or glabrate. Leaves narrowly elliptical (oblanceolate), 5.5–11 × 1.3–3.5 cm; petiole 2– 5 mm; base cunate to narrowly convex or somewhat attenuate; apex narrowly acute (to rounded); margins crenate-serrulate with few shallow, sometimes inconspicuous teeth, sometimes slightly revolute, with glands slightly elongated, in margins inside apex of tooth; adaxial surface drying greenish (to grayish, mottled dark, or when immature blackish brown), abaxial surface brown. Inflorescences paniculate, relatively often racemiform with short branches, lateral, many cauliflorous on large twigs below most distal leaves, sometimes pseudoterminal, 4–15(–20) cm, pilose; flowers pedicellate with pedicels (0.5–)1.5–3.5(–5) mm; bracts ovate to oblong, elliptical or lanceolate, 0.5–3.5 mm, most rapidly caducous. Flowers 8–10-merous; calyx tube funnelform, pilose; sepals linear (very narrowly spatulate) with acute apex, 1.3–3 mm; petals linear (very narrowly spatulate) with acute apex, 1.4–3.5 mm; sepals and petals white to yellowish or the sepals greenish and petals white to greenish white, accrescent, slightly longer than calyx tube to about equal (slightly shorter), ascending (sometimes spreading in a few flowers), long-ciliate, both surfaces pilose; sepal glands pilose; filaments 0.7–1 mm, pilose; ovary pilose; styles 5, 1– 1.4 mm, basally sparsely pilose. Vernacular names. – “Hazomainty ” (Service Forestier 2960); “Hazombato” (Service Forestier 7785); “Jabozahaditro” (Antilahimena 841); “Jambaraohoditry” (Hong-Wa et al. 319). Distribution, ecology and conservation status. – Homalium cauliflorum is native to low-elevation (once to 500 m) humid forests in Antsiranana; it is reported to occur on gneiss and along rivers and in ravines. The Area of Occupancy is estimated at 60 km ² and the Extent of Occurrence at c. 19,700 km ²; the species is found in the Tsaratanana and Marojejy Reserves and more than 10 locations are or have been known. Its conservation status is therefore considered to be “Least Concern” [LC]. Additional material examined. – MADAGASCAR. Prov. Antsiranana: Ampasindava, forêt d’Andranomatavy, 13°41'41"S 47 °59'15"E, 160 m, 27.XI.2009, fl., Ammann et al. 426 (P); Fkt. Ambobake, Ramena River, 13°45'00"S 48°31'18"E, 251 m, 21.XI.2001, fl., Antilahimena 841 (MO); Behefaka, Anjahana, forêt d’Ambinanibekona, 13°21'06"S 49°10'06"E, 124 m, 7.VI.2005, fl., Hong-Wa et al. 319 (MO, P); massif de Marivorahona au SW de Manambato (Haute Mahavavy du Nord, Distr. d’Ambilobe), 500 m, 18–26.III.1951, fl., Humbert & Capuron 25617 (P); Loky Manambato, village Tsinjrano, aux bords de la riv. Manambato, 13°13'40"S 49°52'36"E, 14 m, fl., 12.X.2013, Rakotovao et al. 6490 (MO); Daraina, forêt de Binara, 13°14.61'S 49°38.85'E, 100 m, 1.IV.2004, fl., Ranirison 562 (MO, P); RNI de Marojejy, suivant la riv. d’Androranga, entre Anketsahely et village Antongodriha, 14°18'S 49°42'E, 150–265 m, 3–13.XI.1994, fl., Rasoavimbahoaka 413 (MO, P); District Sambava, 19.VIII.1956, fl., Réserves Naturelles 8421 (P); Marovato, district Ambanja, 12.III.1956, fl., Réserves Naturelles 8611 (P); Antsirabe, Ambanja, 16–30.III.1951, fl., Service Forestier 2960 (P); env. d’Amb[a]kirano, district d’Ambilobe, s.d., fl., Service Forestier 3073 (P [2 sheets]); Marojejy, Andapa, 27.X.1953, fl., Service Forestier 7785 (MO, P); Ambahatra, cours moyen (13°56'S 48°27'E), Bemamboly (rive droite), 140 m, 28.V.2000, fl., Wohlhauser 60273 (MO)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on page 230, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328."]}

12. Homalium perrieri Sleumer in Bull. Jard. Bot. Natl. Belg. 43: 257. 1973. ≡ Homalium buxifolium H. Perrier in Mém. Mus. Natl. Hist. Nat. 13: 286. 1940 [nom. illeg.] [non Däniker, 1933]. Holotypus: MADAGASCAR. Prov. Antsiranana: Massif de Manongarivo, versant de Sambirano, 500 m, V.1909, fl., Perrier de la Bâthie 4620 (P [P04734362]!; isotype: L [L0010989] image seen). Tree to 14 m, 20 cm dbh, or large shrub; young twigs minutely pubescent. Leaves elliptical to broadly elliptical (somewhat ovate or obovate, suborbicular), 0.5 – 2.3 × 0.3 – 1.4 cm; petiole 1 – 1.5(– 2) mm; base rounded to convex; apex acute with rounded tip; margins crenateserrate, with glands elongated in tooth apices, often producing exudate; adaxial surface drying dark brown, abaxial surface brown; domatia occasionally present in axils of secondary veins. Inflorescences racemose, lateral, few-flowered, (0.8 –)1.4 – 3(– 3.8) cm, minutely to short-pubescent; flowers pedicellate with pedicels 0.5– 1.4(–2.5) mm; bracts 0.4–0.8 mm. Flowers 6–7-merous; calyx tube funnelform with base broadening in fruit, sparsely long-pubescent (to sparsely pilose); sepals ligulate (narrowly elliptical) with acute apex, 1–2(–3.1) mm; petals ligulate (narrowly spatulate to narrowly oblanceolate-elliptical) with acute apex, 1.3–2.1(–3.3) mm; sepals and petals pale yellow (to pale beige, white, reddish white), probably moderately accrescent, from substantially longer than calyx tube (especially in fruit) to equal in length, ascending (partly spreading to reflexed), long-ciliate, abaxial surface sparsely long-pubescent (to sparsely pilose), adaxial surface sparsely pilose to glabrate; sepal glands pilose; filaments 1–2.6 mm, sparsely pilose (moderately pilose, glabrous); ovary pilose; styles 4, 1.2–2.5 mm, basally pilose. Vernacular names. – “Beando” (Bernard et al. 72); “Hazombato” (Rakotomalaza 10); “Maroankoditra” (Schatz & Antilahimena 4019). Distribution, ecology and conservation status. – Homalium perrieri is native mostly to mid-elevation humid forest (to low-elevation or moderately high-elevation) in Antsiranana and northern Toamasina. It has been reported to occur on basalt and near water. There are at least six distinct populations, and the range includes the protected areas of Manongarivo and Anjanaharibe Sud, so its conservation status is tentatively estimated as “Least Concern” [LC]. Notes. – Homalium perrieri is among the species with racemose inflorescences and pubescent sepal glands. It is most similar to H. viguieri H. Perrier, which has usually sessile flowers with broad sepals and petals, and H. micranthum, which has usually much larger leaves and usually larger inflorescences. Leaves of pressed specimens of H. perrieri often appear almost distichously arranged. It is easily distinguished from the other very small-leaved species in this group, H. myrtifolium, which has obovate pale-drying leaves with cuneate bases, strongly ascending secondary veins, and small marginal glands that do not produce noticeable exudate. Ravelonarivo & Rabesonina 646, from a very high altitude in Antsiranana, is a particularly problematic specimen. It has much larger leaves (to 3.3 × 1.7 cm) than other specimens, whose leaves are always less than 2.4 cm long, sometimes much less; the sepals and petals are usually reflexed shortly after anthesis. The leaves appear distichous and the inflorescences are very short, like H. perrieri; hence identifying this as a small-leaved H. micranthum is not satisfactory. It might be suspected of being a distinct locally endemic taxon, but there is not enough material available to describe it. Further collections from the area would be desirable. Homalium perrieri was a replacement name for H. buxifolium H. Perrier [nom. illeg.]. As far as is known, there was initially only one duplicate of the type collection, which is the holotype. The “isotype” at L, like most cited type duplicates at L., consists of a few leaves plainly borrowed after the fact from a complete specimen housed elsewhere, and does not have equal standing. Additional material examined. – MADAGASCAR. Prov. Antsiranana: Antalaha, Ambohitralalana (Cap-Est), forêt Tanandavehely, 15°18'S 50°29'E, 0–15 m, 9.XII.1994, fl., Bernard et al. 72 (G, MO, P); E of Ankaramy, Réserve Spéciale of Manongarivo, Bekolosy, 14°02'S 48°19'E, 800–1100 m, 7–12.XII.1992, fl. & fr., Malcomber et al. 1984 (MO, P); E of Ankaramy, Réserve Spéciale de Manongarivo, Antsatrotro, 14°05'S 48°23'E, 600–900 m, 26–30.III.1993, fl. & fr., Malcomber et al. 2326 (MO, P); E of Ankaramy Be, Manongarivo Réserve Spéciale, Bekolosy, 14°02'S 48°18'E, 1000–1200 m, 4.XII.1993, fl. & fr., Malcomber & Rakotomalala 2616 (MO, P); E de Ankaramy-be, Réserve Spéciale de Manongarivo, Bekolosy, 14°04'S 48°17'E, 1000–1480 m, IV.1993, fl., Rakotomalaza 10 (MO, P); Andravinambo, forêt de Tsihomanaomby, 14°06'S 50°00E, 23.I.2014, fl., Rakotonirina et al. 413 (MO); Réserve Spéciale Anjanaharibe-Sud, village de Mandritasrahely, 14°43'10"S 49°27'12"E, 1700 m, 14.II.1995, fl., Ravelonarivo & Rabesonina 646 (G, MO, P); Réserve Spéciale #5, Manongarivo, W slopes of Antsatrotro, 14°08'S 48°21'E, 800 m, 23–25. III.1991, fl. & fr., Schatz 3216 (MO, P). Prov. Toamasina: Maroantsetra, fkt. Anjiahely, forest Ankahaminivola, 15°23'57"S 49°26'08"E, 780 m, 15.XII.2002, fl. & fr., Antilahimena et al. 1490 (MO, P); Ankirindro massif, slopes above village Ambodivato, 15°19'13"S 49°33'29"E, 150 m, 20.XI.2002, fl., Schatz & Antilahimena 4019 (G, K, MO, P); bassin de la Fananehana, massif de l’Androrona, crête terminale, 850 m, II.1954, Service Forestier 9001bis (P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 239-240, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

5. Homalium decaryanum H. Perrier in Mém. Mus. Natl. Hist. Nat. 13: 288. 1940. Lectotypus (designated by SLEUMER, 1973: 255): MADAGASCAR. Prov. Fianarantsoa: Befotaka, [Prov.] de Farafangana, 12.VIII.1926, fl., Decary 4776 (P [P00375089]!; isolectotype: K [K000231498] image seen, L [L0010902] image seen). Tree to 3 m; young twigs minutely pubescent. Leaves narrowly elliptical (oblanceolate), 2.1–3.3 × 0.5–0.7 cm; petiole 1.5– 3 mm; base narrowly cuneate (slightly attenuate); apex acute with rounded tip; margins serrulate, with glands small, rounded, in tooth apices; adaxial surface drying greenish, abaxial surface pale brown. Inflorescences racemose, lateral, sometimes paired, (1.3–) 2.8–3.7 cm, pilose or long-pubescent; flowers pedicellate with pedicels 0.5–1.4 mm; bracts ovate to lanceolate, 1.4–2.5 mm. Flowers 8 –9-merous; calyx tube funnelform, long-pubescent; sepals ligulate with acute apex, 1.7–2.1 mm; petals ligulate with acute apex, 1.7–2.4 mm; sepals and petals white, accrescence unknown, longer than calyx tube, ascending (few spreading to reflexed), ciliate with long soft trichomes, abaxial surface sparsely appressed-pubescent, adaxial surface densely pilose; sepal glands glabrous; filaments 0.6– 0.7 mm, glabrous or glabrate; ovary pilose; styles 3–4, 0.7–0.8 mm, pilose. Vernacular name. – “Kidravy” (Decary 4776). Distribution, ecology and conservation status. – The only known collection is from Befotaka, an area of humid forest near Midongy du Sud in Fianarantsoa, near water. Most of the remaining forest in that area is protected. However, the fact that this species has not been recollected for over 90 years suggests that it is quite rare and of limited distribution, so that all remaining individuals could be affected by a single extreme event such as a typhoon. Its conservation status is therefore suggested to be “Vulnerable” [VU D2]. Note. – SLEUMER (1973) termed the duplicate of Decary 4776 at P the “ holotype ”. That was certainly the specimen used by Perrier de la Bâthie, but he did not overtly state that only material at P was used, and there is a duplicate at K. Therefore these are syntypes (the fragment at L that must have been taken from one of them has a lesser standing) and Sleumer’s declaration actually constitutes an effective lectotypification., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 230-231, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328."]}

13. Homalium retivenium Sleumer in Bull. Jard. Bot. Natl. Belg. 43: 264. 1973. Holotypus: MADAGASCAR. Prov. Toamasina: Env. de Navana (à l’E de Maroantsetra), I.1954, fl. & fr., Service Forestier 8884 (P [P00375174]!; isotype: L [L0011003] image seen, P [P04734404, P04734406]!, TEF [TEF000233] image seen). Tree to 16 m, 12 cm dbh; latex white; young twigs glabrous. Leaves obovate to elliptical, oblanceolate or narrowly elliptical, 2.8–7.5 × 1.3–3.8 cm; petiole 1–3.5 mm; base cuneate to convex; apex rounded-acute to short-acuminate (rounded, cuspidate); margins entire or subentire, usually revolute, with glands absent or inconspicuous; adaxial surface drying grayish (blackish brown), abaxial surface brown to olive. Inflorescences racemose, lateral, often paired, 1.8–6.8(–10) cm, moderately pilose; flowers pedicellate with pedicels 1 – 2.4(– 4.5) mm, apparently early caducous; bracts linear to lanceolate, (0.7–) 1– 1.8 mm. Flowers (7–)8–9-merous; calyx tube short or narrowly funnelform with narrow base, sparsely (to moderately) pilose; sepals linear to narrowly ligulate (narrowly spatulate) with acute apex, 2.5–5.5(–6.4) mm; petals linear to narrowly ligulate with acute apex, 2.5–5.5(–6.8) mm; sepals and petals greenish-white to yellowish, strongly accrescent, much longer than calyx tube, ascending (sometimes becoming spreading, seldom to reflexed), long-ciliate, both surfaces sparsely pilose; sepal glands moderately pilose on apex; filaments 2–3 mm, moderately pilose; ovary sparsely pilose; styles 3–4, (1.6–) 2.5–3.4 mm, basally pilose. Vernacular names. – “Hazombato à p[etits] f[euilles]” (Service Forestier 15758); “Hazombatofotsy” (Service Forestier 2791, 15694); “Marankoditra” (Réserves Naturelles 6193); “Tsilaitra” (Raharimalala 219). Distribution, ecology and conservation status. – Homalium retivenium is native to low-elevation and littoral forests in northeastern Madagascar. There are more than five distinct localities, and the species occurs in several protected areas, including Betampona, Masoala, Nosy Mangabe, and Tampolo. Therefore, a preliminary assessment of its conservation status is “Least Concern” [LC]. Notes. – Among eastern species of sect. Blackwellia with racemose inflorescences and pubescent sepal glands, H. retivenium is notable for its pilose inflorescences and flowers and its relatively numerous, long, narrow sepals and petals, much exceeding the calyx tube (which at anthesis is rather short and narrow). The only specimen included in H. retivenium from the province of Antsiranana, Service Forestier 2791, is morphologically atypical with very long inflorescences and pedicels and unusually broad sepals and petals. Further investigation of this population would be desirable. Additional material examined. – MADAGASCAR. Prov. Antsiranana: Ampavatonampingotra, Andapa, 12.II.1951, fr., Service Forestier 2791 (P). Prov. Toamasina: Maroantsetra, fkt. Ambanizana, 15°37'00"S 49°57'00"E, 10.IV.2002, fl., Antilahimena & Aridy 962 (MO, P); Betampona RNI, fkt. Andratambe, 17°55'S 49°13'E, 250–650 m, I-X.2000, fl., Iambana et al. 268 (G, MO); Forêt d’Ibanda, 2.II.1990, fl., Raharimalala 219 (P); RN I [Betampona], Ambodiriana, Tamatave, 12.II.1954, fr., Réserves Naturelles 6193 (MO, P); Masoala Peninsula c. 6 km NNE of Ambanizana along trail from Ambanizana River to summit of Ambohitsitondroina, 24–25.XII.1989, fl., Schatz & Modeste 2910 (MO, P); Nosy Mangabe in the Bay of Antongil, 15°30'S 49°46'E, 0–330 m, 2–19.I.1990, fl., Schatz & Carlson 2926 (MO, P); Masoala Peninsula, along coast at third major river, approx. 6 km S of Ambanizana, 0–10 m, 26.XII.1990, fl., Schatz & Modeste 3092 (MO, P); Tampolo, Fénérive, 14.II.1956, fl. & fr., Service Forestier 15694 (MO, P [2 sheets]); Vohilengo, Fénérive-Est, 20.II.1956, fl. & fr., Service Forestier 15758 (P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on page 240, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

2. Homalium axillare (Lam.) Benth. in J. Proc. Linn. Soc., Bot. 4: 34. 1859. ≡ Blakwellia axillaris Lam., Encycl. 1: 428. 1785. Lectotypus (designated by SLEUMER, 1973: 255): MADAGASCAR: sine loco, s.d., fl., Commerson s.n. (P-JU 14407 image seen; isolectotype: G [G00018417, G00018418, G00018419] images seen, P [P00624058]!). Tree to 16 m, 26 cm dbh, or large shrub; young twigs glabrous or minutely papillate. Leaves elliptical to narrowly or broadly elliptical (seldom to ovate, obovate, or rarely lanceolate), (2–)2.5–7.5(–9.3) × (0.9–)1.1–3.8(–4.3) cm; petiole (1.5–) 2–8 mm; base convex (rounded, cuneate); apex acute to rounded or obtuse (somewhat acuminate); margins crenateserrate, with small or elongated glands at apices of teeth, to subentire; adaxial surface drying darker, brown to greenish or grayish, abaxial surface pale to medium brown (greenish). Inflorescences racemose, 1.5–14.5 cm, pubescent; flowers pedicellate with pedicels 0.2–1.8(–2.5) mm (subsessile); bracts oblong-ovate to ovate or lanceolate (suborbicular, oblongobovate, deltoid), 0.5– 1.1(–1.6) mm. Flowers 7– 10-merous (most commonly 8-merous); calyx tube narrowly funnelform to basally rounded, pubescent; sepals narrowly oblanceolateoblong to ligulate or narrowly oblong with acute apex, 0.8–3.7 mm; petals narrowly oblanceolate-oblong to ligulate or narrowly oblong with acute apex, 0.8–3.5 mm; sepals and petals whitish to cream or yellow-green (possibly becoming pink or reddish in fruit), accrescent, longer than calyx tube (very rarely similar in length), ascending to irregularly spreading or partly reflexed (most often in fruit), ciliate with long trichomes, both surfaces sparsely short-pubescent to glabrous; sepal glands glabrous; filaments 0.8–2.4(–3.5) mm, sparsely villous (to glabrate); ovary hispid or pilose to long-pubescent; styles 3–4(–5), 1.3–3 mm (apparently accrescent), basally hispid to pilose. Notes. – There is substantial geographic variation in morphology within H. axillare, which is the commonest species in its section. Specimens from the province of Toamasina have usually relatively large, crenate leaves with usually large, elliptical marginal glands that may occupy much of the apical margins of teeth; the leaves usually dry brown. The type of the species, the locality of which is unknown, falls into this group, as do a few specimens from Antananarivo, Fianarantsoa, and eastern Mahajanga. Their inflorescences are usually 4–14.5 cm long and mostly borne near twig apices, so that they often extend much beyond the leaves. (However, rare specimens from this range have short or poorly developed inflorescences.) Most specimens, including all from Toamasina, have filaments 2–3.5 mm long, though outlying specimens sometimes have short filaments H. axillare subsp. delphinense (H. Perrier) Appleq., while the variant native primarily to Toamasina is treated as subsp. axillare . Because there is variation within as well as among subspecies in all of the potentially distinguishing characters, they are presumed not to be reproductively isolated. A group of specimens collected contemporaneously at Manongarivo (McPherson & van der Werff 16356, 16379, 16381, van der Werff & McPherson 13533) show particularly large variation within a single population, including some characters typical of H. axillare subsp. axillare . In northern Madagascar and the province of Fianarantsoa, H. axillare subsp. delphinense generally occurs at moderate to high altitudes in humid forest, whereas in Toliara, most populations are from littoral or low-altitude coastal forests. Most specimens from Toliara have quite small, entire, pale-colored leaves, and the difference in ecological preferences might well suggest that they should be treated as a distinct subspecies. However, coastal specimens are known that have moderately large, prominently crenate, and/or brown-drying leaves, while a very small-leaved specimen from a high altitude in Toliara is known. Since it is not possible to separate ecologically distinct groups using morphology, they have not been formally recognized. However, the likelihood of geographically correlated genetic variation should be noted. SLEUMER (1973) quoted among specimens seen “ Commerson (P, P-LA), ‘ Blackwellia de Madagascar’, P-JU 14407 holotype Blackwellia axillaris ”. Since multiple unnumbered Commerson duplicates exist that have been presumed to be part of the same gathering, this actually constitutes a lectotypification with an erroneous term used. No material corresponding to this collection is locatable in the Lamarck herbarium. There is a second sheet in the main collection in Paris [P00624058], which Sleumer labeled as the “type” of B. axillaris; this is an isolectotype. A second specimen at P [P00624059], which bears a Herb. Mus. Paris, Herbier de A.N. Desvaux label, was determined by Sleumer as a “prob[able] isotype” of B. axillaris. Only the piece on the upper left of that sheet is actually Homalium axillare. The remainder appears to be H. planiflorum (Boivin ex Tul.) Baill. The former portion does resemble the Commerson material but there is no clear evidence that it derives from the same collection; the handwriting on the main labels is similar but does not appear to be identical. Hence that sheet is not considered here to be an isotype. There are also three specimens at G that do bear the name of Commerson in their labeling (which is variable) and that have been considered to be type material. Key to the subspecies of Homalium axillare 1. Leaves mostly elliptical to broadly elliptical, (2.5–)3.6– 7.5(–9.3) × 1.5– 3.8(–4.3) cm; petiole 4–8 mm; margins crenate-serrate (crenulate, subentire), with glands usually elongated in tooth apices; adaxial surface drying darker, brown (greenish), abaxial surface brown (greenish); racemes (2.6–) 4–14.5 cm; filaments (1.3–) 2.1–3.5 mm............................................ 2a. H. axillare subsp. axillare 1a. Leaves mostly elliptical to narrowly (broadly) elliptical, (2 –)2.5–6.5 × (0.9–) 1.1–3.6 cm; petiole (1.5–)2–4(–5) mm; margins crenate-serrulate to subentire, with glands round or slightly elongated; adaxial surface drying darker, usually greenish or grayish, abaxial surface pale to medium brown (greenish); racemes 1.5–8 cm; filaments 0.8–2.4 mm........................................ 2b. H. axillare subsp. delphinense, Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 225-226, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328."]}

14. Homalium thuarsianum (Tul.) Baill., in Bull. Mens. Soc. Linn Paris 1: 574. 1886 [as thouarsianum]. ≡ Blackwellia thuarsiana Tul. in Ann. Sci. Nat., Bot. ser. 4, 8: 60–61. 1857. Holotypus: MADAGASCAR: sine loco, s.d., fl., du PetitThouars s.n. (P [P04679701]!; isotype: L [L0011030] image seen). Tree to 20 m, 41 cm dbh; young twigs glabrous. Leaves oblanceolate to obovate (oblong-oblanceolate, elliptical), (2.8 –)3.5 – 7.8(– 10) × (1.2 –)1.6 – 3.8(– 5.2) cm; petiole (0 –)2– 5(–6) mm; base cuneate to convex; apex rounded to somewhat obtuse or acute with rounded apex (shallowly emarginate, rounded-cuspidate); margins entire or subentire, sometimes revolute, occasionally slightly repand, with glands absent or few and sunk in margins; adaxial surface drying grayish to dark (greenish) brown, abaxial surface brown. Inflorescences paniculate usually with well-developed branches, lateral (pseudoterminal), (4–)6.7–17(–20) cm, moderately to densely pubescent with fine white, usually erect hairs, to short-pilose; flowers pedicellate with pedicels 1–2.7 mm; bracts ovate to broadly elliptical (linear), (0.8–)1.5– 2.5 (seldom foliose, to 22 mm). Flowers 7–9-merous; calyx tube subcylindrical with rounded base, pubescent with erect white hairs to pilose; sepals ligulate with acute apex, 1.5–4.2 mm; petals ligulate with acute apex, 1.6–4.5 mm; sepals and petals variable in color, greenish-white to yellowish, cream or beige, pink to red or purple probably in fruit, accrescent, longer than calyx tube, almost always ascending (some flowers spreading at anthesis, apparently returning to ascending in fruit), ciliate with hairs very long near apex, abaxial surface appressed-pubescent (rarely sparsely pilose), adaxial surface sparsely pilose; sepal glands sparsely pilose to sparsely pubescent; filaments 0.8– 1.7 mm, pilose; ovary sparsely to moderately pilose; styles (4)5, (0.6–) 0.8–1.3 mm, pilose at base. Vernacular names and uses. – “Hazobato” (Service Forestier 2875); “Hazombato” (Lehavana & Zackarie 711, Noyes et al. 960, Rabevohitra et al. 4370, 4610, 4864, 4970, Réserves Naturelles 10340, Service Forestier 4603, 5861, 8314, 9525, 15701, 19526); “Hazombato à g[randes] f[euilles]” (Service Forestier 15701); “Hazombato à p[etites] f[euilles]” (Service Forestier 15172); “Hazombato-mena à p[etites] f [euilles]” (Service Forestier 15702); “Hazondrano-fotsy” (Service Forestier 15704); “Maraokoditra” (Service Forestier 14228); “Tsitakonala” (Service Forestier 4153). Wood of H. thuarsianum is used for construction (Service Forestier 19526). Distribution, ecology and conservation status. – Homalium thuarsianum is native to littoral and sublittoral forests along the eastern coast of Madagascar; it is reported on sand. One specimen (Antilahimena & Felix 7756) was collected at 1053 m altitude, geographically relatively close to the species’ range but far outside its normal altitude range; this specimen cannot be excluded from the species by morphology. Though the habitat is fragmented and highly threatened, this species is relatively widely distributed and occurs in the protected areas of Tampolo, Soanierana-Ivongo, Nosy Mangabe, and Masoala. Therefore its conservation status is estimated as “Least Concern” [LC]. Notes. – Homalium thuarsianum is the most widespread and common of three species with pubescent sepal glands and usually well-developed paniculate inflorescences, the other two being the newly described H. fortunatii and H. martialii. All three share other features in common, including often relatively large bracts, 7–9-merous flowers, and sepals and petals that are usually pubescent on the abaxial surface (in H. thuarsianum, occasionally sparsely pilose) but somewhat pilose to villous on the adaxial surface. They are all native to low-altitude eastern humid forests. This author considers it reasonable to presume that they are a natural group. The distinctions between H. thuarsianum and the other two species of this group are described under those species. Flowers are visited by bees (Lehavana et al. 54, Ludovic & Rakotoarivony 241). Selected material examined. – MADAGASCAR. Prov. Fianarantsoa: Fkt. Ambahy, forêt d’Andohan’I Martin, 20°48'07"S 48°28'56"E, 22 m, 13.XI.2003, fl., Rabehevitra et al. 749 (MO, P); Fkt. Ambahy, 20°46'15"S 48°28'48"E, 21.III.2003, fl., Rabevohitra et al. 4610 (MO, P); Entre Vohipeno et la mer, VII.1954, fl., Service Forestier 9221 (P); Ankijy, Vatomasina, Vohipeno, 13.IX.1956, fl., Service Forestier 14228 (P); Ankazondratana, Nosy Varika, 30.V.1958, fl., Service Forestier 19526 (P [2 sheets]). Prov. Toamasina: Brickaville, fkt. Ambodilendemy, Ankerana, 18°24'58"S 48°47'21"E, 1053 m, 23.III.2011, fl., Antilahimena & Felix 7756 (MO, P); Mananara, 28.IX.1920, fl. & fr., Decary 63 (P); Ambila-Lemaitso, 18°49'S 49°09'E, 2–5 m, 15.XI.1989, fl. & fr., Keating & Miller 2261 (MO, P); Tampolo, 17°17'16"S 49°24'30"E, 16 m, 14.IV.2004, fl. & fr., Lehavana et al. 54 (MO, P); Fkt. Tanambao Ambodimanga, près du gite, 16°47'S 49°43'E, 27.V.2010, fr., Lehavana & Zackarie 711 (MO); Île Sainte-Marie, Lokintsy, forêt d’Ambohidena on NE coast, 16°51'11"S 49°57'10"E, 10 m, 13.V.2003, fr., McPherson et al. 18913 (MO, P); 9.4 km W of Mahavelona village (Foulpointe), 17°38'S 49°30'E, 10 m, 28.II.1992, Noyes et al. 960 (MO); Fkt. Ambohidena, forêt d’Ambohidena, 16°51'05"S 49°57'13"E, 10 m, 10.X.2003, fl., Rabehevitra et al. 654 (MO, P); Cne rurale Manompana, Forêt Antanambao-Ambodimanga, 16°45'40"S 49°42'35"E, 1.II.2003, fl., Rabevohitra et al. 4370 (MO, P); Andranokoditra, Ankanin’ny nofy, forêt de Vohibola, 18°35'42"S 49°14'02"E, 10.II.2003, fl., Rabevohitra et al. 4413 (MO, P); Soanierana-Ivongo, Andrangazaha, 16°52'02"S 49°40'52"E, 29.1.2004, fl., Rabevohitra & Rakotomamonjy 4864 (MO, P); Forêt au NW de Tanambao, 16°45'12"S 49°42'56"E, 31.I.2004, fr., Rabevohitra et al. 4970 (MO, P); Fkt. Ambohidena, forêt d’Ambohidena, 16°51'11"S 49°57'18"E, 18.II.2004, fl., Rabevohitra et al.5016 (MO, P); Tampini, X.1924, fl. & fr., Réserves Naturelles (Dumazer) s.n. (P); RN II [Masoala], Ambohitralalana, district Antalaha, 16.II.1954, fr., Réserves Naturelles 6620 (P); Soanierana-Ivongo, 18.V.1959, fl. & fr., Réserves Naturelles 10340 (P); Nosy Mangabe RS 4–5 km S of Maroantsetra, 15°30'S 49°46'E, 0–330 m, 21.II.1992, fr., Schatz et al. 3243 (MO, P); Gare d’Ambila-Lemaitso, 28.VIII.1951, fl. & fr., Service Forestier 4153 (P); Antetezana, Tamatave, 28.XII.1951, fl. & fr., Service Forestier 4603 (P); Ambila-Lemaitso, 10.X.1953, fl., Service Forestier 5861 (P); S de Tenina, S de Rantabe, IV.1954, fl. & fr., Service Forestier 9168 (MO, P); Tampolo, Fénérive, 29.VIII.1955, fl., Service Forestier 15172 (MO, P); Tampolo, Fénérive, 22.II.1956, fl., Service Forestier 15701 (MO, P); Île Sainte-Marie, forêt d’Ampaniny, 17.V.1969, fr., Service Forestier 28858 (P [2 sheets])., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 240-241, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607

8. Homalium longiracemosum Appleq., spec. nova (Fig. 3). Holotypus: MADAGASCAR. Prov. Toliara: Anosy Rég., Distr. Taolagnaro, Cne Mahatalaky, Fkt. Sainte Luce, forêt littorale d’Analavinaky, située à l’W du village d’Ambandrika au bord de la riv. de Rianabo, 24°43'21"S 47°00'44"E, 143 m, 24.XI.2009, post-fl., Rakotovao 4714 (MO-6481608!; isotype: TAN). Homalium longiracemosum Appleq. is distinguished from other Malagasy species of Homalium sect. Blackwellia in having leaves mostly narrowly lanceolate to narrowly elliptical, 7.3–15 × 2.6–3.9 cm with petiole (6.5–) 9–18 mm, racemose inflorescences 9–25 cm long, and pubescent sepal glands. Tree to 18 m, 45 cm dbh; young twigs glabrous. Leaves narrowly oblanceolate to narrowly elliptical (obovate-elliptical), 7.3–15 × 2.6–3.9 cm; petiole (6.5–) 9–18 mm; base convex to cuneate; apex short-acuminate to cuspidate (acute); margins shallowly repand, undulate or crenate-serrulate, with glands elongated, prominent at tooth apices; adaxial surface drying dark brown, abaxial surface medium brown. Inflorescences racemose, lateral, 9–25 cm, short-pubescent; flowers pedicellate with pedicels 0.5–2(–3.5) mm; bracts linear to narrowly oblong, c. 1.5–2 mm. Flowers 6-merous; calyx tube funnelform, sparsely short-pubescent to glabrate; sepals narrowly oblanceolate (to narrowly oblong) with acute apex, 1.8– 3.8 mm; petals narrowly lanceolate to narrowly oblong with acute apex, 2.5– 3.8 mm; sepals and petals yellowish green, accrescent, moderately longer than calyx tube, in fruit ascending (partly and irregularly reflexed), ciliate with straight well-separated hairs, both surfaces glabrous or glabrate; sepal glands shortpubescent; filaments 1.6–2.5 mm, glabrous; ovary moderately pubescent with thick straight hairs; styles 4, c. 1.6 mm, basally pubescent. Candollea 73, 2018 Homalium sect. Blackwellia (Salicaceae) in Madagascar – 233 Vernacular name. – “Varikanda” (Rakotovao 4714). Distribution, ecology and conservation status. – The two known collections of H. longiracemosum are from a small region in extreme southeastern Madagascar, where they occur in lowaltitude humid forest. Though both are indicated in TROPICOS (2018) to be from the protected area of Tsitongambarika, the very small Area of Occupancy (8 km ²), with two populations in very close proximity, supports a conservation status of “Vulnerable” [VU D2]. Notes. – Homalium longiracemosum is distinguished from other species of sect. Blackwellia with pubescent sepal glands and racemose inflorescences by its long, narrow, long-petioled leaves and long sturdy racemes. Both known specimens, which were collected around the same time but from two different localities, evince an unusually high level of insect damage. Paratypus. – MADAGASCAR. Prov. Toliara: Taolagnaro, Mahatalaky, Farafara Vatanibe, forêt humide à haute futaie à l’W du village de Malama, au bord de la grande riv. Belomotse, 24°51'02"S 47°00'20"E, 135 m, 14–18.XI.2009, fl., Rakotovao et al. 4513 (MO)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 232-235, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["TROPICOS (2018). Missouri Botanical Garden, Saint Louis [http: // www. tropicos. org]."]}

6. Homalium erianthum (Tul.) Baill. in Bull. Mens. Soc. Linn. Paris 1: 574. 1886. ≡ Blackwellia eriantha Tul. in Ann. Sci. Nat., Bot. ser. 4, 8: 62. 1857. Lectotypus (first step designated by SLEUMER, 1973: 257; second step designated here): MADAGASCAR. Prov. Antsiranana: Nossi-bé, bords de la riv. de Djabal, II.1849, fl., Boivin 2125 (P [P00624053]!; isolectotype: BM, G [G00018409] image seen, L [L0010916] image seen, P [P00624052]!). Syntypus: “ Provenit etiam in insula Mauritiana ” [cultivated], Commerson s.n. (P, not seen). Tree to 22 m, 26 cm dbh; bark platanoid; young twigs sparsely short-pubescent to glabrous (moderately pubescent). Leaves elliptical to narrowly (broadly) elliptical (seldom somewhat ovate), 6–13(–15) × 2–6.6 cm; petiole (4–)6–9(–13) mm; base convex (rounded); apex short-acuminate, acute to obtuse, or rounded; margins crenate-serrulate (barely repand), with glands small, in margins of tooth apices; adaxial surface drying darker brown (to near black, paler, or grayish), abaxial surface brown (greenish). Inflorescences paniculate (rarely partly racemose), lateral and pseudoterminal, (3–) 5.5– 15.5 cm, shortpubescent, rarely to villous; flowers pedicellate with pedicels (0.6 –)1 –5(–6) mm; bracts ovate to elliptical, lanceolate or ligulate, 0.8–3.5 mm. Flowers 9–10(–11)-merous; calyx tube long-funnelform with rounded base, moderately to sparsely short-pubescent; sepals ligulate (to narrowly oblong, narrowly deltoid) with acute apex, 1.4–4 mm; petals ligular to narrowly spatulate (narrowly elliptical, oblanceolate) with acute apex, 1.4–4.2 mm; sepals and petals white to pale yellowish green or cream, accrescent, moderately shorter or longer than calyx tube, ascending (seldom partly spreading, then reflexing after anthesis), ciliate mostly towards apex, both surfaces sparsely pubescent (sometimes sparsely villous to pilose adaxially); sepal glands glabrous; filaments 1.5–2.4 mm, moderately pilose to villous; ovary sparsely pilose to villous; styles 5, 1.3–2(–3) mm, basally sparsely pilose to villous. Vernacular names. – “Hompa” (Service Forestier 12037); “Sarigidroa” (Service Forestier 15786); “Zahamena” (Service Forestier 15958). Distribution, ecology and conservation status. – Homalium erianthum is primarily native to low-elevation humid forests in the province of Antsiranana, rarely found farther southwest in Mahajanga. It is reported to grow on gneiss and quartzite. The species may be uncommon, as it has not been recollected in some well-botanized areas for several decades. The Area of Occupancy is estimated at 52 km ² and the Extent of Occurrence at c. 74,100 km ². Though most of the habitat is threatened by ongoing anthropogenic damage, the species has been collected in the protected areas of Montagne d’Ambre and Namoroka. More than ten distinct locations are known. Therefore the species’ conservation status is tentatively assessed as “Least Concern” [LC]; however, this may understate its rarity. Notes. – Among species of Homalium sect. Blackwellia with glabrous glands, H. erianthum is the only species with welldeveloped paniculate inflorescences and has unusually large leaves. SLEUMER (1973) incorrectly cited the material of Boivin 2125 at P as the holotype, which at the time constituted an effective lectotypification. However, since there are actually two duplicates at P, a second-stage lectotypification should select one of them as lectotype. The duplicate chosen has a more complete label and a fragment packet. Randrianasolo et al. 519 has paniculate inflorescences but leaves that, though possibly immature, are crowded and quite small, resembling those of H. axillare; it is possibly a hybrid between H. erianthum and H. axillare. Service Forestier 9384 has unusually small leaves and short sepals and petals that are subequal but barely and weakly ciliate, and glabrous. It is suggested that this may represent a hybrid involving H. erianthum and an unknown second parent, possibly from sect. Odontolobus rather than sect. Blackwellia proper. Additional material examined. – Madagascar. Prov. Antsiranana: Daraina, forêt de Bekaraoka, aux abords du village d’Andranotsimaty, 13°11'S 49°42'E, 130 m, 7.XI.2001, fl., Gautier & Ravelonarivo 4095 (MO, P); Ins. Sakatia prope Nosi-be, II.1880, fl., Hildebrandt 3361 (P); ibid loco, II.1880, fr., Hildebrandt 3361a (P); Vallée de l’Antsahabe, affluent de la Lokoho, 50–300 m, 10–11.III.1949, fl. & fr., Humbert 23351 (MO, P [2 sheets]); Montagne d’Ambre (S de Diego), III.1962, fl., Keraudren 1659 (P); Riv. Ankarakely, Nossibe, XII.1912, fl., Perrier de la Bâthie 6206 [?] (P); Fkt. Manakana, forêt d’Ambondrobe, 13°42'46"S 50°05'25"W, 18.V.2004, fl., Rabehevitra et al. 1062 (MO, P); Plateau de Beankany, Ambanja, 12.XII.1963, fl., Rakotozafy 335 (P); Daraina, forêt d’Ampondrabe, 12°58.33'S 49°42.25'E, 65 m, 13.IV.2004, fl., Ranirison 651 (MO, P); Ankara JB8, Diego-Suarez, 10.XI.1954, sterile, Service Forestier 12037 (P); Antanamanino, Vohémar, 18.VI.1956, fr., Service Forestier 15958 (P); Le long de l’Andrafiamena affluent du haut Rodo, 8.II.1966, fl., Service Forestier 24548 (MO [2 sheets], P [2 sheets]); Prov. de Diego-Suarez, s.d., fl. & fr., Ursch 157 (P). Prov. Mahajanga: R.N. 8, Soalala, 14.V.1951, fl. & fr., Réserves Naturelles 3309 (P); Antanimora, Ambolobozo, Analalava, 20.IV.1956, buds, Service Forestier 15786 (P). Possible hybrid specimens. – MADAGASCAR. Prov. Antsiranana: Imahagaga, Montagne des Français, 12°24'42"S 49°22'22"E, 220 m, 14.II.2005, fl., S . Randrianasolo et al. 519 (MO); Ankara, Diego-Suarez, 11.III.1954, fl. & fr., Service Forestier 9384 (P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 231-232, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328."]}

2b. Homalium axillare subsp. delphinense (H. Perrier) Appleq., comb. & stat. nova. ≡ Homalium microphyllum var. delphinense H. Perrier in Mém. Mus. Natl. Hist. Nat. 13: 289. 1940. Lectotypus (designated here): MADAGASCAR. Prov. Toliara: Fort Dauphin, s.d., fl., Cloisel 199 (P [P00624057]!; isolectotype: BM, L [L0010881] image seen). Syntypi: MADAGASCAR. Prov. Toliara: Fort Dauphin, Cloisel 180 (BM not seen, P not seen); Vinanibe, 23.X.1932, Decary 10864 (P!, BM, PRE [PRE0602194-0] image seen, S [S10-10152] image seen, TAN [TAN000589] image seen); Cap Ivatra, s.d., fl., Decary 10912 (MO [MO- 5587081]!, P [P00624056]!, G [G00018420] image seen, L [L0010882] image seen); vallée moyenne du Mandrare près d’Anadabolava, mont Vohibaria, 800–810 m (sommet), XII.1933, buds, Humbert 12626 (P [P00624051]!); FortDauphin, July, Scott-Elliot 3041 (K [K000231499] image seen, P [P00624055]!). = Homalium humblotii Baill. in Bull. Mens. Soc. Linn. Paris. 1: 574. 1886. Holotypus: MADAGASCAR: “Madag. bor.”, s.d., fr., Humblot 582 (P [P00624054]!; isotype: K [K000231500] image seen, LD [LD1215854] image seen, TAN [TAN000243] image seen, US [US00114722] image seen). Tree to 10 m, 15 cm dbh, or shrub. Leaves elliptical to narrowly (broadly) elliptical (seldom to ovate or rarely lanceolate), (2–)2.5–6.5 × (0.9–) 1.1–3.6 cm; petiole (1.5–)2–4(–5) mm; margins crenate-serrulate to subentire, with glands round or slightly elongated; adaxial surface drying darker, usually greenish or grayish, abaxial surface pale to medium brown (greenish). Inflorescences 1.5–8 cm. Filaments 0.8–2.4 mm. Vernacular names and uses. – “Fotsivony” (Humbert 12626); “Hazofotsy” (Randriatafika & Ramisy 408, Randriatafika & Rabenantoandro 715, Ratovoson 1661); “Hazombahy” (Réserves Naturelles 1812, 2303); “Hazombarorona” (Cours 2801); “Hazombato” (Cours 1898, Gautier et al. 3120); “Hazombatofotsy” (Service Forestier 13711); “Hazombatry” (Réserves Naturelles 2401); “Hazondrena” (Réserves Naturelles 2272); “Laza laza madiniky” (Gautier & Derleth 2563); “Mampay” (Hong-Wa 21); “Marankoditra” (Ramison & Ramisa 110, Randriataf ika & Ramisy 263); “Nefinzaza” (Cloisel 199); “Taimboalavo” (Réserves Naturelles 8936, 10716, Service Forestier 26398); “Tsilastra” (Service Forestier 9979); “Tsivelonatavy” (Service Forestier 6440); “Zora” (Service Forestier 6085, 6396, 7415); “Zore” (Réserves Naturelles 1350). Wood of H. axillare subsp. delphinense is used for construction (Hong-Wa 21, Ratovoson 1661, Réserves Naturelles 1812, Service Forestier 26398) and manufacture of wooden objects (Réserves Naturelles 1350). Distribution, ecology and conservation status. – Homalium axillare subsp. delphinense as herein defined is widespread and its habitat is variable, ranging from littoral forest on sand to mid- or somewhat high-elevation humid forests on gneiss. As noted above, though there is regional variation in morphology, that variation is not entirely consistent with reported habitat or altitude. It is possible that two distinct infraspecific taxa exist within this subspecies, but means of consistently separating them have not been identified. Because of the numerous localities, several of them protected, the subspecies’ appropriate conservation status is “Least Concern” [LC]. Notes. – As noted above, this subspecies remains variable and possibly heterogeneous. Sleumer annotated Cloisel 199 (P) as the lectotype of H. microphyllum var. delphinense but failed to indicate this choice in print (SLEUMER, 1973). Duplicates of three syntypes at BM that are not available through JSTOR are reported based on statements by SLEUMER (1973). Selected material examined. – MADAGASCAR. Prov. Antsiranana: RNI de Tsaratanana, 14°02'51"S 48°46'54"E, 1300 m, 27.XI.2000, fl., Birkinshaw et al. 767 (BR, K, MO); Cne Beramanja, 13°38'40"S 48°40'18"E, 632 m, 22.XI.2007, fl., Callmander et al. 711 (MO, P); Bekolosy (14°02'S 48°18'E), vallon en amont de la chute de la riv. Bekolosy, 1100 m, 15.XI.1994, fr., Gautier & Derleth 2563 (MO, P); vallée inférieure de l’Androranga, affluent de la Bemarivo (NE) aux env. d’Antongondriha, mont Anjenabe, 1000 m, 3–7.XI.1950, fl., Humbert & Capuron 24146 (P); Montagnes au N de Mangindrano jusqu’aux sommets d’Ambohimirahavavy (partage des eaux Mahavavy-Androranga), 1600 m, 19.I-12.II.1951, fl., Humbert & Capuron 24915 (P); Manongarivo Massif above village of Ambodisakoana, 14°05'S 48°20'E, 16.X.1994, fl., McPherson & van der Werff 16356 (MO); Daraina, forêt de Binara, 13°16'S 49°36'E, 960 m, 19.XI.2005, fl., Nusbaumer & Ranirison 1636 (MO, P); Réserve Spéciale de Manongarivo, 14°04'34"S 48°22'29"E, 1100–1350 m, 15.VIII.1994, fl., Randriambololona et al. 121 (K, MO, P); Sainte Luce, 24°44'27"S 47°10'55"E, 50 m, 19.XI.2011, fl., Ratovoson 1661 (MO); forêt de Kalabenono, 13°38'59"S 48°40'41"E, 785 m, 24.XI.2006, fl., Razafitsalama et al. 1117 (MO); Soahita, Ambohimitsinjo, Sambava, 22.X.1957, fr., Réserves Naturelles 9253 (P); Flanc S de l’Anjenabe, vallée de l’Andromanga, c. 900 m, 5.XI.1950, fl., Service Forestier 753 (MO, P); crête S du Betsomanga, bassin de l’Androranga, dct. de Sambava, 900 m, 19.XI.1950, fl., Service Forestier 835 (P); massif de l’Ambohimarahavavy, [Bealanana?] S du plateau de Marofameno, 1500 m, 7.II.1951, fl., Service Forestier 994 (P); Manongarivo massif, 14°05'S 48°20'E, 500–1000 m, 20.X.1994, fl., van der Werff & McPherson 13533 (MO); vallée de l’Ambahatra (cours moyen), Maevatanana (haut de pente), 13°54'51"S 48°29'08"E, 380 m, 19.XI.2000, fl., Wohlhauser et al. 319 (MO, P). Prov. Fianarantsoa: Ambodiriana rive droite du fleuve, 700 m, 14.XII.1944, fl., Cours 1898 (P [2 sheets]); Ambodimanga à Antanambao, 1200 m, 11.X.1945, fr., Cours 2801 (P); Riambavy Waterfall, 22°08'45"S 46°53'31"E, 1760 m, 8.XI.2003, fr., Phillipson & Rakotonandrasana 5689 (BR, CAS, G, K, L, MO, P); Tandrokomby, massif de l’Andringitra, R.N. V, 4.X.1948, fl., Réserves Naturelles 1812 (P [2 sheets]); Ambodirindahy, Sendrisoa, 7.X.1950, fl., Réserves Naturelles 2272 (P [2 sheets]); RN V, Ambalavao, Zazafo, 23.X.1949, fl. & fr., Réserves Naturelles 2401 (P [2 sheets]); Antambohobe, 12.III.1960, fl., Réserves Naturelles 10716 (P); Fenoarivo, Ilakatra, Vohipeno, 20.XI.1952, fl., Service Forestier 6440 (P); Ifanadiana SW, 19.XII.1953, fr., Service Forestier 9979 (P); Ampasinambo, Nosy-Varika, 25.XI.1954, fl., Service Forestier 13711 (P); cTOn Amtambohobe, village le plus proche Ankazomby, Lohony, 15.VIII.1967, fl., Service Forestier 26398 (P); Massif de l’Isalo, à l’W de Ranohira, 20.XII.1968, fr., Service Forestier 28506 (P). Prov. Mahajanga: Fkt. Andranomena, Befandriana Nord, 15°08'14"S 49°21'27"E, 1084 m, 12.II.2008, fl., Ravelonarivo et al. 2793 (G, MO). Prov. Toliara: Réserve Spécial de Kalambatritra, forêt d’Analamaro, 23°28'00"S 46°23'20"E, 1420 m, 6.XI.2004, fl., Andrianjafy et al. 527 (MO, P); District de Fort-Dauphin, Vinanibe, 23.X.1932, fl., Decary 10863 (P); Fort Dauphin, forêt de Mandena, 16.III.1985, fr., Dorr et al. 3978 (MO, P); Petriky Forest, 0–10 m, 25°04'S 46°51'E, 13.III.1989, fl., Gereau & Dumetz 3260 (MO, P); Ampasimena, Mt Tsimokotra-Vohimavo, 24°20'11"S 47°08'55"E, 542–547 m, 30.XI.2001, fl., Hong-Wa 21 (MO); Petriky, 25°05'S 46°52'E, 2.XI.1989, fl., McPherson 14361 (MO, P); Fkt. Sainte Luce, bloc S9, 24°46'28"S 47°09'50"E, 3 m, 4.XI.2003, fl., Rabenantoandro et al. 1558 (MO, P); Fort-Dauphin, forêt au N de Manafiafy, 12.XI.1990, fl., Rabevohitra 2442 (MO, P); Ambohisapa, Thalweg à l’E d’Ambatotsorongorongo, 25°05'11"S 46°46'13"E, 31 m, 23.II.2009, fr., Rakotovao et al. 4402 (MO); Mandromodromotra, 24°55'22"S 47°01'28"E, 9 m, 6.XII.2006, fl., Ramison & Ramisa 110 (MO); Mahatalaky, forêt de Belavenoka, 24°50'S 47°04'E, 17.XI.2007, fr., Ramison 410 (MO, P); Station forestière de Mandena (M.16), 30.VIII.2001, fl., Randriatafika & Ramisy 263 (MO, P); Fkt. Petriky, 25°03'19"S 46°51'55"E, 18.II.2004, fl., Randriatafika & Ramisy 408 (MO); Antsotso, forêt Ivohibe, 24°33'52"S 47°11'43"E, 386 m, XI.2005, fl., Razakamalala et al. 2440 (MO, P); Réserve forestière de Mandena, 16.XI.1948, fr., Réserves Naturelles 1350 (P [2 sheets]); Vinanibe, à l’W de Fort-Dauphin, 7.XII.1968, fl., Service Forestier 28623 (P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 228-229, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328."]}

10. Homalium micranthum (Boivin ex Tul.) O. Hoffm., Sert. Pl. Madagasc.: 18. 1881. , ≡ Blackwellia micrantha Boivin ex Tul. in Ann. Sci. Nat. Bot. ser. 4, 8: 63. 1857. Lectotypus (first step designated by SLEUMER, 1973: 263; second step designated here): MADAGASCAR. Prov. Antsiranana: Île Nossi Bé, 9.XII.1840, fr., Pervillé 389 (P [P04704026]!; isolectotype: K [K000231493, K000231494] images seen, P [P04704030, P04704025]!, TAN [TAN000588] image seen). Tree to 25 m, 70 cm dbh; bark grayish, platanoid; young twigs glabrous or minutely (to short-)pubescent. Leaves elliptical to oblong-elliptical or narrowly oblong (occasionally somewhat obovate to oblanceolate or ovate to lanceolate), (2.4–)3.7– 9.5 × 1.3– 3.4 cm; petiole (1 –)2– 5(–6) mm; base convex (rounded, cuneate); apex acuminate (rounded, acute, cuspidate); margins serrulate to crenate-serrulate on apical half, with glands elongated, at or above tooth apex, often producing exudate; adaxial surface drying dark brown to blackish (greenish), abaxial surface brown (olive). Inflorescences racemose, lateral, often paired (sometimes very numerous), (1.8–) 2.5–10 cm, short-pubescent (in South villous); flowers pedicellate with pedicels (0.2–)0.4 – 1.6(– 2.5) mm; bracts lanceolate to ligulate or ovate, 0.3 – 1.3(– 2) mm. Flowers (6)7(8)-merous; calyx tube funnelform with base broadening in fruit, sparsely pubescent (sometimes becoming glabrous); sepals more or less ligulate with acute apex, 1.0– 3.8 mm; petals ligulate to spatulate (narrowly elliptical) with acute apex, 1.4–4.4 mm; sepals and petals white to pale or yellowish green, sometimes red-tinged, or pink (probably only in fruit), accrescent, moderately to slightly longer than calyx tube, usually mostly ascending in flower, sometimes fully reflexed in early flower, commonly becoming irregularly spreading to reflexed in fruit, ciliate, both surfaces sparsely pubescent to glabrous; sepal glands pubescent; filaments (0.9–) 1.3–2.7 mm, glabrous (sparsely pilose); ovary pubescent; styles 4, 1.4–2.6 mm, pubescent at base. Vernacular names and use. – “Fotsiakara” (Service Forestier 16265 [Taisaka dialect]); “Janganito” (Antilahimena 211, Birkinshaw & Antilahimena 540, Réserves Naturelles 3004, Service Forestier 7718); “ Janganito malandy” (Madiomanana et al. 279, Service Forestier 3881, 11074, 13108); “Zanganito” (Réserves Naturelles 9398); “Zanganito malandy” (Service Forestier 11514). Wood of H. micranthum is used for construction (Service Forestier 16265). Distribution, ecology and conservation status. – Homalium micranthum is mostly native to northeastern Madagascar, primarily from Antsiranana; it has been reported on laterite. As presently circumscribed, it includes several specimens from eastern and southeastern Madagascar. As such, it has a large range and has been collected in several protected areas (Lokobe, Montagne d’Ambre, Manombo, Masoala, Manongarivo) so its conservation status must be estimated as “Least Concern” [LC]. Notes. – Most collections of H. micranthum are from northeastern Madagascar, almost all from Antsiranana. Several collections outside Antsiranana have previously, or now for the first time, been tentatively identified as having affinities to H. micranthum. The geographic range of these specimens is suspiciously broad, but for the most part, though they are all at the extreme of the species’ morphological range in at least one or two characters, they are not clearly distinguished from it. Specimens from eastern littoral forests have moderately villous inflorescences, in contrast to the more usual short-pubescent inflorescences. The few specimens from the southeast (Prov. Tulear) are of particular interest: mature leaves are not available for all, but they seem to share an unusual leaf shape (obovate, rounded-cuspidate apex, sometimes cuneate base) and to have the calyx tube normally, rather than rarely, glabrate at maturity. The bark description in the label data of Service Forestier 16265 from Fianarantsoa is somewhat inconsistent with descriptions from Antsiranana. Since the floral morphology of these specimens is unremarkable, they are not segregated at this time, but the availability of more material in future might allow them to be recognized as distinct. Among the atypical specimens of H. micranthum is Perrier de la Bâthie 18374, originally a syntype of H. viguieri H. Perrier. This specimen seems to fit better in H. micranthum than in H. viguieri, primarily due to its long and relatively narrow petals; its relatively large, short-acuminate leaves and mostly shortpedicellate flowers are more typical of H. micranthum, though not totally unknown in H. viguieri. It has slightly longer bracts than typical of H. micranthum and unusual sparsely pilose filaments, but otherwise resembles typical H. micranthum. Further investigation of any surviving population in the area would be desirable. Another problematic specimen is Service Forestier 23946 (P), from coastal Fianarantsoa between Farafangana and Manombo. This collection resembles H. micranthum from the same area, with some unexpected characters; in particular, the sepal glands are glabrous. It may be a hybrid involving H. micranthum and a species with glabrous glands, most plausibly H. axillare (which is not known from that area, but which has a sufficiently wide range that its presence would not be surprising). The protologue of Blackwellia micrantha cited Pervillé 389 and Boivin 2125. The latter collection provided the holotype of the simultaneously published B. eriantha Tul. A portion of the material of Pervillé 389 at P was wrongly marked in the herbarium with a Boivin label and “2125”. The real Boivin 2125 is very different in appearance, as Homalium micranthum and H. erianthum are not at all confusable. It may be presumed that Tulasne intended to quote label data, including the misused number, and did not intend to include the real Boivin 2125 in Blackwellia micrantha. Therefore, Pervillé 389 constitutes the only original material. SLEUMER (1973) stated that the “ holotype ” was “Pervillé Herb. 389 in Herb. Boivin 2125” at P, with an isotype at K. This would constitute an effective first- step lectotypification, but there are actually three probable duplicates of Pervillé 389 at P, from which one still needs to be selected as lectotype. The sheet chosen here, [P04704026], is preferred because it is the largest fragment and also the least susceptible to confusion, since it is marked only with a printed “Pervillé 1841” label and a handwritten “No. 389”. The second sheet, [P04704030], bears a printed “Boivin 1847–1852” label and hand-marked “no. 2125”, as well as handwritten labeling that says “Pervillé leg. et... comm, sub n. 389” and also “2125. Nossi-bé. Bernier comm. 1846” [i.e., before the date of Boivin’s voyage]. The third sheet, [P04704025], bears only the Boivin label and herbarium number, with a fragment packet marked “ Blackwellia micrantha Tul. ”; it is Homalium micranthum and can be presumed to be unlabeled material of Pervillé 389. Selected material examined. – MADAGASCAR. Prov. Antsiranana: Nosy-Be, RNI Lokobe, 13°24'30"S 48°18'15"E, 0–50 m, 7.XII.1994, fl. & fr., Antilahimena 211 (MO, P); Nossi-be, env. d’Ambatozandry, 50–250 m, 14.XII.1967, fr., Bernardi 11867 (P); Réserve Naturelle Intégrale No. 6, Lokobe, 5 km SE of Hell Ville, 13°25'S 48°18'E, 100 m, 28.XI.1992, fl., Birkinshaw 193 (MO, P); Ramena valley, fkt. Antsahabe, 13°49'30"S 48°46'59"E, 500 m, 10.X.1998, fl., Birkinshaw & Antilahimena 540 (MO, P); Ambahatra cours supérieur, 13°59'S 48°26'E, crête entre les deux bras de l’Ambahatra, 24.V.2000, fr., Gautier & Rakotomamonjy 3649 (MO, P); Nosi-be, IX.1879, fr., Hildebrandt 3178 (P [2 sheets]); Ampasindava, forêt d’Andranomatavy, 13°39'42"S 47°58'41"E, 615 m, 30.XI.2009, fr., Madiomanana et al. 279 (P); Parc National de Montagne d’Ambre, 12°32'S 49°11'E, 990–1100 m, 16–20.XI.1992, fl., Malcomber et al. 1805 (MO, P); Daraina, forêt d’Antsahabe, 13°12'53"S 49°33'19"E, 796 m, 14.I.2004, fr., Nusbaumer 961 (MO, P); vallée du Sambirano, I.1909, fl., Perrier de la Bâthie 4591 (P); H[aut]e Mananjeba, 300 m, IV.1913, fl., Perrier de la Bâthie 6702 (P); Daraina, forêt de Binara, 13°14.06'S 49°37.31'E, 300 m, 31.III.2004, fl. & fr., Ranirison 552 (MO, P); Marotongoza, Nossi-Bé, 12.III.1957, fr., Réserves Naturelles 9398 (P); Ambodimangatelo, Ambanja, 12.X.1951, fr., Service Forestier 3880 (P); Ambaliha, Ambanha, 18.X.1953, fr., Service Forestier 7718 (P); Maromiandra, Ambanja, 6.X.1954, fl., Service Forestier 11074 (MO [2 sheets], P); massif de l’Antsatrotro (Manongarivo), 900 m, XI.1954, fl., Service Forestier 11514 (P [2 sheets]); Bekolosy, Ambanja, 17.XI.1954, fl., Service Forestier 13108 (P); massif de la Montagne d’Ambre, rive droite de la riv. des Makis entre la Station des Roussettes et la grande cascade, 18–20.XI.1958, fr., Service Forestier 20039 (MO, P); Prov. de DiegoSuarez, s.d., fr., Ursch 205 (P). Prov. Fianarantsoa: Distr. Farafangana, cTOn Ihorombé, Manombo, 31.V.1956, fr., Service Forestier 16265 (P). Prov. Toamasina: Ambatovola, Vohitra, 700 m, I.1928, fl. & fr., Perrier de la Bâthie 18374 (P); [Masoala] Peninsula, trail just above Antalavia River on S bank, 1–2 km from coast, 15°45'S 50°02'E, 50–100 m, 19.II.1988, fl. & fr., Schatz et al. 1922 (MO, P). Prov. Toliara: Fort-Dauphin, s.d., fl., Cloisel 188 (MO, P); près de la riv. Antorendrika avant Belavenona, 24°52'S 47°07'E, 0–20 m, 22.III.1989, fl., Rabevohitra 1773 (MO, P); Fkt. Ambandrika, Analavinaky S8, 24°46'29"S 47°08'57"E, s.d., fl. & fr., Randriatafika & Rabenantoandro 412 (MO, P)., Published as part of Applequist, Wendy L., 2018, A revision of the Malagasy species of Homalium sect. Blackwellia (Salicaceae), pp. 221-244 in Candollea 73 (2) on pages 236-238, DOI: 10.15553/c2018v732a7, http://zenodo.org/record/5724607, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Natl. Belg. 43: 239 - 328."]}

9b. Homalium planiflorum subsp. roseiflorum Appleq., subspec. nova (Fig. 5). Holotypus: MADAGASCAR. Prov. Toliara: Anosy, Taolagnaro, Ampasy Nahampoana, for��t littorale d���AmbavaranoMandena, 24��57���S 47��00���E, 5 m, 13.V.2006, fl., Rabenantoandro & Ramisy 1765 (MO-6450859!; iso-: G [G00418622] image seen, P [P00806077]!, TAN). Homalium planiflorum subsp. roseiflorum Appleq. differs from H. planiflorum (Tul.) Baill. subsp. planiflorum in having inflorescences mostly clustered at twig ends and occasionally paniculate (vs scattered along twigs and almost always racemose), and pink to reddish flowers (vs whitish to greenish or yellowish flowers). Leaves narrowly elliptical to oblanceolate (aberrantly obovate), 3.5-7.7(-8.7) �� 1-2.3(-2.9) cm. Inflorescences racemes, often clustered near twig ends, or occasionally panicles with a few long branches. Flowers pink to reddish, commonly 8-merous, less often 7-merous (9-merous). Vernacular names. ��� ���Hazofotsy��� (Faliniana et al. 2, Randriatafika et al. 319, R��serves Naturelles s.n., Soanary 26); ���Zora��� (Rabenantoandro et al. 336, Ratovoson 1731); ���Zora fotsy��� (Ratovoson 1629, 1692). Uses. ��� Wood of H. planiflorum subsp. roseiflorum is reported to be hard and to be used for construction and to make tool handles (Randriatafika et al. 319, Ratovoson 1629, 1692, 1731, R��serves Naturelles s.n.). Distribution, ecology and conservation status. ��� Homalium planiflorum subsp. roseiflorum is confined to coastal forests in southeastern Madagascar. All collections are from a very small portion of Toliara province; most are from the protected areas of Mandena and Sainte Luce, with single collections from four other localities in close proximity (all within ca. 13 km of one of the significant populations). GeoCAT calculates an EOO as c. 320 km 2 and an AOO of 40 km 2. Since there may be six distinct populations, the species cannot be formally assessed as ���Vulnerable��� [VU D2]. However, all localities are so close together that they could easily be affected by a single event such as a typhoon, and habitat outside the rather limited boundaries of the protected areas is at high threat of anthropogenic damage. A status of ���Near Threatened��� [NT] is therefore recommended to call attention to this concern. 1 cm 1 cm Notes. ��� Homalium planiflorum subsp. roseiflorum is distinguished from the typical subspecies by its flower color (pink to red) and, to some extent, by inflorescence morphology (often clustered near twig apices, occasionally paniculate). Its leaves are on average somewhat narrower and more likely to tend to oblanceolate than those of subsp. planiflorum, and the average number of sepals and petals per flower is slightly greater (more often eight than seven, and rarely nine, whereas seven- and eight-petaled flowers are about equally common in subsp. planiflorum). However, these latter characters overlap too much to be taxonomically useful. Paratypi. ��� MADAGASCAR. Prov. Toliara: Mandena, ��tude parcelles campement, 24��57���S 47��00���E, 0-10 m, 17.IV.1989, ster., Dumetz et al. 666 (P); Sainte Luce, 24��47���S 47��10���E, 0-10 m, 20.X.1989, fl., Dumetz et al. 754 (MO, P); Mandena, 24��57���S 47��00���E, 6.XII.1989, fl. & fr., Dumetz & McPherson 1127 (MO, P); Mandena, 0-10 m, 24.I.1990, fr., Dumetz 1236 (MO, P); Sainte Luce, 24��46���46������S 47��10���17������E, 10 m, 15.XII.2000, fl., Faliniaina et al. 2 (MO); Sainte Luce, proposed QIT mining conservation area, 24��46���42������E 47��10���15������E, 0 m, 13.X.2000, fl., Hoffmann et al. 209 (MO [2 sheets]); Near Mandena beyond QIT camp, 24��57���S 47��00���E, 25 m, 31.X.1989, buds, McPherson & Dumetz 14339 (P); ibid. loco, 20 m, 4.XI.1989, fl., McPherson 14369 (MO, P); M7, Taolagnaro, M 7 [Mandena], 24��57���31������S 47��00���02������E, 0-10 m, 6.X.2000, fl., Rabenantoandro et al. 336 (G, MO, P); Sainte Luce, pr��s du campement QMM, 24��46���47������S 47��10���17������E, 3 m, 29.X.2003, buds, Rabenantoandro et al. 1522 (MO, P); Mandena, 24��57���S 47��02���E, 10 m, 13.I.1990, fr., Rabevohitra 2123 (MO, P); Taolagnaro, �� 1 km au N d���Evatra, 24��57���S 47��06���E, 17.I.1990, fl. & fr., Rabevohitra 2153 (MO, P); Sainte Luce, for��t d���Analavinaky �� l���W d���Ambandrika, 24��46���30������S 47��09���00������E, 21 m, 24.XI.2009, post-fl., Rakotovao 4737 (MO); Fkt. Fenoambony, E du village Androangabe, 24��53���S 46��59���E, 58 m, 5.XII.2009, post-fl., Rakotovao et al. 5042 (MO); Sainte Luce, S5 [forest parcel], 24��47���S 47��11���E, 2.II.2008, post-fl., Ramison & Rabehevitra 561 (MO); Sainte Luce, 4-5 km N of Manafiafy along main trail into forest, 24��47���S 47��09���E, 0-10 m, 19.XI.1996, fl., A. Randrianasolo 567 (G, MO, P); Fkt. Ampasy-Nahampona, Station foresti��re de Mandena (M3), 3.X.2001, fl., Randriatafika et al. 319 (MO); Sainte Luce, S 10, 3 km au N d���Ambandrika, 24��44���39������S 47��10���45������E, 50 m, 19.XI.2011, fl., Ratovoson 1629 (MO); Sainte Luce, S11, 3.1 km au N d���Ambandrika, 24��44���26������S 47��10���38������E, 20 m, 21.XI.2011, fl., Ratovoson 1692 (MO); Mahatalaky, Belavenoko, S 5, 1 km E de Belavenoko For��t, 24��49���51������S 47��06���49������E, 20 m, 23.XI.2011, fl., Ratovoson 1731 (MO); Mandena (route Nancy), 6.XII.1949, fl., R��serves Naturelles (Ranjatson) s.n. (P); Sainte Luce, QMM Forestry Station, parcel S9, 24��46���05������S 47��10���16������E, 11 m, 23.I.2006, fl., Rogers et al. 970 (G, MO); for��t de Vinanibe, pr��s de Fort Dauphin, 10.XII.1961, fl., Service Forestier 20513 (P [2 sheets]); Fkt. Ampasinahampoana, for��t de Mandena, block M15, 24��57���09������S 47��00���10������E, 6.XII.2001, fl., Soanary et al. 26 (MO) Intermediate or unidentifiable specimens. ��� MADAGASCAR. Prov. Toliara: vicin. Fort-Dauphin, 15.XI.1967, fl., Bernardi 11517 (P); Fort-Dauphin, s.d., fl., Cloisel 120 (P); Fort-Dauphin, Ambinanibe, 28.II.1973, fl., Debray 1985 (P); Petriky forest, N of NE corner of Lake Andranany, NW of large dune at end of road, 25��03���S 46��53���E, 0-10 m, 12.IV.1989, ster., Gereau et al. 3365 (P); Fort Dauphin, VI.1889, fl., Scott Elliot 2835 (P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 44-46, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136

9. Homalium planiflorum (Tul.) Baill. in Bull Mens. Soc. Linn. Paris 1: 574. 1886. �� Blackwellia planiflora Boivin ex Tul. in Ann. Sci. Nat., Bot. ser. 4, 8: 64. 1857 [nom. conserv. prop.; APPLEQUIST, 2017]. �� Blackwellia gracilis Blume in Mus. Bot. Lugd.-Bat. 2: 26. 1856. Holotypus: MADAGASCAR: ��� Ile Ste. Marie ���, s.d., Richard 297 (L [L0010991] image seen; iso-: G [G00018396] image seen, P [P00375177, P00375178]!). Tree to 30 m tall, 60 cm dbh; bark young twigs glabrous (minutely pubescent when very young). Leaves: petiole (2-)3- 8(-14) mm long, glabrous (to sparsely and minutely pubescent); blade narrowly elliptical to elliptical or oblanceolate (aberrantly obovate), (2.8-)3.5-10.3 �� 1-4 cm; base convex (cuneate to attenuate); margins crenate-serrate to serrulate, at least apically (partly subentire); apex acute to acuminate (to rounded, emarginate, cuspidate). Inflorescences racemose (paniculate with a few long branches, racemiform panicles), sometimes clustered terminally, (2.2-)5-10(-22) cm long, moderately to sparsely short-pubescent; flowers mostly in small clusters, pedicellate with pedicels (0.4-)1-3(-3.5) mm long; bracts broadly ovate-deltoid to transversely ovate; bracteoles ovate to deltoid, small, not thickened, caducous. Flowers 7-8 (-9)-merous, whitish to pale green, pale yellow or creamcolored or pink to reddish; sepals lanceolate to narrowly oblong-lanceolate (ovate), 0.5-0.9(-1.1) mm long; petals oblanceolate (to broadly spatulate, narrowly obovate, to somewhat oblong or narrowly elliptical), 1-1.7 mm long, margins ciliate and both surfaces sparsely pubescent to glabrate, apex acute to rounded; filaments (0.5-)0.6-1.2(-1.4) mm long; upper surface of ovary moderately pubescent; styles 3-4, (0.4-) 0.8-1.4 mm long. Notes. ��� Most specimens assigned to H. planiflorum have white to pale green or yellowish flowers. A group of specimens from southeastern Madagascar that are characterized by pink or reddish flowers are herein segregated as subsp. roseiflorum Appleq. While specimens of subsp. planiflorum have sometimes large leaves and racemes usually borne along the length of the twigs, those of subsp. roseiflorum often have relatively small leaves and relatively large (though not long) distal clusters of racemes, or occasionally paniculate inflorescences borne well below twig apices; however, these characters are not fully consistent. Several specimens from the extreme southeast, where both subspecies occur, are not assigned to either subspecies, though these collections generally have narrow leaves of modest size, typical of subsp. roseiflorum. One is sterile and two are only weakly flowering with one or two racemes, so subspecies identity cannot be confidently assigned without information on flower color. Two other collections are described as having possibly reddish flower color (���terre br��l��e��� according to Cloisel and ���fulvo-rosei (sordid)��� according to Bernardi) but the distribution of their well-developed inflorescences appears to be more consistent with subsp. planiflorum, and flowers may brown as they turn to fruit. The inability to assign identities to some specimens in the zone where these taxa are sympatric supports a view that they are not fully genetically isolated, and that the distinctions between them are not adequate to justify recognition at the species level. Alternatively, the intermediate specimens could be interpreted as hybrids between two distinct species; however, because three fixed differences between the two morphological forms have not been observed, the conservative approach of treating them as conspecific has been preferred. Service Forestier 9952 (Andovolava, Nosy-Varika, 14.XI.1953) is a specimen in poor condition with few flowers; the leaves are very large and broad (up to at least 10 �� 5.5 cm). It may represent a regional variant, an aberrant individual, or an undescribed species; the available material is inadequate to clarify its status. Additional collections from this population would be highly desirable. Blackwellia planiflora was published in 1857, the year after B. gracilis Blume, though its author, Tulasne, was apparently unaware of the latter publication. The citation of Richard 297 in the protologue of B. gracilis explicitly referred only to the duplicate at L, which is therefore its holotype. The protologue of B. planiflora cited two syntype gatherings, Boivin 1847 and Richard 297; three duplicates each of the former are present at P and G, and two of the latter at P. PERRIER DE LA B��THIE (1946: 92) chose Boivin 1847 as ��� type ��� (i.e., the lectotype) without selecting among the available duplicates. Under Art. 52.2 of the ICN (MCNEILL et al., 2012), the citation of Richard 297, which included the holotype of B. gracilis, in the protologue of B. planiflora would appear to make the latter a superfluous and illegitimate name. Past literature has implicitly treated it as being the citation only of duplicates that were available to Boivin and Tulasne, not the duplicate at L that they presumably did not have access to (e.g., SLEUMER, 1973: 248). By that view, B. planiflora need not be treated as illegitimate. However, Art. 9.5 of the ICN states that the citation of a gathering, if not qualified or limited, is equivalent to the citation of all duplicates of that gathering, even those not seen or known of. Tulasne did not indicate that only material at P was used. The informal practice of treating a duplicate in an author���s home institution as a holotype despite the absence of a qualifying statement is seen in the literature (see Homalium lucidum above), but normally in cases that do not involve questions of legitimacy. Hence Blackwellia planiflora is illegitimate due to the citation of the holotype of B. gracile, which is therefore also the obligate type of B. planiflora, regardless of Perrier de la B��thie���s attempt to select another gathering as the (lecto) type. This creates serious problems, because under the ICN, the species under question here would have no legitimate name. Blackwellia gracilis Blume cannot now be transferred to Homalium and retain that epithet because of the existence of H. gracile Briq., a replacement name for Blackwellia gracilis Vieill. [nom. illeg., non Blume]. However, because Homalium gracile had not yet been published when H. planiflorum was described, H. planiflorum as a replacement name credited to Baillon would also be illegitimate (see Art. 58, Note 1, of the ICN; MCNEILL et al., 2012). The publication of a new name would therefore be required. However, since this is the most common species of its section, introducing an entirely unfamiliar name would cause some confusion among botanists. A proposal to conserve Blackwellia planiflora to permit the continued use of Homalium planiflorum (the preferable approach, since Prop. 235 to amend the ICN to permit this (WIERSEMA et al., 2016) was accepted at the 2017 Botanical Congress) has therefore been offered (APPLEQUIST, 2017) but will not be considered by the relevant Committees for some time yet. Key to the subspecies of Homalium planiflorum 1. Inflorescences usually racemose (rarely a racemiform or few-branched panicle), usually borne along much of the length of small twigs; flowers whitish to pale green, pale yellow, or cream-colored; widely distributed............................................................................ 9a. subsp. planiflorum 1a. Inflorescences racemose, usually clustered near twig ends, or paniculate with few long racemoid branches; flowers pink to reddish; confined to southeastern Madagascar.............................................................9b. subsp. roseiflorum, Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 42-43, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136, {"references":["PERRIER DE LA BATHIE, H. (1946). Flacourtiacees. In: HUMBERT, H. (ed.), Fl. Madagascar Comores 140.","MCNEILL, J., F. R. BARRIE, W. R. BUCK, V. DEMOULIN, W. GREUTER, D. L. HAWKSWORTH, P. S. HERENDEEN, S. KNAPP, K. MARHOLD, J. PRADO, W. F. PRUD'HOMME VAN REINE, G. F. SMITH, J. H. WIERSEMA & N. J. TURLAND (2012). International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Veg. 154.","SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","WIERSEMA, J. H., W. GREUTER & J. MCNEILL (2016). Some proposals to resolve problems relating to the conservation or rejection of names, suppression of works, and binding decisions. Taxon 65: 642 - 646.","APPLEQUIST, W. L. (2017). Proposal to conserve the name Blakwellia (' Blackwellia') planiflora (Homalium planiflorum) (Salicaceae). Taxon 66: 206 - 207."]}

6. Homalium moniliforme H. Perrier in M��m. Mus. Natl. Hist. Nat. 13: 291. 1940. Lectotypus (designated by SLEUMER, 1973: 308): MADAGASCAR. Prov. Fianarantsoa: Vondrozo, province de Farafangana, s.d., fl., Decary 5469 (P [P04679983]!; isolecto-: K [K000231481] image seen, L [L0010965] image seen). Syntypus: MADAGASCAR. Prov. Toamasina: Tampina, 25.VIII, fr., Louvel 53 (P [P04679987]!). Tree to 13 m tall, 30 cm dbh; young twigs moderately pubescent with minute hairs or glabrous (glabrate). Leaves: petiole 1-7(-10) mm long, glabrous (minutely pubescent); blade narrowly elliptical to elliptical (oblanceolate) or obovate to elliptical, (1.8-)2.7-6.8 �� 0.7-2.6(-3) cm; base cuneate to attenuate or convex; margins serrulate to crenate-serrate, shallowly wavy or subentire, sometimes slightly revolute; apex rounded-acute to rounded (slightly acuminate, rounded-cuspidate, aberrantly emarginate). Inflorescences spicate, (2-)4-7(-10) cm long, minutely pubescent or glabrous; flowers in severalflowered glomerules, sessile; bracts transversely oblong, often irregular; bracteoles reduced to minute teeth, rapidly caducous. Flowers 6-7(-8)-merous, greenish (corolla pale yellow); sepals deltoid to ovate, 0.1-0.3 mm long; petals transversely (very broadly) ovate (very broadly oblong-ovate), 0.5-0.7(-0.8) mm long, glabrous, apex rounded (rounded-obtuse); filaments 0.2-0.4(-0.6) mm long; upper surface of ovary densely shortpubescent; styles (2-)3, 0.2-0.3(-0.5) mm long. Notes. ��� Homalium moniliforme displays geographic variation in its vegetative morphology and ecological preferences. Collections from the southern provinces of Toliara and Fianarantsoa occur consistently at low altitudes on sand and have small, short-petioled, often narrowly elliptical to elliptical leaves and minutely pubescent twigs. This form extends to coastal forests in Toamasina province. Inland collections from moderate-elevation humid forests, which occur on more diverse substrates, range from Antsiranana to Toamasina provinces and rarely Fianarantsoa province. These have somewhat larger leaves that are longer-petioled and usually mostly obovate, less often elliptical; the twigs are glabrous or glabrate, rarely papillate at the extreme apices. The two geographically distinct forms are usually readily distinguishable. It therefore seems appropriate to recognize them as subspecies. The lectotype is from Fianarantsoa province at a low-moderate elevation, in a region where no other material of the species has been collected. Its morphology is somewhat intermediate between the two forms, with mostly narrow, oblanceolate leaves, and the leaves dry relatively pale, which is much more common in the small-leaved subspecies (though that is not a fully reliable character in any related group observed to date). However, it is overall more similar to material from the moderate-elevation populations, for which it would represent a substantial range extension. These populations are therefore herein recognized as subsp. moniliforme, and the coastal subspecies is described below as subsp. littoralis Appleq., to which the other syntype (Louvel 53) belongs. The original publication of Homalium moniliforme did not explicitly cite specimens, but did enumerate the vernacular names provided on the labels of both Decary 5469 and Louvel 53, which were the only two collections cited in the Flora treatment six years later (PERRIER DE LA B��THIE, 1946). SLEUMER (1973) therefore termed them syntypes. That interpretation could be questioned, since they were only indirectly cited, but they are certainly original material, and he appropriately designated the P duplicate of Decary 5469 as lectotype. Key to the subspecies of Homalium moniliforme 1. Young twigs glabrous (glabrate, minutely papillate at the apex); petiole 2-7(-10) mm long; leaf blade obovate to elliptical, narrowly elliptical, or oblanceolate, 3-6.8 �� (1.1-)1.3- 2.6(-3) cm; mid- to low-elevation humid forests....................................................................... 6a. subsp. moniliforme 1a. Young twigs moderately minutely pubescent; petiole 1-3 mm long; leaf blade narrowly elliptical to elliptical (oblanceolate), (1.8-)2.7-4.7(-5.6) �� 0.7-1.6(-1.9) cm; littoral or low-elevation coastal forests.... 6b. subsp. littorale, Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 35-37, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136, {"references":["PERRIER DE LA BATHIE, H. (1940). Revision des Flacourtiacees de Madagascar et des Comores. Mem. Mus. Natl. Hist. Nat. 13: 261 - 301.","SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","PERRIER DE LA BATHIE, H. (1946). Flacourtiacees. In: HUMBERT, H. (ed.), Fl. Madagascar Comores 140."]}

6b. Homalium moniliforme subsp. littorale Appleq., subspec. nova (Fig. 3). Holotypus: MADAGASCAR. Prov. Fianarantsoa: Fiv. Farafangana, firaisam-pokontany Mahabo Mananivo, fkt. Nosy ala, for��t d���A[g]nalazaha, 23��10���13������S 47��43���27������E, 22 m, 26.VII.2003, fl., Rabehevitra et al. 516 (MO-04810771!; iso-: G [G00418523] image seen, P [P04705768]!, TEF). Homalium moniliforme subsp. littorale Appleq. differs from H. moniliforme H. Perrier subsp. moniliforme in having young twigs moderately pubescent with minute trichomes (vs glabrous to glabrate or minutely papillate at apex), and leaves with the petiole 1-3 mm (vs 2-7(-10) mm) and the blade usually narrowly elliptical to elliptical, (1.8-)2.7-4.7(-5.6) �� 0.7-1.6(-1.9) cm. Young twigs moderately pubescent with minute hairs. Leaves: petiole 1-3 mm; blade narrowly elliptical to elliptical (oblanceolate especially when young), (1.8-)2.7-4.7(-5.6) �� 0.7-1.6(-1.9) cm; margins serrulate to crenate-serrate or subentire; apex rounded-acute to rounded (slightly acuminate with rounded tip). Vernacular names. ��� ���Fotsiakara��� (Ludovic et al. 870); ���Fotsiakara mainty��� (Ludovic 649); ���Fotsiakara minty��� (Ludovic et al. 696); ���Fotsikara minty��� (Ludovic 761); ���Hazontsindrano��� (Louvel 53). Uses. ��� Wood is used for construction (Ludovic et al. 696, 870). Distribution, ecology and conservation status. ��� Homalium moniliforme subsp. littorale is native to littoral or low-elevation coastal forests from Toamasina to Toliara provinces; it is reported to occur on sand. Coastal forest, where not protected, is one of the most threatened habitat types in Madagascar. The habitat is in fact quite fragmented; three locations are within the protected area network (i.e., Mahabo/Agnalazaha, Mandena, and Ste. Luce) but continuing decline in habitat area and quality can be expected for the remainder. There are only seven distinct locations, three of which (from Tulear) are very close together. The EOO is estimated by GeoCAT as c. 10.5 km 2, and the AOO as 56 km 2. The preliminary assessment of conservation status suggests that a status of ���Vulnerable��� [VU B 1ab(iii)+ B 2ab(iii)] would be appropriate, though one could argue for ���Endangered��� [EN] on the grounds that the habitat might be considered ���severely fragmented.��� Paratypi. ��� MADAGASCAR. Prov. Fianarantsoa: Fkt. Vohimasy, for��t d���Agnalazaha, bloc Agnanto, 23��10���09������S 47��41���51������E, 28 m, 23.III.2004, fl., Ludovic 649 (MO); Fkt. Nosiala, for��t d���Agnalazaha, bloc de Betaindambo, 23��08���42������S 47��42���18������E, 31 m, 22.IV.2004, fl., Ludovic 761 (MO); Fkt. Nosiala, bloc forestier d���Ampandramadinika, 23��12���13������S 47��42���41������E, 21 m, 8.IV.2004, fl., Ludovic et al. 696 (MO); Nosiala, for��t d���Ampitavananima, 23��11���10������S 47��43���02������E, 30 m, 5.VIII.2004, fr., Ludovic et al. 870 (MO); MahaboMananivo, for��t de Mahabo, 23��10���20������S 47��42���23������E, 29 m, 23.IX.2002, fr., Rabenantoandro et al. 961 (MO, P); ibid. loco, 23��10���37������S 47��43���E, 10 m, 25.IX.2002, fl., Rabenantoandro et al. 990 (MO, P); Trail E of Vohimasy Ankorabe, W edge of Ampanasanay forest, 23��10���56������S 47��42���00������E, 24 m, 19.VI.2004, fr., A. Randrianasolo et al. 853 (MO, P); Mahabo-Mananivo, Sahalava, 4 km �� l���E du gite MBG, 23��11���27������S 47��42���39������E, 4 m, s.d., fl., Razafitsalama et al. 1004 (G, MO, P). Prov. Toamasina: Fkt. Tanambao Ambodimanga, for��t de Menagisy 500 m du village Antanandava Martin, 16��47���S 49��43���E, 26.V.2010, fl., Lehavana & Zackarie 692 (MO); Tampina, s.d., fr., Louvel 53 (P); F��n��rive, for��t �� Mahambo, 12.VIII.1961, fl., Peltier & Peltier 3419 (P); Vohibola, �� l���W de Tampina, 12-14.VIII.1957, fl., Service Forestier 18072 (P); Mahambo, au S de F��n��rive, 30.VIII.1957, fl. & fr., Service Forestier 18146 (MO, P [3 sheets]). Prov.Toliara: Mandena, 24��57���S 47��02���E, 10 m, 13.I.1990, fl., Dumetz 1184 (MO, P); for��t au NE de la rivi��re Antorendrika avant Belavenona, 24��52���S 47��07���E, 0-20 m, 22.III.1989, fl., Rabevohitra et al. 1781 (MO, P); Forest of Ste. Luce (Manafiafy), 2 km NW of Ambandrika village, 24��46���S 47��12���E, 0-10 m, 9.VI.1994, fl., A. Randrianasolo 343 (MO, P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on page 38, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136

5. Homalium masoalense Appleq., spec. nova (Fig. 2). Holotypus: MADAGASCAR. Prov. Toamasina: Masoala Peninsula, trail along coast 3-6 km S of Ambanizana, 15��39���S 49��58���E, 0-20 m, 30.X.1992, fl., Schatz et al. 3367 (MO-6703444!; iso-: BR!, G!, K!, P!, TAN, USMS). Homalium masoalense Appleq. differs from H. longistaminum H. Perrier in having leaves mostly ovate to lanceolate (vs elliptical to broadly elliptical, less often ovate or seldom lanceolate), with the apex usually acuminate (vs acute to obtuse or rounded, rarely short-acuminate) and the petiole shorter, 3-5.5 vs 4-11 mm long; inflorescences longer, 5-11 cm long (vs (2.2-) 4-6.5 cm long), moderately pubescent with the surface visible (vs at least partly canescent), the flowers sessile or subsessile with pedicels to 0.5 mm long (vs pedicellate with pedicels 0.3-1.4 mm long); sepals oblong-lanceolate to narrowly oblong, 1.2-1.4 mm long (vs deltoid to ovate, at most 0.6 mm long); petals narrowly oblong, 1.5-1.7 mm long, only modestly longer than sepals, with the apex acute (to narrowly rounded) (vs oblong to somewhat obovate, 0.8-1.2 mm long, much longer than sepals, with the apex rounded to obtuse). Tree to 16 m tall; young twigs glabrous. Leaves: petiole 3-5.5 mm long, glabrous; blade ovate to lanceolate (elliptical), (3.5-)5-9.1 �� (1.8-) 2-4.2 cm; base convex to rounded with the extreme base attenuate; margins irregularly crenulate to slightly undulate, usually for most of their length, or subentire; apex acuminate (acute, emarginate). Inflorescences racemose, 5-11 cm long, moderately pubescent with short erect trichomes that are often relatively dense but leave the surface visible; flowers mostly in clusters of up to 4, sessile or subsessile with pedicels to 0.5 mm; bracts transversely oblong to transversely oblong-ovate; bracteoles rapidly caducous (or possibly absent?). Flowers (6-)7-merous, greenish yellow; sepals oblong-lanceolate to narrowly oblong, 1.2-1.4 mm long; petals narrowly oblong, 1.5-1.7 mm long, margins ciliate, abaxial surface pubescent, mostly glabrous adaxially, apex acute (to narrowly rounded); filaments (0.7-) 1.5-1.9 mm long; upper surface of ovary moderately pubescent with long trichomes; styles 3-4, (1.1-) 1.3-1.6 mm long. Distribution, ecology and conservation status. ��� Homalium masoalense is known from a single collection, made in coastal forest of the Masoala Peninsula in northwestern Madagascar just outside the Masoala protected area in an area with human activities. Since accessible coastal areas of Masoala have been relatively well collected, the new species is likely to be rare. The new species should be considered ���Vulnerable��� [VU D2] based on its rarity and its probable occurrence within the Masoala protected area even if no collection has yet been made there. Notes. ��� Homalium masoalense and another new species, H. ovatifolium Appleq., which is discussed below, have clear affinities to H. longistaminum, previously placed in Homalium sect. Blackwellia , which likewise has unusually long stamens and styles compared to most species of Homalium sect. Odontolobus . All three have ciliate petals that are relatively long compared to most species of Homalium sect. Odontolobus. Homalium masoalense and H. ovatifolium are most readily distinguished from H. longistaminum by having usually ovate leaves (in the former species frequently to lanceolate) with the apex usually acuminate or long-acute. Homalium masoalense is distinguished from both H. longistaminum and H. ovatifolium by having larger flowers and in particular, much longer sepals both proportionately and in absolute measurements (1.2-1.4 mm long, more or less narrowly oblong, versus at most 0.7 mm long and deltoid to ovate in the other two species). The flowers are thereby more similar in proportion to those of H. brachystylis, which has smaller, more or less elliptical leaves and solitary flowers, than to those of any other species in Homalium sect. Odontolobus. Homalium longistaminum and H. ovatifolium both have thickened pubescent bracteoles; no bracteoles were observed on the type collection of H. masoalense., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on page 35, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136

7. Homalium ovatifolium Appleq., spec. nova (Fig. 4). Holotypus: MADAGASCAR. Prov. Antsiranana: Ambilobe, Marivorahona, village le plus proche Betsimiranja, for��t d���Andohan���Antsohihy, Ambohibe, RN6 �� 1 km au SE d���Ambilomagodra, 13��02���31������S 49��09���19������E, 100 m, 17.VII.2005, fl., Randrianaivo et al. 1206 (MO-6082912!; iso-: CNARP, P [P04679126]!, TAN). Homalium ovatifolium Appleq. differs from H. longistaminum H. Perrier in having leaves larger, (4.2-)6.2-10.2 �� (2.1-) 3.1-5.2 cm (vs (3.2-)3.5-7.5 �� 1.6-3.3(-4.8) cm), ovate (to elliptical) (vs irregularly elliptical to broadly elliptical, ovate or rarely lanceolate) with the base rounded (vs convex to rounded), margins less toothed, sometimes subentire (vs irregularly crenate apically to crenate-serrate for most of their length), apex acute to acuminate (vs acute to obtuse or rounded, rarely shortacuminate), and petals narrowly oblong (vs oblong to obovate), the abaxial surface pubescent and the adaxial surface largely glabrous (vs both surfaces pubescent, the abaxial surface densely so and often the base of the adaxial surface densely so). It differs from H. masoalense Appleq. in having possibly broader leaves, ovate or rarely elliptical (vs often lanceolate), with the petiole longer, 5-11 mm long (vs 3-5.5 mm long); inflorescences shorter, (2.2-) 4-6.5 cm long (vs 5-11 cm long), often canescent (vs moderately pubescent), the flowers more often pedicellate with pedicels 0.5-1 mm long (vs absent or up to 0.5 mm long); sepals ovate to deltoid or narrowly deltoid, (0.3-)0.4-0.5(-0.7) mm long (vs oblong-lanceolate to narrowly oblong, 1.2-1.4 mm long); petals 0.8-1.2 mm long (vs 1.5-1.7 mm long), much longer than sepals (vs only moderately longer). Candollea 73, 2018 Homalium sect. Odontolobus (Salicaceae) in Madagascar - 39 1 cm 1 cm 1 cm Tree to 10 m tall, 50 cm dbh; young twigs glabrous. Leaves: petiole 5-11 mm long, glabrous; blade ovate (elliptical), (4.2-) 6.2-10.2 �� (2.1-) 3.1-5.2 cm; base rounded; margins irregularly crenate-serrulate, sometimes with only 1 tooth per side, to slightly wavy or subentire; apex acute to acuminate. Inflorescences racemose, (2.2-) 4-6.5 cm long, canescent to moderately pubescent; flowers often in clusters of 2 or 3, pedicellate, with pedicels 0.5-1 mm long, or subsessile; bracts broadly ovate to deltoid or transversely oblong; bracteoles very thick, to subterete at base, densely pubescent, often caducous. Flowers 7-8-merous; sepals ovate to deltoid or narrowly deltoid, (0.3-) 0.4-0.5(-0.7) mm long; petals narrowly oblong, 0.8-1.2 mm long, margins long-ciliate, abaxial surface pubescent, mostly glabrous adaxially, apex rounded; filaments 1.4-1.9 mm long; upper surface of ovary densely pubescent to moderately so in fruit; styles 3-4, 1.3-1.8 mm long. Vernacular name. ��� ���Taindalitra��� (Guittou et al. 160). Distribution, ecology and conservation status. ��� Homalium ovatifolium is known from only two collections and a single location in deciduous forests at low elevation in extreme northern Madagascar in the western edge of the Andrafiamena Andavakoera protected area. Even if the only known location is within the protected area network, the new species is under threat and can be affected by a single event (e.g., fire), so its conservation status should be considered ���Vulnerable��� [VU D2]. Notes. ��� Homalium ovatifolium has clear affinities to H. longistaminum, which likewise has a canescent or mostly densely pubescent inflorescence, fleshy, densely pubescent bracteoles and long filaments and styles. It differs in its larger leaves, (4.2-) 6.2-10.2 �� (2.1-) 3.1-5.2 cm (vs (3.2-)3.5-7.5 �� 1.6- 3.3(-4.8) cm), which are more usually ovate with a rounded base, often long-acute to acuminate apex and few-toothed, shallowly wavy or subentire margins, and its narrow, less pubescent petals. Interestingly, the two taxa are found at the opposite extremes of latitude, H. ovatifolium being confined to the extreme north of Madagascar and H. longistaminum to the southeast. It also is likely to be closely related to H. masoalense, which is from northeastern Madagascar and shares the unusual character of often ovate, acuminate leaves, as well as long filaments and styles. Homalium masoalense has often lanceolate, short-petiolate leaves and larger flowers (petals 1.5-1.7 mm long) with the sepals more similar to the petals in length (1.2- 1.4 mm long) and narrowly oblong to oblong-lanceolate. Paratypus. ��� MADAGASCAR. Prov. Antsiranana: Ambilobe, Tanambao Marivorahona, Betsimiranja, Andohan���Antsohy, 4 km au NE de Betsimiranja, 13��02���32������S 49��09���24������E, 76 m, 2.VII.2005, fl. & fr., Guittou et al. 160 (MO, P, TAN)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 38-41, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136

9a. Homalium planiflorum subsp. planiflorum Leaves narrowly elliptical to elliptical (oblanceolate, aberrantly obovate), (2.8-)3.8-10.3 �� (1.1-) 1.3-4 cm. Inflorescences almost always racemose (rarely a racemiform or few-branched panicle), usually borne along length of twigs. Flowers whitish to pale green, pale yellow or cream-colored, 7-8-merous. Vernacular names. ��� ���Fotsakara��� (Service Forestier 15282); ���Fotsakara (petits feuilles)��� (Service Forestier 2719); ���Haraka��� (Louvel 68); ���Hazombalovary��� (Louvel 146); ���Hazombarorana��� (Cours 2965); ���Hazombato��� (Ludovic & Rakotoarivony 213, Rabevohitra et al. 3876, R��serves Naturelles s.n., Service Forestier 1577, 1789, 5693, 5858, 5866, 16862, 17810); ���Hazombato mena (lahy)��� (R��serves Naturelles [Dumazer] s.n.); ���Hazomboangy��� (Service Forestier 11070); ���Hazondrano��� (Service Forestier 2861); ���Marimbody��� (Service Forestier 10155); ���Maroankoditra��� (Service Forestier 8916); ���Masonambatsy��� (Service Forestier 14449, 17999); ���Menavony ��� (R��serves Naturelles 8807); ���Moranga ala��� (R��serves Naturelles 3344); ���Rodrano��� (Service Forestier 14404); ���Tsimahamasatsokina��� (Service Forestier 16259); ���Tsofofonjahana��� (Bernard 126); ���Voangiala��� (Service Forestier 15183); ���Zanganitofotsy��� (Service Forestier 8090); ���Zora��� (Rakotovao 4713). Uses. ��� Wood is used for construction (Service Forestier 16259, 17810, 17999). Distribution, ecology and conservation status. ��� Homalium planiflorum subsp. planiflorum occurs most commonly in littoral and coastal eastern forests on sand, though it also occurs in low-elevation humid forests and rarely in mid-elevation forests (to 706 m) or dry forests. It is often described as occasional, but is common enough to be frequently collected. The preliminary assessment of its conservation status is ���Least Concern��� [LC]. Note. ��� Homalium planiflorum subsp. planiflorum has been reported to be visited by bees (Ludovic & Rakotoarivony 213). Selected material examined. ��� MADAGASCAR. Prov. Antsiranana: Ambohitralalana, Masoala PN, 15��18���45������S 50��20���30������E, 75-150 m, 24.XI.1995, fl. & fr., Bernard 126 (G, K, MO, P); Parc Masoala, for��t Ambodipont, 15��44���45������S 50��19���25������E, 0-10 m, 3.X.1996, fl., Bernard 361 (G, MO, P); vall��e inf��rieure de l���Androranga, affluent de la Bemarivo (NE) aux env. d���Antongondriha, 100-250 m, 1-24.XI.1950, fl., Humbert & Capuron 23979 (P); Between Mandena and the Reserve Integral de Marojejy, 14��27���S 49��17���E, 100- 200 m, 30.IX.1988, fl., J. Miller et al. 3311 (MO [2 sheets], P); Sambava, au N d���Antalaha, XI.1912, fr., Perrier de la B��thie 6699 (P); Andravinambo, for��t de Tsihomanaomby, 14��06���S 50��01���E, 225 m, 3.II.2014, fl., Rakotonirina et al. 446 (MO); Sambava, for��t d���Ambodivohitra, 14��24���42������S 49��51���49������E, 148 m, 20.X.2010, fl., Ravelonarivo & Raharivelo 3541 (MO, P); vall��e de l���Ampalohandrano, affluent de l���Anohoranga, aux env. d���Antongondriha, 150 m, 9.XI.1950, fl., Service Forestier 794 (P); Antsambalahy, Voh��mar, 2.XII.1955, fl., Service Forestier 15228 (P); env. S d���Analamanara (pr��s de Tsaratanana), entre Sambava et Antsirabe-Nord, 2-7.XII.1966, fl., Service Forestier 27184 (P); massif du Bezavona, entre la Fanambana et la Manambery, basse vall��e de l���Andilana, 13.XII.1966, fl., Service Forestier 27226 (P). Prov. Fianarantsoa: route Farafangana-Vangaindrano, XII.1963, fl., Bosser 18580 (P); Province de Mananjary, zone c��tiere, III-IV.1909, fl., Geay 7855 (P); Manombo RS, parcelle n�� 2, 23��02���31������S 47��46���24������E, 20 m, 18.VIII.1995, fl., Rakotomalaza et al. 410 (MO, P); Fkt. Marohita, for��t d���Alimamba, 21��28���04������S 48��17���43������E, 13 m, X.2004, fl., Razakamalala et al. 1646 (G, MO, P); Mananjary, Rindrimbolo, 23.XII.1954, fl., Service Forestier 14404 (MO); Analamarina, Fort-Carnot, 15.X.1955, fl., Service Forestier 15282 (P); Evato, village le plus proche Loharano, for��t d���Analavory, 10.XI.1955, fl. & fr., Service Forestier 16259 (P). Prov. Toamasina: Fkt. Ambanizana, between Andranobe & Tampolo, 15��42���12������S 49��57���38������E, 1 m, 27.X.2004, fl. & fr., Antilahimena 2993 (G, K, MO, P); 0-2 km E of Andavakimenarana (location of ferry to cross Pangalanes), 18��52���34������S 49��08���12������E, 1-10 m, 14-16. II.1999, fl., Birkinshaw et al. 559 (MO, P); Ile Ste. Marie, X.1849, Boivin 1847 (P [3 sheets]); Maroantsetra, IX.1923, fr., Louvel 68 (P); Masoala peninsula, Antalavia, coastal path leading S, 15��46���S 50��01���E, 25 m, 17.IV.1987, fl., Nicoll et al. 553 (MO, P); Pointe �� Larr��e, for��t Menagisy, 16��46���39������S 49��41���32������E, XI.2008, fl., Nikolov 1815 (MO); bassin inf��rieur du Mangoro, X.1922, fl., Perrier de la B��thie 18262 (P); entre Bedinta et Andranobe, 15��39���30������S 49��57���30������E, 350-500 m, 24-26.XI.2001, fr., Rabevohitra 4006 (P); Tampolo forestry station, 17��17���S 49��23���E, 10 m, 11.X.1999, fl., A. Randrianasolo et al. 603 (G, K, MO, P); Ambila-Lemaitso, W of Brickaville-Ambila road, 18��49���S 49��08���E, 0-5 m, 11.XI.1999, fl., A. Randrianasolo & Ranivojaona 649 (G, K, MO, P); trail to Andranokoditra from Ambodivonanto, 18��33���39������S 49��14���33������E, 27 m, 28.X.2002, fr., A. Randrianasolo et al. 746 (G, MO, P); Fkt. Ambalavontaka, for��t d���Antaimby, 20��22���21������S 48��33���15������E, 13 m, 27.IX.2004, fl., Razakamalala et al. 1518 (G, MO, P [3 sheets]); RN II [Masoala], cton Ambohitralana [na], 21.I.1953, fr., R��serves Naturelles 4932 (P); [Ile Ste. Marie], s.d., fl., Richard 260 (P); Masoala Peninsula, N of Antalavia, 15��47���S 50��02���E, 200-380 m, 13-16.XI.1989, fl. & fr., Schatz et al. 2780 (MO, P); poste forestier de Mananara-Nord, 4.XII.1953, fl., Service Forestier 8090 (P); Tenina, au S de Rantabe, I.1954, fl. & fr., Service Forestier 8916 (P); Tampina, s.d., fr., Ursch s.n. (P). Prov.Toliara: massif de l���Andohahel[a] (SE), vall��e de Ranohela, 300-1200 m, 18-26.X.1928, fl. & fr., Humbert 6244 (P [2 sheets]); Sainte Luce, for��t de Beboaka, 24��40���25������S 47��12���00������E, 13 m, 25.XI.2011, fl., Miandrimanana 535 (MO); Ivorona, for��t de Mamoareny, 24��49���S 46��56���E, 706 m, 30.XI.2009, Razakamalala et al. 5021 (MO); Fort Dauphin, VI.1889, fl., Scott Elliot 2824 (P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 43-44, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136

6a. Homalium moniliforme subsp. moniliforme Young twigs glabrous (glabrate, minutely papillate only at extreme apex). Leaves: petiole 2-7(-10) mm; blade obovate to elliptical, narrowly elliptical, or oblanceolate, 3-6.8 �� (1.1-)1.3-2.6(-3) cm; margins shallowly wavy to crenateserrate or subentire, often slightly revolute; apex rounded to acute (rounded-cuspidate, slightly acuminate, aberrantly emarginate). Vernacular names. ��� ���Hazombato��� (Razakamalala et al. 55); ���Mafaikoditre��� (Antilahimena et al. 2171); ���Matrambody��� (Razanatsima & C��l��stin 712); ���Menahihy��� (Iambana & Ars��ne 208, Iambana 270); ���Rohitra��� (Decary 5469); ���Tamehaka madinidravina��� (Andrianjafy et al. 189); ���Tendrompony��� (Razanatsima 859). Uses. ��� Wood is used for construction (Andrianjafy et al. 189). Distribution, ecology and conservation status. ��� Homalium moniliforme subsp. moniliforme is native to mid- to low-elevation humid forests in Antsiranana, Toamasina, and rarely Fianarantsoa provinces. It is reported to occur on laterite, humus soil with rocky substrate, and on ferricrete, sometimes near water. Because it is relatively widespread and occurs in four protected areas (i.e., Zahamena, Masoala, Betampona, Makirovana-Tsihomanaomby), the preliminary assessment of its conservation status is ���Least Concern��� [LC]. Notes. ��� Homalium moniliforme subsp. moniliforme has on average larger leaves than subsp. littorale, but its leaves usually do not exceed 6.8 �� 2.6(-3) cm. Collections from lower-elevation sites, e.g., in Betampona, seem more often to have acute or cuspidate leaves while those at higher elevations have more rounded leaf apices and the leaves are more strongly coriaceous; at the highest elevations the margins are generally revolute. Two specimens from northern Madagascar with very large leaves (to 9.7 �� 4.1 cm) (Service Forestier 28807 and Razakamalala et al. 55) are tentatively placed with the large-leaved, usually southern H. densispicatum but may have affinities to H. moniliforme. Selected material examined. ��� MADAGASCAR. Prov. Antsiranana: Ambodivoapaza, for��t de Makirovana-Tsihomanaomby, 14��08���33������S 49��55���06������E, 593 m, 23.V.2009, post-fl., Randrianarivony et al. 154 (MO); Antanandava, Makirovana, 14��10���01������S 49��57���12������E, 685 m, 4.V.2010, fr., Razakamalala et al. 5447 (MO). Prov. Toamasina: Zahamena PN, 17��38���27������S 48��52���32������E, 700-1040 m, 4.X.2001, fr., Andrianjafy et al. 189 (MO, P); Ampokafo, village Ambatoledama, 15��17���S 50��00���E, 15.VIII.2003, fr., Antilahimena 2046 (G, MO); Fkt. Anjiahely, 15��24���19������S 49��30���28������E, 360 m, 28.VI.2004, fl., Antilahimena 2597 (P); Masoala NP, Ambohitsitondroina Mahalevona, 15��25���37������S 49��57���24������E, 1158 m, 10.X.2003, fr., Antilahimena et al. 2171 (P); Fkt. Anjian���i Madirano, 15��54���08������S 49��27���44������E, 829 m, 22.VII.2007, post-fl., Antilahimena et al. 5616 (MO); Betampona RNI, piste principale 2 �� 4 km, 17��55���S 49��13���E, 21.IV.1999, fr., Iambana & Ars��ne 208 (MO, P); Vatomandry, 19��09���16������S 48��34���56������E, 686 m, 22.XI.2004, old fls., A. Randrianasolo et al. 947 (MO); Brickaville, vers le sommet d���Ankerana, 18��25���24������S 48��47���06������E, 896 m, 22.III.2011, fl., Ravelonarivo et al. 3785 (MO); Fkt. Ambatolampy, for��t d���Ankarana, 18��23���51������S 48��47���53������E, 948 m, 25.I.2012, fl., Ravelonarivo et al. 4194 (MO); Sahanionaka, for��t de Vohibe, 19��10���49������S 48��32���27������E, 763 m, 2.VI.2010, post-fl., Razanatsima 859 (BR, G, K, MO, WAG); Ambinanindrano II, for��t de Vohibe, 19��09���19������S 48��35���04������E, 655 m, 5.IV.2009, fl., Razanatsima & C��l��stin 712 (MO [2 sheets]); Fkt. Ambatolampy, for��t d���Ankerana massif de Beanjada (N de la presqu���ile Masoala), 1000 m, I.1954, fr., Service Forestier 8825 (P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 37-38, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136

8. Homalium parkeri Baker in J. Linn. Soc., Bot. 20: 150. 1883. Lectotypus (designated by SLEUMER, 1973: 309): MADAGASCAR. Prov. Antananarivo: Andrangolaoka, s.d., Parker s.n. (K [K000231480] image seen). Syntypus: MADAGASCAR. Prov. Antananarivo: forests of Imerina, s.d., Baron 1295 (K [K000231479] image seen, TAN [TAN000592] image seen). Tree to 20 m tall, 36 cm dbh, or shrub; young twigs glabrous (glabrate). Leaves: petiole 2.5-7(-8) mm long, glabrous (glabrate); blade elliptical to narrowly elliptical (to narrowly obovate, ovate or obovate), (2.8-)4-7.3(-9.3) �� 1.3-3.1 (-3.4) cm; base cuneate (moderately convex, attenuate); margins serrate with rounded tooth apices; apex acute (rounded, aberrantly emarginate). Inflorescences spicate, (1.8-) 3-7(-11.5) cm long, minutely pubescent (glabrate); flowers clustered, sessile; bracts broadly to transversely deltoid; bracteoles deltoid, minute, caducous. Flowers 5-7-merous, pale green to yellowish; sepals deltoid (ovate), 0.2-0.4(-0.5) mm long; petals ovate (oblong-ovate), 0.7-1.5 mm long, glabrous (sparsely pubescent on abaxial surface), apex acute to rounded; filaments 0.3-0.6 mm long; upper surface of ovary shortpubescent, usually densely; styles (2-)3(-4), 0.2-0.5 mm long. Vernacular names. ��� ���Hazombato��� (Anonymous 19, Hong-Wa et al. 392, Lehavana et al. 487, Service Forestier 16807, 19957, 28755); ���Hazombatofotsy��� (Louvel 43); ���Hazombatomainty��� (Service Forestier 16010, 16822); ���Hazomby��� (Service Forestier 595, 1043, 3926, 6006, 10370, 15990); ���Hazompoza��� (Service Forestier 7592); ���Moara��� (R��serves Naturelles 9460); ���Ramaindafa��� [?] (R��serves Naturelles 10472); ���Ranga��� (Service Forestier 15873). Distribution, ecology and conservation status. ��� Homalium parkeri is widespread in humid forests of eastern Madagascar, extending to high-elevation montane moss forest. A preliminary assessment of its conservation status should be ���Least Concern��� [LC]. Selected material examined. ��� MADAGASCAR. Prov. Antananarivo: Ambohitantely RS, 1620 m, 18��11���52.5������S 47��17���03������E, 7.III.2004, fl., Almeda et al. 8645 (MO); Angavokely (Carion), V.1956, fl., Bosser 9575 (P); PK 138 de la route Tananarive-Majunga, 8.VII.1971, post-fl., Cremers 1636 (MO); W Imerina, Andrangolaoka, III.1881, fl. & fr., Hildebrandt 4102 (P [2 sheets]); Firarazana, for��t de Manjato, 18��06���19������S 47��14���43������E, 1447 m, 6.VII.2005, fl., Hong-Wa et al. 392 (MO, P); 7 km E of Anjozorobe, 18��22���S 48��00���E, 1350 m, 2.IV.1988, fl., Lowry & Randrianasolo 4419 (MO, P); Ambatolaona, 1700 m, VII.1914, fl., Perrier de la B��thie 6720 (P); Tsinjoarivo, Ambatotsipihina, 22.XI.1949, fl., Service Forestier 1043 (P); Antsahambavy, Manjakandriana, 14.V.1956, fl., Service Forestier 15873 (P); Ambohidraondriana, Ankazobe, 3.V.1956, fl., Service Forestier 15990 (P); Ambatondradama au N d���Ambohimanga, 30.IV.1957, fl., Service Forestier 18027 (P); Tsiazompaniry, for��t d���Ambohimangakely, 6.IV.1961, fl., Service Forestier 19894 (P); Ibity massif W, Vohipisaka, 20��09���42������S 46��58���54������E, 1423 m, 29.II.2004, fl., Skema et al. 22 (MO); SW Andranofeno Sud village, 18��04���59������S 47��10���26������E, 1407 m, 5.IV.2004, fl., Skema et al. 54 (MO). Prov. Antsiranana: Manongarivo RS, 14��05���S 48��23���E, 1470-1570 m, 14-15.IV.1992, fl., Malcomber et al. 1495 (MO, P); SW edge of Anjanaharibe-Sud Reserve, 14��48���15������S 49��26���45������E, 1000-1100 m, 6.VIII.1997, fr., McPherson 17257 (MO); Anjanaharibe-Sud, 14��47���45������S 49��27���54������E, 1161 m, 22.V.1995, fl., Ravelonarivo & Rabesonina 811 (G, K, MO, P); Andranomilolo, 13 km �� l���W du village d���Andranopositra, 14��19���16������S 49��17���56������E, 1462 m, 10.XI.2006, fl., Ravelonarivo et al.2015 (MO, P); Befingotra, Andranotsarabe, 14��45���11������S 49��28���49������E, 762 m, 21.IV.1997, fl., Razafindramora et al. 26 (MO). Prov. Fianarantsoa: Andringitra, 26.II.1938, fl., Herb. Jard. Bot. 3141 (P [2 sheets]); Ranomafana, W side of Namorona riv., 21��16���S 47��21���E, 1080 m, 6.III.1992, fl., Malcomber et al. 1317 (MO, P); Ambalamanakana, 30 km S d���Ambositra sur RN 7, 20��44���06������S 47��11���37������E, 1650 m, 26.III.1996, fl, Rakotomalaza et al. 678 (MO); Ranomafana, parcelle n�� 3, 21��15���30������S 47��25���E, 900-1100 m, 14.VI.1994, fr., J. Randrianasolo et al. 65 (MO). Prov. Mahajanga: Ambalotsangana, for��t de Makira, 15��31���S 49��05������E, 1169 m, s.d., fl., Lehavana et al.487 (MO, P). Prov.Toamasina: Fkt. Ampita[m]be, Ambatovy, Ampanatovana forest, 18��51���17������S 48��19���07������E, 1085 m, 9.VI.2008, fl., Antilahimena et al. 6277 (G, MO); Andapanomby, pr��s de la riv. Ampandisanana, Makira NW, 15��21���17������S 49��06���25������E, 25.IV.2007, fr., Bernard & Birkinshaw 456 (G, MO, P); Fkt. Ambohibato, for��t de Rianan���i Galy, 18��39���32������S 47��57���56������E, 1291 m, 10.IV.2005, fl., Raharijaona et al. 72 (MO); Mantadia PN, Andranomanamponga, 18��50���23������S 48��26���31������E, 1110 m, 19.III.2013, fl., Ramahenina et al. 224 (G, MO, P); Zahamena PN, campement Analalentitra, 17��32���47������S 48��44���21������E, 1235-1400 m, 29.IX.2001, fr., Randrianjanaka et al. 629 (MO); Fkt. Ampitambe, Ambatovy, for��t d���Analamay, 18��49���07������S 48��19���26������E, 1138 m, 5.IX.2011, fl., Ravelonarivo 3950 (G, MO); Ambatondrazaka, Manaka Est, 27.VII.1960, fl., R��serves Naturelles 10472 (P); Bekiritsika [?], Lakato, 8.X.1953, fl., Service Forestier 7592 (P [2 sheets]); Perinet, km 7 Antaniditra, 5.V.1954, fl., Service Forestier 10370 (P); W du massif de l���Ampahana, �� l���E de Fierenana, 950-1300 m, 10-16.III.1969, fl., Service Forestier 28755 (MO, P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 41-42, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328."]}

4. Homalium lucidum Scott-Elliot in J. Linn. Soc., Bot. 29: 24. 1891. Lectotypus (designated by SLEUMER, 1973: 307): MADAGASCAR. Prov. Toliara: Fort Dauphin, V.1889, fl., Scott Elliot 2661 (K [K000231842] image seen; isolecto-: P [P04704037]!). Tree to 30 m tall, 60 cm dbh; bark dark gray-brown, sometimes slightly fissured or peeling, with a bright yellow slash; young twigs glabrous. Leaves: petiole 3-10(-19) mm long, glabrous (glaucous, peeling); blade elliptical to lanceolate (narrowly lanceolate, aberrantly broadly elliptical), 3.5-11.5 �� 1.5-3.8(-4.3) cm; base convex; margins serrulate to crenateserrulate (shallowly crenate, subentire); apex acute to acuminate (emarginate, rounded-cuspidate). Inflorescences racemose, (2.5-)4-10(-12) cm long, glabrous; flowers proximally often in clusters of 2 or 3, sessile (pedicellate with pedicels to 2 mm long); bracts ovate to broadly ovate; bracteoles ovate, minute, caducous. Flowers 6-7(-8)-merous, pale green to white, cream or yellowish, or brown with age (once described as red); sepals deltoid, 0.3-0.4(-0.5) mm long; petals lanceolate (narrowly ovate) to oblong-lanceolate or narrowly elliptical (elliptical), 1.5-2.5(-3.1) mm long, ciliolate, otherwise glabrous (or adaxial surface sparsely short-pubescent), apex acute; filaments 0.3-0.7 mm long; upper surface of ovary sparsely to moderately short-pubescent; styles 2-3, 0.3-0.5 mm long. Vernacular names. ��� ���Fotsakara (g[rands] f[euilles])��� (Service Forestier 2718, 2737); ���Hazofotsy��� (Antilahimena et al. 4835); ���Hazokoaky��� (Service Forestier 13942); ���Mafaikoditra��� (Antilahimena et al. 1139); ���Maroampotolia��� (R��serves Naturelles 3668); ���Menavahatra��� (Antilahimena et al. 7110); ���Ramisaona��� (Service Forestier 14569, 14608). Distribution, ecology and conservation status. ��� Homalium lucidum is widely distributed in humid regions of Madagascar and occurs from near sea level to over 1400 m elevation. The species is widely distributed from Antananarivo province in the center to Toliara province in the south and is relatively common with many localities within the protected area network (i.e., Andohahela, Anjozorobe, Betampona, Makira, Nosy Mangab�� and Tsitongambarika). Homalium lucidum is therefore assigned a preliminary assessment of conservation status of ���Least Concern��� [LC]. Note. ��� Homalium lucidum was described from only one collection. As for other species published by Scott Elliot, the duplicate at K, where he worked, has been informally considered to be the ��� holotype.��� However, as other duplicates of his own collections were certainly available to him, and the publication did not specify that specimens cited were held at K, the two known duplicates of Scott Elliot 2661 should be regarded as syntypes under Art. 9.5 of the ICN (MCNEILL et al., 2012). SLEUMER���S (1973) subsequent statement that the K and P duplicates were the ��� holotype ��� and ���isotype���, respectively, was an effective selection of the former as lectotype, with the use of ��� holotype ��� being a correctable error under Art. 9.9. Selected material examined. ��� MADAGASCAR. Prov. Antananarivo: 3.0 km NW of Ambohitsaratelo-Bebao (NW of Tsiroanomandidy), 1100-1200 m, 16.I.1985, fl., Dorr et al. 3587 (MO, P); Bongolava, 1300 m, VII.1974, fl. & fr., Morat 4639 (MO [2 sheets], P [4 sheets]); Fkt. Ambaravaranala, 18��25���S 45��40���E, 1190-1200 m, 25.VI.1997, fr., Rakotomalaza et al. 1344 (G, K, MO); for��t de Besofina �� 8 km au S de Betatao, 18��15���10������S 47��53���27������E, 1386 m, 9.III.2000, fl., Randrianaivo et al. 517 (MO, P); 7 km E of Anjozorobe, 18��22���S 48��00���E, 1300 m, 11.V.1987, fl., Schatz et al. 1380 (MO, P). Prov. Fianarantsoa: Fkt. Iabomaro, 23��11���07������S 47��42���32������E, 24 m, 19.VI.2004, fr., A. Randrianasolo et al. 878 (P); Vangaindrano, 20.I.1950, fl., Service Forestier 2737 (P); Manakara, Vakoary, 11.V.1955, fr., Service Forestier 14569 (P); Manakara, for��t Belambo, 11.V.1955, Service Forestier 14608 (P). Prov. Mahajanga: 9.6 km NW of Ambohistaratelo-Bebao, 1200 m, 14.I.1985, fl., Dorr et al. 3535 (MO, P); 5 km �� l���E de Bandabe, 15��31���08������S 49��06���00������E, 3696���, 25.IV.2007, fr., Ravelonarivo et al. 2253 (MO, P). Prov. Toamasina: 5 km du village d���Anony, vers Ankarefobe, 17��12���26������S 48��33���13������E, 1164 m, 22.IV.2005, fl., Andriamijaharivo et al. 577 (MO); Fkt. Andranofotsy, 15��25���14������S 49��48���51������E, 140 m, 14.VII.2002, fr., Antilahimena et al. 1139 (MO, P); Fkt. Ampitambe, Ambatovy, Andranovery, 18��52���43������S 48��17���38������ W, 1010 m, 10.VI.2009, fl., Antilahimena et al. 7110 (G, K, MO, P); Andapanomby, pr��s de la rivi��re Ampandisanana, Makira NW, 15��21���48������S 49��07���06������E, 25.IV.2007, fr., Bernard & Birkinshaw 462 (G, MO, P); Nosy Mangab��, 15��30���S 49��45���E, 17.IV.1988, fl. & fr., Leeuwenberg 13933 (MO, P); confluence de l���Onive et du Mangoro, 600 m, IX.1926 [?], fl., Perrier de la B��thie 17012 (P); entre Moango et Antanamarina, 17��33���57������S 48��53���58������E, 740 m, 27.X.2000, fr., Randrianjanaka et al. 586 (MO, P); entre Manakambahiny Est et Nonokambo, 17��45���34������S 48��42���04������E, 7.III.2001, fl. & fr., Randrianjanaka et al. 606 (MO, P); RN I [Betampona], 20.III.1952, fl., R��serves Naturelles 3668 (MO, P [2 sheets]); Nosy Mangabe, 15��30���S 49��46���E, 0-330 m, 13-23. IV.1988, fl., Schatz & Gentry 2055 (MO, P); Bassin de la Manonga (effluent [rive gauche] de la Rantabe), env. de Sahajinja, 700 m, 5.III.1954, fl., Service Forestier 9107 (P). Prov. Toliara: Fkt. Antsotso, Ivohibe forest, 24��34���10������S 47��12���37������E, 41 m, 24.V.2006, fr., Antilahimena et al. 4835 (P); Andohahela [NP], parcelle 1, Eminiminy, 24��40���S 46��48���E, 200-700 m, 4-24.V.1993, fr., Randriamampionona 309 (MO, P); ibid. loco, 4-24.V.1993, fl., Randriamampionona 376 (MO, P); Imonty, 3.VIII.1955, fr., Service Forestier 13942 (P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on page 34, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","MCNEILL, J., F. R. BARRIE, W. R. BUCK, V. DEMOULIN, W. GREUTER, D. L. HAWKSWORTH, P. S. HERENDEEN, S. KNAPP, K. MARHOLD, J. PRADO, W. F. PRUD'HOMME VAN REINE, G. F. SMITH, J. H. WIERSEMA & N. J. TURLAND (2012). International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Veg. 154."]}

1. Homalium brachystylis (Tul.) Baill. in Bull. Mens. Soc. Linn. Paris 1: 573. 1886 [as brachystylum]. �� Blackwellia brachystylis Tul. in Ann. Sci. Nat., Bot. ser. 4, 8: 59-60. 1857. Lectotypus (designated by SLEUMER, 1973: 246): MADAGASCAR. Prov. Antsiranana: For��ts �� l���entr��e de la Baie de Voh��mar, 1837, fl. & fr., Richard 63 (P [P04679017]!; isolecto-: G [G00018411] image seen, L [L0010889, L0010890] images seen). Syntypi: MADAGASCAR. Prov. Antsiranana: Voh��mar, 1846, fl. & fr., Richard 110 [= Boivin 2568] (P [P 04734119]!). Sine loco: ��� Sud de Madagascar [?]���, s.d., fr., Richard 561 (P [P 04734120]!); Herb. Galldich [cl. Jaubert] 40 (not found). = Homalium humbertii H. Perrier in M��m. Mus. Natl. Hist. Nat. 13: 290. 1940 [as humberti]. Lectotypus (first step designated by SLEUMER, 1973: 247; second step designated here): MADAGASCAR. Prov. Toliara: bassin de r��ception de la Mananara, affluent du Mandrare, pentes occidentales des montagnes entre l���Andohahela et l���Elakelaka, au Vatazo (S d���Imonty), 900-950 m, II.1934, fl., Humbert 14077 (P [P04734112]!; isolecto-: B [B100153995] image seen, G [G00018412] image seen, P [P04734111, P04734113, P04734114]!, PRE [PRE0297345 -0] image seen, TAN [TAN000242] image seen). Syntypi: MADAGASCAR. Prov. Toliara: vall��e moyenne du Mandrare pr��s d���Anadabolava, Mont Vohitrotsy, 700-850 m, XII.1933, fl., Humbert 12655 (G [G 00018413, G 00018414, G 00018415] images seen, P [P 04734115, P 04734116, P 04734117, P 04734118]!, S [S 10-10143] image seen). Tree to 12 m tall, 15 cm dbh; bark sometimes described as platanoid; young twigs glabrous or sparsely pubescent, later becoming glabrous. Leaves: petiole (1-)1.5-2.5(-4) mm long, glabrous (sparsely pubescent); blade narrowly elliptical to narrowly oblong-elliptical or elliptical (somewhat obovate), 2.5-7.7 �� 0.5-2(-3.4) cm; base convex to cuneate; margins shallowly serrulate to crenulate, usually with few teeth, or subentire; apex acute to rounded (slightly acuminate, obtuse). Inflorescences racemose, 2-7.7 cm long, glabrous or with small patches of pubescence near the point of insertion of flowers (rarely sparsely and minutely pubescent on much of the inflorescence or with small patches of yellowish scaly indument around flowers); flowers solitary, sessile; bracts broadly to transversely ovate, thick-textured; bracteoles transversely to broadly ovate, broadly elliptical, or irregularly transversely oblong, thick-textured. Flowers (5-)6-7(-8)-merous, white to cream-colored, yellow or yellowish green (browning after anthesis); sepals narrowly deltoid to oblong-lanceolate or ligulate, 0.8-1.8 mm long; petals narrowly oblong to narrowly oblong-elliptical or oblong-lanceolate (to somewhat oblanceolate, in north), 1.1-1.8(-2.3) mm long, margins ciliate, otherwise glabrous (adaxial surface rarely pubescent), apex acute (to rounded); filaments 0.3-0.6 mm long; upper surface of ovary moderately villous or pubescent; styles 2(-3), 0.3-0.6 mm long. Vernacular names. ��� ���Ampolilahy��� (Service Forestier 14092); ���Mantsapoty��� (Service Forestier 9872, 10080); ���Somontsohihy��� (Be et al. 42); ���Tsohontsobe��� (Humbert 19161). Distribution, ecology and conservation status. ��� Homalium brachystylis is widespread in dry seasonal low to moderateelevation forests and is reported from limestone, sand and silica substrates and in acid soil. It is noteworthy that the single specimen (Andriamihajarivo et al. 293) from the high-elevation quartzite massif [Itremo] near the town of Ambatofinandrahana has much larger leaves than other specimens in the southern part of this species��� range. This massif is an unusual locality that is home to a relatively large number of endemic species, e.g., Andropogon itremoensis Voronts. (VORONTSOVA et al., 2013), Buxus itremoensis G.E. Schatz & Lowry (SCHATZ & LOWRY, 2002), Streptocarpus lanatus MacMaster (MACMASTER et al., 2005), Xerochlamys diospyroidea (Baill.) F. G��rard and X. itremoensis Hong-Wa et al. (HONG-WA, 1999). Thus this specimen may be suspected of representing a genetically distinct population that merits further investigation. The species is widely distributed from the Antsiranana province in the north to the Toliara province in the south and is relatively common with many localities within the protected area network. Homalium brachystylis is therefore assigned a preliminary assessment of conservation status of ���Least Concern��� [LC]. Notes. ��� In most of its range, including much of the northern province of Antsiranana, H. brachystylis usually has narrowly elliptical or oblong-elliptical leaves, seldom to elliptical or aberrantly obovate, which are 0.5-2 cm broad (after the exclusion of one large-leaved specimen from Ambatofinandrahana), and has petals that are usually less than 1.8 mm long. A few older collections made in the Voh��mar region of northeastern Madagascar have elliptical leaves at maturity, which are 1.9-3.4 cm broad, and unusually large flowers, with petals to 2.3 mm long. They also have relatively narrow bracteoles, short pubescence on the calyx cup, and small patches of indument on the inflorescence rachis above flowers, all features that likewise occur in some narrow-leaved material, though the latter is rare. The protologue of Blackwellia brachystylis indicated that the original material came from the Voh��mar region; not all available syntypes are labeled as such (one was later, in another hand and probably erroneously, labeled as ���Sud de Madagascar���), but their morphology is consistent with the claim. All original material of Homalium humbertii, which SLEUMER (1973) synonymized with H. brachystylis, is from southern Madagascar and has narrow leaves. The possibility therefore had to be considered that H. brachystylis and H. humbertii should be distinguished from one another at some rank. However, no recent collections from around Voh��mar reflect the extreme morphology of the older broadleaved specimens; a couple of recent collections have elliptical leaves and relatively narrow bracteoles, but the flowers are not large, nor is the calyx cup pubescent. Therefore, it appears that there is no clear evidence of a persistent, clearly distinguished taxon endemic to the Voh��mar region, and the unusual older specimens represent local genetic variation that does not warrant formal recognition. The epithet of this species, once transferred to Homalium, is usually given incorrectly as brachystylum. Epithets such as brachystylis should properly be treated as nouns in apposition, which are not declined as adjectives according to the gender of the genus name (STEARN, 1992: 96-97). Since many authors do treat them as adjectives, the decision to do so is not correctable. However, since the author of the basionym (Blackwellia brachystylis Tul.) did choose to use a form correct for a noun in apposition, that choice should be preserved. SLEUMER (1973) designated Humbert 14077 in the Paris herbarium as the lectotype of Homalium humbertii; there are in fact four sheets of this collection at P, and he made no selection among them. The International Code of Nomenclature [ICN] (MCNEILL et al., 2012: Art. 9.17) recommends that a second-stage lectotypification be published to designate a single sheet as the lectotype. The selected sheet is one of the better duplicates, in terms of the size and completeness of the material, and it bears a complete original label. Selected material examined. ��� MADAGASCAR. Prov. Antsiranana: env. de RS d���Analamera, 12��40���40������S 49��32���43������E, 127 m, 17.I.1995, fl., Andrianantoanina et al. 738 (BR, G, K, MO, P, WAG); massif de l���Ankarana, 12��57���09������S 49��09���01������E, 120 m, 26.I.2003, fl. & fr., Bardot-Vaucoulon et al. 1325 (MO); Sahafary, 12��34���30������S 49��27���32������E, 200 m, 25.VII.2004, fl., Be et al. 42 (MO, P); Analamera, 50-400 m, I.1938, buds, Humbert 19161 (P [3 sheets]); for��t d���Antsahabe, 10 km �� l���W du village d���Ankarafa, 13��12���38������S 49��33���29������E, 471 m, 29.X.2005, Rakotondrafara et al. 335 (MO); Sahafary be, 3 km �� l���E de Saharenana, 12��35���09������S 49��27���34������E, 235 m, 11.IX.2004, fr., Rakotondrajaona et al. 336 (MO, P); Ankarongana, for��t d���Andranomadiro, 12��36���18������S 49��26���34������E, 258 m, 7.XI.2006, fl., Ranaivojaona et al. 1519 (MO); Vohemar, for��t de Binara, 13��14���09������S 49��37���22������E, 300 m, 31.III.2004, fl., Ranirison 551 (MO); Fkt. Saharenana, for��t d���Andranomadiro, 12��36���18������S 49��26���35������E, 300 m, 14.II.2005, fl., Schatz et al. 4264 (MO); Di��go-Suarez, Ankara JB8, 27.VII.1954, buds, Service Forestier 10426 (P); plateau de l���Ankarana pr��s d���Ambondromifehy, 4-6.X.1954, Service Forestier 11257 (MO); Di��go-Suarez, Ankara, 24.III.1955, fl., Service Forestier 13208 (P); Ankarana �� l���W d���Ambondromifehy, 24.IV.1963, fl., Service Forestier 22695 (MO, P [3 sheets]); rebord S du plateau de Mahory, au SW de Marotaolana (Anivorano-Nord), 3.IV.1964, fl., Service Forestier 23376 (P); massif de l���Ankitakona, au S de la Baie d���Ambararata, 25-265 m, 25.IV.1966, fl., Service Forestier 24673 (P); massif du Bezavona entre la Fanambana et la Manambery, pentes inf��rieures de la rive droite de l���Andilana, 20.III.1967, fl., Service Forestier 27537 (MO, P). Prov. Fianarantsoa: Ambatofinandrahana, 20��30���03������S 46��50���51������E, 1451 m, 20.III.2004, fl., Andriamihajarivo et al. 293 (MO); haute vall��e de la Menarahaka a l���E d���Ihosy, 700-800 m, 28.I.1955 & 10.IV.1955, fl., Humbert 28538 (MO, P); between Ankazobetroka and Ambararata, ca. 25 km NE of Ihosy, 22��15���S 46��15���E, 750 m, 11.IV.1971, fl., Mabberley 910 (P); Ihosy, 800 m, VI.1953, fl. & fr., Perrier de la B��thie 19272 (P [2 sheets]); for��t de Kitranga, �� 12 km au N d���Ihosy, I.1955, fl., Service Forestier 11611 (P [2 sheets]); Zasafotsy, Ihosy, 21.VII.1954, buds, Service Forestier 14482 (P); Kitranga, entre Ambararata et Ivandrika, 22.II.1970, fl., Service Forestier 29063 (P). Prov. Mahajanga: Bemanevika, for��t d���Analafaly, 17��13���50������S 46��59���58������E, 640 m, 9.V.2005, fr., Andrianjafy et al. 1038 (MO, P). Prov. Toliara: Andohahela RNI parcelle 2, path E to Ambohibory massif, 24��56���23������S 46��38���40������E, 120 m, 9-12.V.1997, fr., Birkinshaw et al. 427 (MO); W du Tsimelahy, Andohahela RNI, 24��56���S 46��35���E, 345 m, 22.III.1995, fl., Eboroke 985 (G, K, MO); for��t de Zombitsy (Sakaraha), 600-850 m, 26-29. III.1955, fl. & fr., Humbert et al. 29567 (P); Andohahela, Parcelle 3, 25��04���S 46��41���E, 100-300 m, 8-10.IV.1993, fr., Randriamampionona 302 (MO, P); RN XI [Andohahela], 18.I.1951, fl., R��serves Naturelles 2756 (MO, P [2 sheets]); Tulear, for��t d���Analavelona, 7.IV.1954, fr., Service Forestier 9872 (MO, P); Mahaboboha, for��t d���Ihera, 9.IV.1954, fr., Service Forestier 10080 (P); for��t de Zombitsy, 700 m, III.1955, fl., Service Forestier 11912 (P); Andranolava, for��t Vohibasia, 27.V.1955, fl., Service Forestier 14092 (P); For��t de Zombitsy, 700-800 m, 20.VI.1958, fl. & fr., Service Forestier 18593 (P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 30-31, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136, {"references":["SLEUMER, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","VORONTSOVA, M. S., G. RATOVONIRINA & T. RANDIAMBOAVONJY (2013). Revision of Andropogon and Diectomis (Poaceae: Sacchareae) in Madagascar and the new Andropogon itremoensis from the Itremo Massif. Kew Bull. 68: 1 - 15.","MACMASTER, G., M. MOLLER, M. HUGHES, T. J. EDWARDS & D. U. BELLSTEDT (2005). A new species of Streptocarpus (Gesneriaceae) endemic to Madagascar. Adansonia ser. 3, 27: 131 - 136.","STEARN, W. T. (1992). Botanical Latin. Ed. 4. David & Charles, Newton Abbot.","MCNEILL, J., F. R. BARRIE, W. R. BUCK, V. DEMOULIN, W. GREUTER, D. L. HAWKSWORTH, P. S. HERENDEEN, S. KNAPP, K. MARHOLD, J. PRADO, W. F. PRUD'HOMME VAN REINE, G. F. SMITH, J. H. WIERSEMA & N. J. TURLAND (2012). International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Veg. 154."]}

3. Homalium longistaminum H. Perrier in M��m. Mus. Natl. Hist. Nat. 13: 287. 1940. Lectotypus (designated here): MADAGASCAR. Prov. Toliara: vall��e de la Manambolo, rive droite (bassin du Mandrare) aux env. d���Isomono (confluent de la Sakamalio), 400-900 m, XII.1933, buds & fl., Humbert 12914 (P [P04679979]!; isolecto-: L [L0010953] image seen, P [P04679980]!). Tree to 12 m tall, 30 cm dbh; young twigs glabrous or minutely pubescent. Leaves: petiole 4-11 mm long, glabrous (sparsely, minutely pubescent); blade irregularly elliptical to broadly elliptical or ovate (lanceolate), (3.2-)3.5-7.5 �� 1.6-3.3(- 4.8) cm; base convex to rounded (attenuate at extreme base); margins irregularly crenate apically to crenate-serrate for most of their length; apex acute to obtuse or rounded (short-acuminate). Inflorescences racemose, (2.2-) 4-6.5 cm long, canescent or in part moderately pubescent; flowers often borne in clusters of 2 or 3, on pedicels 0.3-1.4 mm long; bracts transversely to broadly deltoid; bracteoles very thick, to subterete at base, densely pubescent (absent or rapidly caducous). Flowers 6- or 7-merous, pale or yellowish green; sepals deltoid to ovate, (0.3-)0.4-0.5(-0.6) mm long; petals oblong to somewhat obovate, 0.8-1.2 mm long, margins short-ciliate and both surfaces pubescent, densely so abaxially and usually at base adaxially, apex rounded or obtuse; filaments 1.1-1.7 mm long; upper surface of ovary canescent; styles (3-)4(-5), 1.1-1.7 mm long. Vernacular names. ��� ���Tsimalagnilamba��� (Eboroke 980); ���Tsimalamba��� (Humbert 12914). Distribution, ecology and conservation status. ��� Homalium longistaminum is confined to a limited area in southeastern Madagascar, where it occurs mostly in low-elevation humid forests. The fact that very few collections are available, although both Andohahela and Ranopiso are relatively well-collected, suggests that the species is rare in its native range. Four collections are known, pertaining to only three distinct localities, and giving an AOO of no more than 12 km 2. Two of these three are in the area of Andohahela RNI, hence protected. However, the species should be considered ���Vulnerable��� [VU B1ab(iii)+B2ab(iii)] based on its small AOO (Note. ��� In publishing this species, PERRIER DE LA B��THIE (1940) did not note that there were two sheets of the type collection at P; Sleumer incorrectly marked both as ��� holotype ���. The sheet numbered P04679980 has a determination label by Perrier, while the sheet numbered P04679979 has notations on the label in the same handwriting that read ��� Homalium (Blackwellia) longistaminum H. Perr. ���, with the addition of ���n. sp.���, and ��� Type ��� (double underlined). The latter sheet is in better condition, having a fragment packet that contains partially open flowers; it would certainly be a more suitable type and Perrier de la B��thie probably intended it as such. However, since both sheets were used by Perrier de la B��thie and no distinction between them was made in the protologue, they are regarded as syntypes and it is therefore necessary to designate a lectotype. Additional material examined. ��� MADAGASCAR. Prov. Toliara: RNI d���Andohahela, parcelle n�� 2, 24��49���49������S 46��32���15������E, 30-50 m, 16.III.1994, Andrianarisata et al. 101 (MO [2 sheets]); Ranopiso, Evasia, 24��46���S 46��41���E, 350 m, s.d., post-fl., Eboroke 980 (G, MO); Andohahela, parcelle n�� 2, Ihazofotsy, 24��50���S 46��33���E, 50-200 m, 2-5.III.1993, fr., Randriamampionona 152 (MO, P)., Published as part of Applequist, Wendy L., 2018, A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar, pp. 27-48 in Candollea 73 (1) on pages 33-34, DOI: 10.15553/c2018v731a4, http://zenodo.org/record/5722136, {"references":["PERRIER DE LA BATHIE, H. (1940). Revision des Flacourtiacees de Madagascar et des Comores. Mem. Mus. Natl. Hist. Nat. 13: 261 - 301."]}

4. Homalium sect. Eumyriantheia Warb. in Engler & Prantl, Nat. Pflanzenfam. III(6a): 36. 1893. �� Myriantheia Thouars, Gen. Nov. Madagasc.: 21. 1806. �� Homalium [unranked as ��] Myriantheia (Thouars) Baill., Bull. Mens. Soc. Linn. Paris 1: 576. 1886 [as Myrianthea]. �� Homalium sect. Myriantheia (Thouars) Kuntze in Post & Kuntze, Lex. Gen. Phan., Prop.: 285. 1903 [as Myriantheja]. Typus: Homalium laxiflorum (Tul.) Baill. (�� Myriantheia laxiflora Tul.) (designated by Sleumer, 1973: 315). Stipules axillary, free. Inflorescences racemose or paniculate; bracts ovate to deltoid, sometimes broadly, of small to moderate size; bracteoles usually 2 per flower, often borne on pedicel (1 per flower, or absent or rapidly caducous). Flowers pedicellate with pedicels articulated (in H. trigynum sessile or short-pedicellate); perianth 4-5(-6)-merous. Sepals oblong to elliptical, ovate or obovate, sometimes narrowly (oblanceolate), very little accrescent; calyx tube funnelform, in fruit becoming convex to cup-shaped or nearly hemispherical (in H. trigynum short-cylindrical or turbinate becoming broadly ellipsoid in fruit); sepal glands usually large, oblong to elliptical or irregularly trapezoid (semicircular), sometimes elevated. Petals obovate to oblanceolate (in H. trigynum to spatulate), similar to sepals in length or up to twice as long, spreading or ascending at anthesis, very little accrescent; sepals and petals not ciliate (appearing ciliolate in conjunction with overall surface indument). Stamens 3 (aberrantly 4, in H. boinense H. Perrier 5) per petal, inserted between glands; anthers dorsifixed, broadly oblong-elliptical with oblong-elliptical locules and a large connective, the slits of dehiscence nearly parallel. Upper surface of ovary conical, usually shallowly, to nearly flat in flower, in fruit sometimes becoming convex; styles 3-5, free to base or near base. Locule of fruit subglobose (in H. trigynum ellipsoid and partly filled with spongy tissue), glabrous (sparsely pubescent); seeds sometimes 1 (to 2) per fruit, large and subglobose to ellipsoid (possibly at least 4 in H. nobile Baill.). Distribution. ��� Madagascar. Species included. ��� Homalium boinense, H. brevipedunculatum Scott-Elliot, H. capuronii Sleumer, H. dorrii Appleq., H. graciliflorum Sleumer, H. laxiflorum, H. maringitra H. Perrier, H. nobile, H. oppositifolium (Tul.) Baill., H. pseudoboinense Appleq., H. pulchrum Sleumer, H. randrianasoloi Appleq., H. ranomafanicum Appleq., H. schatzii Appleq., H. trigynum. Notes. ��� Sleumer (1973) wrongly corrected the name of Homalium sect. Eumyriantheia to ���sect. Myriantheia ��� and presumed it to have been based upon Thouars��� Myriantheia. The two were not nomenclatural synonyms, but because he explicitly treated them as such, he effectively designated the same type for both simultaneously. Homalium trigynum has some unusual features that are similar to features common in sect. Blackwellia, which also occurs in Madagascar (see discussion above); molecular data regarding its relationships would be of interest. A recent revision describes five new species in the section (Applequist, 2016)., Published as part of Applequist, Wendy L., 2016, A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae), pp. 231-256 in Candollea 71 (2) on pages 252-253, DOI: 10.15553/c2016v712a9, http://zenodo.org/record/5721656, {"references":["Sleumer, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328.","Applequist, W. L. (2016). A revision of the Malagasy species of Homalium sect. Eumyriantheia Warb. (Salicaceae). Candollea 71: 33 - 60. DOI: http: // dx. doi. org / 10.15553 / c 2016 v 711 a 7"]}

8. Homalium sect. Polyanthera Warb. in Engler & Prantl, Nat. Pflanzenfam. III(6a): 35, 36. 1893. Lectotypus (designated here): Homalium deplanchei (Vieill.) Warb. (�� Blackwellia deplanchei Vieill.) Stipules axillary, free. Inflorescences racemose (to spicate) or paniculate (usually short-branched); bracts usually small or narrow, bracteoles small, at least the bracts usually persistent past anthesis. Flowers pedicellate with pedicels articulated to subsessile or sessile; perianth 5-12(-13)-merous. Sepals ligulate to linear or narrowly elliptical (ovate to lanceolate, narrowly oblong; seldom much reduced), usually modestly accrescent; calyx tube turbinate to narrowly funnelform or tubular, at least in fruit (rarely funnelform in flower), usually prominently ridged; sepal glands usually small and round, rarely broad. Petals similar in size and shape to sepals or narrowly elliptical to spatulate or oblanceolate (narrowly obovate, oblong-ovate, narrowly deltoid) and markedly longer than sepals, ascending to partly spreading (seldom nearly erect or reflexed after anthesis), usually modestly accrescent; sepals and petals usually ciliate, usually with long and/or stiff trichomes. Stamens 2-10(- 12) per petal, inserted between glands; anthers dorsifixed, broadly oblong-elliptical with oblong-elliptical locules and a large connective, the slits of dehiscence nearly parallel, or broader than long, usually very small, with subglobose locules diverging at a broad angle, slits of dehiscence often toward the apex. Upper surface of ovary conical in flower and fruit (in a few species, nearly flat and narrow, the ovary almost entirely inferior); styles 3-5(-7), free to near base. Locule of fruit much taller than broad, apex shaped like that of the ovary, pubescent at least in lower portion; seeds usually several per fruit, small, confined to apical portion of ovary. Distribution. ��� Southern Asia, Malesia, Pacific to Australia. Species included. ��� Homalium acuminatum Cheeseman, H. alnifolium Thwaites & F. Muell., H. aneityense Guillaumin, H. austrocaledonicum Seem., H. betulifolium D��niker, H. bracteatum Benth., H. buxifolium D��niker, H. caryophyllaceum (Zoll. & Moritz) Benth., H. celebicum Koord., H. circumpinnatum, H. dasyanthum, H. decurrens (Vieill.) Briq., H. dentrecasteauxense, H. deplanchei, H. foetidum, H. francii Guillaumin, H. guillainii (Vieill.) Briq., H. intermedium (Vieill.) Briq., H. juxtapositum Sleumer, H. kanaliense (Vieill.) Briq., H. kunstleri, H. laurifolium A.C. Sm., H. leratiorum Guillaumin, H. maneauense Craven, H. mathieuanum, H. moto H. St. John, H. moultonii, H. mouo H. St. John, H. myriandrum Merr., H. nitens Turrill, H. oblongifolium Merr., H. pallidum A.C. Sm., H. polystachyum (Vieill.) Briq., H. protectum Sleumer, H. ramosii Merr., H. rivulare (Vieill.) Briq., H. rubiginosum (Vieill.) Briq., H. rubrocostatum Sleumer, H. samarense Merr., H. schlichii Kurz, H. spathulatum Ridl., H. streimannii Craven, H. subcordatum Craven, H. tatambense Sleumer, H. taypau H. St. John, H. tongaense H. St. John, H. travancoricum Bedd. H. undulatum King, H. velutinum Craven, H. villarianum S. Vidal, H. vitiense Benth., H. whitmeeanum H. St. John. Notes. ��� The original publication of Homalium sect. Polyanthera did not designate a type, and included two species, Homalium deplanchei and H. polyandrum Warb. Because H. polyandrum is now considered to be a junior synonym of H. mathieuanum (Sleumer, 1974; Lescot, 1980), H. deplanchei is preferred as type. Some species-level taxonomic problems exist in this group, perhaps especially in the Pacific island taxa most recently dealt with by St. John (1977). Whistler (2004) tentatively treats H. tongaense as a synonym of H. whitmeeanum (described from Samoa) and implies that H. taypau and H. mouo are likely to be synonymous with H. acuminatum and H. moto respectively., Published as part of Applequist, Wendy L., 2016, A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae), pp. 231-256 in Candollea 71 (2) on page 254, DOI: 10.15553/c2016v712a9, http://zenodo.org/record/5721656, {"references":["Sleumer, H. (1974). A concise revision of the Flacourtiaceae of New Caledonia and the Loyalty Islands. Blumea 22: 123 - 147","Lescot, M. (1980). Flacourtiaceae. Fl. Nouvelle-Caledonie & Dependances 9: 3 - 134.","St. John, H. (1977). Revision of the Polynesian species of Homalium (Flacourtiaceae). Mitt. Deutsch. Dendrol. Ges. 69: 35 - 47.","Whistler, W. A. (2004). Rainforest trees of Samoa. A guide to the common lowland and foothill forest trees of the Samoan Archipelago. Isle Botanical, Honolulu."]}

5. Homalium sect. Nisa (Noronha ex Thouars) Baill. ex Warb. in Engler & Prantl, Nat. Pflanzenfam. III(6a): 36. 1893. �� Nisa Noronha ex Thouars, Gen. Nov. Madagasc.: 24. 1806. �� Homalium [unranked as ��] Nisa (Noronha ex Thouars) Baill. in Bull. Mens. Soc. Linn. Paris 1: 575. 1886. Typus: Homalium nudiflorum (DC.) Baill. (�� Nisa nudiflora DC.) (designated by Sleumer, 1973: 295). Stipules opposite the petiole base and fused (in H. intercedens Sleumer axillary, free). Inflorescences spicate; bract and bracteoles large, broad, thick-textured, persistent. Flowers sessile (rarely subsessile); perianth 5-6(-7)-merous. Sepals obovate to oblanceolate (to nearly ligulate), oblanceolateoblong or narrowly elliptical, or ovate to oblong-ovate), spreading, accrescent; calyx tube short, broadly funnelform to cup-shaped, or in fruit hemispherical; sepal glands large, elliptical to elongated oblong-elliptical or roughly trapezoid. Petals ovate to oblong, usually smaller than sepals and curving over fruit (in H. louvelianum similar in size and spreading), accrescent; sepals and petals not ciliate or ciliolate in conjunction with overall surface pubescence). Stamens 1 per petal, inserted between glands; anthers dorsifixed, broadly oblongelliptical with oblong-elliptical locules and a large connective, the slits of dehiscence nearly parallel. Upper surface of ovary often nearly flat in flower, in fruit becoming hemispherical or broadly conical; styles fused basally into a short but distinct column (rarely fused for most of length), the branches 3-4. Locule of fruit subglobose or vertically compressed, glabrous (short-pubescent); seeds 1 per fruit, subglobose, largely filling the locule (or several small immature seeds). Distribution. ��� Madagascar. Species included. ��� Homalium intercedens, H. louvelianum, H. nudiflorum, H. stelliferum H. Perrier., Published as part of Applequist, Wendy L., 2016, A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae), pp. 231-256 in Candollea 71 (2) on page 253, DOI: 10.15553/c2016v712a9, http://zenodo.org/record/5721656, {"references":["Sleumer, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328."]}

1. Homalium sect. Homalium = Racoubea Aubl., Hist. Pl. Guiane 1: 589, tab. 236. 1775. �� Homalium sect. Racoubea (Aubl.) C.B. Clarke in Hooker, Fl. Brit. India 2: 597. 1879. Typus: Homalium guianense (Aubl.) Oken. (�� Racoubea guianensis Aubl.). = Napimoga Aubl., Hist. Pl. Guiane 1: 592, tab. 237. 1775. Typus: Napimoga guianensis Aubl. Stipules axillary, free. Inflorescences racemose (to spicate) or paniculate; bracts small or narrow, usually persistent, bracteoles small, usually borne high on pedicel. Flowers pedicellate to subsessile (in part sessile), with pedicels articulated, 5-7- merous. Sepals ovate (to somewhat oblong or oblong-lanceolate), often moderately accrescent; calyx tube funnelform, in fruit to nearly hemispherical or broadly turbinate; sepal glands quite large, rounded to oblong-elliptic. Petals ovate to elliptical (to somewhat oblong, in H. fulviflorum Sleumer broadly spatulate), similar to sepals in size or longer, usually spreading at anthesis, often moderately accrescent; sepals and petals not ciliate or minutely ciliate. Stamens (2-)3-7(-8) per petal, inserted between glands; anthers dorsifixed, broadly oblongelliptical with oblong-elliptical locules and a large connective, the slits of dehiscence nearly parallel. Upper surface of ovary conical (to nearly flat) in flower, conical (sometimes very prominently) in fruit; styles fused basally into a short column (less often free to near base, or fused for most of length), the branches 3-4(-5). Locule of fruit ovoid to ellipsoid, apically conical, pubescent; seeds where known 1-2 per fruit, often ellipsoid, mostly filling the locule (sometimes several immature seeds present, possibly not maturing). Distribution. ��� Africa, Central and South America. Species included. ��� Homalium abdessammadii, H. angustifolium Sm., H. fulviflorum, H. guianense, H. racemosum, H. senarium Mo��. & Sess�� ex DC., Published as part of Applequist, Wendy L., 2016, A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae), pp. 231-256 in Candollea 71 (2) on pages 250-251, DOI: 10.15553/c2016v712a9, http://zenodo.org/record/5721656

2. Homalium sect. Antinisa Warb. in Engler & Prantl, Nat. Pflanzenfam. III(6a): 35. 1893. �� Nisa Noronha ex Thouars [unranked] Antinisa Tul. in Ann. Sci. Nat., Bot. ser. 4, 8: 73. 1857. �� Homalium [unranked as ��] Antinisa (Tul.) Baill. in Bull. Mens. Soc. Linn. Paris 1: 576. 1886. Typus: Homalium involucratum (DC.) O. Hoffm. (�� Nisa involucrata DC.). Stipules axillary, free. Inflorescences spicate with flowers mostly in few-flowered clusters; bracts very large, reniform and usually completely concealing flowers, persistent; bracteoles 2 per flower, broad, ciliolate, persistent. Flowers sessile; perianth 5-8-merous. Sepals lanceolate to narrowly lanceolate, modestly accrescent; calyx tube broad-based funnelform, densely sericeous; sepal glands irregularly rounded, unusually prominent. Petals lanceolate, becoming longer than sepals after anthesis, erect or nearly so, more accrescent than sepals and sometimes curving over fruit; sepals and petals ciliolate in conjunction with surface pubescence. Stamens 1 per petal, inserted between glands; anthers basifixed, the locules very small, attached by a deep bulbous connective (hence anther is somewhat rounded-deltoid in shape viewed from above) diverging at a relatively narrow angle, with slits of dehiscence lateral. Upper surface of ovary conical (sometimes shallowly) in flower, in fruit conical; styles 3-6, free to near base. Locule of fruit narrowly ellipsoid, densely pubescent in upper portion; seeds 1-4 per fruit, ellipsoid, largely filling the locule. Distribution. ��� Madagascar. Species included. ��� Homalium involucratum. Notes. ��� The description of Homalium sect. Antinisa includes fruit morphology not seen in this study but mentioned in literature (Sleumer, 1973)., Published as part of Applequist, Wendy L., 2016, A reconsideration of the infrageneric classification of Homalium Jacq. (Salicaceae), pp. 231-256 in Candollea 71 (2) on page 251, DOI: 10.15553/c2016v712a9, http://zenodo.org/record/5721656, {"references":["Sleumer, H. (1973). Revision du genre Homalium Jacq. (Flacourtiacees) en Afrique (y compris Madagascar et les Mascareignes). Bull. Jard. Bot. Natl. Belg. 43: 239 - 328."]}