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1 May 2018 A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar
Wendy L. Applequist
Author Affiliations +

Applequist, W.L. (2018). A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar. Candollea 73: 27–48. In English, English and French abstracts. DOI:

Homalium sect. Odontolobus Warb. (Salicaceae) is endemic to Madagascar and formerly included three species. In this revision, nine species are recognized, three of them newly described (Homalium densispicatum Appleq., Homalium masoalense Appleq., Homalium ovatifolium Appleq.) and three transferred from Homalium sect. Blackwellia Benth. Homalium densispicatum has highly reduced flowers like those of Homalium moniliforme H. Perrier, but the leaves are larger and the inflorescences thicker, with flowers in elongated many-flowered glomerules. Homalium masoalense and Homalium ovatifolium are northern species that have frequently ovate leaves and long styles and filaments; the flowers of Homalium masoalense are larger, with longer sepals, while Homalium ovatifolium has often canescent inflorescences. Two new subspecies are described: Homalium moniliforme subsp. littorale Appleq. and Homalium planiflorum subsp. roseiflorum Appleq. A key for identification of the species of Homalium sect. Odontolobus is provided.


Homalium Jacq. is a pantropical woody genus traditionally placed in Flacourtiaceae; along with a large portion of that formerly recognized family, its proper affinities are now known to be with the taxa placed in an expanded Salicaceae (Chase et al., 2002). The genus includes over 150 currently recognized species (see Applequist, 2013). Characters these species have in common include hermaphroditic flowers with two perianth whorls, semi-inferior ovaries, oppositipetalous stamens, single conspicuous large glands at the base of each sepal, and very small fruits with few maturing seeds; in other floral characters, they encompass an unusual degree of diversity.

Madagascar is perhaps the primary center of diversity of Homalium. Warburg (1894) published a complex classification of the genus that recognized two subgenera and nine sections. With some emendations and additions such as the publication of Homalium sect. Rhodonisa (Tul.) Sleumer, that classification has been mostly maintained since. The subgenera, defined by whether stamens were only one per petal (Homalium subg. Blackwellia (Benth.) Warb.) or fasciculate (Homalium subg. Homalium), were implicitly known by early authors to be artificial and were apparently accepted and maintained for convenience.The present author (Applequist, 2016) has proposed to emend this classification to recognize ten sections and omit recognition of subgenera, since the relationships among some sections are unclear. Of those ten sections, six are found in Madagascar and five are endemic there.

One of these endemic Malagasy sections is Homalium sect. Odontolobus Warb., which traditionally is characterized by a reduction of the sepals to tiny deltoid teeth; the flowers are usually sessile and quite small, in two species much reduced and borne in clusters, and the filaments and styles are unusually short. Warburg (1894) initially included two species, H. parkeri Baker and H. lucidum Scott-Elliot, in this section. Perrier de la Bâthie (1940) did not recognize Homalium sect. Odontolobus; he included these two species within Homalium sect. Blackwellia Benth. and described another species in that section, H. moniliforme H. Perrier, that was later transferred to Homalium sect. Odontolobus by Sleumer (1973), author of the most recent treatment of the Malagasy species of Homalium. Recent study of existing herbarium material has indicated that several additional, unrecognized taxa are assignable to this section. Moreover, three species traditionally placed in Homalium sect. Blackwellia were more similar to members of Homalium sect. Odontolobus in overall morphology, though lacking the extreme sepal reduction. A modernized treatment of Homalium sect. Odontolobus was thus needed and is herein presented.

Material and methods

Herbarium specimens at P and MO (herbarium acronyms according to Index herbariorum, 2018) were examined, as were then-undistributed duplicates for exchange available at those herbaria and images available through JSTOR Global Plants website [] of types held by other institutions. Most recent collections have duplicates at TAN or TEF, including all those derived from the Missouri Botanical Garden's Madagascar program. As these were not seen they are not listed among material examined; however, the locations of many such duplicates are available through Tropicos (2018).

Species were defined according to a taxonomic species concept (e.g., Grant, 1981), the most widely used and usually most practical approach in plant taxonomy, with three morphological features differing among groups of populations considered sufficient to warrant recognition at the species level. Flowers undergo little expansion in fruit, but are usually caducous at or before fruit maturity. Because fruits with mature seeds are rarely seen, only flowering characters are used in species delimitation. In descriptions of taxa, character states or size ranges given in parentheses are uncommon.

Known distributions and habitat are described for each species or subspecies. To save space, locality data are not provided for all specimens seen of common taxa, and those that are provided are edited for brevity, especially when full label data are available from Tropicos (2018). “Fkt.” is used throughout as an abbreviation for “fokontany”. A complete index of specimens seen is provided as an appendix. Maps of georeferenced specimens may also be generated within the Madagascar Catalogue (2018), which is continually updated with new determinations and specimens.

A preliminary, unofficial assessment of conservation status using the categories and criteria of IUCN (2012) is provided for each taxon recognized. In instances when the extent of occurrence (EOO) and area of occupancy (AGO) might affect the assessed status, GeoCAT (Bachman & Moat, 2012) was used to estimate those values. Geographic coordinates were taken from label data or from the Missouri Botanical Garden's gazetteer (Tropicos, 2018).

Taxonomic treatment

Homalium sect. Odontolobus Warb, in Engl. & Prantl, Nat. Pflanzenfam. III(6a): 35.1894.

  • Typus (designated by Sleumer, 1973: 306): Homalium parkeri Baker.

  • Trees (occasionally shrubby in H.parkeri). Stipules axillary, free. Inflorescences spicate to racemose (partly paniculate with racemoid branches in H. planiflorum subsp. roseiflorum) with most flowers borne in small clusters or glomerules (singly in H. brachystylis); bracts small, often broad, usually persistent; bracteoles caducous or persistent (rapidly caducous or possibly absent in H. masoalense), minute to broad and larger than bracts, usually thick-textured, in 2 species fleshy, approaching subterete at base. Leaves alternate (sometimes opposite or subopposite), glabrous (occasionally glabrate with few trichomes on midrib or sparsely pubescent when young). Flowers sessile or short-pedicellate, pedicels not articulated; perianth 5-8-merous. Sepals reduced to minute deltoid teeth or small and ligulate to lanceolate-oblong, not at all accrescent; calyx tube broadly funnelform (to narrowly so in H. lucidum), in fruit becoming nearly hemispherical; sepal glands rounded (to elliptical), densely pubescent (to sparsely so, usually with age). Petals ovate, sometimes broadly to transversely so, or narrowly oblong-lanceolate to oblong-elliptical, sometimes quite small but always larger than sepals, spreading, not or very little accrescent; sepals and petals lacking cilia, ciliolate, or ciliate with sometimes long, wavy, fine trichomes. Stamens 1 per petal, inserted between glands (filaments usually short); anthers basifixed, broader than long, very small, with subglobose locules diverging at a wide angle, dehiscent by short slits at or near the apex. Upper surface of ovary broad and nearly flat (to convex or broadly conical) in flower, in fruit becoming convex to hemispherical; styles 2-4(-5), free, often quite short. Locule of fruit subglobose to obovoid, sparsely pubescent to glabrate; seeds 1 per fruit, subglobose, occupying the entire locule (sometimes several-seeded, the seeds then possibly not maturing).

  • Distribution. — Homalium sect. Odontolobus is endemic to Madagascar, where its members occur throughout much of the island.

  • Notes. — Homalium sect. Odontolobus has been expanded to include three species formerly placed in Homalium sect. Blackwellia (Applequist, 2016). It is characterized by its small open flowers borne in spicate or racemose inflorescences (or seldom panicles with racemiform branches), which are sessile to short-pedicellate and have a usually relatively short, broad calyx tube, a broad upper ovary surface that is nearly flat (sometimes to convex, but not narrowly conical) at anthesis, sometimes highly reduced sepals, and small anthers dehiscing towards the apex. The styles and filaments are often short; the locule of the ovary is subglobose to obovoid, with little or no internal pubescence, and there is usually only one large seed per fruit, though mature fruit is rarely seen.

  • Key to the species of Homalium sect. Odontolobus

    1. Sepals reduced to small teeth, barely visible from above 2

    1a. Sepals from nearly as long as petals to less than half as long, sometimes quite small but readily visible from above 5

    2. Inflorescences canescent; most flowers borne in elongated, many-flowered clusters inserted close together on a thick rachis 2. H. densispicatum

    2a. Inflorescences glabrous or sparsely or minutely pubescent; flowers in few-flowered clusters or subglobose moniliform clusters mostly separated on rachis 3

    3. Flowers mostly in groups of 2 or 3, sessile or petiolate; petals lanceolate (narrowly ovate) to oblong-lanceolate or narrowly elliptical (elliptical), 1.5-2.5(-3.1) mm long, margin ciliolate, apex acute 4. H. lucidum

    3a. Flowers in subglobose, often moniliform clusters, sessile; petals ovate (or somewhat oblong) to transversely ovate, not over 1.5 mm long, margin not ciliolate, apex acute or rounded (to obtuse) 4

    4. Petals transversely ovate (seldom very broadly ovate or oblong-ovate), 0.5-0.7(-0.8) mm long, apex rounded (rounded-obtuse) 6. H. moniliforme

    4a. Petals ovate (oblong-ovate), 0.7–1.5 mm long, apex acute to rounded 8. H.parkeri

    5. Flowers solitary; filaments and styles 0.3–0.6 mm long 1. H. brachystylis

    5a. Flowers at least partly in clusters of 2 or more; filaments and styles (0.4-)0.6-1.9 mm long 6

    6. Inflorescences canescent (to in part moderately pubescent); bracteoles densely pubescent, fleshy to subterete at the base; petals 0.8–1.2 mm long; sepals (0.3-)0.4-0.5(-0.6) mm long, deltoid to ovate or narrowly deltoid 7

    6a. Inflorescences moderately to sparsely short-pubescent; bracteoles in 1 species glabrous or minutely pubescent, flat; petals 1–1.7 mm long; sepals 0.5-0.9(-1.1) or 1.2–1.4 mm long, oblong-lanceolate to narrowly oblong or lanceolate (ovate) 8

    7. Leaves 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 length; apex acute to obtuse or rounded (short-acuminate); petals oblong to somewhat obovate, pubescent on both surfaces, densely so abaxially and usually at base adaxially; SE Madagascar 3. H. longistaminum

    7a. Leaves 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; petals narrowly oblong, abaxially pubescent, mostly glabrous adaxially; N Madagascar 7. H. ovatifolium

    8. Leaf blade ovate to lanceolate (elliptical) with acuminate (acute, emarginate) apex, convex to rounded base; flowers sessile or subsessile with pedicels to 0.5 mm long; petals 1.5–1.7 mm long, sepals 1.2–1.4 mm long; Masoala Peninsula 5. H. masoalense

    8a. Leaf blade narrowly elliptical to elliptical or oblanceolate with acute to acuminate (rounded, emarginate, cuspidate) apex, convex (cuneate, attenuate) base; flowers pedicellate with pedicels (0.4-)1.3-3(-3.5) mm long; petals 1–1.7 mm long, sepals 0.5-0.9(-1.1) mm long 9. H.planiflorum


    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: Forêts à l'entrée de la Baie de Vohémar, 1837, fl. & fr., Richard 63 (P [P04679017]!; isolecto-: G [G00018411] image seen, L [L0010889, L0010890] images seen). Syntypi: Madagascar. Prov. Antsiranana: Vohémar, 1846, fl. & fr., Richard 110 [= Boivin 2568] (P [P04734119]!). Sine loco: “Sud de Madagascar [?]”, s.d., fr., Richard 561 (P [P04734120]!); Herb. Galldich [cl.Jaubert] 40 (not found).

  • = Homalium humbertii H. Perrier in Mém. 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 réception 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: vallée moyenne du Mandrare près d'Anadabolava, Mont Vohitrotsy, 700–850 m, XII.1933, fl., Humbert 12655 (G [G00018413, G00018414, G00018415] images seen, P [P04734115, P04734116, P04734117, P04734118]!, S [S10-10143] 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 ob ovate), 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. Gérard 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 Vohémar 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 Vohémar 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 Vohémar 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 Vohémar 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]); forêt d'Antsahabe, 10 km à 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 à 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, forêt d'Andranomadiro, 12°36′18″S 49°26′34″E, 258 m, 7.XI.2006, fl., Ranaivojaona et al. 1519 (MO); Vohemar, forêt de Binara, 13°14′09″S 49°37′22″E, 300 m, 31.III.2004, fl., Ranirison 551 (MO); Fkt. Saharenana, forêt d′Andranomadiro, 12°36′18″S 49°26′35″E, 300 m, 14.11.2005, fl., Schatz et al. 4264 (MO); Diégo-Suarez, Ankara JB8, 27.VII.1954, buds, Service Forestier 10426 (P); plateau de l'Ankarana près d'Ambondromifehy, 4-6.X.1954, Service Forestier 11257 (MO); Diégo-Suarez, Ankara, 24.III.1955, fl., Service Forestier 13208 (P); Ankarana à 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 inférieures 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 vallée de la Menarahaka a l'E d'Ihosy, 700–800 m, 28.1.1955 & 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 Bâthie 19272 (P [2 sheets]); forêt de Kitranga, à 12 km au N d'Ihosy, 1.1955, fl., Service Forestier 11611 (P [2 sheets]); Zasafotsy, Ihosy, 21.VII.1954, buds, Service Forestier 14482 (P); Kitranga, entre Ambararata et Ivandrika, 22.11.1970, fl., Service Forestier 29063 (P). Prov. Mahajanga: Bemanevika, forêt 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); forêt 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], 18.1.1951, fl., Réserves Naturelles 2756 (MO, P [2 sheets]); Tule ar, forêt d'Analavelona, 7.IV.1954, fr., Service Forestier 9872 (MO, P); Mahaboboha, forêt d'Ibera, 9.IV.1954, fr., Service Forestier 10080 (P); forêt de Zombitsy, 700 m, III.1955, fl., Service Forestier 11912 (P); Andranolava, forêt Vohibasia, 27.V.1955, fl., Service Forestier 14092 (P); Forêt de Zombitsy, 700–800 m, 20. VI.1958, fl. & fr., Service Forestier 18593 (P).

  • 2. Homalium densispicatum Appleq., spec, nova (Fig. 1).

  • Holotypus: Madagascar. Prov. Toliara: Anosy, Fort Dauphin, Iabokoho, Antsotso, forêt d'Ivohibe, 24°34′14″S 47°12′10″E, 230 m, 2.IV2008, fl., Rabenantoandro et al. 1884 (MO-6474628!; iso-: P [P06171672]!,TAN).

  • Homalium densispicatum differs from H. moniliforme H. Perrier in having larger leaves, (5.5-)7-15 (-17) × (1.5-)2.1-6(-6.9) cm (vs (1.8-)2.7-6.8 × 0.7-2.6(-3) cm); inflorescences canescent (vs minutely pubescent or glabrous); and flowers borne in elongated many-flowered glomerules along most of the rachis (vs in well-separated, moniliform glomerules).

  • Tree to 12 m tall; young twigs glabrous. Leaves: petiole 2.5-20(-30) mm long, glabrous; blade narrowly elliptical to elliptical (oblanceolate to oblong-oblanceolate, obovate), (5.5-)7-15(-17) × (1.5-)2.1-6(-6.9) cm; base cuneate to attenuate or convex, usually attenuate at extreme base; margins crenate-serrulate (subentire, crenate-serrate or crenulate), often slightly revolute; apex rounded to acute or cuspidate (shallowly emarginate, short-acuminate). Inflorescences spicate with a thick rachis, 1.4–7 cm long, canescent; flowers in elongated manyflowered clusters, sessile; bracts transversely (to very broadly) oblong, deltoid or ovate, often irregular; bracteoles reduced to minute teeth, caducous. Flowers (6-)7(-8)-merous, white or greenish; sepals ovate to oblong or broadly ovate-deltoid, 0.2-0.3(-0.4) mm long; petals broadly (transversely) oblongelliptical to oblong-ovate, (0.4-)0.5-0.8 mm long, glabrous (pubescent abaxially but not ciliate), apex rounded; filaments 0.1-0.4(-0.5) mm long; upper surface of ovary densely shorttomentose; styles (2-)3, 0.1–0.4 mm long.

  • Distribution, ecology and conservation status. — Homalium densispicatum is primarily native to humid forests in southeastern Madagascar at low (seldom to moderate) elevations. As herein circumscribed, it includes one disjunct population from northern Madagascar. Only eight distinct locations are known for H. densispicatum (if the Antsiranana collection is included). Most low-elevation forest in Madagascar is now badly fragmented. However, most collections are from two protected areas (Manombo, Tsitongambarika) where the species seems not to be rare and further decline in habitat extent or quality may be avoidable. Therefore it is suggested that an appropriate preliminary assessment of conservation status would be “Least Concern” [LC].

  • Notes. — Homalium densispicatum shares extremely reduced, tightly clustered flowers with H. moniliforme, and some specimens have previously been assigned to that species. However, the leaves are much larger, with numerous veins and sometimes much longer petioles, the inflorescences are much more densely pubescent, and the flowers are borne on a thick rachis in elongated many-flowered clusters that at their termini are mostly close to one another on alternating sides of the rachis, giving the inflorescence a uniformly thick, rather than moniliform appearance.

  • The distribution is primarily southeastern. One smallleaved collection (Razakamalala et al. 55) is known from Antsiranana, far to the north; though this distribution is suspicious, the specimen is overall morphologically consistent with H. densispicatum and not with H. moniliforme, e.g., in having thick inflorescences with elongated, nearly contiguous glomerules. A second northern collection, Service Forestier 28807 from Ile Sainte-Marie (partie S de la forêt de Kalalao, 16.V.1969, fr., P [2 sheets]), has leaves to 9.7 × 4.1 cm, yet in other features it seems to resemble H. moniliforme more than H. densispicatum, it is therefore treated as incertae sedis. Specimens from Fianarantsoa province have smaller leaves than those from Toliara; one of the latter, Razafimandimbison et al. 223, has unusually narrow and strongly toothed leaves and might represent a distinctive local variant.

  • Paratypi.Madagascar. Prov. Antsiranana: Fiv. Antalaha, Fkt. Sahafary, 15°17′34″S 50°22′07″E, 173 m, 18.11.2001, fl., Razakamalala et al. 55 (MO). Prov. Fianarantsoa: Ranomafana, XII.1963, fl., Chabonis s. n. (P); Fkt. Manombo, Réserve Spéciale de Manombo, 23°01′19″S 47°43′56″E, 30 m, 14.XI.2001, fr., et al. 780 (MO); forêt de Manombo, à 30 km au S de Farafangana, J.B. 16,26. VI.1954, fl. & fr., Service Forestier 9205 (MO, P [7 sheets]); Ihorombe, Manombo, 21.VII.1955, fr., Service Forestier 15240 (P). Prov. Toliara: Fkt. Antsotso, Ivohibe forest, 24°34′10″S 47°12′37″E, 41 m, 24. V.2006, fl., Antilahimena et al. 4846 (P); Fkt. Iaboakoho, forêt d'Ampasina, 24°34′30″S 47°08′35″E, 119 m, 25.1.2015, fl., Randrianarivony et al. 623 (MO); village d' Antsotso, forêt de Bemangily, 24°35′33″S 47°12′52″E, 22.V.2006, fl., Randriatafika et al. 672 (MO, P); forêt de Manantantely à 15 km au NW de Fort-Dauphin, 24°59′S 46°55′E, 50–100 m, 24.VI.1996, fl., Razafimandimbison 223 (BR, G, K, 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. 4113 (MO, P). Sine loco: s.d., fl., Service Forestier 71-R-176 (P).

  • Fig. 1.

    Homalium densispicatum Appleq. A. Flowering branch; B. Flower cluster.

    [Rabenantoandro et al. 1884, TAN] [Drawing: R.L. Andriamiarisoa]


    3. Homalium longistaminum H. Perrier in Mém. Mus. Natl. Hist. Nat. 13:287.1940.

  • Lectotypus (designated here): Madagascar. Prov. Toliara: vallée 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 km2. 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 (< 20 km2) and number of locations (≤ 5).

  • Note. — In publishing this species, Perrier de la Bâthie (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 Bâthie probably intended it as such. However, since both sheets were used by Perrier de la Bâthie 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).

  • 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” (Réserves 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 Mangabé 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, 16.1.1985, 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); forêt de Besofina à 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, 20.1.1950, fl., Service Forestier 2737 (P); Manakara, Vakoary, 11.V.1955, fr., Service Forestier 14569 (P); Manakara, forêt Belambo, 11.V.1955, Service Forestier 14608 (P). Prov. Mahajanga: 9.6 km NW of Ambohistaratelo-Bebao, 1200 m, 14.1.1985, fl., Dorr et al. 3535 (MO, P); 5 km à 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, 73., Antilahimena et al. 7110 (G, K, MO, P); Andapanomby, près de la rivière Ampandisanana, Makira NW, 15°21′48″S 49°07′06″E, 25.IV.2007, fr., Bernard & Birkinshaw 462 (G, MO, P); Nosy Mangabé, 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 Bâthie 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., Réserves 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).

  • 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.

  • 6. Homalium moniliforme H. Perrier in Mém. 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 re volute; 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 Bâthie, 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.

  • Fig. 2.

    Homalium masoalense Appleq. A. Flowering branch; B. Flowers.

    [Schatz et al. 3367, TAN] [Drawing: R.L. Andriamiarisoa]


    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

    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 & Céléstin 712); “Menahihy” (Iambana & Arsène 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 latente, 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, forêt 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 à 4 km, 17°55′S 49°13′E,21.IV.1999, fr., Iambana & Arsène 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 a'Ankerana, 18°25′24″S 48°47′06″E, 896 m, 22.III.2011, fl., Ravelonarivo et al. 3785 (MO); Fkt. Ambatolampy, forêt d'Ankarana, 18°23′51″S 48°47′53″E, 948 m, 25.I.2012, fl., Ravelonarivo et al. 4194 (MO); Sahanionaka, forêt 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, forêt de Vohibe, 19°09′19″S 48°35′04″E, 655 m, 5.IV.2009, fl., Razanatsima & Céléstin 712 (MO [2 sheets]); Fkt. Ambatolampy, forêt d'Ankerana massif de Beanjada (N de la presqu'ile Masoala), 1000 m, I.1954, fr., Service Forestier 8825 (P).

  • 6b. Homalium moniliforme subsp. littorale Appleq., subspec. nova (Fig. 3).

  • Holotypus: Madagascar. Prov. Fianarantsoa: Fiv. Farafangana, firaisam-pokontany Mababo Mananivo, fkt. Nosy ala, forêt 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., Mababo/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 km2, and the AOO as 56 km2. The preliminary assessment of conservation status suggests that a status of “Vulnerable” [VU Blab(iii)+B2ab(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, forêt d'Agnalazaha, bloc Agnanto, 23°10′09″S 47°41′51″E, 28 m, 23.III.2004, fl., Ludovic 649 (MO); Fkt. Nosiala, forêt 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, forêt d'Ampitavananima, 23°11′10″S 47°43′02″E, 30 m, 5.VIII.2004, fr., Ludovic et al. 870 (MO); Mahabo-Mananivo, forêt 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 à 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, forêt de Menagisy 500 m du village Antananflava Martin, 16°47′S 49°43′E, 26.V.2010, fl., Lehavana & Aackarie 692 (MO); Tampina, s.d., fr., Louvel 53 (P); Fénérlve, forêt à Mahambo, 12.VIII.1961, fl., Peltier & Peltier 3419 (P); Vohibola, à l'W de Tampina, 12-14.VIII.1957, fl., Service Forestier 18072 (P); Mahambo, au S de Fénérlve, 30.VIII.1957, fl. & fr., Service Forestier 18146 (MO, P [3 sheets]). Prov. Toliara: Mandena, 24°57′S 47°02′E, 10 m, 13.1.1990, fl., Dumetz 1184 (MO, P); forêt au NE de la rivière 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).

  • 7. Homalium ovatifolium Appleq., spec, nova (Fig. 4).

  • Holotypus: Madagascar. Prov. Antsiranana: Ambilobe, Marivorahona, village le plus proche Betsimiranja, forêt d'Andohan'Antsohihy, Ambohibe, RN6 à 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).

  • 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).

  • Fig. 3.

    Homalium moniliforme subsp. littorale Appleq. A. Flowering branch; B. Flowers; C. Inflorescence; D. Leaf (upper surface); E. Leaf (lower surface). [Rabehevitra et al. 516, TEF] [Drawing: R.L. Andriamiarisoa]


    Fig. 4.

    Homalium ovatifolium Appleq. A. Flowering branch; B. Flowers.

    [Randrianaivo et al. 1206, TAN] [Drawing: R.L. Andriamiarisoa]


    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” (Louvel43); “Hazombatomainty” (Service Forestier 16010, 16822); “Hazomby” (Service Forestier 595, 1043, 3926, 6006, 10370, 15990); “Hazompoza” (Service Forestier 7592); “Moara” (Réserves Naturelles 9460); “Ramaindafa” [?] (Réserves 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 Concer”n [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 Imerma, Andrangolaoka, III.1881, fl. & fr., Hildebrandt 4102 (P [2 sheets]); Firarazana, forêt 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 Bâthie 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, forêt d'Ambohimangakely, 6.IV.1961, fl., Service Forestier 19894 (P); Ibity massif W, Vohipisaka, 20°09′42″S 46°58′54″E, 1423 m, 29.11.2004, 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 à 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, 26.11.1938, 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, forêt 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. 62 77 (G, MO); Andapanomby, près 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, forêt 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, 19.111.2013,11., 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, forêt 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., Réserves 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, à l'E de Fierenana, 950–1300 m, 10-16.III.1969, fl., Service Forestier 28755 (MO, P).

  • 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.; Appleq,uist, 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 ob ovate), (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 ob ovate, 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 brûlée” according to Cloisel and “fulvo-rosei (sordid)” according to Bernard!) 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 Bâthie (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 Bâthie'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

    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, Réserves Naturelles s.n., Service Forestier 1577, 1789, 5693, 5858, 5866, 16862, 17810); “Hazombato mena (lahy)” (Réserves 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” (Réserves Naturelles 8807); “Moranga ala” (Réserves 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, 24X1.1995, fl. & fr., Bernard 126 (G, K, MO, P); Parc Masoala, forêt Ambodipont, 15°44′45″S 50°19′25″E, 0–10 m, 3.X.1996, fl., Bernard 361 (G, MO, P); vallée inférieure 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 Bâtbie 6699 (P); Andravinambo, forêt de Tsihomanaomby, 14°06′S 50°01′E, 225 m, 3.II.2014, fl., Rakotonirina et al. 446 (MO); Sambava, forêt d'Ambodivohitra, 14°24′42″S 49°51′49″E, 148 m, 20.X.2010, fl., Ravelonarivo & Raharivelo 3541 (MO, P); vallée de l'Ampalohandrano, affluent de l'Anohoranga, aux env. d'Antongondriha, 150 m, 9.XI.1950, fl., Service Forestier 794 (P); Antsambalahy, Vohémar, 2.XII.1955, fl., Service Forestier 15228 (P); env. S d'Analamanara (près 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 vallée 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 côtiere, 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. Marchita, forêt 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, forêt d'Analavory, 10.XI.1955, fl. 8c 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. 11.1999, 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 à Larrée, forêt Menagisy, 16°46′39″S 49°41′32″E, XI.2008, fl., Nikolov 1815 (MO); bassin inférieur du Mangoro, X.1922, fl., Perrier de la Bâthie 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, forêt 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., Réserves 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, 1.1954, fl. & fr., Service Forestier 8916 (P); Tampina, s.d., fr., Ursch s.n. (P). Prov. Toliara: massif de l'Andohahel[a] (SE), vallée de Ranohela, 300–1200 m, 18-26.X.1928, fl. & fr., Humbert 6244 (P [2 sheets]); Sainte Luce, forêt de Beboaka, 24°40′25″S 47°12′00″E, 13 m, 25.XI.2011, fl., Miandrimanana 535 (MO); Ivorona, forêt 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).

  • 9b. Homalium planiflorum subsp. roseiflorum Appleq., subspec. nova (Fig. 5).

  • Holotypus: Madagascar. Prov. Toliara: Anosy, Taolagnaro, Ampasy Nahampoana, forêt littorale d'Ambavarano-Mandena, 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, Réserves 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 (Randriataflka et al. 319, Ratovoson 1629, 1692, 1731, Réserves 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 ofToliara 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 km2 and an AOO of 40 km2. 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.

  • Notes. — Homalium planifiorum subsp. roseifiorum 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. planifiorum, 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, étude 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, 24.1.1990, 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, près 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, 13.1.1990, fr., Rabevohitra 2123 (MO, P); Taolagnaro, à 1 km au N d'Evatra, 24°57′S 47°06′E, 17.1.1990, fl. & fr., Rabevohitra 2153 (MO, P); Sainte Luce, forêt d'Analavinaky à 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 forestière de Mandena (M3), 3.X.2001, fl., Randriatafika et al. 319 (MO); Sainte Luce, S10, 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, S5, 1 km E de Belavenoko Lorêt, 24°49′51″S 47°06′49″E, 20 m, 23.XI.2011, fl., Ratovoson 1731 (MO); Mandena (route Nancy), 6.XII.1949, fl., Réserves Naturelles (Ranjatson) s.n. (P); Sainte Luce, QMM Forestry Station, parcel S9,24°46′05″S 47°10′16″E, 11 m, 23.1.2006, fl., Rogers et al. 970 (G, MO); forêt de Vinanibe, près de Fort Dauphin, 10.XII.1961, fl., Service Forestier 20513 (P [2 sheets]); Fkt. Ampasinahampoana, forêt 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, 28.11.1973, 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).

  • Fig. 5.

    Homalium planiflorum subsp. roseiflorum Appleq. A. Flowering branch; B. Flower clusters; C. Leaf (lower surface).

    [Rabenantoandro & Ramisy 1765, TAN] [Drawing: R.L. Andriamiarisoa]



    The author thanks the Muséum national d'Histoire naturelle for permitting study of collections at P; Sovanmoly Hul, Peter Phillipson, Simon Verlynde, and Jacques Florence for assistance during my visit and useful discussions; Porter P. Lowry II for valuable comments on a draft manuscript and help processing image files; Robbie Hart for help with an image file; Roger Lala Andriamiarisoa for drawing the figures; George Schatz for advice on conservation assessments; and Martin Callmander for editing and advice regarding conservation assessments and for providing images of type material at G.



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    Index to collectors. Collections are listed alphabetically by first collector's last name, with determinations indicated by numbers corresponding to those of species in the taxonomic treatment; types are indicated in boldface.

    Almeda 8645 (8); Andriamihajarivo 293 (1), 577 (4);Andrianantoanina 738 (1); Andrianarisata 101 (3); Andrianjafy 189 (6a), 1038 (1); Antilahimena 1139 (4), 2046 (6a), 2171 (6a), 2597 (6a), 2993 (9a), 4835 (4), 4846 (2), 5242 (4), 5616 (6a), 6277 (8), 7110 (4).

    Bardot-Vaucoulon 1325 (1); Baron 1295 (8), 5301 (8), 5928 (8), 5966 (9a); Be 42 (1); Bernard 126 (9a), 361 (9a), 456 (8), 462 (4); Bernardi 11517 (9); Birkinshaw 279 (8), 427 (1), 559 (9a); Boivin s.n. (9a), 1847 (9a); Bosser 9575 (8), 18580 (9a).

    Carlson 192 (4); Chaboniss.n. (2); Cloisel 120 (9), 188 (9a); Cours 2468 (4), 2965 (9a), 4875 (9a); Cremers 1636 (8).

    Debray 1985 (9); Decary 5469 (6a); Dorr 3535 (4), 3587 (4); Dumetz 666 (9b), 754 (9b), 1127 (9b), 1184 (6b), 1236 (9b).

    Eboroke 980 (3), 985 (1).

    Faliniaina 2 (9b).

    Geay 7855 (9a); Gereau 3365 (9); Giraudy s.n. (9a); Guittou 160 (7).

    Harder 1533 (4); Herb. Jard. Bot. 3141 (8); Hildebrandt 4102 (8); Hoffmann 209 (9b); Hong-Wa 392 (8); Humbert 6244 (9a), 12655 (1), 12914 (3), 14077 (1), 19161 (1), 23979 (9a), 28538 (1), 29567 (1).

    Iambana 181 (4), 208 (6a), 270 (6a).

    Lam s.n. (9a); Lantz s.n. (9a); Le Thomas 122 (9a); Leeuwenberg 13933 (4); Lehavana 487 (8), 692 (6b); Louvel 43 (8), 55 (6b), 64 (9a), 65 (9a), 146 (9a); Lowry 4419 (8); Ludovic 46 (8), 184 (9a), 213 (9a), 649 (6b), 696 (6b), 761 (6b), 870 (6b).

    Mabberley 910 (1); Malcomber 1317 (8), 1495 (8), 1514 (4); McPherson 14339 (9b), 14369 (9b), 17257 (8); Miandrimanana 253 (8), 555 (9a); Miller 3311 (9a); Morat 4639 (4).

    Nicoll 553 (9a); Nikolov 1815 (9a).

    Parker s.n. (8); Peltier 3419 (6b); Perrier de la Bâthie 6699 (9a), 6720 (8), 14073 (9a), 17012 (4), 15262 (9a), 19272 (1); Poncy 1543 (9a).

    Rabehevitra 516 (6b); Rabenantoandro 336 (9b), 753 (9a), 780 (2), 885 (9a), 961 (6b), 990 (6b), 1522 (9b), 1765 (9b), 1884 (2); Rabevohitra 1781 (6b), 2125 (9b), 2155 (9b), 3876 (9a), 4006 (9a); Raharijaona 72 (8); Rakotomalaza 410 (9a), 675 (8), 1555 (8), 1344 (4); Rakotondrafara 335 (1); Rakotondrajaona 122 (8), 336 (1); Rakotonirina 446 (9a); Rakotovao 4737 (9b), 5042 (9b); Ramahenina 224 (8); Ramison 561 (9b), 584 (4); Ranaivojaona 365 (1), 431 (9a), 1519 (1); Randriamampionona 152 (3), 302 (1), 309 (4), 376 (4); Randrianaivo 517 (4), 1206 (7); Randrianarivony 154 (6a), 623 (2); Randrianasolo 65 (8), 151 (8), 343 (6b), 567 (9b), 603 (9a), 649 (9a), 746 (9a), 853 (6b), 878 (4), 947 (6a); Randrianjanaka 586 (4), 606 (4), 629 (8); Randriatafika 319 (9b), 672 (2); Ranirison 551 (1); Ratovoson 1208 (1), 1220 (1), 1629 (9b), 1692 (9b), 1731 (9b); Ravelonarivo 811 (8), 1925 (8), 2015 (8), 2253 (4), 3541 (9a), 3785 (6a), 3950 (8), 4194 (6a); Razafimandimbison 223 (2); Razafindramora 26 (8); Razafitsalama 1004 (6b); Razakamalala 55 (2), 1518 (9a), 1646 (9a), 4113 (2), 5021 (9a), 5447 (6a); Razanatsima 712 (6a), 859 (6a); Reserves Naturelles s.n. (9a), s.n. [Dumazer] (9a), s.n. [Ranjatson] (9b), 1526 (9a), 2756 (1), 3344 (9a), 3668 (4), 4932 (9a), 8807 (9a), 9460 (8), 10472 (8); Richard 63 (1), 87 (1), 110 (1), 260 (9a), 297 (9a), 561 (1), 593 (1), 2974 (9a); Rogers 970 (9b), 2006 (9a).

    Sauquet 96 (9a); Schatz 1380 (4), 2055 (4), 2349 (4), 2542 (4), 2694 (4), 2780 (9a), 3094 (9a), 3367 (5), 3377 (9a), 4264 (1); Scott Elliot 2661 (4), 2824 (9a), 2555 (9); Service Forestier 71-R-176 (2), 595 (8), 794 (9a), 865 (9a), 1043 (8), 1577 (9a), 1789 (9a), 2715 (4), 2719 (9a), 2737 (4), 2756 (1), 2861 (9a), 3926 (8), 4226 (9a), 5693 (9a), 5858 (9a), 5866 (9a), 6006 (8), 7592 (8), 8090 (9a), 8420 (8), 8825 (6a), 8916 (9a), 9107 (4), 9205 (2), 9872 (1), 10080 (1), 10155 (9a), 10370 (8), 10426 (1), 11070 (9a), 11257 (1), 11611 (1), 11912 (1), 12000 (8), 13208 (1), 13728 (1), 13942 (4), 14092 (1), 14404 (9a), 14449 (9a), 14482 (1), 14569 (4), 14605 (4), 15183 (9a), 15228 (9a), 15240 (2), 15282 (9a), 15873 (8), 15990 (8), 16010 (8), 16259 (9a), 16807 (8), 16822 (8), 16862 (9a), 17810 (9a), 17999 (9a), 18027 (8), 18072 (6b), 18146 (6b), 18593 (1), 18767 (8), 19894 (8), 19957 (8), 20513 (9b), 22695 (1), 23376 (1), 23627 (9a), 24673 (1), 27184 (9a), 27226 (9a), 27537 (1), 28755 (8), 28807 (aff 2?), 29063 (1); Skema 22 (8), 54 (8); Soanary 26 (9b).

    Turk 545 (8), 547 (8).

    Ursch s.n. (9a).

    Wendy L. Applequist "A revision of Homalium sect. Odontolobus (Salicaceae) endemic to Madagascar," Candollea 73(1), 27-48, (1 May 2018).
    Published: 1 May 2018
    Homalium sect. Odontolobus
    new species
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