Research Article |
Corresponding author: Neil Snow ( nsnow@pittstate.edu ) Academic editor: Peter de Lange
© 2015 Neil Snow, Martin Callmander, Peter B. Phillipson.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Snow N, Callmander M, Phillipson PB (2015) Studies of Malagasy Eugenia – IV: Seventeen new endemic species, a new combination, and three lectotypifications; with comments on distribution, ecological and evolutionary patterns. PhytoKeys 49: 59-121. https://doi.org/10.3897/phytokeys.49.9003
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Seventeen new endemic species of the genus Eugenia L. (Myrtaceae) are proposed from Madagascar, including: E.andapae N. Snow, E.barriei N. Snow, E.bemangidiensis N. Snow, E.calciscopulorum N. Snow, E.delicatissima N. Snow, Callm. & Phillipson, E.echinulata N. Snow, E.gandhii N. Snow, E.hazonjia N. Snow, E.iantarensis N. Snow, E.malcomberi N. Snow, E.manomboensis N. Snow, E.obovatifolia N. Snow, E.ranomafana N. Snow & D. Turk, E.ravelonarivoi N. Snow & Callm., E.razakamalalae N. Snow & Callm., E.tiampoka N. Snow & Callm., and E.wilsoniana N. Snow, and one new combination, Eugeniarichardii (Blume) N. Snow, Callm. & Phillipson is provided. Detailed descriptions, information on distribution and ecology, distribution maps, vernacular names (where known), digital images of types, comparisons to morphologically similar species. Preliminary assessment of IUCN risk of extinction and conservation recommendations are provided, including Vulnerable (4 species), Endangered (2 species), and Critically Endangered (4 species). Lectotpyes are designated for Eugeniahovarum H. Perrier, Eugenianompa H. Perrier, and E.scottii H. Perrier respectively.
Dix-sept nouvelles espèces endémiques du genre Eugenia L. (Myrtaceae) sont proposées pour Madagascar, incluant E.andapae N. Snow, E.barriei N. Snow, E.bemangidiensis N. Snow, E.calciscopulorum N. Snow, E.delicatissima N. Snow, Callm. & Phillipson, E.echinulata N. Snow, E.gandhii N. Snow, E.hazonjia N. Snow, E.iantarensis N. Snow, E.malcomberi N. Snow, E.manomboensis N. Snow, E.obovatifolia N. Snow, E.ranomafana N. Snow & D. Turk, E.ravelonarivoi N. Snow & Callm., E.razakamalalae N. Snow & Callm., E.tiampoka N. Snow & Callm., et E.wilsoniana N. Snow, ainsi qu’une nouvelle combination, E.richardii (Blume) N. Snow, Callm. & Phillipson. Des descriptions détaillées, des informations sur la distribution et l’écologie, des cartes de distribution, des noms vernaculaires (quand ils sont connus), des images digitalisées des types, des comparaisons avec les espèces morphologiquemnet proches et une évaluation préliminaire du risque d’extinction selon l’UICN ainsi que des recommendations de conservation sont fournies. Des lectotypes sont désignés pour Eugeniahovarum H. Perrier, Eugenianompa H. Perrier et Eugeniascottii H. Perrier.
Biogeography, conservation, Eugenia , Eulemur fulvus , leaf cutter bees, Madagascar, Myrtaceae , new species, systematics, vernacular names
This paper is the fourth in a series devoted to clarifying the systematics of Malagasy Eugenia L. (
Type specimens of all nearly described species of Eugenia from Madagascar have been examined. All Malagasy specimens of Eugenia housed at MO (abbreviations following
Species descriptions include all information that can be interpreted with confidence and, given available information, are more or less in parallel. The species concept and criteria follow previous applications (
Hotlinks to holotypes on Tropicos® are included in lieu of illustrations of the new species. In addition, scanned images for all barcoded specimens from P are available online through the MNHN Vascular Plant Database (http://science.mnhn.fr/institution/mnhn/collection/p/item/search/form). Photos of living material are included when available. The conservation status of each species was assessed following IUCN Red List Categories and Criteria (
Frutex usque ad 9 m altus; ramuli complanati, glabri; folia usque ad 19 × 11.5 cm, obovata vel late elliptica, basi cordata vel rotundata, nervo medio supra sulcato, nervis secundariis infernis elevatis.
Madagascar. Prov. Antsiranana: Sud-Ouest d’Andapa, Réserve Spéciale d’Anjanaharibe-Sud. Ambodisatrana, aux environs des sommets, 14°32'45"S, 49°35'15"E, 809–1364 m, 25 May–3 June 1994, D. Ravelonarivo 206 + Raymond & Bekamisy (holotype: MO-6277713!; isotypes: KSP [KSP000041]!, P [P05097480]!, TAN).
Shrubs or trees 4–12 m tall; bark of main bole unknown. Vegetative and reproductive parts mostly glabrous except as noted. Branchlets laterally compressed, the terminal internode sometimes with a distal sulcus but becoming rounded, smooth, minutely and sparsely short-sericeous but soon glabrous, oil glands faint and moderately common but soon fading; emerging (youngest) internodes sometimes bearing 1–2 pairs of opposite to broadly ovate bracts 1.5–4.0 mm long. Leaves opposite, thinly coriaceous (dried material cracking with only moderate pressure), discolorous, matte above and below, venation brochidodromous. Axillary colleters absent. Petioles 3–4 mm long, slightly striate below, flattened above, epunctate. Leaf blades (15–)18.5–25 × (7–)13.5–14 cm, broadly elliptic to broadly obovate, base cordate and somewhat clasping (or rounded), surface and margin flat, apex obtuse to broadly acute, tip acuminate (or rarely retuse), secondary veins more or less straight, 11–15 per side protruding prominently below and arising at 25–55° angles and connected by slightly arching (but also prominent) inner marginal veins, tertiary veins relatively well-spaced but projecting only slightly (dried material); adaxial surface glabrous, oil glands faint (use magnification), sparse to common, more or less flush and darkish (dried), midvein sulcate; abaxial surface glabrous, midvein projecting prominently throughout and punctate (especially proximally) or epunctate, secondary veins projecting prominently, straight or only curving slightly towards margin, the secondaries connected at their ends by moderately arching connecting veins, tertiary veins projecting but less so than secondaries, intramarginal vein of same thickness as tertiaries, 1.5–5 mm from margin at midpoint of blade. Inflorescence (material scant) a monad; flowers cauliflorous, arising from short brachyblasts (< 3 mm long) above nodes on naked branches. Pedicels 8–32 mm long (possibly elongating after fertilization), 0.7–2.0 mm wide, somewhat compressed laterally, longitudinally striate, somewhat flexuous (bending with light touch), habit unknown, moderately glandular (glands faint), anthopodium and metaxyphylls absent. Bracteoles narrowly to broadly ovate, 1.5–2.5 × 0.5–1 mm, minutely and sparsely hairy dorsally and apically (hairs clear or whitish with some reddish). Hypanthium campanulate, 3.0–3.3 mm long, 3–4 mm wide at base of calyx lobes, densely but very shortly sericeous in proximal half (hairs reddish-brown) but glabrous distally; ovary apex glabrous. Calyx lobes 4 and often tearing irregularly towards hypanthium, up to 3.5 mm long × 4.5 mm broad (at base), irregularly hemispherical, glabrous on both faces apart from occasional minute hairs, evidently reflexed irregularly in anthesis. Petals 4 (material scant), 5.5–19.0 × ca. 10 mm, narrowly to broadly obovate, glabrous, epunctate, rose to violet. Staminal ring 3.5–4.0 mm in diameter (rounded or somewhat squarish), sparsely short hairy (hairs whitish). Stamens ca. 140 (estimated from scars on ring), multiseriate; filaments up to 10 mm long; anthers globular, ca. 0.8 mm long. Fruit 23–35 × 21–50 mm, depressed globular to globose, glabrous, base and apex rounded or apex crowned by calyx lobes, pinkish-reddish.
Flowering in February and March; fruiting March through November.
The species occurs in the Andapa Basin.
Known in northeast Madagascar in and around the Anjarahabe-Sud and Marojejy protected areas (Figure
Humid forests, riparian areas and near summit of Ambodisatrana; ca. 200–1540 m.
With an Extent of Occurrence (EOO) of 1586 km2, and Area of Occupancy (AOO) of 45 km2 and five subpopulations, two of which are situated within the protected area network (Anjanaharibe-Sud, Marojejy), Eugeniaandapae is assigned a preliminary risk of extinction of “Vulnerable” [VU B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (
The calyx lobes of this species tear towards the hypanthium during anthesis (see also E.lacerosepala N. Snow and E.ambanizanensis N. Snow). In this regard Eugeniaandapae is similar to taxa first described by
MADAGASCAR. Prov. Antsiranana: Vallée inférieure de l’Androranga, affluent de la Bemarivo (NE), aux environs d’Antongondriha, à la base du massif du Betsomanga, [14°15'30"S, 49°44'00"E], 200 m, 17-20 Nov. 1950, H. Humbert 24234 + R. Capuron (P [P05208578]); Quartier d’Ambodisatrana, SW d’Andapa, Réserve Spéciale d’Anjanaharibe-Sud, suivant la piste au bord de la rivière de Beamalona, vers la chaîne d’Anjanaharibe dans la réserve, 14°38'30"S, 49°25'30"E, 1235 m, 23 Mar. 1995, D. Ravelonarivo 694 + R. Rabesonina (MO). Andapa, Anjialavabe, Ankiakabe, 14°09'50"S, 49°22'47"E, 952 m, 11 Feb. 2007, R. Razakamalala 3234 + D. Ravelonarivo, C. Rakotovao, Jacky & José (G, K, MO-6175410, P [P04885355]). Prov. Mahajanga: Amparihy, Ruisseau d’Andasinanantsomanga, 14°55'38"S, 49°25'50"E, 1199 m, 23 Feb. 2008, P. Bernard 860 + J. Ramiadana & J. Jocelyn (MO-6432613).
Haec species a congeneris madagascariensibus pedicellis gracilibus delicatis, foliis tenuiter coriaceis, floribus minutis atque hypanthio dense villoso distinguitur.
Madagascar. Prov. Mahajanga: Fiv. Port Bergé, Marosely, Bongolava, 15°38'58"S, 47°35'03"E, 217 m, 17 Nov. 2004, R. Razakamalala 1735 + R. Ramananjanahary & A. Rabezafy (holotype: MO-4849778!).
Shrubs to 3 m tall; bark of main bole unknown. Vegetative and reproductive parts (where indicated) bearing a moderately dense, shortish indumentum, the individual trichomes dibrachiate or not, appressed to somewhat reflexed (appearing villous), frequently irregularly contorted, whitish or reddish. Branchlets laterally compressed but becoming rounded, smooth, moderately short villous (hairs mostly reflexed and not dibrachiate) becoming glabrous, oil glands common and prominent (after indumentum falls away). Leaves opposite, mostly occurring in 2–4 pairs along seasonal growth of branchlet, thinly coriaceous, venation brochidodromous (invisible to obscure), discolorous, somewhat glossy above but matte below. Axillary colleters absent. Petioles 1.7–2.5 mm long, broadly sulcate above, moderately hairy towards base adaxially in sulcus. Leaf blades 0.9–2.0 (–2.7) × 0.6–0.9 cm, narrowly elliptic or elliptic to narrowly obovate, base cuneate, surface flat to slightly and irregularly (but broadly) sinuous on drying, margin flat or drying slightly revolute, apex obtuse; abaxial surface glabrescent, oil glands common (use magnification) and drying brownish and slightly sunken, midvein flush and becoming imperceptible towards apex; abaxial surface sparsely glabrescent, oil glands relatively sparse to moderate and somewhat less prominent than adaxially, secondary veins few and barely perceptible, the secondaries connected at their ends by a slightly arching pseudo-intramarginal vein 0.3–0.8 mm from leaf margin (i.e., lacking an intramarginal vein distinct from pseudo-intramarginal vein). Inflorescence a monad; the base of the flowering branchlets each with (2–)4–6 flowers arising alternately, each flower subtended by a short, hairy and somewhat ovate to broadly triangular caducous bract. Pedicels (5–)10–15(–20) mm long, 0.3–0.5 mm wide, round in transverse section, stiff, ascending, sparsely hairy (especially near base) to nearly glabrous, moderately glandular throughout, anthopodium present or absent. Bracteoles 2, linear, 1.0–1.2 × 0.3–0.5 mm, sparsely hairy. Hypanthium cupulate 2.0–2.5 mm long, 1.4–1.8 mm wide at base of calyx lobes, densely short-hairy, oil glands absent or sparse (and obscured by hairs); ovary apex glabrous. Calyx lobes 4, 1.5–1.9 mm, broadly ovate to rounded, glabrous on both faces apart from sparse apical hairs (white or reddish), strongly reflexed in athesis. Petals 4 (material scant), ca. 2.5 mm × 2 mm, obovate to widely obovate, glabrous on both faces apart from sparse apical hairs (contorted irregularly), oil glands absent. Staminal region (i.e., lacking a well-defined staminal ring) 1.6–1.8 mm diameter in anthesis, sparsely hairy (trichomes simple); stamens 35–45; filaments 1.5–2.5 mm; anther sacs 0.5–0.7 mm long, globose, basifixed, eglandular. Style 2.5–2.8 mm, glabrous or sparsely hairy basally; stigma narrow and only slightly capitate. Fruit unknown.
The specific epithet honors Dr. Fred Barrie (b. 1948) of the Missouri Botanical Garden in recognition of his contributions to our knowledge of Eugenia and other genera of Mesoamerican Myrtaceae (e.g.,
Flowering confirmed only for the middle of November; fruiting likely late November through December.
With only one collection known from Central-western Madagascar collected in an unprotected and threathened dry forest, Eugeniabarriei is assigned a preliminary risk of extinction of “Critically Endangered” [CR A3c] following the IUCN Red List Categories and Criteria (
The type specimen of Eugeniabarriei initially was determined as E.tropophylla H. Perrier. The latter species and the varieties described by
Among taxa from southeastern Africa, Eugeniabarriei resembles some specimens of E.capensissubsp.gracilipes. In particular, the slender pedicels of a specimen from Malawi (Chapman 6570 [MO]) have a similar but less dense indumentum on the branchlets, pedicels and hypanthium. Other differences of the Chapman specimen include longer and more densely and prominently punctate leaves, and inflorescences that mostly arise from ramiflorous brachyblasts.
Eugeniabarriei also has gross morphological similarity to the widespread and relatively common west African species E.leonensis Engler & Brehmer (e.g., D.K. Harder 3372 et al. [MO] from Ghana), but it differs from that species by the generally glabrous aspect of the latter. Likewise, E.barriei somewhat resembles E.mufindiensis Verdc. by virtue of the indumentum of the branchlets, but the latter differs by its much more densely punctate leaves (above and below), the glabrous hypanthium, and a narrower and more deeply sulcate petiole (e.g., M.A. Mwangoka 5945 + H. Mgalla [MO]).
Haec speciesEugeniaewilsonianae N. Snow similis, sed ab ea inflorescentiis fasciculatis caulifloris distinguitur; etiam ad altitudines inferiores crescit.
MADAGASCAR. Prov. Toliara: Anosy, Taolagnaro, Iaboko, Antsotso Avaratra, 24°34'35"S, 47°12'28"E, 25 m, 13 Dec. 2007, R. Razakamalala 4056 + D. Rabehevitra, M. Maka, Roger & B. Mara (holotype: MO-6335452!; isotypes: P [P06490265]!, TAN).
Shrubs 3–4 m; bark of main bole more or less smooth but somewhat flaking, brown to gray. Foliage and reproductive parts glabrous except where noted. Branchlets terete, light brown (dried), smooth, oil glands sparse to moderate (but indistinct). Leaves mostly concentrated near branch tips, coriaceous, slightly discolorus, surfaces matte. Axillary colleters lacking. Petioles 3–4(–5) mm, slightly sulcate above, elgandular. Leaf blades 6–10 × 1.7–3.5 cm, narrowly elliptic to narrowly ovate, base rounded, apex acute to acuminate or a few falcate, margin flat, oil glands not seen (dried material); adaxial midvein broadly but shallowly sulcate, secondary veins indistinct to prominent (with magnification), intramarginal vein 0.5–1.8 mm from edge at midpoint of leaf blade. Inflorescences cauliflorous, arising from amorphous knobby protrusions (short shoots?) at or near nodes; flowers triads, monads, or fascicled (and possibly a few short botryoids); pedicels 1.5–6 mm, flexuous. Bracteoles 2 or absent, ca. 0.5 × 0.5 mm, broadly triangular to broadly rounded, ascending to divergent, stiff. Hypanthium 2.5–4.0 mm, cupuliform, oil glands common but small. Sepals 4, 2–3.0 × 3.5–4 mm, semi-circular to broadly elliptic or oblate, apex broadly rounded to obtuse, lobes very sparsely and minutely ciliate distally, moderately glandular; persistent and crowning (at least) young fruits. Petals 4 (material limited), 5–6 × 3–4 mm, widely elliptic to obovate, glabrous to sparsely and minute ciliate distally, glands sparse to moderate but faint (in dried material). Stamens 50–75, multiseriate; staminal disk short-hairy; ovary apex glabrous but distinctly punctate; filaments 2–5 mm; anthers 0.5–0.8 mm, elliptic, sub-basifixed, connective apex eglandular. Styles 6–7 mm; stigma narrow (scarcely if at all capitate). Fruits ca. 10–15 × 10–15 mm, (material possibly not fully mature) subglobose to globose, base sometimes sharply tapered, glabrous, dark bluish-black (dried).
The epithet is derived from the place name Bemangidy, a biologically rich region within the provisional Protected Area of Tsitongambarika.
Flowering commencing November and December; fruiting likely commencing by late December or early January (unconfirmed).
Known only from southeast Madagascar in Toliara Province, ca. 55 km northeast of Taolognaro, in Tsitongambarika (Fig.
Label information is sparse, but based on recent satellite imagery, evidently occurring in moderately to highly disturbed hilly areas that retain thin remnants of forest in ssome drainages and on steeper slopes, the type gathering located some 0.5 km east of much less disturbed primary forests that occur on steeper slopes, the paratype gathering in highly disturbed sites; elevation ca. 25–110 m.
With only three collections known, an AOO of 9 km2 and one subpopulation, which is situated outside the current protected area network, Eugeniabemangidiensis is assigned a preliminary risk of extinction of “Critically Endangered” [CR A3c+B1ab(iii)] following the IUCN Red List Categories and Criteria (
The leaf morphology of Eugeniabemangidiensis resembles E.wilsoniana (see discussion above). However, the short-pedicellate, fasciculate, and cauliflorous infloresences of E.bemangidiensis differ from the axillary, long-pedicellate flowers of E.wilsoniana. The type gathering at ca. 25 m elevation of E.bemangidiensis is approximately 645 km south of the most southerly collections of E.wilsoniana, the latter of which occur at middle elevations of ca. 980–1100 meters.
An indetermined fruiting specimen (Rakotovao 241 [MO-6437191]) from Near Ivohibe approximately 260 km north of the type gathering of E.bemangidiensis, which appears to have ramiflorous (and possibly cauliflorous) inflorescences, may be related, although it occurs at 1210 m, far above the known occurrences of E.bemangidiensis.
MADAGASCAR. Prov. Toliara: Iaboko, Antsotso, forêt Ivohibe, 24°33'52"S, 47°14'5"E, 26 Nov. 2005, R. Razakamalala 2326 + E. Ramisa & B. Mara (MO-6433204, P [P06490266]); ibid. loc., 24°34'16"S, 47°12'06"E, 8 Dec. 2007, R. Razakamalala 3799 + D. Rabehevitra, M. Maka, Roger & B. Mara (MO-6308460, P [P04827649]).
Haec species quoad faciem habitumqueEugeniaerichardii (Blume) N. Snow et al. et E. hazonjia N. Snow subsimilis, sed ab eis lamina foliari grosse sinuata, lobis calycinis roseoalbidis atque petalis subroseis distinguitur.
MADAGASCAR. Prov. Antsiranana: Mahavanona, Andranomanitra, 12°23'27"S, 49°20'01"E, 11 Dec. 2004, S. Rakotonandrasana 883 + R. Randrianaivo, A. Rakotondrafara, C. Christian & J. Be (holotype: MO-6245566!; isotypes: CNARP, KSP [KSP000016]!, P [P04885354]!, TAN).
Trees to 6 m; dbh 13 cm; bark of main bole unknown. Herbage glabrous except as noted. Branchlets rounded; oil glands sparse to common but indistinct; first epidermal layer soon flaking irregularly and drying grayish; later-forming bark smooth, light brown or light gray, frequently cracking horizontally. Leaves coriaceous, mostly concentrated near branch tips; opposite to disjunct opposite, discolorous, surfaces matte. Foliar colleters absent. Petioles 3–4 mm, terete or somewhat sulcate proximally when young, broadening and flattening somewhat distally at base of blade. Leaf blades (1.7–)2.0–4.3 × 1.8–3.0 cm, broadly elliptic (mostly) or a few ovate, base rounded to slightly cuneate, apex obtuse to somewhat acute, margins more or less flat, surface overall broadly sinuous; adaxial midvein sulcate lower 1/3–1/2 becoming flush distally, oil glands common to dense but becoming faint with maturity; abaxial surface oil glands sparse to common but faint at maturity and relatively small, secondary veins relatively indistinct, intramarginal vein 1–2 mm from margin of leaf at midpoint of blade, faint. Inflorescence terminal or axillary, of 1–3 monads per leaf axil. Pedicels 7–15 mm, relatively stiff. Bracteoles 2; ca. 0.5–1.0 mm, ovate, stiff, sparsely short-pubescent distally (trichomes ca. 0.1 mm long). Hypanthium 1.5–2.0 mm, cupuliform to obconic, densely glandular. Calyx lobes 4, 2–3 mm, elliptic to oblong, apex rounded, whitish to pink when fresh, oil glands common and prominently. Petals 4 (material scant), 6–11 × 5–7 mm, elliptic to widely elliptic, sparsely short-ciliate apically, pinkish, oil glands common. Stamens ca. 70–90, multiseriate; filaments 5–8 mm, whitish; anthers 0.6–0.8 mm, globose to elliptic, sub-basifixed, yellowish, the connective bearing a single apical oil gland; staminal disk squarrish, ca. 3 mm across, sparsely short-hairy. Styles 7–9 mm; stigma narrow. Fruits unknown.
Calci and scopulorum combine to form a genitive plural masculine noun, in reference to the locality at the base of a calcareous (limestone) cliff.
Flowering December; fruiting most likely late December through January (unconfirmed).
Known only from the type gathering in north-eastern Madagascar in Antsiranana, in the Montagne des Français limestone massif (Figure
Degraded dry forest at the foot of a cliff over calcareous substrates; ca. 410 m.
The species was said to be abundant at the type gathering. However, with only one collection known, an AOO of 9 km2 and one subpopulation, which is situated within a proposed protected area that currently holds a temporary protection status (Montagne des Français), Eugeniacalciscopulorum is assigned a preliminary risk of extinction of “Critically Endangered” [CR A3c+B1ab(iii)] following the IUCN Red List Categories and Criteria (
See Eugeniarichardii for similar species.
Haec species a congeneris madagascariensibus lamina foliari tenuiter coriacea irregulariter sinuosa atque pedicello gracili delicato distinguitur.
MADAGASCAR. Prov. Mahajanga: Tsaratanana massif, N of Mangindrano, along trail from camp at Mahatsabory to Be Pia, 14°07'40"S, 48°58'50"E, 2350 m, 20 Oct. 2001, P.P. Lowry II 5438 + R. Razakamalala & R. Lala (holotype: MO-6224858!; isotypes:P [P05208430]!, TAN).
Shrubs to treelets 2–4 m tall; bark of main bole smooth, light brownish or grayish. Plants glabrous or (where noted) sericeous (hairs dibrachiate, reddish). Branchlets laterally compressed, sparsely hairy upon emergence but becoming glabrous, oil glands faint, sparse or occasional and of uniform size, flush or only slightly protruding, smooth, light brown or gray. Leaves evenly distributed along branchlets, thinly coriaceous, venation brochidodromous, strongly discolorous, matte above and below. Axillary colleters if present then soon deciduous. Petioles 4.0–6.5 mm long, slightly to deeply sulcate above, sparsely glabrescent, eglandular. Leaf blades (18–)30–52 mm × (8–)18–25 mm, narrowly elliptic to elliptic (or occasionally broadly elliptic), base cuneate to strongly cuneate, margin flat to slightly revolute (dry), surface slightly irregularly wavy (sometimes including along midvein), apex acute to acuminate, tip obtuse to mostly acute or acuminate or slightly falcate; adaxial surface glabrous, oil glands faint, dense and of uniform size, midvein sulcate but becoming flush distally; abaxial surface glabrous, oil glands faint, sparse to occasional and of different sizes, vein connecting tips of secondaries near margin slightly arching arching only slightly, 0.5–0.9 mm from margin at midpoint of leaf blade. Inflorescence terminal, axillary or ramiflorous, consisting of short brachyblasts with 1–3 pairs of monads, often 2 or more brachyblasts per axil or point of insertion (on naked branches), or monads arising in axils. Pedicels 3–11 mm long, 0.3–0.4 mm thick, stiff to somewhat flexuous, ascending to sometimes reflexed, glabrous or very sparsely short-hairy, lightly striate, round, oil glands sparse and faint, anthopodia and metaxyphylls absent. Bracteoles 0.8–1.3 mm long, less than 0.5 broad (at base), very narrowly triangular to ovate, normally persisting in anthesis, sparsely hairy (or merely ciliate). Hypanthium obconic, (0.8–)1.2–1.3 mm long, 0.9–1.0 mm broad at apex beneath base of calyx lobes, sparsely hairy to glabrous, smooth, oil glands sparse to common and protruding. Calyx lobes 4, 1.3–1.9 mm long, broadly ovate to rounded, sparsely ciliate, sparsely glandular, greenish. Petals 4, 2.8–4.4 × 1.6–1.8 mm, obovate to broadly elliptic, glabrous or sparsely ciliate apically, white, oil glands sparse to common, of uniform size and somewhat protruding below. Staminal disk ca. 1.8 mm in diameter, sparsely hairy. Stamens 15–25; filaments 3–4 mm long; anther sacs ca. 0.5 mm long, globose to subcylindrical, dorsifixed, brown, connective with an apical gland. Style 4–5 mm long, glabrous; stigma narrow. Fruit 0.9–1.4 × 0.8–1.6 cm, subglobose to globose, texture smooth, rounded or tapered at base, glabrous, greenish but drying blackish, calyx lobes reflexed flat against apex.
A superlative of delicata in Latin, in reference to the thin, almost thread-like pedicels.
Rotramadinika (Rakoto 294).
Flowering in late October through at least early November; fruiting November.
Known from the northern mountains of Madagascar in and around the Manongarivo and Tsaratanana protected areas and in the Sorata region (Fig.
High elevation montane forests with bamboo; 1100–2350 m.
With an EOO of 3278 km2, an AOO of 36 km2 and four subpopulations, two of which are situated within the protected area network (Manongarivo, Tsaratanana), Eugeniadelicatissima is assigned a preliminary risk of extinction of “Vulnerable” [VU B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (
Eugeniadelicatissima is part of an apparent species group characterized by thinly coriaceous, sinuous leaf blades and slender, often elongate and delicate pedicels.
The thinly coriaceous, sinuous leaf blades of Eugeniadelicatissima are somewhat suggestive of E.echinulata, but the fruits of E.delicatissima are smooth (e.g. Randriambololomamonjy 270), unlike the highly irregular (warty to echinate) texture of of E.echinulata.
The slender pedicels are similar to those of Eugeniatropophylla H. Perrier, a highly heterogenous species that
An indetermined specimen of Eugenia from Mayotte in the Comoros (Barthelat 992 [G, K, MO]) also has slender pedicles, but they are much shorter than those of E.delicatissima. The leaf blades of the Barthelat specimen are longer and the bases rounded, unlike the cuneate bases of E.delicatissima. Taxa from eastern and southern Africa with delicate pedicels include E.capensissubsp.gracilipes F. White (e.g., P. van Wyk BSA 2844 [MO] from Zimbabwe and J.D. Chapman 6570 [MO] from Malawi). Another species from Africa with delicate pedicels that differs from E.delicatissima in various traits is E.congolensis De Willd & T. Durand (e.g., J. Madidi 642 et al. [MO] and F. Bujo 602 [MO] from Congo). None of these, however, are good morphological matches for E.delicatissima.
MADAGASCAR. Prov. Antsiranana: Massif du Manongarivo, [13°59'24"S, 48°22'12"E], c. 1600 m, 1909, Perrier de la Bâthie 6503 (P [P05258587]); Sommet de Beampoko, Ambohimirahavavy, 14°13'55"S, 49°08'23"E, 2137 m, 21 Nov. 2005, C. Rakotovao 2565 (G, MO-6174672); Vohemar, Andrafainkona, Ampsarahina, forêt de Maromaniry située à 5 km au N d’Ampiarahina, 13°38'49"S, 49°32'13"E, 1177 m, 7 Nov. 2007, O. Randriambololomamonjy 270 + R. Razakamalala & Jaowind (MO-6186659); Massif du Tsaratanana, crête (et ses abords) séparant les bassins du Sambirano et de la Mahavavy, entre l’Andohanisamborano et la cote 2362, 9-10 Nov. 1966, Service Forestier 27019 (P [P05097488]).
Haec species inter congeneros madagascarienses fructibus maturis valde echinulatis distinguitur.
Madagascar. Prov. Toamasina: zone d’occupation controlée d’Antenina, sur piste entre Antenina et Ankosy, à 2 km du village d’Antenina, 17°30'00’’–17°29'47"S, 48°46'19"E–48°45'35"E, 917 m, 1 Feb. 2002, L.M. Randrianjanaka 702 + N.M. Andrianjafy (holotype: MO-5786207!; isotypes: P, TAN).
Shrubs or trees 1.5–4 m tall; dbh 4–5 cm; bark of main bole brown, somewhat cracking. Foliage and flowers glabrous except as noted. Branchlets terete to laterally compressed, drying light brown to gray, smooth, oil glands occasional but barely visible. Leaves slightly discolorous, more or less evenly distributed, margins coarsely sinuous, venation brochidodromous, surfaces matte. Axillary colleters absent. Petioles 4–6 mm long, slightly suclate, eglandular. Leaf blades (3.0–)3.8–6.5 × (1.0–)1.6–3.5 cm, narrowly elliptic or narrowly ovate to elliptic, base cuneate, apex acute to acuminate; adaxial midvein sulcate proximally but becoming flush distally, oil glands dense but faint, sometimes protruding slightly on dried material; abaxial surface midvein prominently raised, oil glands common to dense and of uniform size but faint, sometimes slightly protruding (dried material). Inflorescence ramiflorous or axillary. Flowers solitary or clustered into short brachyblasts; brachyblasts 2–5 mm long, densely short-bracteolate, sparsely covered with twisted, whitish-maroon trichomes. Pedicles 5–7.5 mm, glabrous. Flower material scant. Hypanthium ca. 3 mm, coarsely large-pustullate (warty), light green or yellowish-green in bud. Calyx lobes 4 (rose-colored in bud), 3–4 mm, elliptic to broadly ovate or rounded, glabrous, mostly persisting and crowning fruit. Petals not seen; said to be pink (Antilahimena 8858). Stamens not seen; anthers said to be yellow with white filaments (Ahtilahimena 8858). Style not seen. Berry 1.3–1.8 × 1.0–1.5 cm, globose to subglobose, coarsely rugose-warty when fresh but drying echinulate (with with short sharp points), densely glandular when younger, green when immature, drying nearly black. Seeds in mature fruit 1; ca. 10–11 mm, globular, outer seed coat elgandular.
The specific epithet comes from the Latin echinulatus (having very small prickles) in reference to the prickly, dried, mature fruits that are unique for the genus in Madagascar.
Rotra (Antilahimena 6841).
Flowering December; fruiting January and February.
Known from the eastern escarpment of Madagascar in the Toamasina Province around Analamazaotra and Ambatovy and in the highlands at Anjozorobe in Antananarivo Province (Fig.
Dense humid evergreen forests; ca. 915–1350 m.
One collection indicated the species as being abundant in its area of collection (Randrianaivo 1463 et al.). However, with an EOO of 4,372 km2, an AOO of 72 km2 and three subpopulations, two of which are situated within the protected area network (Analamazaotra, Zahamena), Eugeniaechinulata is assigned a preliminary risk of extinction of “Vulnerable” [VU B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (
MADAGASCAR. Prov. Antananarivo: Reg. Analamanga, Anjozorobe, Ambongamarina, Ampamoa, à 10 km au Nord-Est d’Anjozorobe, prolongement N de la fôret d’Antsahabe, 18°23'19"S, 47°55'49"E, 1352 m, 15 Feb. 2007, R. Randrianaivo 1463 + L. Vary, François, Jacques & Rakoto (MO-6440448). Prov. Toamasina: Alaotra-Mangoro, Fkt: Menalamba, Ambatovy, 1168 m, 18°50'11"S, 48°18'44"E, 22 Jan. 2007, P. Antilahimena 5170 et al. (MO-5786207, KSP [KSP000006], P [P00730617]); ibid. loc., 18°52'25"S, 48°20'44"E, 979 m, 4 Oct. 2008, P. Antilahimena 6566 et al. (MO, P, TAN); ibid. loc., 18°52'28"S, 48°17'37"E, 1010 m, 10 Nov. 2008, P. Antilahimena 6841 + M. Ratolojanahary, B.A. Ratodimanana, M. Randrianarivony, M. Ratovomanana & F. Edmond (MO, P, TAN); ibid. loc., 18°52'30"S, 48°17'35"E, 970 m, 10 Nov. 2008, P. Antilahimena et al. 6854 (MO, P, TAN); ibid. loc., 18°55'22"S, 48°25'39"E, 1013 m, 21 Dec. 2013, Antilahimena 8858 + Rabarison, Honoré, Félix, Randrimitantsoa Jean & V. Razafindrahaja (MO, P, TAN); Phelps Dodge project site, ca. 15 air-km NE of Moramanga, ca. 11 km E of Antanambao. Ambatovy, SE valley (Ambohimanga), 18°51'34”S, 48°18'25”E, 1050 m, P.J. Rakotomalaza 1208a (MO, P [P00730620]); Analamazaotra, Ambatovy, 18°56'10”S, 48°25'33”E, 11 Dec. 2013, Ramahenina 276 et al. (MO); Ambatovy, forêt de Savihara, 18°52'06"S, 48°16'43"E, 22 Feb. 2010, D. Razafimelison 10 et al. (P [P00730618]); Alaotra-Mangoro, Moramanga, Andasibe, Réserve Spécial d’Analamazaotra, Circuit Indri 1, 18°56'20"S, 48°25'09"E, 947 m, 14 Dec. 2013, H.M.J. Rasoazanany 541 + H. Razafindraibe, B. Ramandimbisoa & Rakotondravelo (MO, P, TAN).
Eugeniaechinulata is unknown in flower apart from one remnant hypanthium on the holotype. However, the coarsely sinuate leaf blades and acuminate apices, coupled with the warty mature fruits, are diagnostic and unknown in combination elsewhere among Malagasy congeners. Whereas E.muscicola H. Perrier has narrowly elliptic leaves, its leaf margins are more or less planar. A number of similar sterile collections from this area may be this taxon.
The texture of the fruit is coarsely rugulose-warty (see link above to image of living material), but the warty protrusions collapse to a large extent and upon drying and then assume a sharper, echinulate texture.
Haec species a congeneris madagascariensibus lamina foliari tenuiter coriacea elliptica usque obovata dense sed inconspicue glandulosa apice acuta acuminatave distinguitur.
MADAGASCAR. Prov. Toamasina: Nosy Mangabe, a 520 ha island in the Bay of Antongil, 5 km from Maroantsetra, 15°30'S, 49°46'E, 0–330 m, 9 Jan. 1989, G.E. Schatz 2482 + J.S. Miller (holotype: MO-4805069!; isotypes: MO-3708875!, P [P05260197]!, TAN).
Eugeniaarthroopodavar.ambalavensis H. Perrier in Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 4(2): 177. 1953. Nom. inval. (lacking Latin diagnosis, ICN, Art. 39.1,
Shrubs to trees, 3–12 m. Trunk 3–8 cm dbh; outer bark of main bole reddish, papery to flakey and peeling. Foliage and reproductive parts glabrous except where noted. Branchlets terete to laterally compressed, reddish when fresh but drying brownish or grayish, glabrous but prominently glandular, the epidermis smooth but soon peeling (dried specimens) and often thin and narrow grayish strips or even threads. Leaves opposite, medium green (fresh), slightly to strongly discolorous, venation brochidodromous, thinly coriaceous (easily cracking when dried), surfaces matte. Axillary colleters lacking. Petioles 2.8–6 mm, slightly sulcate distally. Leaf blades (5.5–)9–16 × 2.5–6.0 cm, elliptic to occasionally obovate, base cuneate, apex acute to mostly acuminate or caudate, margins flat; adaxial surface glabrous, densely punctate (glands small and best seen with magnification); abaxial surface similar, lateral veins indistinct to prominent; intramarginal vein 1–3.5 mm from margin at midpoint of blade. Inflorescence terminal, axillary, or ramiflorous, of soliltary monads or in fascicles of 2–3 flowers. Pedicels 9–29 mm long × 0.8–1.2 mm thick, stiff, moderately glandular, ascending to erect, green. Bracteoles narrowly ovate to broadly rounded, 0.5–1.3 × 1.0–1.2 mm, stiff but thin, glabrous to minutely and sparsely sericeous on margin, often persistent in fruit. Hypanthium 4–5 mm long, cupuliform, densely glandular (some glands larger proximally). Calyx lobes 4, 6–9 × up to 11 mm, broadly elliptic to rounded, cream-colored (fresh), reflexed in flower, persistent and often crowning the fruit. Petals 4, 11–14 × up to 12 mm, widely elliptic to widely obovate (narrowly greatly at base), sparsely and minute ciliate marginally, densely glandular (glands of variable size but typically large proximally). Stamens 250–300, multiseriate, exserted; staminal disk short-hairy (trichomes slightly ferrugineous); filaments 8–15 mm, white; anthers globose, 0.3–0.5 mm, brownish, bearing a single large apical gland. Ovary apex glabrous but somewhat glandular. Style 8–10 mm; stigma narrow (scarcely if at all swollen). Berries 2.5–4.0 × 3.0–4.0 cm, subglobose to globose, purplish or violet (drying nearly black). Locules 2; placentation axile. Seeds up to 20 mm long and wide (available material possibly not fully mature), up to 6 per fruit, evidently often somewhat flattened, cotyledons and hypocotyl barely if at all differentiated.
The species honors Dr. Kanchi N. Gandhi (b. 1948) of Harvard University Herbaria, North American editor of the International Plants Names Index, and an expert of botanical nomenclature who has assisted colleagues with the proper usage of botanical Latin and interpretation of the Code over many years.
Gavoala (Ravelonarivo 103); gavoalabe.
The name “guaavaala” appears on one specimen, but native speakers inform us that it is a nonsensical name based on a combination of the common English name guava (for members of Psidium, a Neotropical genus) and ala, meaning locally “of the forest”.
Flowering January through June; fruiting March through November.
Endemic to northeast Madagascar, recorded from the Island of Nosy Mangabe, the Masoala Peninsula, and in and around the Anjaniharibe Reserve (Figure
Humid and typically dense forests; 0–1424 meters.
With an “Extent Of Occurrence” (EOO) of 9,504 km2, an “Area Of Occupancy” (AOO) of 45 km2 and five subpopulations, three of which are situated within the protected area network (Anjanaharibe-Sud, Marojejy and Nosy Mangabe), Eugeniagandhii is assigned a preliminary status of “Least Concern” [LC] following the IUCN Red List Categories and Criteria (
Perrier de la Bâthie’s E.arthroopodavar.ambalavensis H. Perrier was based on a single collection from the lowlands just to the south of Marojejy Reserve, but we have little doubt that it is conspecific with the material we include in E.gandhii, which ranges from the nearby Anjaniharibe Reserve to the southwest of Marojejy to the Masoala peninsula and Nosy Mangabe farther south.
The thinly coriaceous leaf blades of Eugeniagandhii are suggestive of Eugeniavatomandrensis H. Perrier, which thus far is known as a coastal species from near Vatomandry north towards Soanierana (
MADAGASCAR. Prov. Antsiranana: Vallee de la Lokoho (Nord-est), près d’Amalavanonio, 9-10 January 1949, H. Humbert + G. Cours 22804 (P [P00118089, P00118090]!); Réserve Spéciale d’Anjanaharibe-Sud, aux environs du sommet, 1161–1424m, 14°46'15"S, 49°28'00"E, 21 Mar.–7 Apr. 1994, D. Ravelonarivo 103 et al. (KSP [KSP000010, KSP000011], MO-6135429, P [P04884879]); Réserve Spéciale Anjanaharibe-Sud, Ambodisatrana, aux environs des sommets, 14°32'45"S, 49°35'15"E, 809–1364 m, 25 May–3 Jun. 1994, D. Ravelonarivo et al. 174 (KSP [KSP000013]); Réserve Spéciale d’Anjanaharibe-Sud, village d’Andranotsarabe, suivant la route Nationale d’Andapa-Bealanana de la piste vers à l’W, Ambatomainty, Camp No. 2, 14°44'42"S, 49°27'42"E, 1185–1335 m, 3 Nov. 1994, D. Ravelonarivo 516 + P. Rabesonina (KSP [KSP000012], P[P04885352]); Réserve Spéciale Anjanaharibe-Sud, suivant la piste pour Ambalaheva, haute rivière d’Andramonta, 14°36'40"S, 49°24'12"E, 628–1879 m, 22 Feb. 1996, D. Ravelonarivo et al. 929 (KSP [KSP000015, KSP000014], MO-6135429, P [P05208458]). Toamasina: Maroantsera Tampolo, Péninsule Masoala, env. 1 km NW du camp (site du posé du radeau), 15°42'77"S, 48°58'25"E, 100–200 m, 1 Nov. 2001, O. Poncy 1548 + S. Rapanarivo (K, P [P00373064], TEF); Nosy Mangabe, 5 km from Maroantsetra in the Bay of Antongil, 15°30'S, 49°46'E: all sterile specimens collected by G.E. Schatz + A. Gentry from 13–23 April 1988: 2010 (MO-3599534), 2020 (MO-3599524), 2100 (MO-3598203), 2137 (MO-3598167; P [P05156045]), 2197 (MO-3596402), 2210 (MO-3596403), 2261 (MO-3597989), 2294 (MO-3597957), 2296 (MO-3597954).
Haec species a congeneris madagascariensibus ramificationis ordinatione V-formi, lamina foliari elliptica vel late elliptica basi cuneata apice rotundata usque late acuta, costa adaxialiter plana atque bracteolis cupulatis base latis distinguitur.
MADAGASCAR. Prov. Antsiranana: Fiv. Vohemar. Fir: Nosibe. Fkt: Anjiabe, 13°04'42"S, 49°54'13"E, 25 m, 2 Nov. 2002, J. Rabenantoandro 1089 + R. Rabevohitra, G. McPherson, H. Ranarivelo, C. Claude & M. Sola (holotype: MO-6038336!, isotype: P [P05208261]!).
Shrubs or small trees 2–5 m; dbh 5 cm; bark of main bole unknown. Herbage glabrous except as noted. Branchlets strongly laterally compressed on emergence but becoming terete, drying light brown to light gray, sometimes slightly striate, oil glands sparse. Leaves mostly evenly distributed, coriaceous, venation brochidodromous, discolorous, surfaces matte and sometimes slightly undulate. Axillary colleters absent. Petioles 1–3 mm, flattened adaxially, sometimes slightly sulcate basally. Leaf blades (2.0–)2.5–4.0 × (1.0–)1.5–3.3 cm, elliptic to broadly elliptic, base cuneate, apex rounded or obtuse to (occasionally) broadly acute, margins usually slightly revolute; adaxial surface oil glands common to dense but faint, midvein flush; abaxial surface midvein flush except proximally, oil glands common to dense (sometimes faint), secondary veins indistinct, intramarginal vein 1–2 mm from edge at midpoint of blade. Inflorescence of terminal or axillary monads, these solitary, paired, or (especially at apex) fasciculate. Pedicels 8–18 mm, ascending, stiff. Bracteoles 2, 1.0–1.5 mm, broadly ovate and cuplike (sometimes keeled), rigid, minutely and sparsely ciliate proximally (use magnification). Hypanthium 2.5–3.0 mm, cupuliform, oil glands dense. Calyx lobes 4, 3–5 mm, broadly rounded, apex obtuse, sparsely short ciliate, prominently glandular, greenish. Petals 4, 8–11 × 6–7 mm, obovate to broadly obovate, glabrous, pink. Stamens >150, multiseriate, staminal disk densely short-hairy; filaments 5–9 mm; anthers 0.6–0.9 mm, ellipsoid, basifixed, connectives with a single (but faint) apical gland. Styles up to 8 mm; stigma narrow and scarcely if at all capitate. Fruits unknown.
Named after the local vernacular name hazonjia, which is recorded on two specimen labels.
Flowering September through December; fruiting unknown.
Known from Antsiranana Province in north-eastern Madagascar approximately 30 km northwest of Vohemar and in the Daraina region (Fig.
From littoral forest on sand and dry deciduous forest; 15–1015 m.
With an EOO of 39 km2, an AOO of 27 km2 and two subpopulations, one of which is located within in the temporarily protected Loky-Manambato area, Eugeniahazonjia is assigned a preliminary risk of extinction of “Endangered” [EN B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (
Eugeniahazonjia somewhat resembles E.urschiana H. Perrier, but the latter has longer petioles and shorter pedicels. It resembles even more closely a specimen (Rabevohitra 5044 et al.) form the eastern side of Ile Sainte Marie approximately 420 km south, but which has a moderately dense indumentum on the hypanthium and pedicels of dibrachiate, ferrugineous trichomes, and a staminal disk with significantly longer and more erect trichomes.
MADAGASCAR. Prov. Antsiranana: Vohemar, Nosibe, Fkt: Anjiabe, Anaborano, Fôret littorale d’Analabe, Mosorolava, 13°05'41"S, 49°54'21"E, 24 Sept. 2004, H. Manjakahery 17 (MO-6308382); Vohemar, Nosy-Be, Anjiabe, forêt de Lac Sahaka, Anaborano, 13°04'49"S, 49°54'14"E, 9 Dec. 2007, R. Randrianaivo 1505 + S. Randrianasolo, R. Rakotondrajaona, V. Beninjara, C. Claude, Cyprien & M. Sola (G, MO-6250496); Loky Manambato, Daraina, forêt d’Antsahabe, 13°12'59”S, 49°32'47”E, 1015 m, 1 Dec. 2004, Gautier 4799 (G, P [P05094974]); ibid. loc., 900 m, 6. Dec. 2004, L. Gautier 4842 + L. Nusbaumer (G [G00019226], P [P04827946], TEF n.v.).
Haec species a congeneris madagascariensibus foliis abaxialiter secus costae partem proximalem glandulis oleiferis manifestis ornatis, fructu permagno atque seminibus testa externa profunde sulcata convolutaque distinguitur.
MADAGASCAR. Prov. Fianarantsoa: Andringitra, Camp I, ca. 45 km S of Ambalavao, east bank of Iantara River, along Ambalamanenjana-Ambatomboay trail, edge of Andringitra Reserve, 22°13'20"S, 47°01'29"E, 20 m, 15–21 Nov. 1993, B. Lewis 776 + J. Raharimalala, G. Rahajasoa, M. Randriambololona & J. McDonagh (holotype: MO-6224855!]; isotypes: KSP [KSP000052]!, P, TAN).
Shrubs or trees, 3–9 m tall; dbh 6–10 cm; bark of main bole unknown. Vegetative and reproductive parts glabrous or (as noted) moderately to densely sericeous (trichomes dibrachiate and ferrugineous). Branchlets somewhat compressed laterally but becoming terete, becoming irregularly wrinkled in age, oil glands common, prominent and somewhat protruding initially but fading with age, grayish brown. Leaves concentrated mostly at branchlet tips, only 1–2 leaves produced during seasonal growth, the pairs ca. 1–1.5 cm distant; blades coriaceous, venation brochidodromous, surface and margin flat or slightly sinuous, somewhat discolorous, matte above and below. Axillary colleters absent. Petioles 6.5–10.5 mm long, rounded above, eglandular, shedding an outer (grayish or light brownish) epidermal layer with age. Leaf blades (9)11.5–22 × (3.4)5–8.7 cm, elliptic (or infrequently narrowly elliptic, narrowly obovate or obovate), base cuneate, apex obtuse; adaxial surface oil gland dense, easily seen with magnification, midvein sulcate but becoming flush apically, secondary veins protruding slightly; abaxial surface sparsely to moderately sericeous (hairs dibrachiate, ferrugineous) but becoming glabrous, oil glands common (notably less dense than adaxial), midvein protruding (and with punctate glands, especially proximally), punctate, striate, secondary veins somewhat protruding (moreso than on adaxial surface), intramarginal vein connecting tips of secondaries arching moderately, 1.5–3.3 mm from margin at laminar midpoint. Inflorescence of terminal, axillary or ramiflorous monads, solitary to paired or occasionally in few-flowered fascicles. Pedicels 8–21 mm long (in fruit), 1.8–3.5 mm wide, round in transverse section, rigidly stiff, ascending, sparsely hairy (use high magnification) or evidently glabrous, eglandular, somewhat striate, the epidermis reddish and eventually cracking in small irregular flakes, anthopodia and metaxyphylls lacking. Bracteoles (only one seen) evidently broadly triangular, ca. 2 mm long. Calyx lobes 4, 5.5–8.8 mm long, broadly rounded, apex obtuse, hairy towards based but otherwise more or less glabrous adaxially, moderately to densely hairy abaxially, bifacially glandular, persisting in fruit and tearing somewhat beneath the based as fruit matures, green to cream-colored or magenta above. Petals violet (Ratolojanahary 88). Staminal disk (in mature fruit) 7–16 mm wide, glabrous. Stamens (estimated from basal scars) 150+. Style (material scant) ca. 9 mm long, glabrous; stigma narrow (scarcely if at all capitate). Ovary 2-locular; placentation axile; ovules up to 5 per locule and radiating from central position. Berries 3.2–4.1 × (1.6)2.4–5.0 cm, irregularly globose and often with 3-numerous irregular lobes (from maturing seeds), densely shortly sericeous, pale green immature becoming grayish-brown when fresh but drying light rusty-brown by virtue of indumentum, outer wall embedded with numerous straight, more or less evenly spaced vertical veins extending between base and apex. Seeds 3–15 per fruit (up to 8 per locule), 17–25 mm long, up to ca. 15 mm thick (shrinking considerably away from seed coat during drying), radiating out from placenta (narrowed towards attachment point); seed coat thickly membranous to somewhat leathery, eglandular, separating easily from embryo, surface drying irregularly bullate, light reddish brown; cotyledons evidently completely fused.
The specific epithet is a Latinization of the Iantara River, location of the type gathering.
Rotramena vaventiravina (Kotozafy 922); rotra (Antilahimena 6914).
Flowering March through November; fruiting November and December.
Known from the eastern escarpment of Madagascar in Finarantsoa and Toamasina provinces around and in the Andringitra, Ranomafana and Analamazaotra Special Reserve (Fig.
Evidently along or near rivers in humid forest, sometimes in disturbed areas; from 770–1210 m. The type collection indicates the plant, presumably the large fruits, is used by the Common brown lemur, Eulemurfulvus.
With an EOO of 4,892 km2, an AOO of 81 km2 and three subpopulations, all of which are situated within the protected area network (Andringitra, Ranomafana and Analamazaotra), Eugeniaiantarensis is assigned a preliminary status of “Least Concern” [LC] following the IUCN Red List Categories and Criteria (
The flowering and early fruiting material (Toamasina Province) and mature fruiting material (Finarantsoa) are disjunct and the flowering material is somewhat incomplete (no petals), but apart from slightly shorter leaves in Toamasina the specimens match well.
Mature fruits are among the largest yet known among Malagasy Eugenia, and the large number of seeds per fruit (up to 13) is considerably greater than that commonly seen for Eugenia worldwide or in Madagascar. The testa in dried material becomes highly furrowed-convoluted to resemble the surface of a human brain, likely due only in part to shrinkage from dessication. A similar testal morphology was recently described in Eugeniaalletiana Baider & Florens from Mauritius (
MADAGASCAR. Prov. Fianarantsoa: Parc National de Ranomafana, 21°15'S, 47°25'E, 1045 m, 5 Dec. 2007, J. Renoult Ma07-225 (P [P05208539]); Parc National de Ranomafana, Parcelle II d’Ambodiamontana, Parcelle II de Talatakely, 21°16'S, 47°26'E, 950 m, 14 Mar. 1995, A. Kotozafy 922 (KSP [KSP000050, KSP000051], MO-04940615, P [P05208460]); Andringitra RN. Ivohibe, à l’W d’Ambarongy, au NE du campement no. 3, sur la piste à l’E du campement no. 3 vers le campement no. 4, 22°13'22"S, 46°58'18"E, 1210–1625 m, 8–11 Dec. 1994, C. Rakotovao 287 (MO-6224856, MO-5474705 [carpo.], P [P04884880]). Prov. Toamasina: Ampitambe, Ambatovy, Sahaviara forest, 18°52'19"S, 48°16'43"E, 1035 m, 9 Nov. 2005, P. Antilahimena 4143 & F. Edmond (MO, P [P05094984]); Ampitambe, Ambatovy, 18°51'39"S, 48°16'27"E, 943 m, 15 Dec. 2005, P. Antilahimena 4384 (MO, P [P00730616]); Ambatovy, Antaralava Besalampy forest, 18°53'15"S, 48°16'09"E, 950 m, 15 Nov. 2008, Antilahimena 6905 (MO-6447003, P [P00730615]); Ambatovy, Sahaevo, 18°50'26"S, 48°16'3"E, 982 m, 23 Nov. 2008, P. Antilahimena 6914 + B.A. Ratodimanana, D. Ravelonarivo, E. Félix & M. Ratovomanana (MO-6447031); Fkt: Ambohibolakely, forêt d’Amparihy, corridor Forestier Analamay Mantadia, 18°47'28"S, 48°22'46"E, 1001 m, 23 Apr. 2012, C. Rakotovao 5783 (P [P00730613]); Ampitambe, Ambatovy, forêt d’Ampadidifanantsy, 18°51'46"S, 48°16'32"E, 975 m, 27 Oct. 2008, M. Ratolojanahary 88 (MO-6453743, P [P04680790]); Forêt d’Analamazaotra, [18.56'S, 48.26'E], 3 Dec. 1934, Ursch 20 (P [P05208538]).
Haec species a congeneris madagascariensibus folia grandia habentibus foliis late ellipticis ad basem rotundatis plerumque plus quam 15 cm longis distinguitur.
MADAGASCAR. Prov. Toliara: NW of Tolagnaro, Reserve Naturelle Integrale #11 (Andohahela) parcelle I, NW of Eminiminy, beside River Itrotroky, 24°38'S, 46°46'E, 500–1000 m, 6–13 Feb. 1993, S. Malcomber 2117 + H. van der Werff, C. Hemingway, M. van Bergen, S. Rapanarivo, P.J. Rakotomalaza, O. Andrianantoanina & B. Randriamampionona (holotype: MO-6277745!; isotypes: KSP [KSP000046]!, P [P05131986]!, TAN).
Trees to 6 meters tall; bark of main bole unknown. Indumentum (where noted) of vegetative and reproductive material sericeous (hairs dibrachiate and typically reddish or brownish). Emerging (youngest) branchlets laterally compressed, sparsely hairy, glandular, color uncertain; older branchlets becoming terete, soon glabrous, mostly eglandular, becoming brownish-gray, smooth. Leaves evenly distributed along branchlets (nodes greater than 9 cm apart), coriaceous, venation brochidodromous, strongly discolorous, matte above and below. Axillary colleters present (type A;
The new species honors Dr. Simon Malcomber (b. 1967), who collected the type material and nearly two thousand specimens from Madagascar in the early 1990s.
Flowering in early February; fruiting probably late February through at least mid-March.
Known only from the type gathering in Toliara in parcelle I of the Reserve Naturelle Integrale #11 northwest of Taolognaro, in south-western Madagascar. This region is near the southern terminus of the humid forests that occur east of the main escarpment that run latitudinally along much of the length of Madagascar (Fig.
The species was collected in a riverine habitat in a rainforest. Some of the (undistributed) type material is heavily clothed in epiphytic mosses, Hymenophyllaceae, and lichens.
Eugeniamalcomberi occurs in the Andohahela protected area (parcel I). No indication was provided on the label about its relative abundance at the time it was collected. Google™ Earth imagery (April 2013) shows a more or less continuous band of primary forest extending ca. 42 km south and ca. 135 km northeast, and ca. 10 km wide (east to west) at comparable elevations from the type locality. Considering its occurence in a nature reserve in fairly rugged terrain with considerable topographic relief, but also because it is known only from a single subpopulation with an AOO of 9 km2, Eugeniamalcomberi is assigned a preliminary risk of extinction of “Vulnerable” [VU D2] following the IUCN Red List Categories and Criteria (
The petals during anthesis are quite large relative to the size of the flower buds, suggesting that they enlarge rapidly during early anthesis.
Haec species a congeneris madagascariensibus petiolo longo, lamina foliari grandi elliptica, pedicello crasso atque fructu grandi globulari laevi distinguitur.
MADAGASCAR. Prov. Fianarantsoa: Région Atsimo-Atsinanana, Dist. Farafangana, comm. Ankarana, Réserve Spéciale de Manombo, parcelle I, forêt d’Anaviavy, 23°00'S, 47°44'E, 12 m, 17 Sep. 2005, R. Razakamala 2136 (holotype: MO-2590168! isotypes: KSP [KSP000043]!, P [P04776395]!, TAN).
Trees to 8 m tall. Foliage glabrous and without evident oil glands except as noted. Branchlets round to somewhat laterally compressed, drying brown to light brown; bark smooth but cracking slightly longitudinally. Leaves stiffly coriaceous, opposite or sometimes three per node, evenly distributed along branchlets, discolorous, somewhat glossy adaxially, matte abaxially. Axillary colleters absent. Petioles 25–35 mm, terete or slightly sulcate. Leaves 16–18 × 6.5–10.5 cm, elliptic, base slightly cuneate to mostly rounded, apex obtuse, surface and margin more or less flat; adaxial surface with impressed midvein in lower ½ to 2/3, becoming flush distally; abaxial surface with dense but small oil glands (faint, use high magnification); secondary veins indistinct, arising at 40–50° angle but mostly too faint to count with confidence; intramarginal vein indistinct to barely visible, 1.5–2.5 mm from margin at midpoint of blade. Inflorescence up to 7 cm, structure uncertain but evidently of triads or botryoids, these solitary in leaf axils. Pedicels 4–14 mm, stiff and thick. Flowers mostly unknown. Calyx lobes (from dried fruit) 4, 3–5 mm, broadly rounded (much broader than long in fruit), persistent in fruit. Fruit 25–30 × 25–30 mm, globose.
The specific epithet is derived from the Réserve Spéciale de Manombo, the only known occurrence of this prominent species.
Fruiting mid September; likely flowering by August (unconfirmed).
Known only from the type gathering in south-eastern Madagascar in Fianarantsoa from Réserve Spéciale de Manombo, near the coast (Fig.
Dense, humid, low-altitude forest over lateritic soils.
Eugeniamanomboensis occurs in the Réserve Spéciale of Manombo. Considering its occurrence in a natural reserve, but also because it is known only from single subpopulation (AOO of 9 km2), Eugeniamalcomberi is assigned a preliminary risk of extinction of “Vulnerable” [VU D2] following the IUCN Red List Categories and Criteria (
Large-leaved but indetermined speicmens of Malagasy Eugenia with large fruits have not been matched closely with flowering material in some cases. However, given that none of the larger-leaved species have petioles as thick or as long as Eugeniamanomboensis, no others are known from near the type locality, and no large-leaved species present globular fruits of this size, it appears that this species has not been named previously.
It seems likely that the large fruits of this tree are consumed by local widelife.
Haec species a congeneris madagascariensibus foliis anguste obovatis atque inflorescentiis glabris saepe ramifloris distinguitur.
MADAGASCAR. Prov. Toliara: Fokontany: Sainte Luce, 24°46'15"S, 47°10'15"E, 5 m, 5 Nov. 2003, J. Rabenantoandro 1592 + A. Monja (holotype: MO-6038332!; isotypes: P [P04885376]!, TAN).
Trees 5–14 m; dbh 14–28 cm; bark of main bole unknown. Indumentum (where indicated) of short, ferrugineous, dibrachiate hairs that become increasingly white or grayish. Branchlets rounded laterally compressed, densely short-hairy becoming glabrescent; light brown to light gray (dried). Leaves concentrated near branch tips, coriaceous, surfaces matte. Axillary colleters absent. Petioles 4–8 mm, terete, pubescent when emerging. Leaf blades (3.0–)4.5–6.5 × 1.5–2.5 cm, narrowly obovate (or infrequently ovate); base narrowly cunate; apex obtuse or somewhat acute; margins flat but drying somewhat revolute; adaxial surface glabrous or moderately hairy (especially proximally), oil glands dense (but small; use magnification) on emergence but becoming less visible, midvein slightly sulcate to flush; abaxial surface moderately hairy, oil glands dense but faint; secondary veins 8–15, alternating along midvein, faint; intramarginal vein not visibile. Inflorescences ramiflorous, less than 1.5 cm long, solitary or more commonly fasciculate, of monads, triads, or short botryoids. Pedicels 1–5 mm, stiff, ascending, sparsely hairy to glabrous. Bracteoles 2, ovate, 0.5-1.0 × < 0.5 mm, rigid, sparsely hairy on margins. Hypanthium 2–3 mm, cuplate to obconic, glabrous to densely ferrugineous-hairy, densely glandular (glands relatively large). Calyx lobes 4, 1.5–2 mm, two each relatively large and small, broadly rounded (much broader than long), apex obtuse, sparsely minutely hairy marginally near apex, otherwise glabrous, greenish to cream colored. Petals 5–8 × 5–6 mm, elliptic to ovate, glabrous, whitish to pink or violet, oil glands sparse but easily visible. Stamens ca. 30–50, multiseriate; staminal disk short-hairy, ca. 1.5 mm diameter; filaments 3–5 mm, white; anthers ca. 0.5 mm, globular, basifixed, eglandular, light yellow. Styles 5–7 mm, glabrous; stigma narrow. Berries not seen, but labels indicate pale green (probably immature).
In reference to the shape of the leaf blades.
Ropsay (Rabenantoandro 366).
Flowering September, November, and January; fruiting August through December.
East-central Madagascar in Toamasina Province and south-eastern in Toliara Province from Mandena to Sainte Luce (Fig.
Littoral forest over sand; elevation ca. 5–53 m.
With an EOO of 8,333 km2, an AOO of 63 km2 and four subpopulations, none of which are situated within the protected network, Eugeniaobovatifolia is assigned a preliminary risk of extinction of “Endangered” [EN 2ab(i, iii, iv)] following the IUCN Red List Categories and Criteria (
Eugeniaobovatifolia has a general resemblance to E.arenicola H. Perrier, E.hazompasika H. Perrier, and E.cloiselii H. Perrier given its ramiflorous, more or less fasciculate inflorescences. The four species also grow in relative proximity in southeastern and eastern Madagascar, often in sandy littoral forests, and may form a species complex that ranges approximately 1300 km between Taolognaro and Sainte Luce along the coast and somewhat inland to Vatomandry.
Eugeniaobovatifolia differs from E.arenicola by its more or less glabrous leaves, whereas those of the latter are densely ferrugineous-hairy upon emergence and retain much of the indumentum on the abaxial surface. Eugeniaobovatifolia differs each from E.cloiselii and E.hazompasika by virtue of the glabrous (or nearly so) inflorescences, which contrast with the hairy inflorescences of the latter two.
Several indetermined specimens cannot be confidently placed yet given variation in leaf morphology and indumentum (e.g., Razanatsima 31 + Ranaivojaona; Ranaivojaona 1177 + Razanatsima; Razanatsima 711 + Céléstin; Rakotovao 3755 et al. [all at MO]). However, including all of these taxa and specimens into one highly heterogenous species likely would equate to the inappropriate lumping of several independently evolving lineages.
MADAGASCAR. Prov. Toamasina: Fokontany: Andravokoditra, 18°34'53"–18°34'20"S, 49°14'42"–49°14'54"E, 7 Jan. 2003, R. Ludovic 228 & F. Rakotoarivony (MO-6038324); Hotel Pangalan on Lake Ampitabe, west of Andranokoditra, 18°35'S, 49°14'E, S.K. Pell 607 + J.D. Mitchell & A. Randrianasolo (MO-5887243); Fokontany: Andranonkoditra Vohibola du côté de l’hôtel Pangalane, 18°35'32"S, 49°14'02"E, 11 Feb. 2003, J. Rabenantoandro 1248 + R. Rabevohitra, P. Lowry, R.Razakamalala & S. Lowry (MO-6038325); Ambinaninony, Andranokoditra, Akinin’ny nofy, 18°34'12"S, 49°14'18"E, 3 Aug. 2003, R. Razakamalala 664 & D. Rabehevitra (MO-6038323); Ambila-Lemaitso, 18°49'S, 49°08'E, 14 Dec. 1967, Service Forestier 28034 (B, BR, G, K, MO-6321087, NY, WAG). Prov. Toliara: Mandena, Jardin Botanique, M16-QMM, Ampasy, Taolagnaro, 24°57'05"S, 47°00'11"E, 26 Sep. 2000, J. Rabenantoandro 241 + P. Lowry, R. Rabevohitra, L. Randrihasipara, & E. Ramisy (MO-5598572); Saint-Luce (Manafiafy), S9, Mahatalaky, Taolagnaro, 24°46'30"S, 47°10'20"E, 16 Dec. 2000, J. Rabenantoandro 366 + F. Lucien & E. Ramisy (MO-5728973); Manambaro, Petriky, 25°03'43"S, 46°52'06"E, 15 Nov. 2006, Ramison 33 (MO-6427490).
Haec species a congeneris madagascariensibus ramulis in sectione transversali non profunde sed late sulcatis, foliis saepe 3- vel 4-verticillatis atque hypanthio sub anthesi discoideo distinguitur.
MADAGASCAR. Prov. Fianarantsoa: Ranomafana PN, Ifanadiana, à 60 km au Nort-Est de la ville de Fianarantsoa, le long de la piste Cabine de Recherche, Vatoaranana, 10 Oct. 1996, 21°13'37"S, 47°22'11"E, 1500–1600 m, R. Randrianaivo 12 et al. (holotype: MO-6224857!; isotypes: KSP [KSP000044, KSP000045]!; P [P04885335]!, TAN).
Shrubs or trees, 3–8 m tall; dbh 2.8 cm; bark of main bole unknown. Foliage and flowers glabrous except as noted below. Branchlets laterally compressed to terete but sometimes trigonal in cross section, and irregularly and broadly and prominently sulcate on one or more sides, flaring distally below nodes but becoming rounded; smooth but sometimes striate, sparsely serciceous (trichomes reddish and dibrachiate) but soon glabrous, epidermis reddish or maroon on emergence but becoming irregularly and finely brown-gray mottled (dried material). Leaves opposite to disjunct opposite or whorled in 3s or 4s, concentrated near branch tips; coriaceous; venation brochidodromous; surfaces concolorous, matte; lateral veins 20–32 on each side, arising at an angle of 65–75˚ from the midrib. Axillary colleters lacking. Petioles 3–7(–12) mm, deeply sulcate adaxially, prominently striate-rugose (especially abaxially), eglandular. Leaf blades (4.4– )6.0–16.0 × (1.9)2.8–3.3 cm, narrowly elliptic, often conduplicate (dried material) in lower ca. 20%, base mostly rounded (sometimes cuneate), apex obtuse but sometimes slightly falcate, margin and surface somewhat sinuous; adaxial midvein deeply sulcate throughout, eglandular; abaxial surface eglandular except on midvein (glands moderately dense and somewhat indistinct); secondary veins consisting of (16–)25–32 pairs, indistinct but slightly raised, diverging at wide angle from midvein; intramarginal vein indistinct, 1.5–2.5 mm from margin of leaf at midpoint of blade. Inflorescence of solitary, paired, or fasciculate and ramiflorous monads. Pedicels (4.5–)8–17 mm, stiff but thin, terete to slightly compressed. Bracteoles 2, 0.5–0.7 × ca. 0.2–0.5 mm, broadly ovate, apex rounded, horizontal to reflexed in fruit, occasionally a few short trichomes. Hypanthium 0.5–1.5 mm long; initially cupuliform but becoming discoid (prominently flattened from above) during and after anthesis, glabrous, oil glands common, cream-colored (fresh). Calyx lobes 4 (but of unequal lengths), the two shorter lobes broader than long, ca. 1.5 × 2.2 mm broad at base, the two longer lobes hemispherical, ca. 2 × 2 mm, glandular, pale green (fresh), persisting or deciduous in fruit, ascending or crowing mature fruit. Petals 4, ca. 4 mm × 3.2 mm, broadly obovate to broadly elliptical, pink, with 9 veins arising from the base. Staminal disk diameter (in fruit) 3.5–5.0 mm; glabrous. Stamens ca. 60–80; filaments (1.5–)4–6 mm, cream-colored; anther sacs ca. 0.5 mm, subglobose to globose, connective apex eglandular; staminal disk glabrous. Style ca. 4–6 mm; stigma narrow. Fruit 11–18 × 8–23 mm, subglobose with irregular lobes (but not costate), base rounded, green (young fresh) but drying dark bluish-black, locules 2, placenta capitate, placentation axile, ovules radiating from center of placenta. Seeds 3–4 per fruit; outer coat leathery, highly contorted, drying to a light yellowish-brown (“fawn”) or dull brownish-yellow (“tawny”,
From Ranomafana National Park.
Fruiting October and November; flowering May.
Known from three collections in Ranomafana National Park in Fianarantsoa Province, southeast-central Madagascar (Fig.
Growing in the undergrowth or subcanopy in humid forests, ca. 950–1600 m. The collection by Turk notes its occurrence with trees of the genera Weinmannia L., Tambourissa Sonn., Decarydendron Danguy, Ephippiandra Decne., Ocotea Aubl. and Cryptocarya R. Br., and understory with Psychotria L. and Oncostemum A. Juss.
With only three collections known, an AOO of 9 km2 and one subpopulation, which is situated within the protected area network (Ranomafana), Eugeniaranomafana is assigned a preliminary risk of extinction of “Vulnerable” [VU D2] following the IUCN Red List Categories and Criteria (
This species was first recognized as undescribed by
With its narrowly elliptic leaf blades with mostly rounded leaf bases, Eugeniaranomafana resembles mostly closely Eugeniawilsoniana, the nearest occurrence of which is ca. 280 km northeast. Eugeniawilsoniana differs by its shorter stature, leaf margins being flat to only slightly sinuous, longer and thinner (on average) pedicels, presence of axillary colleters, and a hairy staminal disk. In addition, E.ranomafana differs by its often trigonous branchlets and the frequent occurrence of 3–4 leaves per node.
MADAGASCAR. Prov. Fianarantsoa: Ranomafana National Park, parcelle 3, S of National Road 25 at 7 km W of Ranomafana; Talatakely trail system, 21°15'30"S, 47°25'00"E, 950–1150 m, 4 May 1993, D. Turk 436 + J. Randrianasolo, J. Solo, & D. Randriamanatenta (MO, KSP); Ranomafana National Park, Talatakely parcel, Trail System F-TBT-B, 21°15'S, 47°25'E, P. Fritsch 1643 et al. (CAS, MO-6287616).
Haec species a congeneris madagascariensibus foliis apice abrupte cupidatis caudatisve atque inflorescentiis ramifloris dense fasciculatis distinguitur.
MADAGASCAR. Prov. Antsiranana: SW d’Andapa, Réserve Spéciale Anjanaharibe-Sud, suivant la piste pour Ambalaheva, haute rivière d’Andramonta, 14°36'40"S, 49°24'12"E, 628–1879 m, 22 Feb. 1996, D. Ravelonarivo et al. 930 (holotype: MO-6135426!; isotypes: KSP [KSP000007, KSP000008]!, P [P04885351]!, TAN).
Shrubs or trees, 3–12 meters; bark of main bole unknown. Herbage glabrous except as noted. Branchlets laterally comprssed and shallowly 2-grooved on each side when young below petioles, oil glands common but indistinct, epidermis smooth, green drying light brown. Leaves evenly distributed along branchlets, opposite to disjunct opposite, thinly coriaceous to thickly membranous, discolorous, surfaces matte. Axillary colleters absent. Petioles 7–10 mm, narrowly and deeply sulcate (especially distally). Leaf blades (5.5–)7.5–14 × 3.3–4.8 cm, mostly elliptic but some broadly elliptic or narrowly oblong, base cuneate, apex obtuse, or acuminate and often falcate, margins flat but slightly undulate (dried); adaxial surface midvein deeply but narrowly sulcate to nearly the apex, secondary veins numerous but faint, oil glands absent; abaxial surface lakcking oil glands, secondary veins faint, slightly raised, diverging at ca. 80°; intramarginal vein 0.5–1.5 mm from margin at midpoint of blade. Inflorescences of moderately dense fascicles of monads arising from short brachyblasts, these evidently ramiflorous or cauliflorous (uncertain given that all are detached on herbarium sheets); pedicels 9–15 mm, rigid, irregularly glandular. Bracteoles 2, ca. 0.5–0.8 mm, triangular to ovate, often obscure. Hypanthium ca. 2–2.5 mm, cupuliform to discoid (somewhat flattened) in anthesis, prominently glandular. Calyx lobes 4, 2.0–5.0 mm, broadly oblong to oblate or rounded, often dimorphic (2 shorter, 2 longer), apex broadly rounded, petaloid, oil glands sparse (and mostly abaxial). Petals 4 (material scant), up to 5 mm; indicated as pinkish on a paratype. Staminal ring more or less square, ca. 4 mm per side, sparsely short hairy (use magnification) and glandular. Stamens (material scant) numerous; anthers ca. 0.5 mm, ellipsoid. Styles 6–11 mm, thin, sparsely glandular (especially proximally); stigma narrow. Berries 24–35 × 30–50 mm (dried), subglobular, greenish (mature or nearly so); seeds 1–3, 16–22 × 25–30 mm, subglobular; embryo globular, hypocotyl and cotyledons not differentiated; testa thinnish, drying light brown.
Flowering January through April; fruiting March through November.
In the northern mountains of Madagascar in Antsiranana and Mahajanga provinces (Fig.
In mid-elevation, wet eastern forests often over granite or gneiss; elevation approximately 630–1880 m but needing confirmation.
It is a pleasure to name this species in honor of Désiré Ravelonarivo (b. 1966), a prolific collector of specimens in Madagascar. Désiré is native to the Andapa basin, where he collected the type specimen of Eugeniaandapae and the only known fruiting specimen of this new species (D. Ravelonarivo 489 & R. Babesonina).
Gavoalehely (Ravelonarivo 930); gavoala (Ravelonarivo 103).
With an EOO of 436 km2, an AOO of 36 km2 and three subpopulations, two of which are situated within the protected area network (Anjanaharibe-Sud), Eugeniaravelonarivoi is assigned a preliminary risk of extinction of “Vulnerable” [VU D2] following the IUCN Red List Categories and Criteria (
The label of the type gathering indicates “rougeâtres” (reddish) for petals, but this is doubtful and probably was meant for pinkish, which accords with information on the label of a paratype.
Among existing species, the leaf morphology of Eugeniaravelonarivoi resembles that of E.alatroensis H. Perrier and E.vatomandrensis H. Perrier, but the bases of their leaves are subsessile to sessile, which contrasts with the distinctly petiolate leaves of E.ravelonarivoi. In addition, the often cuspidate-caudate aspect of the leaf apex of new species is much more pronounced than of those two species, whose tips are barely (if at all) cuspidate.
Eugeniaantongilensis H. Perrier has cuspidate apices on the blade, but it is a much longer leaf with shorter petioles. Eugeniamusicola H. Perrier also resembles E.ravelonarivoi, especially given the membranous to weakly coriaceous texture of the leaf blades; however, the pedicels of the former are much thinner and more lax, and its petioles are shorter and much less (if at all) sulcate adaxially. Eugeniadiospyroides differs with its generally larger flowers and leaves, and leaves having a rounded apex.
The new species also closely resembles Eugeniaradiciflora in leaf morphology, but that species is said to have solitary flowers with pedicels only 2–3 mm long, which does not match the fasciculate brachyblasts of E.ravelonarivoi. The leaves of Eugeniaravelonarivoi also superficially resemble those of E.gandhii, but the latter has densely punctate leaf blades and solitary flowers. Finally, Eugeniaravelonarivoi resembles closely some evidently undescribed taxa
Dr. David Gordon, an entomologist at Pittsburg State University, indicates that the visible damage on the leaves of some specimens may be from leaf cutter bees (Family Megachilidae), which often damage leaves in this manner to obtain material for the construction of their nests.
MADAGASCAR. Prov. Antsiranana: SW d’Andapa, Réserve Spéciale d’Anjanaharibe-Sud, aux enrivons de sommet, 14°46'15"S, 49°28'00"E, 1112–1424 m, 21 Mar.–7 Apr. 1994, Ravelonarivo 103 + F. Rasoavimbahoaka, B.T. Rafaliarimanana, H. Rasitefarinina & Motera (P [P04884879]); Massif de l’Anjanaharibe (pentes et sommet N) a l’W d’Andapa (Haut Andramonta, Bassin de Lokoho), [14°37'S, 49°25'E], 900 m, 10 Dec. 1950–9 Jan. 1951, H. Humbert 24576 + R. Capuron & G. Cours (P [P05208600]); Sud-Ouest d’Andapa, Réserve Spéciale d’Anjanaharibe-Sud, village d’Andranotsarabe, suivant la route Nationale d’Andapa-Bealanana de lat piste vers à l’oueste, Ambatoomainty, Camp No. 2, 14°44'22”S, 49°27'42”E, 3 Nov. 1994, 1185–1335 m, D. Ravelonarivo 489 & R. Babesonina (KSP [KSP003115, KSP003116], P [P05208456]). Prov. Mahajanga: Ankaizinana, [14°30'S, 48°55'E], 1400 m, 20 Apr. 1923, R. Decary 2009 (P [P00118106]).
Haec species a congeneris madagascariensibus lamina foliari rigide coriacea late elliptica usque obovata utrinque ad paginas (nec ad margines tantum) nonnihil sinuosa atque glandulis densis folii paginam abaxialem pedunculos hypanthium lobos calycinos petala necnon fructum ornantibus distinguitur.
Madagascar. Prov. Antsiranana: Fokontany: Anjia, Ambato, 15°17'26"S, 50°20'28"E, 37 m, 25 Feb. 2001, R. Razakamalala 67 + S. Rakotoarisoa, A. Rasolohery & P. Antilahimena (holotype: MO-6277747!; isotypes: KSP [KSP000003]!; P, TAN).
Trees to 10 m; dbh up to 13 cm; bark of main bole unknown. Plants glabrous throughout. Branchlets laterally compressed; youngest epidermis drying light brown to light gray but soon peeling and flaking to reveal reddish-brown color (dried); oil glands of epidermis prominent and common. Leaves stiffly coriaceous, mostly concentrated near branch tips, medium green and slightly discolorous, irregularly sinuous, surfaces matte. Axillary colleters absent. Petioles 3–5 mm, becoming slightly sulcate distally and broadening towards apex. Leaf blades 3.5–9.0 × 2.5–5.0 cm, broadly elliptic to obovate or broadly obovate, base cuneate, apex obtuse to somewhat acute; adaxial surface with midvein broadly but shallowly sulcate in the lower half, becoming flush distally; oil glands absent; abaxial surface oil glands common to dense but small and faint and fading with maturity (visible only with magnification), secondary veins somewhat indistinct, intramarginal vein indistinct and ca. 0.5–1.0 mm from margin at midpoint of blade. Inflorescence terminal or axillary, monads 1–3 in each axil. Pedicels 15–25 mm, relatively thick (1.2–2.0 mm at maturity), ascending to erect, rigid. Bracteoles 2, 1–2 mm, ovate, stiff, persistent and more or less divergent in fruit. Hypanthium 3.5–5 mm, cupulate, oil glands common to dense. Calyx lobes 4, 5–8 mm, broadly rounded, apex obtuse, densely glandular, persistent and crowning the fruit, stramineous or rufous-beige (fresh). Flowers unknown. Berries 1.5–1.8 × 2.0–2.3 cm, subglobular to globular, densely glandular, slightly verrucose (dried), yellowish (possibly not fully mature).
It is a pleasure to name this species in honor of Richard Razakamalala (b. 1962), a Malagasy botanist who has collected over seven thousand specimens of plants from Madagascar since 2001. His prolific collecting efforts have contributed significantly to our increased knowledge of the Malagasy flora. Four of his Eugenia collections are designated as type specimens in this publication.
Fruiting late February.
Known only from the type gathering in Antsiranana Province, north-eastern Madagascar, about 14 km west of the coast in the Masoala Peninsula (Fig.
In dense humid lowland forests over quartzite from ca. 35 m elevation.
With only one collection known from just outside the Masoala protected area, Eugeniarazakamalalae is assigned a preliminary risk of extinction of “Endangered” [EN 2ab(i, iii, iv)] following the IUCN Red List Categories and Criteria (
The leaf morphology of Eugenia razakamalalae resembles that of Eugeniaarthroopodavar.ambalavensis H. Perrier, an invalid name given the lack of a Latin diagnosis in the protologue (
Jossiniarichardii Blume in Mus. Bot. 1: 123. 1851.
MADAGASCAR. Prov. Antsiranana: forêt à Vohémar, [13°21'30"S, 50°00'30"E], comm. 1837, J.M.C. Richard 119 (holotype: L [L0009467]!; isotype: P [P00118150]!).
Shrubs 2–4 m tall; ca. 4 cm dbh; bark of main bole rough, grayish-brown. Branchlets terete, lightish gray-brown (dried); internodes short, (0.5–)1.5–2.3 cm; epidermis smooth but becoming fissured and flaking on older internodes, indumentum moderately dense of minutely dibrachiate trichomes; oil glands common, and somewhat prominent, and protruding above surface. Leaves mostly evenly distributed along branchlets, coriaceous, slightly discolorous, surfaces matte. Axillary colleters absent. Petioles 1–2 mm, slightly sulcate but mostly flush adaxially, glabrous. Leaf blades 2.5–4.5 × 1.6–2.5 cm, elliptic; base rounded to somewhat cuneate; apex broadly obtuse to rounded; margins slightly revolute; adaxial surface glabrous, oil glands dense upon emergence but fading as blade thickens, midvein broadly but slightly sulcate proximally; abaxial surface resembling adaxial, secondary veins indistinct or not visible, intramarginal vein 1.0–1.5 mm from margin at midpoint of blade. Inflorescence a monad arising in terminal leaf axils. Anthopodia (short internode between base of hypanthium and subtending bracteoles) present, 1–2 mm. Pedicels 1.0–1.4 mm, firm but slightly bowed, glabrous. Bracteoles 2, 0.8–1.0 × ca. 1.0 mm, ovate, minutely hairy on margins near apex. Hypanthium 4–5 mm, cupuliform to obconic, glabrous but evidently exuding salt, oil glands common to dense; ovary apex densely short-hairy. Calyx lobes 4, ca. 4 × 4 mm (material limited), oblong, apex obtuse, glabrous, greenish. Petals 9–10 × 6–9 mm, widely elliptic to widely obovate, sparsely short-ciliate on margins, thin (nearly transluscent), pinkish, oil glands sparse to common but prominent. Stamens ca. 70–100, multiseriate; anthers ellipsoid, 0.6–0.8 mm, sub-basifixed, connective with a single apical gland. Filaments 3–9 mm, white; ovary apex densely short-hairy. Styles 9–10 mm, glabrous; stigma narrow. Berries unknown.
Flowering August; fruiting January and likely commencing in late August (unconfirmed).
Known from the north-eastern coastal region of Madagascar in Antsiranana Province, from Orangéa to about 140 km along the coast near Vohemar (Fig.
Littoral forests over sand, inland from the ocean shore less than 1 km; elevation 10 m.
With an EOO of 358 km2, an AOO of 36 km2 and three subpopulations, one of which is situated within a proposed protected area that currently holds a temporary protection status (Oronjia), Eugeniarichardii is assigned a preliminary risk of extinction of “Endangered” [EN B1ab(iii)+ B2ab(iii)] following the IUCN Red List Categories and Criteria (
Eugeniarichardii most closely resembles two species, including Eugeniavanwykiana N. Snow, which differs by its larger leaves with cuneate bases (
Eugeniarichardii also resembles E.quadriflora H. Perrier, which is known from a single collection in 1961 from east of Lake Alaotra. Both species have thickly coriaceous, densely glandular leaf blades with irregularly undulating surfaces and moderately to strongly recurved margins, and simple flowers arising terminally. However, the leaves of E.quadriflora are sessile with strongly and irregularly recurved blades (although this may reflect to some degree its condition when pressed and dried); in contrast, the leaf blades of of E.richardii are distinctly but shortly petiolate and slightly recurved.
Elsewhere, the leaf morphology of Eugeniarichardii somewhat resembles that of Eugenianeofasciculata Bennet from Mauritius (formerly E.fasciculata Guého & A.J. Scott). Based on comparison digitally with an isotype (Lorence DL 1214 (P [P00390098]!), the species are similar by virtue of stiffly coriaceous, densely punctate, obovate-elliptic leaf blades with slightly undulating surfaces and recurved margins that arise from relatively short petioles. However, whereas the adxial petiolar surface of E.richardii is flat, that of E.neofasciculata is somewhat sulcate. In addition, the inflorescence of E.richardii is a terminal pair of flowers with one arising in each leaf axil, whereas that of E.neofasciculata is conspicuously fasciculate (mostly) on naked branches or within leaf axils.
MADAGASCAR. Prov. Antsiranana: Vohémar, [12°16'24"S, 49°23'20"E], s.d., L.H. Boivin 2697 (=J.M.C. Richard 134) (P [P00118153]); Orangéa, [12°14'S, 49°22'E], 1–100 m, 22 Jan. 1960, G. G. Cours 5394 (P [P05097489]; Env. de Diégo-Suarez, Orangéa, [12°14'S, 49°22'E], 1–100 m, 22 Jan. 1960, H. Humbert 32245 + G. Cours (P [P05576991]); Ramena, Baie des Sakalava, 3.5 km à l’E du Fokontany Ankorikihely, 12°16'24"S, 49°23'20"E, 17 Aug. 2004, J.B. Leopold 27 + L.J. Razafitsalama, R. Guittou, J. Be, & V. Benjara (CNARP, KSP [KSP000005], MO-6245567, P [P05208542], TAN); Andrafiabe, presqu’île entre Ambolobozobe et Ambolobozokely, 12°29'39"S, 49°34'04"E, 9 Feb. 2005, F. Ratovoson 965 + R. Guittou & D. Elifara (G, MO-6028974, P [P05208572]); Vohémar, [13°21'30"S, 50°00'30"E], s.d., J.M.C. Richard 75 (P [P00118148, P00118149]).
Haec species Eugeniae thouvenotianae H. Perrier simillima, sed ab ea lamina foliari glandulis oleiferis punctatis abaxialiter ornata atque pedicellis multo brevioribus distinguitur.
MADAGASCAR. Prov. Antsiranana: Doany, en aval des deux côtes de la rivière Ambalanirano. Ambohimirahavavy, 14°13'43"S, 49°08'07"E, 2011 m, 19 Nov. 1995, C. Rakotovao 2511 + Jaovazaha & Tsarajery (holotype: MO-6174870!; isotypes: G [G00341423]!, P [P05260190]!, TAN).
Trees 8–18 m; dbh 12–60 cm; bark of main bole unknown. Indumentum of foliage and floral parts dibrachiate, short and appressed, ferrugineous, denser upon emergence but becoming (often) nearly glabrous (or as indicated below). Branchlets rounded to laterally compressed, greenish fresh but drying light brown to light gray; epidermis thin and peeling away in thin strips or flakes, oil glands absent to sparse and indistinct. Leaves discolorous, margin sometimes strongly revolute and irregularly sinuous, venation reticulate, emerald and somewhat glossy adaxially, citrine and matte abaxially, concentrated near branch tips. Axillary colleters occasionally present. Petioles (2.5–)5–10, narrowly and deeply sulcate adaxially, elgandular. Leaf blades 4.0–6.8 × 2.3–.8 cm, elliptic to obovate, base cuneate, apex obtuse or occasionally retuse; adaxial surface remaining at least sparsely sericeous when fully emerged (trichomes closely appressed), oil glands common to dense but faintly visible, midvein deeply and narrowly suclate proximally becoming flush distally; abaxial surface indumentum as above, oil glands not as dense but much more prominent, secondary veins indistinct to somewhat prominent, intramarginal vein 1.0–1.5 mm from margin at midpoint of leaf blade (but often obscured by revolute margin). Inflorescence 2–5 cm long, mostly axillary or ramiflorous, sometimes terminal, consisting of triads, botryoids, or less commonly metabotryoids or pseudo-umbels of 4–5 flowers apically clustered on peduncle; peduncles mostly solitary but often opposite in leaf axils or on naked branches, mostly ascending and somewhat stiff but also sometimes flexuous. Pedicels 1–2 mm long. Bracteoles absent; bract-like structures surrounding triads of flowers when inflorescence is emerging but these soon deciduous. Hypanthium 1.5–2.5 mm, obconic, densely hairy and densely punctate. Calyx lobes 4, (1.2–)1.5–1.8 mm, broadly rounded, apex obtuse, mostly glabrous above to sparsely sericeous, moderately sericeous dorsally and on margins below, deciduous in fruit, light green when fresh. Petals 4, 4–7 mm, obovate, minutely and sparsely ciliate apically (use magnification) but otherwise glabrous, oil glands sparse to moderate. Stamens 20–45, exserted; staminal disk glabrescent; ovary apex glabrous; filaments 4–7 mm, yellowish-white; anthers 0.4–0.6 mm, globose to subelliptic, basifixed, eglandular; style 4–5 mm, stigma narrow and scarcely if at all capitate. Fruit (material limited), ca. 2.2 × 2.2 cm, globose, glabrate, green when young drying nearly black; outer layer leathery and prominently glandular. Seed 1 at maturity (material scant), round from above but laterally compressed (due to pressing?), ca. 16 × 16 × 9 mm; embryo not differentiated into distinct cotyledons or epicotyl.
The specific epithet is derived from elements of the native language of northern Madagascar, in which ampoka means mosses. Eugeniatiampoka thus means the “Eugenia that likes mosses”.
Flowering October through February; fruiting by early April, likely commencing by March and extending through May.
Northern Madagascar in Antsiranana Province in the northern mountains (Fig.
Mid- to high-altitude humid forests laden with mosses, sometimes along river banks; 1990–2300 m in the northern highlands of Madagascar. A team of botanists lead by MC conducted a series of field expeditions between 2005 and 2008 to explore the flora and vegetation of this long-neglected region, during which more than 4,400 collections were made, yielding material of several plant species new to science (
With an EOO of 71 km2, an AOO of 45 km2 and two subpopulations, neither of which is within the protected area network, Eugeniatiampoka is assigned a preliminary risk of extinction of “Endangered” [EN B1ab(iii)+2ab(iii)] following the IUCN Red List Categories and Criteria (
The leaves reportedly are aromatic (Birkinshaw 950, Rakotovao 2398). One specimen is said to have a nectariferous disk (Randriarivelo 350). A specimen of 30 m stature (Ravelonarivo 647) from the same general area and also said to be aromatic, but growing at 1700 m, has a slightly different aspect to the leaves, also may be this species.
The morphological gestalt of the leaves and inflorescences resemble most closely those of E.thouvetiana H. Perrier, which however lacks the punctate oil glands of the abaxial leaf surface, and which has much longer pedicels subtending the individual flowers than those of E.tiampoka. The type locality of E.thouvetiana is located in Analamazaotra-Andasibe in Toamasina Province, some 525 km south of the southernmost known collection of E.tiampoka. Eugeniatiampoka also somewhat resembles Eugeniahazompasika H. Perrier, the latter of which has much longer, broader and stiffer leaves with a less abruptly cuneate base.
MADAGASCAR. Prov. Antsiranana: 13 km N of Mangindrano, Tsaratanana Massif, SW ridge of Andomanisambiraro, 14°08'41”S, 48°57'53”E, 2150 m, 17 Oct. 2001, C. Birkinshaw 950 + R. Lala (KSP [KSP000048, KSP000049], P [P05156041]); Crête E d’Ambohimirahavavy, 14°12'15”S, 49°05'54”E, 2278 m, 3 Nov. 2005, C. Rakotovao 2398 + Jaovazaha & Torize (G, MO-6202095); Sommet de Beampoko, Ambohimirahavavy, 14°13'55”S, 49°08'23”E, 2137 m, 21 Nov. 2005, C. Rakotovao 2566 + Jaovazaha & Tsarajery (G, MO-6174671, P [P05260203]); Doany, forêt d’altitude d’Ambohimirahavavy, 14°13'41”S 49°08'14”E, 1991 m, 18 Nov. 2005, C. Randrianarivelo 350 + J. Randriantiavina & Torize (G, MO-6081166), KSP [KSP000047], P [P05260189]); Fokontany: Ampanompy, Ampanompy, 14°08'31”S, 48°58'04”E, 2050–2300 m, 5 Apr. 2001, R. Razakamalala 99 + R. Ranaivojaona, F. Ratovoson, A. Rasolohery, A. Andriamaniry & Mahavory (KSP [KSP000047]!).
Haec speciesEugeniaebemangidiensi N. Snow simillima, sed ab ea inflorescentia axillari longipedicellata distinguitur; etiam ad altitudines superiores crescit.
MADAGASCAR. Prov. Toamasina: Alaotra Mangoro Reg., Moramanga, Ambohibary, Ampitambe, 18°48'55"S, 48°16'37"E, 1103 m, 14 Oct. 2008, R. Rakotondrajaona 649 (holotype: MO-6419537!); isotypes: P, TAN).
Shrubs or trees, (1–)2–6 m. Trunk dbh ca. 2 cm (measurements few); bark of main bole fissured, maroon. Indumentum where present of highly asymmetric and short, ferrugineous, dibrachate hairs. Branchlets laterally compressed, smooth, sparsely puberulous, drying light brown to greenish, punctate glands absent. Leaves opposite or disjunct opposite, mostly concentrated near tips of branches; venation reticulate; blades thinly coriaceous, dark green above and lighter green below. Axillary colleters obscure when present. Petioles 3–5 mm, deeply and narrowly sulcate adaxially, laterally compressed, elgandular, glabrescent (especially abaxially), longitudinally striate initially but thickening and becoming somewhat latitudinally striate with age. Leaf blades 4.0–12.5 × 1.4–3.0 cm, narrowly elliptic to narrowly ovate, base rounded and slightly constricted-conduplicate above petiole, apex acute, margins flat; adaxial surface glabrous, eglandular, midvein narrowly but deeply sulcate lower 2/3–4/5; abaxial surface like adaxial except: midvein protruding, sometimes prominently glandular and longitudinally striate, secondary veins faint to prominent, arising only 10–20° from midvein, tertiary veins faint to nearly as prominent as secondaries; intramarginal vein 1.0–1.5 mm from leaf edge at midpoint of blade. Inflorescence terminal, axillary, or arising on naked branch of current year’s growth. Flowers solitary, or occasionally as up to four arising from short brachyblasts; pedicels 6–12 × ca. 0.5 mm, glabrous, strongly laterally compressed (especially distally), longitudinally striate, sparsely to moderately glandular. Bracteoles 2, 0.6–1.1 mm, narrowly ovate to ovate, sparsely hairy abaxially and apically, rigid, somewhat ascending to appressed against base of hypanthium. Hypanthium 2.5–4.0 mm, cupuliform, sparsely hairy becoming glabrous, sparsely to moderately (but only faintly) glandular. Calyx lobes 4, 4–5 × 3–5 mm, oblate to broadly elliptic, apex broadly obtuse to rounded, minutely and sparsely ciliate in upper half, prominently glandular, greenish when fresh. Petals 4, 4–5 × 2–3 mm, widely elliptic to ovate, sparsely short-ciliate upper 1/2–1/3, faintly and somewhat sparsely glandular, whitish or pinkish. Stamens exserted, staminal disk ca. 4 mm diameter, more or less square from above, short-hairy, adjacent ovary apex glabrous; filaments 2–3 mm; anthers 0.8–1.0 mm, elliptic, yellow, sub-basifixed, apical gland between connectives lacking. Styles 7–8 mm, glabrous; stigma narrow and scarcely if at all capitate. Berries (immature?) 12–15 × 12–15 mm, globular, greenish.
In honor of Dr. Peter G. Wilson (b. 1950) of the Royal Botanic Gardens in Sydney, Australia; colleague, collaborator, and long-time student of Myrtaceae (e.g.,
Hazompasina (Antilahimena 4935); Rotra (Ravelonarivo 3148).
Flowering mid-October through mid-November; fruiting October and November.
East-central Madagascar in Toamasina Province, occurring near Analamazaotra National Park (Fig.
In humid, sometimes disturbed forests at middle altitudes from ca. 980–1103 m.
With five collections known, an AOO of 9 km2 and one subpopulation that is situated outside the protected area, Eugeniawilsoniana is assigned a preliminary risk of extinction of “Critically Endangered” [CR A3c+B1ab(iii)] following the IUCN Red List Categories and Criteria (
The leaf and floral morphologies of Eugeniawilsoniana are suggestive of Eugeniadiospyroides H. Perrier. However, E.wilsoniana has much shorter leaves and a hairy staminal ring, in contrast with the much larger leaves and glabrous staminal ring of E.diospyroides (e.g., Randriatifika 118 et al. [KSP]). This new species resembles even more closely another newly described species herein, E.bemangidiensis, which differs by its cauliforous inflorences occurring in tight, relatively short fascicles, secondary veins on the leaf blades arising at steeper angles, and occurring in a different habitat at much lower elevations some 750 km southeast from the presently known occurrences of E.wilsoniana. Moreover, the foliage and floral parts of E.bemangidiensis are entirely glabrous, the abaxial laminar midvein protrudes less prominently, the adaxial laminar sulcus is not as deep or narrow, and its adaxial petiolar sulcus is broader and shallower than those of E.wilsoniana.
MADAGASCAR. Prov. Toamasina: Ambatovy, Sahaevo forest, 18°50'26"S, 48°16'33"E, 11 Nov. 2006, P. Antilahimena 4935 & F. Edmond (MO-6338238); ibid. loc., 18°50'26"S, 48°16'34"E, 23 Nov. 2008, P. Antilahimena 6912 + B.A. Ratodimanana, D. Ravelonarivo, E. Félix & M. Ratvomanana (MO-6447030); Ambatovy, Antaniditra, 18°49'11"S, 48°16'53"E, 13 Oct. 2008, R. Bernard 1154 (MO); Ampitambe, Ambatovy, 18°49'06"S, 48°17'06"E, 1041 m, 13 Oct. 2008, R. Rakotondrajaona 642 + M. Ratolojanahary (MO-6419531); Mararano, Marovoay, 18°48'10"S, 48°17'59"E, 14 Nov. 2008, D. Ravelonarivo 3148 (MO-6447573).
Many species of Eugenia in Madagascar described by
Lectotypification is particularly important for Eugeniascottii, since the protologue included twelve syntype collections that clearly included more than one taxon. Eugeniahovarum H. Perrier and Eugeniascottii H. Perrier were described for relatively small-leaved species occuring mostly in the mountainous parts of Madagascar, and both were based on multiple syntypes (
Eugeniascottii H. Perrier in Mém. Inst. Sci. Mad., Sér. B, Biol. Vég. 4(2): 168-169. 1953. Lectotype (designated here): MADAGASCAR. Prov. Antsiranana: Massif du Tsaratanana et haute vallée du Sambirano, Mt. Andilabe, 2300 m, Nov.-Dec. 1937, H. Humbert 18501 (lectotype: P [P00118145]!; isolectotype: P [P00118146]!).
Among the specimens that we exclude from Eugeniascottii is material that can be accommodated in E.cassinoides Lam., a species from the littoral forest in the south-east (Perrier de la Bâthie 6486); E.richardii (e.g., Boivin 2697, Richard 75, Richard 119), a species from low elevation dry forest in the north-east (the needed new combination of which is provided in this article); E.petrikiensis N. Snow & Randriat. (Scott Eliott 2847) from Fort Dauphin; and a new, as yet undescribed species from the mountains of the south-east at mid elevation in the Mandrare basin (Humbert 6503, 6685, 6835), and probably also includes the last syntype, Humbert 11965, from Kalambatritra.
Additional specimens examined: MADAGASCAR. Prov. Antsiranana: Réserve Naturelle Intégrale du Marojejy, versant W et SW du sommet de Marojejy, 14°27'S, 49°44'E, 2040 m, 15 Nov. 1996, Rakotomalaza 918 et al. (G, MO, P [P05208453]); Massif du Tsaratanana, hauts sommets de l’Amboabory à l’Antsianongatalata, 2300-2609 m, Nov.-Dec. 1937, H. Humbert 18389 (P [P00118147]); Sommet oriental du Massif du Marojejy, à l’W de la Haute Manantenina, affluent de la Lohoho, 1850-2137 m, 26 March–2 Avril. 1949, H. Humbert 23709 + G. Cours (P [P00118154]).
Excluded syntypes: MADAGASCAR. Prov. Antsiranana: Vohemar, L.H. Boivin 2697 (= J.M.C. Richard 134) (syntype: P [00118153]!); Vohémar, [12°16'24"S, 49°23'20"E], s.d., J.M.C. Richard 75 (syntypes: P [P00118148, P00118149]!); Vohémar, [12°16'24"S, 49°23'20"E], comm. 1837, J.M.C. Richard 119 (syntypes: L [L0009467]!, P [P00118150]!). Prov. Fianarantsoa: massif de Kalambatitra (centre S), Mt. Analatsitendrika, Nov. 1933, H. Humbert 11965 (syntypes: P [P00118151, P00118152]!); près de l’embouchure du Matitana (SE), Oct. 1911, H. Perrier de la Bâthie 6486 (syntype: P [P00118144]!). Prov. Toliara: Haute vallée du Mandrare (SE), 600-900 m, 8–15 Nov. 1928, H. Humbert 6503 (syntypes: P [P00118141, P00118142]!); Bassin sup. du Mandrare (SE), du col de Vavara à la vallée du Manambolo, 700–1200 m, 20–22 Nov. 1928, H. Humbert 6685 (syntype: P [P00118140]!); Bassin sup. du Mandrare (SE), Mt. d’Amboahangy près d’Esira, 1000–1150 m, 25 Nov. 1928, H. Humbert 6835 (syntypes: P [P00118139, P00118138]!); Fort Dauphin, s.d., Scott Eliott 2847 (syntype: P [P00118137]!).
Eugeniahovarum H. Perrier in Mém. Inst. Sci. Mad., Sér. B, Biol. Vég. 4(2): 170. 1953. Lectotype (designated here): MADAGASCAR. Prov. Antananarivo: env. de Manjakandriana, comm. 1932, Perrier de la Bâthie 14007 (lectotype: P [P00118032]!; isolectotype: P [P00118031]!). Syntype: MADAGASCAR. Prov. Antananarivo: Imerina, comm. 20 Nov. 1889, Campenon s.n. (P [P00118033]!).
Additional specimen examined: MADAGASCAR. Prov. Antsiranana: Parc National de Ranomafana, ca. 4 km E of Vohiparara, 21°13'32”S 47°26'43”E 1170 m, 5 Nov. 1993, Schatz 3531 (MO, P [P05208543]).
Eugenianompa H. Perrier in Mém. Inst. Sci. Mad., Sér. B, Biol. Vég. 4(2): 181. 1953. Lectotype (designated here): MADAGASCAR. Prov. Antsiranana: Dist. Ambanja, Beangona (haut Sambirano), [14°03'00"S 48°42'15"E], 24 Oct. 1947, Réserves Naturelles 1275 (lectotype: P [P00118008]!; isolectotype: P [P00118007]!). Syntypes: MADAGASCAR. Prov. Antsiranana: env. du Mt. Tsaratanana, Nov. 1912, Perrier de la Bâthie 5481 (P [P00118004, P00118005, P00118006]!).
The species concept for Eugenianompa H. Perrier has been uncertain. Because the syntypes were clearly designated by
Additional specimen examined: MADAGASCAR. Prov. Antsiranana: District Ambanja, Canton: Marovato. Réserve Naturelle 4 (Tsaratanana), 21 Sept. 1953, Réserve Naturelle 5750 (P [P05097466]).
The eighteen species of Eugenia newly proposed here increases the known number of native species in Madagascar from approximately 46 to 68, all of which are endemic. Of these, 32 (approximately half) have been described in the last fifteen years (
Many of the original descriptions by
Malagasy Eugenia have a relative paucity of easily diagnostic, qualitatively different morphological character states compared to some baccate genera of Myrtaceae (e.g.,
Despite additional new species of Eugenia remaining to be described and other gaps in knowledge of Malagasy taxa, tentative patterns of distribution, ecological preferences and evolution are beginning to emerge (
Many species of Malagasy Eugenia, including several proposed here, are Threatened, Endangered, or Critically Endangered following IUCN Criteria and Recommendations (
We believe that at least ten or more additional species remain to be described, and stress here that more material of other species would enhance our knowledge of taxonomy, distribution, and conservation priorities. Duplicate material of Malagasy Eugenia sent to the first author at the T.M. Sperry Herbarium (KSP) for future studies would be greatly appreciated.
Corrigendum: The correct collection number for the paratype specimen of Eugeniavanwykiana N. Snow is Razakamalala 1234 et al. The protologue incorrectly cited the collection number as 1324 (
The first author thanks colleagues at Missouri Botanical Garden in St. Louis (MO) for loans of specimens, access to collections, and general assistance; in particular, Zachary Rogers, Jennifer Kuhl, and James Solomon. We also thank MO for permission to use holotype images on Tropicos® (http://www.tropicos.org). MC is grateful to Laurent Gautier and Fabienne Thonney for their help at G. Tabita Randrianarivony assisted with formulation of the specific epithet for Eugeniatiampoka, and Lalao Andriamahefarivo provided valuable advice on Malagasy vernacular names. Peter Wilson (NSW), Kanchi Gandhi (GH) and especially Roy Gereau (MO) wrote or advised on the Latin descriptions and epithets. General discussions of Malagasy Myrtaceae with James Byng and George Schatz have been helpful. David Gordon (Pittsburg State University) examined leaves in the context of insect grazing. NS thanks the Department of Biology and the Graduate School and Continuing Education at Pittsburg State University for financial support.