Studies of Malagasy Eugenia – IV: Seventeen new endemic species, a new combination, and three lectotypifications; with comments on distribution, ecological and evolutionary patterns

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Studies of Malagasy Eugenia – IV: Seventeen new endemic species...


Studies of Malagasy Eugenia – IV: Seventeen new 

endemic species, a new combination, and three 

lectotypifications; with comments on distribution, 

ecological and evolutionary patterns

Neil Snow


, Martin Callmander


, Peter B. Phillipson


1 T.M. Sperry Herbarium, Department of Biology, Pittsburg State University, 1701 S. Broadway, Pittsburg, 

KS 66762 USA Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, USA Conser-

vatoire et Jardin botaniques de la Ville de Genève, case postale 60, 1292 Chambésy, Switzerland 4 Institut de 

systématique, évolution, et biodiversité (ISYEB), Unité mixte de recherche 7205, Centre national de la recher-

che scientifique/Muséum national d’Histoire Naturelle/École pratique des hautes études, Université Pierre et 

Marie Curie, Sorbonne Universités, CP 39, 57 rue Cuvier, F-75231 Paris cedex 05, France

Corresponding author: Neil Snow (


Academic editor:

 Peter de Lange  |  Received 21 November 2014  |  Accepted 27 March 2015  |  Published 28 April 2015


 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. 

doi: 10.3897/phytokeys.49.9003


Seventeen new endemic species of the genus Eugenia L. (Myrtaceae) are proposed from Madagascar, in-

cluding: 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

Eugenia richardii (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 Eugenia hovarum H. Perrier, Eugenia 

nompa H. Perrier, and E. scottii H. Perrier respectively.

PhytoKeys 49: 59–121 (2015)

doi: 10.3897/phytokeys.49.9003

Copyright Neil Snow et al. 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.


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Neil Snow et al.  /  PhytoKeys 49: 59–121 (2015)



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 éva-

luation préliminaire du risque d’extinction selon l’UICN ainsi que des recommendations de conservation 

sont fournies. Des lectotypes sont désignés pour Eugenia hovarum H. Perrier, Eugenia nompa H. Perrier 

et Eugenia scottii 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 Eu-

genia L. (Snow 2008, 2011; Snow et al. 2012), which until the work of Miller (2000) 

and Labat and Schatz (2002) had been dormant for five decades since early treatments 

by Perrier de la Bâthie (1953a,b). Its purpose is to propose seventeen new species, 

make a new combination, and lectotypify three names. It also provides detailed spe-

cies descriptions with comparisons to morphologically similar taxa, distribution maps, 

vernacular names, digital images of types, and preliminary conservation assessments 

following IUCN Red List Categories and Criteria (2012).

Materials and methods

Type specimens of all nearly described species of Eugenia from Madagascar have been 

examined. All Malagasy specimens of Eugenia housed at MO (abbreviations following 

Thiers (2015)) have been viewed by NS (through May 2014), as have all duplicates 

housed at his institution (KSP). Specimens housed at G and P have been studied by 

MC and PP. Duplicates of many specimens housed at KSP also have been consulted by 

NS. Specimens cited here also were compared against all material of Eugenia from the 

Comoros and Mascarenes housed at MO through May 2014. All specimens housed 

at MO should have duplicates at P and TAN given existing exchange agreements, al-

though none at TAN have been seen by the authors. Accession numbers of specimens 

(where indicated) follow the herbarium acronym with a hyphen, whereas barcode 

numbers are enclosed in square brackets.

Studies of Malagasy Eugenia – IV: Seventeen new endemic species...


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 (Snow 1997; Snow et al. 2003). Terminology large-

ly follows earlier works for Malagasy Eugenia (e.g., Snow et al. 2012) but with greater 

consistency regarding general terminology (Beentjie 2010). Terms specific to Myrtaceae 

have been minimized, but may include anthopodium, metaxyphyll, brachyblast, bracte-

ole, monad, and triad, given their descriptive utility in Myrtaceae (see Briggs and John-

son 1979). The collection number cited for each specimen follows immediately after the 

senior collector, in accordance with most database systems now in use. Geocoordinates 

in square brackets were determined retrospectively by various workers.

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 on-

line through the MNHN Vascular Plant Database (

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 (IUCN 2012). The calculations of Area of Occupancy (AOO), 

Extent of Occurrence (EOO) and number of subpopulations follow the methodology 

of Callmander et al. (2007), using a grid cell size to calculate AOO of 3 × 3 km.

Results and taxonomic treatments

Eugenia andapae

 N. Snow, sp. nov.

holotype (Figure 1):

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 (holo-

type: MO-6277713!; isotypes: KSP [KSP000041]!, P [P05097480]!, TAN).


 Shrubs or trees 4–12 m tall; bark of main bole unknown. Vegeta-

tive and reproductive parts mostly glabrous except as noted. Branchlets laterally com-

pressed, 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 oppo-

site, thinly coriaceous (dried material cracking with only moderate pressure), discolor-

ous, matte above and below, venation brochidodromous. Axillary colleters absent. 

Petioles 3–4 mm long, slightly striate below, flattened above, epunctate. Leaf blades 

Neil Snow et al.  /  PhytoKeys 49: 59–121 (2015)


Figure 1. Holotype specimen of Eugenia andapae (MO).

Studies of Malagasy Eugenia – IV: Seventeen new endemic species...


(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 pro-

jecting prominently, straight or only curving slightly towards margin, the secondaries 

connected at their ends by moderately arching connecting veins, tertiary veins pro-

jecting 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 mon-

ad; 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, 



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 7).

Habitat and ecology. 

Humid forests, riparian areas and near summit of Ambodi-

satrana; ca. 200–1540 m.

Conservation status. 

With an Extent of Occurrence (EOO) of 1586 km


, and 

Area of Occupancy (AOO) of 45 km

and five subpopulations, two of which are situ-

ated within the protected area network (Anjanaharibe-Sud, Marojejy), Eugenia anda-

pae is assigned a preliminary risk of extinction of “Vulnerable” [VU B1ab(iii)+2ab(iii)] 

following the IUCN Red List Categories and Criteria (IUCN 2012).

Neil Snow et al.  /  PhytoKeys 49: 59–121 (2015)


Figure 2. Distribution of new Eugenia species in Madagascar with selected Protected Areas (hatched): 

E. bemangidiensis (crosses), E. razakamalalae (triangle), E. richardii (squares), E. tiampoka (stars), and 

E. wilsoniana (circles).

Studies of Malagasy Eugenia – IV: Seventeen new endemic species...



 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  Eugenia andapae is similar to taxa first described by Scott (1979) from the 

Mascarenes in Monimiastrum A.J. Scott, which were reduced to synonymy under Eu-

genia (Snow 2008). The relatively long and broadly elliptic to obovate leaf blades with 

cordate bases, and the relatively straight and projecting abaxial secondary veins are 

diagnostic among other species of Eugenia  in Madagascar. On herbarium material 

the abaxial tertiary veins also protrude slightly despite being thin. The label of the 

type specimen indicates that the fleshy fruits typically are crowned by the calyx lobes, 

although this was not true for the specimen from the Beamalona River.

Specimens examined.

 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, Anjia-

lavabe, Ankiakabe, 14°09'50"S, 49°22'47"E, 952 m, 11 Feb. 2007, R. Razakama-

lala 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).

Eugenia barriei

 N. Snow, sp. nov.

holotype: (Figure 3):

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. Ra-

mananjanahary & A. Rabezafy (holotype: MO-4849778!).


 Shrubs to 3 m tall; bark of main bole unknown. Vegetative and re-

productive parts (where indicated) bearing a moderately dense, shortish indumentum, 

the individual trichomes dibrachiate or not, appressed to somewhat reflexed (appear-

ing 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 

Neil Snow et al.  /  PhytoKeys 49: 59–121 (2015)


Figure 3. Holotype specimen of Eugenia barriei (MO).

Studies of Malagasy Eugenia – IV: Seventeen new endemic species...


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 sur-

face 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-intramargin-

al 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); sta-

mens 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., Barrie 2004, 2005).


 Flowering confirmed only for the middle of November; fruiting likely 

late November through December.


 Known only from near Port Bergé in Mahajanga Province (Figure 4).

Conservation status.

 With only one collection known from Central-western 

Madagascar collected in an unprotected and threathened dry forest, Eugenia barriei 

is assigned a preliminary risk of extinction of “Critically Endangered” [CR A3c] fol-

lowing the IUCN Red List Categories and Criteria (IUCN 2012). In the absence of 

effective protection and the high human pressure on these forests, it is unlikely that the 

forest will persist beyond 3 generations of Eugenia barriei (ca. 30 years).


 The type specimen of Eugenia barriei initially was determined as E. 

tropophylla H. Perrier. The latter species and the varieties described by Perrier de la 

Bâthie (1953a,b) do not represent a single taxon, which even a cursory glance at the 

numerous syntypes (at P) will reveal.

Neil Snow et al.  /  PhytoKeys 49: 59–121 (2015)


Figure 4. Distribution of new Eugenia species in Madagascar with selected Protected Areas (hatched): 

E. barriei (star), E. delicatissima (squares), E. malcomberi (triangle), E. ranomafana (circles), and E. rave-

lonarivoi (crosses).

Studies of Malagasy Eugenia – IV: Seventeen new endemic species...


Among taxa from southeastern Africa, Eugenia barriei resembles some specimens 

of E. capensis subsp. 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.

Eugenia barriei also has gross morphological similarity to the widespread and rela-

tively 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]).

Eugenia bemangidiensis 

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