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Taxonomic treatment of Abrahamia Randrian. & Lowry, a new genus of Anacardiaceae BOISSIERA from Madagascar Armand Randrianasolo, Porter P. Lowry II & George E. Schatz 71 boissiera

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Boissiera 71 was published on 27 December 2017 © Conservatoire et Jardin botaniques de la Ville de Genève boissiera Systematic Botany Monographs vol.71 Boissiera is indexed in: BIOSIS ® ISSN 0373-2975 / ISBN 978-2-8277-0087-5 Taxonomic treatment of Abrahamia Randrian. & Lowry, a new genus of Anacardiaceae from Madagascar

Armand Randrianasolo Porter P. Lowry II George E. Schatz Addresses of the authors AR William L. Brown Center, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A. [email protected]

PPL Africa and Madagascar Program, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A.

Institut de Systématique, Evolution, Biodiversité (ISYEB), UMR 7205, Centre national de la Recherche scientifique/Muséum national d’Histoire naturelle/École pratique des Hautes Etudes, Université Pierre et Marie Curie, Sorbonne Universités, C.P. 39, 57 rue Cuvier, 75231 Paris CEDEX 05, France.

GES Africa and Madagascar Program, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A. Taxonomic treatment of Abrahamia (Anacardiaceae) 7

Abstract he Malagasy endemic genus Abrahamia Randrian. & Lowry (Anacardiaceae) is T described and a taxonomic revision is presented in which 34 species are recognized, including 19 that are described as new. The previously known taxa from Mad- agascar have historically been treated in the genus Protorhus Engl. along with the type species from southern Africa, Protorhus longifolia (Bernh.) Engl. However, recent morphological and molecular phylogenetic studies have shown that the African species is more closely related to Ozoroa Delile and the Malagasy taxa comprise a distinct clade. Therefore the Malagasy taxa could no longer be accommodated in Protorhus, necessi- tating the description a new genus, described as Abrahamia. In the present revision, species of Abrahamia are recognized and distinguished from one another on the basis of morphological characters, primarily of the leaves, inflorescences, flowers and fruits, in combination with eco-geographic features. A total of 15 new combinations are made for taxa from Madagascar previously placed in Protorhus, and lectotypes are designated for 11 names. A key to the species is provide in English and French, and a full description is presented for each species, along with a list of material examined and a distribution map; an illustration is also included for each of the new species. Preliminary conservation assessments following the IUCN Red List criteria indicate that 17 species, representing 50% of the total, are threatened, of which 5 are considered “Endangered” [EN] and 12 “Vulnerable” [VU].

Résumé e genre Abrahamia Randrian. & Lowry (Anacardiaceae), endémique de Madagascar, L est décrit et une révision taxonomique est proposée avec 34 espèces reconnues, dont 19 sont nouvellement décrites. Les espèces précédemment décrites ont été jusqu’alors attribuées au genre Protorhus Engl., qui a pour espèce type, Protorhus longifolia (Bernh.) Engl., d’Afrique australe. Des études morphologiques et de phylogénie moléculaire récentes ont montré que l’espèce africaine est plus proche du genre Ozoroa Delile. Les représentants malgaches appartiennent à un clade distinct et donc ne peuvent plus être inclus dans Protorhus, nécessitant la description d’un nouveau genre, décrit ici sous le nom Abrahamia. Dans le cadre de la présente révision, les espèces d’Abrahamia sont décrites et distinguées les unes des autres à partir des caractères morphologiques, prin- cipalement des feuilles, des inflorescences, des fleurs et des fruits, en combinaison avec des caractères éco-géographiques. Un total de 15 combinaisons nouvelles sont faites pour les taxons de Madagascar précédemment placés dans Protorhus et des lectotypes sont désignés pour 11 noms. Une clé d’identification des espèces est fournie en anglais et en français, et une description complète est présentée pour chaque espèce, accom- pagnée d’une liste des échantillons examinés et d’une carte de distribution. Une illustration est également proposée pour chacune des nouvelles espèces. Une évaluation provisoire du statut de conservation des espèces suivant les critères de la Liste Rouge de l’UICN indique que 17 espèces, soit 50% du total, sont menacées, dont 5 sont considé- rées comme “En Danger” [EN] et 12 comme “Vulnérables” [VU].

Keywords ANACARDIaceae – Abrahamia – Protorhus – Madagascar – Conservation – New Species 8 Boissiera 71

Introduction nacardiaceae are a moderately sized family of c. 800 species belonging to 81 cur- A rently recognized genera (Pell et al., 2011). Widely known for economically important plants such as cashew nuts and mangos, the family is most diverse in humid tropical regions, especially in Asia, South America and Africa, although some members occur in temperate areas. Anacardiaceae are also well represented in Madagascar, where 14 native genera and an estimated total of 94 species are found (Madagascar Catalogue, 2017). The family was treated in the Flore de Madagascar et des Comores by Perrier de la Bâthie (1946; see also Perrier de la Bâthie, 1944), who recognized a total of just 38 species in 13 genera. This remained the only available treatment for more than five decades until the first author began a series of investigations aimed at updating the taxonomy of the entire family in Madagascar. Since then the amount of available herbarium material has grown tremendously, primarily through the efforts of Madagascar’s Service Forestier during the 1960s and 70s, and more recently from field work carried out by staff of the Missouri Botanical Garden and other research institutions starting in the 1970s. This has provided the basis for recent revisions of several genera, including Campnosperma Thwaites (Randrianasolo & Miller, 1998), Micronychia Oliv. (Randrianasolo, 2000; Randrianasolo & Lowry, 2009), and Operculicarya H. Perrier (Randrianasolo & Lowry, 2006), along with the description of one new genus, Poupartiopsis Capuron ex J.D. Mitch. & Daly (Mitchell et al., 2006) and new species in several genera, viz. Campno- sperma (Randrianasolo & Lowry, 2004), Operculicarya (Randrianasolo & Lowry, 2015), Poupartia Comm. ex Juss. (Randrianasolo & Miller, 1999), and Spondias L. (Mitchell et al., 2012). Among the genera of Anacardiaceae present in Madagascar that remain to be revised, the largest comprises species long referred to the genus Protorhus Engl. This genus, described 135 years ago (Engler, 1881), was originally circumscribed to include eight species, one of which, P. longifolia (Bernh.) Engl., occurs in southern Africa and the remainder in Madagascar. Several decades later a second African species, P. namaquensis Sprague, was added (Sprague, 1913) but was later transferred to Ozoroa Delile by von Teichman & van Wyk (1994). Perrier de la Bâthie (1944, 1946) described 8 new species from Madagascar, bringing the total number for the island to 15, and more recently Randrianasolo (1998) recognized a total of 19 Malagasy species.

While Madagascar has been regarded as the center of diversity of Protorhus (von Teichman, 1991a, 1991b; von Teichman & van Wyk, 1994, 1996; Randrianasolo, 1998), questions have been raised regarding whether the single African species should be treated as belonging to the same genus as the species present in Madagascar (von Teich- man, 1991a, 1991b; Randrianasolo, 1998; Pell, 2004). Several striking morphological differences distinguish the taxa from these two areas, especially fruit characters. In particular, Malagasy Protorhus have ellipsoidal, symmetrical fruits with ruminate seeds and inseparable cotyledons (with the exception of one species), whereas the fruits of P. longifolia are ovoid to oblong and distinctly asymmetrical, with non-ruminate seeds and separable cotyledons. In a recent molecular phylogenetic study, Pell et al. (2008) tested the monophyly of Protorhus and confirmed that the genus, as historically circumscribed, indeed comprises two distinct clades, one containing all 11 taxa from Mada- Taxonomic treatment of Abrahamia (Anacardiaceae) 9

gascar included in their sample, which is sister to Heeria argentea (Thunb.) Meisn. and Micronychia macrophylla H. Perrier (the only representatives sampled of these genera of 1 and 10 species, respectively), and another in which Protorhus longifolia from Africa is sister to the tropical African species Ozoroa insignis Delile, the only sampled member of this genus of c. 40 species (Kokwaro, 1986).

While Engler (1881) did not indicate a type when he described Protorhus, 70 years later Phillips (1951) designated P. longifolia as the lectotype. In light of the results of the molecular phylogenetic work of Pell et al. (2008), Protorhus, as previously circumscribed, is polyphyletic and the name thus cannot be applied to the Malagasy species. As no other generic name is available for the members of the clade comprising species in Madagascar, it must therefore be recognized as a new genus. In the present study we formally describe Abrahamia Randrian. & Lowry to accommodate these species and provide a comprehensive taxonomic revision in which a total of 34 species are recognized, 19 of which are newly described. We also discuss additional morphological characters that, in combination, can be used to distinguish between the Malagasy taxa assigned to Abrahamia and Protorhus, which is henceforth restricted to a single African species.

Materials and methods

Herbarium studies he taxonomic revision of Abrahamia presented here is based on two phases of her- and data management T barium studies, conducted prior to and after field work. The first phase involved the examination of herbarium specimens available at the Missouri Botanical Garden herbarium (MO) and received on loan from other institutions with important holdings from Madagascar (in particular G, K, and P; acronyms according to Index Herbariorum, 2017). The goal of this phase was to identify and delimit those species for which ade- quate material was available and to identify gaps and/or problems that would require field work to generate additional specimens and make further observations. The second phase, which took place following field work (see below), was aimed at carrying out more detailed morphological analyses that incorporated newly collected material as well as observations made both in the field and through the consultation of herbarium specimens deposited at the two main herbaria in Madagascar, TAN and TEF. This work provided a robust basis for re-assessing species limits, including the delimitation of new species and the preparation of detailed descriptions and an identification key. The terminology used follows Hickey (1973), Radford et al. (1974), Weberling (1989), Harris & Harris (1994), and Ellis et al. (2009).

Data on all collections examined during this study were captured in Tropicos (2017) and are available on-line, along with distribution maps, photos of selected species, and other information, through the Madagascar Catalogue (2017). For collections lacking geo-coordi- nates, these were assigned, when possible, using the on-line gazetteer of Malagasy place names (Schatz & Lescot, 2003) and are indicated in square brackets in the specimens cited under each species. Distribution maps were generated from the data in Tropicos (2017) projected on a base map of the bioclimatic zones of Madagascar (after Cornet, 1974; see Schatz, 2000). 10 Boissiera 71

Field studies Field work, conducted in Madagascar between 1996 and 2013, initially focused on collecting material needed to fill gaps identified during the first phase of herbarium studies, and was later aimed at sampling as many taxa as possible, including those that appeared to be new, in order to provide a robust understanding of morphological and geographic variation within the group. The collection of herbarium specimens followed standard MBG protocols [www.mobot.org/MOBOT/molib/fieldtechbook/pdf/handbook.pdf]. Whenever possible, a minimum of five duplicate specimens were prepared for each collection, two of which were deposited in Madagascar (at TAN and TEF) and the remaining were sent to MO, where the majority of herbarium studies were conducted (a single sheet was mounted at MO, and additional duplicates have been distributed). Fresh material (leaves, inflorescence, flower buds, flowers, and fruits) was also collected and preserved in FAA, and deposited in the spirit collection of the Missouri Botanical Garden. Leaf samples preserved in silica gel were prepared for as many species as possible for molecular studies (see Pell et al., 2008). Finally, mature seeds were collected and provided to the horticulture depart- ment of the Parc Botanique et Zoologique de Tsimbazaza in Antananarivo, Madagascar.

Risk of extinction The conservation status of each species was assessed using the IUCN Red List Categories assessments and Criteria (IUCN, 2012). Data from the collection records in the Madagascar Catalogue (2017) were used to calculate geographic range parameters (area of occupancy AOO, extent of occurrence EOO, and number of localities), and the number of locations (sensu IUCN, 2012) was determined for each species with respect to the most serious threats.

Morphology number of morphological features have proven to be of value for recognizing and taxonomically A and distinguishing species of Abrahamia. These characters are briefly informative characters presented below.

Habit Two major growth forms can be discerned in Abrahamia. All wet forest species are can- opy or midstory trees with straight trunks and conical crowns. By contrast, those occurring in dry areas (i.e., Madagascar’s western slopes and in the far South) or in highly impacted or degraded habitats (in particular on the high plateau) are usually short and/ or scrubby trees (e.g. A. ibityensis (H. Perrier) Randrian. & Lowry and A. itremoensis Randrian. & Lowry). Also, the color of exudate in the bark, observable when a slash is made on the trunk, varies from milky white or translucid/watery white to red.

Leaves All species of Abrahamia have leaves that are entire and distinctly petiolate. Phyllotaxy, however, varies from alternate and subopposite to opposite or verticillate. Most species are evergreen, with the exception of A. humbertii (H. Perrier) Randrian. & Lowry, which is deciduous. Several additional foliar characters are also of taxonomic importance, as follows: Taxonomic treatment of Abrahamia (Anacardiaceae) 11

Blade shape and size The size and shape of the leaf blade vary significantly inAbrahamia and are useful for recognizing and identifying species. The smallest leaves occur in A. minutifolia Randrian. & Lowry and A. louvelii (H. Perrier) Randrian. & Lowry, ranging from 1.8-4 cm long, whereas in A. latifolia (Engl.) Randrian. & Lowry and A. viguieri (H. Perrier) Randrian. & Lowry they are 8-37 cm long. Some species have narrow leaves, such as A. grandidieri (Engl.) Randrian. & Lowry, in which the blade is oblanceolate and at least 3 times as long as wide, while in other species the length to width ratio is closer to 1.5-2 and the shape therefore ranges from widely elliptic or circular to broadly ovate (in A. humbertii) to broadly obovate (as in A. pauciflora (Engl.) Randrian. & Lowry).

Venation All species of Abrahamia have pinnately veined leaves in which the secondary (lateral) veins are either craspedodromous or semicraspedodromous. According to the classification pro- posed by Ellis et al. (2009), in craspedodromous leaves the major secondary veins terminate at the margin whereas in semicraspedodromous leaves they branch near the margin and only one of the branches reaches the margin. Only five members of the genus have semicraspedodromous venation (viz. A. antongilensis Randrian. & Lowry, A. latifolia, A. lecomtei (H. Perrier) Randrian. & Lowry, A. lenticellata Randrian. & Lowry and A. pauciflora), whereas all others have craspedodromous venation. In addition to the overall venation type, three more venation characters are also of taxonomic value – the number and the shape of the secondary veins, and the distance between them – all three of which show considerable variation among species. When used in combination, these characters make it possible to distinguish, for example, species with parallel and very closely spaced (0.5-3 mm) secondary veins (e.g., A. grandidieri, A. itremoensis, A. louvelii and A. sericea (Engl.) Randrian. & Lowry) from those with arcuate and more distantly spaced (2-15 mm) secondary veins (such as P. thouvenotii (Lecomte) Randrian. & Lowry, and P. viguieri). Another useful venation character in Abrahamia is the pattern of tertiary veins. Two groups of species can be distinguished, one with prominent and visible tertiary venation on at least one of the leaf surfaces and the other in which it is impressed or inconspicuous reticulate.

Leaf surface The surface of leaf blades in Abrahamia species is glabrous or it can have indument com- indument prising various types of trichome. The presence or absence of indument has proven to be particularly useful in delimiting species. Six general indument types are found within the genus, in addition to the glabrous condition. The definitions of these types, as presented below, are based in large part on Harris & Harris (1994). We use ‘pubescent’ in the narrow sense (Harris & Harris, 1994) rather than as a general term to refer to all types of indument comprising trichomes of various orientation, shape, texture.

1. Lanate: densely covered with long and tangled trichomes (e.g., A. viguieri). 2. Pubescent: with various types of short and soft trichomes (e.g., A. turkii Randrian. & Lowry). 3. Puberulent (puberulous): minutely pubescent (e.g., A. itremoensis). 4. Velutinous: with short, soft, spreading trichomes. This type is only found on the abaxial leaf surface of a single species, A. antongilensis. 12 Boissiera 71

5. Strigose: very short and stiff appressed trichomes. 6. Scabrous: very short, stiff, erect trichomes. This type occurs on the leaves of A. deflexa (H. Perrier) Randrian. & Lowry.

Petiole The length of the petiole is also useful for distinguishing species within Abrahamia. For example, A. buxifolia (H. Perrier) Randrian. & Lowry and A. itremoensis have short petioles (2-5 mm and 1-3 mm, respectively) while the petiole is very long in A. longipetiolata (H. Perrier) Randrian. & Lowry (15-30 mm) and A. sambiranensis Randrian. & Lowry (20-58 mm).

Inflorescences In Anacardiaceae as a whole, various inflorescence characters such as structure, size, and type can be taxonomically informative (Wannan & Quinn, 1991; Kearns & Inouye, 1993). Barfod (1988) hypothesized phylogenetic trends among South American Anacardiaceae using inflorescence morphology, andW annan & Quinn (1992) examined the affinities of the genusLaurophyllus Thunb. using inflorescence structure. InAbra - hamia, inflorescence structure, length, type, and position are useful for recognizing and delimiting species. The terminology used here is that of Weberling (1989). The inflorescences of all species ofAbrahamia are compound and erect, and are either panicles or thyrses. Most species (such as A. antongilensis and A. deflexa) have a terminal inflorescence while in some others (e.g.,A. elongata Randrian. & Lowry, A. latifolia, A. lecomtei and A. littoralis Randrian. & Lowry) it is axillary. However, a few species (such as A ditimena (H. Perrier) Randrian. & Lowry and A. turkii) have both axillary and terminal inflorescences in the same individual and often even on the same branch.

Flowers The flowers ofAbrahamia are actinomorphic, small (2-4 mm long), white to yellow in color, 4-, 5-, or rarely 6-merous, and borne on a short and articulate pedicel. Two floral characters are of particular taxonomic value.

Corolla While the petals in Abrahamia are consistently alternisepalous, their size, which is significantly larger than the sepals, is useful for separating certain species. For example, A. ditimena and A. delphinensis Randrian. & Lowry, which have small leaves, velutinous fruits, and overlapping distributions, can be separated by the size of their petals, which are 2.5-3 × 1.5-2 mm in A. ditimena vs 2 × 1 mm in A. delphinensis.

Disk The presence or absence of trichomes on the surface of the disk is also helpful for distinguishing species of Abrahamia. For example, both A. buxifolia and A. itremoensis have small leaves and similar ranges, but the former has a glabrous disc while that of the latter has evident indument. Taxonomic treatment of Abrahamia (Anacardiaceae) 13

Fruits The fruits of Abrahamia are always drupes with a resiniferous mesocarp and thin papyraceous endocarp, but several characters are very useful for delimiting species.

Shape Fruits in Abrahamia can vary from globose or ellipsoid to ovoid or obovoid, depending on the species.

Size Fruit dimensions are also useful for delimiting species within Abrahamia, varying from 1-1.5 × 0.5-0.8 cm in A. minutifolia, which has the smallest fruits, to 3-3.5 × 1.5-1.8 cm in A. ellipticarpa Randrian. & Lowry, which has the largest fruits in the genus.

Surface indument The fruits of Abrahamia are either glabrous or velutinous, and this character is very useful for distinguishing species. Eight species have fruits with indument, viz. A. betamponensis Randrian. & Lowry, A. buxifolia, A. delphinensis, A. ditimena, A. itremoensis, A. lenticellata, A. minutifolia, and A. sericea, while those of the remaining 25 species are glabrous (fruits of A. pauciflora are unknown).

Taxonomic treatment Abrahamia Randrian. & Lowry, gen. nov.

Typus: Abrahamia ditimena (H. Perrier) Randrian. & Lowry (≡ Protorhus ditimena H. Perrier).

Diagnosis Dioecious trees or shrubs. Leaves simple, persistent, opposite, subopposite, verticillate or alternate, chartaceous, subcoriaceous or coriaceous, lateral veins parallel, arcuate, forming an obtuse to acute angle with the midrib toward the base of the blade. Inflorescence erect, a thyrse or panicle. Flowers (4-)5(-6)-merous; pedicel short, articulated; sepals much smaller than petals; stamens alternipetalous, filament broadened at the base (sterile in female flowers, forming staminodes), anthers dorsifixed, introrse, glabrous; ovary pubescent or glabrous (rudimentary or absent in male flowers), with 1 locule containing a single subpendulous or pendulous ovule. Fruits drupaceous, ellipsoidal or ovoid, symmetrical or slightly asymmetrical, resiniferous; seeds ruminate; cotyledons usually inseparable (except in Abrahamia ibityensis (H. Perrier) Randrian. & Lowry, in which the resiniferous canals are only peripheral).

Description Dioecious trees or shrubs; bark with white, translucent or red resinous latex. Leaves alternate, opposite or subopposite (verticillate in one species), simple, coriaceous, subcoriaceous or chartaceous, persistent (deciduous in A. humbertii). Venation pinnate, craspedodromous or semicraspedromous, midvein usually flat, rarely very slightly raised above, prominently raised below (rarely on both surfaces), secondary veins parallel or nearly so, straight or arcuate, prominent below or on both surfaces, forming an obtuse to acute angle with the midrib toward the base of the blade. Inflorescence terminal and/ or axillary, an erect thyrse or panicle, or intermediate and forming a paniculiform thyrse. Flowers usually 5-merous (sometimes 4- or 6-merous); sepals much smaller than petals; 14 Boissiera 71

stamens 5 or 4 (rarely 6), alternipetalous, filaments straight, flattened, broadened at the base, inserted basally on the outer surface of the disk, anthers dorsifixed, introrse, dehis- cent with longitudinal slits (stamens reduced in size and sterile in female flowers, forming staminodes); disk usually cupuliform and crenulate in male flowers, annular and cupuliform in female flowers; ovary with 1 locule containing a single pendulous or subpendulous or ovule on a short funiculus, placentation subapical (ovary rudimentary or absent in male flowers); style one, short, apically trifid or unbranched with 3 capitate stigmatic lobes. Fruits drupaceous, ellipsoidal or ovoid, symmetrical or slightly asymmetrical, exo- carp usually with longitudinal striations on surface, glabrous or velutinous, mesocarp resinous, endocarp thin, papyraceous; seed solitary, with resiniferous canals throughout (only peripheral in A. ibityensis), ruminate; cotyledons usually inseparable (separable in A. ibityensis).

Etymology The genus is named in honor of the late Jean Prosper Abraham (1936-1996), a well- known Malagasy forester with a remarkable knowledge of the woody plants of Madagas- car and who contributed much to our knowledge of the island’s rich and diverse flora.

Notes Abrahamia includes a total of 34 species, all endemic to Madagascar, including 15 species formerly placed in Protorhus (e.g., by Perrier de la Bâthie, 1944, 1946), to which 19 additional species are newly described here. With the recognition of Abrahamia as a separate genus, Protorhus is now restricted to a single species in Africa, P. longifolia. The two genera differ in several flower and fruit characters, as indicated byP ell (2004) and Pell et al. (2011), and summarized in Table 1.

Table 1. Abrahamia Protorhus Morphological features distinguishing Abrahamia Fruit shape Ellipsoidal or ovoid, Ovoid, mango-shaped or Randrian. & Lowry from Protorhus Engl. radially symmetrical transversely oblong, asymmetrical

Seed Resiniferous canals present, Resiniferous canals lacking, making the seed ruminate; seed not ruminate; cotyledons inseparable cotyledons separable

Number of styles 1 (apically trifid or with 3 3, slightly connate at the capitate stigmas) base or completely free Taxonomic treatment of Abrahamia (Anacardiaceae) 15

Abrahamia is represented in all of Madagascar’s major vegetation types (Moat & Smith, 2007) and broad bio-climatic zones (Cornet, 1974; Schatz, 2000). Species can be found in humid forest in the eastern and Sambirano regions (A. antongilensis, A. sambiranensis), wooded grassland and bushland in the center of the island (A. ibityensis, A. buxifolia), western dry forest (A. deflexa, A. humbertii), and southwestern subarid spiny forest-thicket (A. phillipsonii Randrian. & Lowry). Members of the genus occur on many types of substrate and at a wide range of elevations, from littoral forest on sand at the sea level (A. nitida (Engl.) Randrian. & Lowry, A. suarezensis Randrian. & Lowry) to humid forest on lateritic soil at altitudes up to 2000 m (A. buxifolia, A. itremoensis).

Species of Abrahamia were described by Perrier de la Bâthie (1946) as polygamo-dioecious, indicating that he interpreted them as having both unisexual and hermaphro- ditic flowers on the same individual. However, careful observation has shown that all members of the genus are functionally dioecious. Female flowers have rudimentary, sterile stamens that are reduced in size, whereas male flowers either have an ovary that is rudimentary and non-functional or lack an ovary altogether. 16 Boissiera 71

Key to the species of Abrahamia [Numbers in parentheses indicate the previous couplet leading to the present one]

1. Leaves completely glabrous on both surfaces (occasionally with extremely small indument visible only under a dissecting microscope on or adjacent to the midvein on the abaxial surface)...... 2 1a. Leaves with at least a few evident trichomes, sometimes restricted to the midvein beneath...... 20

2. Leaves usually verticillate (3 per node, sometimes one leaf broken off but leaving a scar); E coast littoral forest (occasionally to 200 m elevation)...... 23. A. nitida 2a. Leaves alternate or opposite to subopposite...... 3

3. Largest leaf blade at least 15 cm long...... 4 3a. Largest leaf blade less than 15 cm long (rarely to 17 cm in A. oblongifolia and A. littoralis)...... 6

4. Petiole stout, c. 4-5 mm in diam.; young branches with corky bark; littoral and low elevation humid forest, Maroantsetra S to Tampolo...... 15. A. latifolia 4a. Petiole slender, no more than 2(-2.5) mm in diam.; young branches without corky bark; humid forest...... 5

5. Largest leaf with c. 50-60 pairs of secondary veins; fruits ellipsoid, covered with dense, short indument; Betampona...... 2. A. betamponensis 5a. Largest leaf with c. 30-45 pairs of secondary veins; fruits obovoid, glabrous; Lokobe and Tsaratanana...... 19. A. lokobensis

6 (3). Leaves elliptic to ovate (rarely a few slightly obovate), broadest at the middle or in the proximal part of the blade...... 7 6a. Leaves obovate to oblanceolate, nearly all broadest in the distal part of the blade...... 11

7. Leaves narrowly elliptic to narrowly ovate (rarely a few slightly obovate), at least (2.5-) 3 times as long as wide; dry forest, Isalo to Andohahela...... 27. A. phillipsonii 7a. Leaves elliptic to ovate, sometimes broadly so (rarely a few slightly obovate), no more than 1.5-2 times as long as wide...... 8

8. Fruits covered with dense short indument; flowers 4-merous (rarely 5-merous); humid forest, Marojejy and Anjanaharibe-Sud S to Ranomafana and Ivohibe...... 17. A. lenticellata 8a. Fruits glabrous; flowers 5-merous (unknown inA. darainensis)...... 9 Taxonomic treatment of Abrahamia (Anacardiaceae) 17

9 (8). Leaves orbicular to broadly elliptic (rarely a few broadly ovate), blades no more than 2 times as long as wide, subcoriaceous, apex rounded (rarely broadly acute); dry forest, calcareous substrates, Ankarana, Bemaraha...... 12. A. humbertii 9a. Leaves elliptic to ovate (occasionally a few broadly ovate or obovate), most blades at least 2 times as long as wide, chartaceous, apex acute to acuminate...... 10

10. Blade of largest leaf to c. 6.5 cm long, often discolorous, the upper surface dark green to chocolate brown, apex emarginate or rounded to acute; dry western forest (Bemaraha, Ketsepy, Antsalova, Marovoay, Mahajanga)...... 6. A. deflexa 10a. Blade of largest leaf c. 8 cm long, concolorous, the upper surface drying grayish green, apex acuminate; humid forest, Daraina...... 5. A. darainensis

11 (6). Largest leaf with 40-70 pairs of secondary veins; humid eastern forest ...... 30 A. sericea 11a. Largest leaf with 10-40 pairs of secondary veins ...... 12

12. Inflorescences all axillary; leaves alternate; E coast littoral forest, Tampolo S to Mahabo ...... 18. A. littoralis 12a. Inflorescences terminal and sometimes also axillary; leaves opposite to subopposite ...... 13

13. Tertiary venation obscure on abaxial surface ...... 14 13a. Tertiary venation visible on abaxial surface, at least toward the margins ...... 16

14. Secondary veins closely spaced, 6-9 pairs per cm in median portion of largest leaf, blade distinctly glaucous on adaxial surface, nearly always strongly discolorous when dry, often chocolate brown above (sometimes obscured by glaucescence); fruit velutinous (sometimes glabrescent); dry forest, central high plateau, Itremo, Ibity, Ambatofinandrahana ...... 3. A. buxifolia 14a. Secondary veins widely spaced, 3-5 pairs per cm in median portion of largest leaf, blade neither glaucous (sometimes weakly so in A. suarezensis) nor strongly discolorous when dry, various shades of green but not chocolate brown above; fruit glabrous...... 15

15. Largest leaf with 15-30 pairs of secondary veins, margins distinctly revolute; trees 15-20 m tall; littoral and dry forest, Mt. d’Ambre, S of Vohémar and Masoala...... 28. A. revoluta 15a. Largest leaf with 12-15 pairs of secondary veins, margins slightly thickened but not revolute; trees to 10(-12) m tall; N dry and coastal forests, Antsiranana, Vohémar, Daraina...... 31. A. suarezensis

16 (13). Leaves with venation craspedodromous, secondary veins all clearly reaching the margin (rarely a few terminating away from the margin in A. oblongifolia)...... 17 16a. Leaves with venation semicraspedodromous, at least some secondary veins branching and not reaching the margin...... 19 18 Boissiera 71

17 (16). Largest leaf with 40-70 pairs of secondary veins, largest blade 9-15 cm long ...... 30. A. sericea 17a. Largest leaf with 15-33 pairs of secondary veins, largest blade 2.5-7.5(-8.5) cm long...... 18

18. Blade of largest leaf at least (4.5-) 5 cm long, coriaceous, rigid; mature fruit 2-3 × 1.4-1.6 cm ...... 8. A. ditimena 18a. Blade of largest leaf no more than 4-(4.5) cm long, subcoriaceous, not rigid; mature fruit 1-1.5 × 0.5-0.8 cm ...... 21. A. minutifolia

19 (16). Largest leaf blade 8-14(-17) cm long, petiole c. 2 mm in diam.; fruits glabrous; dry to humid forest, Nossi Be, Antsiranana area ...... 24. A. oblongifolia 19a. Largest leaf blade 3.5-6 cm long, petiole c. 0.5-1 mm in diam.; fruits with short dense indument; littoral forest and low-elevation humid forest, Tolagnaro area ...... 7. A. delphinensis

20 (1). Blade of largest leaf at least 10 cm long...... 21 20a. Blade of largest leaf no more than 10 cm long...... 34

21. Largest leaves with at least 40 pairs of secondary veins...... 22 21a. Largest leaves with fewer than 40 pairs of secondary veins...... 26

22. Inflorescences all axillary...... 23 22a. Inflorescences terminal and sometimes also axillary...... 24

23. Leaves with evident tertiary venation, apex flat; fruits glabrous; humid forest, Marojejy, Anjanaharibe-Sud, Masoala, Rantabe, Ambalabe ...... 10. A. elongata

23a. Leaves lacking evident tertiary venation, apex usually folded in pressed material; fruits pubescent when young, glabrescent at maturity; humid forest, Ambilobe, Ampondrabe, Ambato, Kalabenono-Galoka, Ambanja, Andavakoera ...... 29. A. sambiranensis

24 (22). Secondary veins in median portion of leaf blade closely spaced, c. 5-6 per cm; fruit with dense indument; humid eastern forest...... 30. A. sericea 24a. Secondary veins in median portion of leaf blade more widely spaced, c. 3 per cm; fruit glabrous or with evident indument...... 25

25. Leaves with an acute to short acuminate apex (rarely a few rounded); fruits with dense, short indument; flowers 4-merous (rarely 5-merous); humid forest...... 17. A. lenticellata 25a. Leaves with a broadly acute to rounded or retuse apex; fruits glabrous; flowers 5-merous; dry forest...... 11. A. grandidieri Taxonomic treatment of Abrahamia (Anacardiaceae) 19

26 (21). Inflorescences all axillary...... 27 26a. Inflorescences terminal and sometimes also axillary...... 29

27. Abaxial surface of leaves with dense, erect to spreading light brown lanate indument...... 34. A. viguieri 27a. Abaxial surface of leaves glabrous or with spare, short indument...... 28

28. Longest petiole 5-10 mm; eastern humid, low to mid-elevation forest.... 16. A. lecomtei 28a. Longest petiole (18-)20-30 mm; E coast littoral forest, Tampolo S to Mahabo...... 18. A. littoralis

29 (26). Leaf apex long acuminate to nearly caudate, blade chartaceous...... 30 29a. Leaf apex acute to retuse or emarginate, sometimes short-acuminate (acumen < 1 cm long), blade chartaceous to subcoriaceous or coriaceous...... 32

30. Abaxial surface of leaves densely covered with reddish-brown lanate indument, venation craspedodromous; eastern littoral forest to premontane forest on sand or laterite, Maroantsetra, Toamasina, Zahamena PN, Fort-Carnot...... 34. A. viguieri 30a. Abaxial surface of leaves glabrous or with scattered, white sericeous indument, venation semi-craspedodromous ...... 31

31. Adaxial surface of leaves always glabrous, widest blade to 3.5(-4) cm; fruit velutinous; eastern humid forest ...... 17. A. lenticellata 31a. Adaxial surface of leaves glabrous or with scattered white appressed indument, widest blade at least 4 cm; fruit glabrous; north-eastern humid forest, Masoala, Maroantsetra ...... 1. A. antongilensis

32 (29). Leaves with 10-20 pairs of widely spaced secondary veins, only 1 or 2 per cm in median portion of blade; eastern and north-eastern humid forest, Maroantsetra, Manongarivo, Andapa, Sambava, Ranomafana, Mantadia, Zahamena ...... 32. A. thouvenotii 32a. Leaves with 20-40 pairs of narrowly spaced veins, 2-6 per cm in median portion of blade ...... 33

33. Leaves obovate, never more than 2.5 times as long as wide, coriaceous; inflorescences terminal and sometimes also axillary; fruits covered with dense indument; humid eastern forest...... 8. A. ditimena 33a. Leaves oblanceolate (sometimes narrowly elliptic), at least 3 times as long as wide, chartaceous to coriaceous, flexible; terminal and axillary; fruits glabrous; along rivers and streams, western and southern Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy...... 11. A. grandidieri

34 (20). Largest leaves with at least 40 pairs of secondary veins...... 35 34a. Largest leaves with fewer than 40 pairs of secondary veins...... 36 20 Boissiera 71

35 (34). Secondary veins in median portion of leaf blade closely spaced, c. 5-6 per cm; fruit with dense indument; humid eastern forest...... 30. A. sericea 35a. Secondary veins in median portion of leaf blade more widely spaced, c. 3 per cm; fruit glabrous; along rivers and streams, western and southern Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy...... 11. A. grandidieri

36 (34). Largest leaves with no more than 20 pairs of secondary veins...... 37 36a. Largest leaves with more than 20 pairs of secondary veins...... 46

37. Leaves elliptic to orbicular or slightly ovate, broadest at or below the middle...... 38 37a. Leaves obovate to oblanceolate, broadest beyond the middle...... 40

38. Leaves orbicular to broadly elliptic (rarely a few broadly ovate), no more than 2 times as long as wide; dry forest, Ankarana, Bemaraha...... 12. A. humbertii 38a. Leaves elliptic to narrowly elliptic, at least 2 times as long as wide...... 39

39. Largest leaves 2.5-3.6 cm wide; humid and transition forest, Manambato, Daraina, Androranga River valley near Sambava...... 4. A. capuronii 39a. Largest leaves 1.5-2(-2.2) cm wide; dry forest, Isalo to Andohahela...... 27. A. phillipsonii

40 (37). Inflorescences all axillary, female inflorescences with no more than 5 flowers (male inflorescences unknown); humid forest, Nosy Be, Ampasindava, Ambongo-Boina...... 26. A. pauciflora 40a. Inflorescences terminal and sometimes also axillary, with 8 to numerous flowers (rarely only 5 in female inflorescences ofA. suarezensis)...... 41

41. Blade of largest leaf 2-3(-4) cm long; fruits 1-1.5 cm long...... 22. A. minutifolia 41a. Blade of largest leaf at least (3.5-)4 cm long; fruits 1.5-2.5 cm long...... 42

42. Fruit densely covered with short indument; secondary veins semicraspedodromous, at least some branching and not reaching the margin; littoral forest, Tolagnaro area...... 7. A. delphinensis 42a. Fruit glabrous; secondary veins craspedodromous, all clearly reaching the margin.....43

43. Petiole stout, (1.5-)2-3 mm in diam. on largest leaves, tertiary venation distinctly raised on abaxial surface; eastern and north-western humid forest, Ambanja, Marojejy, Anajanaharibe Sud, Ranomafana, Analamazaotra, Zahamena ...... 32. A. thouvenotii 43a. Petiole slender, no more than 1.5 mm in diam. on largest leaves, tertiary venation obscure to weakly raised on abaxial surface...... 44 Taxonomic treatment of Abrahamia (Anacardiaceae) 21

44 (43). Secondary veins closely spaced, 5-6 pairs per cm in median portion of largest leaf; humid eastern forest, Analamazaotra, Ambalabe ...... 21. A. louvelii 44a. Secondary veins widely spaced, 3-4 pairs per cm in median portion of largest leaf.....45

45. Inflorescences 1-3(-3.5) cm long; leaf margins undulate, secondary veins c. 10-12(-14) pairs; northern dry and coastal forest, Antsiranana, Vohémar, Daraina...... 31. A. suarezensis 45a. Inflorescences 4-9 cm long; leaf margins plane or only weakly undulate, secondary veins 14 or more pairs; humid forest, Ankazobe, E of Fianarantsoa, Ranomafana, Andohahela (Parcel #1), Ivohibe...... 33. A. turkii

46 (36). Venation semicraspedodromous, at least some secondary veins branching and not reaching the margin; fruit with evident indument ...... 17. A. lenticellata 46a. Venation craspedodromous, secondary veins all clearly reaching the margin; fruit glabrous or with evident indument ...... 47

47. Petiole of the largest leaves at least 10 mm long ...... 48 47a. Petiole of the largest leaves less than 10 mm long ...... 53

48. Inflorescences all axillary; east coast littoral forest, Tampolo S to Mahabo...... 18. A. littoralis 48a. Inflorescences terminal and sometimes also axillary...... 49

49. Tertiary veins of leaves barely visible, almost appearing absent; corolla of female flowers glabrous on outer surface...... 20. A. longipetiolata 49a. Tertiary veins of leaves visible at least on one surface; corolla of female flowers with indument on outer surface...... 50

50. Leaves no more than 2.5 times as long as wide; fruit with evident indument ...... 51 50a. Leaves at least 3 times as long as wide; fruits glabrous ...... 52

51. Leaves chartaceous; littoral forest, Tolagnaro area...... 7. A. delphinensis 51a. Leaves coriaceous; humid eastern forest...... 8. A. ditimena

52 (50). Leaves obovate to elliptic; secondary veins 18-33 pairs, 1-4 mm apart; NE humid and sub-humid transitional forest (Daraina, Vohémar)...... 4. A. capuronii 52a. Leaves oblanceolate; secondary veins 35-40-(47) pairs, 0.5-5 mm apart; dry and sub- humid forest, W and SW...... 11. A. grandidieri

53 (47). Secondary venation weakly raised on upper leaf surface; fruits glabrous ...... 54 53a. Secondary venation distinctly raised on upper leaf surface; fruits with evident indument...... 60 22 Boissiera 71

54 (53). Fruits 3-3.5 cm long, ellipsoid; infructescence axes stout, 1.5-2 mm in diam.; largest leaves 8-9 × 2.5 cm, apex acute; littoral forest, Tolagnaro area...... 9. A. ellipticarpa 54a. Fruits 1.5-2.6 cm long, ovoid or ellipsoid; infructescence axes slender, to c. 1(-1.2) mm in diam.; largest leaves either smaller than 8 × 2 cm or with a rounded or emarginate apex...... 55

55. Leaves obovate to oblanceolate...... 56 55a. Leaves elliptic to oblong, rarely a few slightly ovate or obovate...... 59

56. Largest leaves with at least 35 pairs of secondary veins; dry forest..... 11. A. grandidieri 56a. Largest leaves with no more than 25(-28) pairs of secondary veins; humid forest...... 57

57. Secondary veins widely spaced, 3-4 pairs per cm in median portion of largest leaf blade; humid forest, Ankazobe, E of Fianarantsoa, Ranomafana, Andohahela (Parcel #1), Ivohibe ...... 33. A. turkii 57a. Secondary veins closely spaced, 5-6 pairs per cm in median portion of largest leaf blade ...... 58

58. Blade of largest leaf to 4(-4.2) cm long; Analamazaotra, Ambalabe; humid forest...... 21. A. louvelii 58a. Blade of largest leaf at least (4.8-)5 cm long; semi-deciduous and evergreen humid forest, Daraina ...... 25. A. patrickii

59 (55). Petiole slender, < 1 mm in diam.; cotyledons fused; trees c. 3-12 m tall; dry forest, Isalo to Andohahela ...... 27. A. phillipsonii 59a. Petiole stout, 1.2-1.5 mm in diam.; cotyledons separable; shrubs to small trees < 2 m tall; sclerophyllous forest, Ibity...... 13. A. ibityensis

60 (53). Leaves glaucous on adaxial surface, nearly always strongly discolorous when dry, often chocolate brown adaxially (sometimes obscured by glaucescence); dry forest, central high plateau, Itremo, Ibity, Ambatofinandrahana...... 3. A. buxifolia 60a. Leaves not glaucous on adaxial surface, uniform in color or weakly discolorous...... 61

61. Leaves puberulous on adaxial surface; Itremo area...... 14. A. itremoensis 61a. Leaves glabrous on adaxial surface...... 62

62. Leaves chartaceous; littoral forest, Tolagnaro area...... 7. A. delphinensis 62a. Leaves coriaceous; eastern humid forest...... 8. A. ditimena Taxonomic treatment of Abrahamia (Anacardiaceae) 23

Clé des espèces d’Abrahamia [Les chiffres entre parenthèses renvoient au couplet de provenance]

1. Feuilles complètement glabres sur les deux faces (parfois avec un indument très petit sur la surface inférieure, visibles seulement sous microscope, ou de part et d’autre de la nervure médiane)...... 2 1a. Feuilles avec au moins quelques trichomes évidents, parfois restreints à la nervure médiane...... 20

2. Feuilles généralement verticellées (trois par nœud, parfois une feuille manquante mais laissant toujours une cicatrice); forêts littorales orientales (parfois jusqu’à 200 m d’altitude)...... 23. A. nitida 2a. Feuilles alternes ou opposées à subopposées...... 3

3. Limbe foliaire le plus grand mesurant au moins 15 cm de long...... 4 3a. Limbe foliaire le plus grand de moins de 15 cm long (rarement jusqu’à 17 cm chez A. oblongifolia et A. littoralis)...... 6

4. Pétiole robuste, d’env. 4-5 mm de diam.; jeunes rameaux avec une écorce liégeuse; forêts littorales et de basse altitude, de Maroantsetra au nord à Tampolo au sud...... 15. A. latifolia 4a. Pétiole grêle, ne dépassant pas 2(-2.5) mm de diam.; jeunes rameaux sans écorce liégeuse; forêts humides...... 5

5. Feuille la plus grande avec env. 50-60 paires de nervures secondaires; fruit ellipsoïde, couvert d’un indument dense et court; Betampona...... 2. A. betamponensis 5a. Feuille la plus grande avec env. 30-45 pairs de nervures secondaires; fruit obovoïde, glabre; Lokobe et Tsaratanana...... 19. A. lokobensis

6 (3). Feuilles elliptiques à ovées (rarement quelques unes légèrement obovales), plus larges dans la partie proximale...... 7 6a. Feuilles obovales à oblancéolées, presque toutes plus larges dans la partie distale...... 11

7. Feuilles étroitement elliptiques à étroitement ovées (rarement quelques unes obovales), au moins (2.5-)3 fois plus longues que larges; forêts sèches, de l’Isalo à Andohahela...... 27. A. phillipsonii 7a. Feuilles elliptiques à ovées, parfois largement (rarement quelques unes légèrement obovales), pas plus de 1.5-2 fois plus longues que larges...... 8 24 Boissiera 71

8 (7). Fruits couverts d’un indument court et dense; fleurs 4-mères (rarement 5-mères); forêts humides, Marojejy et Anjanaharibe-dans le nord jusqu’à Ranomafana et Ivohibe dans le sud...... 17. A. lenticellata 8a. Fruits glabres; fleurs 5-mères (inconnues chezA. darainensis) ...... 9

9. Feuilles orbiculaires à largement elliptiques (rarement quelques unes largement ovées), limbes pas plus de 2 fois plus longs que larges, subcoriaces, apex arrondi (rarement aigu); forêts sèches sur substrats calcaires, Ankarana, Bemaraha...... 12. A. humbertii 9a. Feuilles elliptiques à ovées (parfois quelques une largement ovées ou obovales), la plupart des limbes au moins 2 fois plus longs que larges, chartacées, apex aigu à acuminé...... 10

10. Limbe de la feuille la plus grande mesurant env. 6.5 cm de long, souvent discolore, la surface supérieure vert foncé à brun-chocolat, apex émarginé ou arrondi à aigu; forêts sèches occidentales (Bemaraha, Ketsepy, Antsalova, Marovoay, Mahajanga)...... 6. A. deflexa 10a. Limbe de la feuille la plus grande env. 8 cm de long, concolore, la surface supérieure gris-vert sur le sec, apex acuminé; forêts humides, Daraina...... 5. A. darainensis

11 (6). Feuille (la plus grande) avec 40-70 paires de nervures secondaires; forêt humide orientale...... 30. A. sericea 11a. Feuille (la plus grande) avec 10-40 paires de nervure secondaires...... 12

12. Inflorescences exclusivement axillaires; feuilles alternes, forêts littorales orientales, de Tampolo au nord jusqu’à Mahabo au sud...... 18. A. littoralis 12a. Inflorescences terminales mais quelquefois aussi axillaires; feuilles opposées à subopposées...... 13

13. Nervures tertiaires invisibles sur la surface inférieure...... 14 13a. Nervures tertiaires visible sur la surface inférieure, du moins vers la marge du limbe.....16

14. Nervures secondaires étroitement espacées, 6-9 paires par cm au niveau de la partie médiane de la plus grande feuille, limbe glauque sur la face supérieure, presque toujours fortement discolore sur le sec, souvent brun-chocolaté sur la surface supérieure (parfois caché par la glaucescence); fruit velu (parfois glabrescent); forêts sèches, partie centrale des hauts plateaux, Itremo, Ibity, Ambatofinandrahana...... 3. A. buxifolia 14a. Nervures secondaires largement espacées, 3-5 paires par cm au niveau de la portion médiane de la feuille la plus grande, limbe ni glauque (sauf un peu chez A. suarezensis) ni fortement discolore sur le sec, divers tons de vert mais jamais brun-chocolaté sur la surface supérieure; fruit glabre...... 15 Taxonomic treatment of Abrahamia (Anacardiaceae) 25

15 (14). Feuille (la plus grande) avec 15-30 paires de nervures secondaires, à marge distinctement révolutée; arbres de 15-20 m de haut; forêts littorales et sèches, Mt. d’Ambre, sud de Vohémar et Masoala ...... 28. A. revoluta 15a. Feuille (la plus grande) avec 12-15 paires de nervures secondaires, à marge légèrement épaissie mais pas révolutée; arbres de 10(-12) m de haut; forêts sèches et côtières du Nord, Antsiranana, Vohémar et Daraina ...... 31. A. suarezensis

16 (13). Feuilles à nervures craspédodromes, toutes les nervures secondaires atteignant la marge (rarement quelques unes n’atteignant pas la marge chez A. oblongifolia)...... 17 16a. Feuilles à nervures semi-craspédodromes, au moins quelques nervures secondaires ramifiées et n’atteignant pas la marge ...... 19

17. Feuille (la plus grande) avec 40-70 paires de nervures secondaires, limbe le plus grand de 9-15 cm long ...... 30. A. sericea 17a. Feuille (la plus grande) avec 15- 33 paires de nervures secondaires, limbe le plus grand de 2.5-7.5(-8.5) cm long...... 18

18. Limbe de la feuille la plus grande d’au moins (4.5-)5 cm de long, coriace, rigide; fruits mûrs de 2-3 × 1.4-1.6 cm ...... 8. A. ditimena 18a. Limbe de la feuille la plus grande ne dépassant pas 4-(4.5) cm de long, subcoriace, non rigide; fruits mûrs 1-1.5 × 0.5-0.8 cm...... 22. A. minutifolia

19 (16). Limbe de la feuille la plus grande de 8-14(-17) cm de long, pétiole env. 2 mm de diam.; fruits glabres; forêts sèches à humides, Nossi Be, Rég. d’Antsiranana...... 24. A. oblongifolia 19a. Limbe de la feuille la plus grande de 3.5-6 cm long, pétiole env. 0.5-1 mm de diam.; fruits à indument dense et court; forêts littorales et humides de basse altitude, Rég. de Tolagnaro...... 7. A. delphinensis

20 (1). Limbe de la feuille la plus grande d’au moins 10 cm de long...... 21 20a. Limbe de la feuille la plus grande ne dépassant pas 10 cm de long...... 34

21. Feuilles les plus grandes avec au moins 40 paires de nervures secondaires...... 22 21a. Feuilles les plus grandes avec moins de 40 paires de nervures secondaires...... 26

22. Inflorescences exclusivement axillaires...... 23 22a. Inflorescences terminales et parfois aussi axillaires...... 24

23. Nervation tertiaire évidente, apex de la feuille plan; fruits glabres; forêts humides, Marojejy, Anjanaharibe-Sud, Masoala, Rantabe, Ambalabe ...... 10. A. elongata 23a. Nervation tertiaire absente, apex de la feuille généralement plié sur les spécimens séchés; fruits jeunes pubescents, glabrescents; forêts humides, Ambilobe, Ampondrabe, Ambato, Kalabenono-Galoka, Ambanja, Andavakoera, Tsaratanana...... 29. A. sambiranensis 26 Boissiera 71

24 (22). Nervures secondaires de la partie médiane du limbe étroitement espacées, env. 5-6 par cm; fruit à indument dense; forêts humides orientales...... 30 A. sericea 24a. Nervures secondaires de la partie médiane du limbe largement espacées, env. 3 par cm; fruit glabre ou à indument clairement visible...... 25

25. Apex de la feuille aigu à courtement acuminé (rarement quelques feuilles à apex arrondi); fruits à indument dense et court; fleurs 4-mères (rarement 5-mères); forêts humides...... 17. A. lenticellata 25a. Apex de la feuille obtus à arrondi ou rétus; fruits glabres; fleurs 5-mères; forêts sèches ...... 11. A. grandidieri

26 (21). Inflorescences exclusivement axillaires...... 27 26a. Inflorescences terminales et parfois aussi axillaires...... 29

27. Surface inférieure des feuilles densément laineuse, à indument brun clair, érigé à étalé ...... 34. A viguieri 27a. Surface inférieure des feuilles glabres ou à indument court, épars...... 28

28. Pétiole le plus long mesurant 5-10 mm; forêts humides orientales de basse et moyenne altitude ...... 16. A. lecomtei 28a. Pétiole le plus long mesurant (18-)20-30 mm; forêts littorales orientales, de Tampolo au nord jusqu’à Mahabo au sud...... 18. A. littoralis

39 (26). Apex de la feuille longuement acuminé, presque caudé, limbe chartacé...... 30 29a. Apex de la feuille aigu à rétus ou émarginé, parfois courtement acuminé (acumen < 1 cm de long), limbe chartacé à subcoriace ou coriace...... 32

30. Surface inférieure des feuilles densément laineuse à indument brun rougeâtre, nervures craspédodromes; forêts littorales orientales sur sable ou forêt de piémont sur latérite, Maroantsetra, Toamasina, Zahamena PN, Fort-Carnot...... 34. A. viguieri 30a. Surface inférieure des feuilles glabres ou à indument blanc soyeux et épars, nervures semi-craspédodromes...... 31

31. Surface supérieure des feuilles toujours glabres, limbe le plus large de 3.5(-4) cm de largeur; fruit velu; forêts humides orientales...... 17. A. lenticellata 31a Surface supérieure des feuilles glabres ou à indument blanc apprimé, limbe le plus large d’au moins de 4 cm de largeur; fruit glabre; forêts humides du nord-est, Masoala, Maroantsetra...... 1. A. antongilensis

32 (29). Feuilles à 10-20 paires de nervures secondaires largement espacées, seulement 1 ou 2 par cm sur la partie médiane du limbe; forêts humides de l’est et du nord-est Maroantsetra, Manongarivo, Andapa, Sambava, Ranomafana, Mantadia, Zahamena...... 32. A. thouvenotii Taxonomic treatment of Abrahamia (Anacardiaceae) 27

32a. Feuilles à 20-40 paires de nervures étroitement espacées, 2-6 par cm au niveau de la partie médiane du limbe ...... 33

33. Feuilles obovales, jamais plus de 2.5 fois plus longues que larges, coriaces; inflorescences terminales et parfois aussi axillaires; fruits à indument dense; forêts humides orientales...... 8. A. ditimena 33a. Feuilles oblancéolées (parfois étroitement elliptiques), au moins 3 fois plus longues que larges, chartacées à coriaces, flexibles; inflorescences terminales et axillaires; fruits glabres; le long des cours d’eau, ouest et sud de Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy...... 11. A. grandidieri

34 (20). Feuilles les plus grandes avec au moins 40 paires de nervures secondaires...... 35 34a. Feuilles les plus grandes avec moins de 40 paires de nervures secondaires...... 36

35. Nervures secondaires au niveau de la partie médiane du limbe étroitement espacées, env. 5-6 par cm; fruit à indument dense; forêts humides orientales...... 30. A. sericea 35a. Nervures secondaires au niveau de la partie médiane du limbe largement espacées, env. 3 par cm; fruit glabre; le long des cors d’eau, ouest et sud de Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy...... 11. A. grandidieri

36 (34). Feuilles les plus grandes avec un maximum de 20 paires de nervures secondaires...... 37 36a. Feuilles les plus grandes avec plus de 20 paires de nervures secondaires...... 46

37. Feuilles elliptiques à orbiculaires ou légèrement ovées, la plus grande largeur au niveau de la partie médiane ou en-deça...... 38 378a. Feuilles obovales à oblancéolées, la plus grande largeur au-delà de la partie médiane....40

38. Feuilles orbiculaires à largement elliptiques (rarement quelques feuilles largement ovées), pas plus de 2 fois plus longues que larges; forêt sèche, Ankarana, Bemaraha...... 12. A. humbertii 38a. Feuilles elliptiques a étroitement elliptiques, au moins 2 fois plus longues que larges....39

39. Feuilles les plus grandes de 2.5-3.6 cm de large; forêts humides de transition, Manambato, Daraina, vallée de l’Androranga près de Sambava...... 4. A. capuronii 39a. Feuilles les plus grandes de 1.5-2(-2.2) cm de large; forêts sèches, de l’Isalo à Andohahela...... 27. A. phillipsonii

40 (37). Inflorescences exclusivement axillaires, inflorescences femelles avec un maximum de 5 fleurs (inflorescences males inconnues); forêts humides, Nosy Be, Ampasindava, Ambongo-Boina...... 26. A. pauciflora 40a. Inflorescences terminales et parfois aussi axillaires, fleurs 8-nombreuses (rarement seulement 5 fleurs dans les inflorescences femelles deA. suarezensis)...... 41 28 Boissiera 71

41 (40). Limbe des feuilles les plus grandes 2-3(-4) cm de long; fruits 1-1.5 cm de long...... 22. A. minutifolia 41a. Limbe des feuilles les plus grandes (3.5-)4 cm de long; fruits 1.5-2.5 cm de long...... 42

42. Fruits densément couverts d’un indument court; nervures secondaires semi- craspédodrome, au moins quelques nervures ramifiées et n’atteignant pas la marge; forêts littorales, Rég. de Tolagnaro...... 7. A. delphinensis 42a. Fruit glabres, nervures secondaires craspédodromes, toutes atteignent clairement la marge...... 43

43. Pétiole robuste, (1.5-)2-3 mm de diam. sur les feuilles les plus grandes, nervures tertiaries distinctivement surélevées sur la surface inférieure; forêts humides de l’est et du nord-ouest, Ambanja, Marojejy, Anajanaharibe Sud, Ranomafana, Analamazaotra, Zahamena ...... 32. A. thouvenotii 43a. Pétiole grêle, pas plus de 1.5 mm de diam. sur les feuilles les plus grandes, nervures tertiaries indistinctes à faiblement surélevées sur la surface inférieure ...... 44

44. Nervures secondaires étroitement espacées, 5-6 paires par cm au niveau de la partie médiane de la feuille la plus grande; forêts humides orientales, Analamazaotra, Ambalabe...... 20. A. louvelii 44a. Nervures secondaires largement espacées, 3-4 paires par cm au niveau de la partie médiane de la feuille la plus grande ...... 45

45. Inflorescences de 1-3(-3.5) cm de long; feuille à marge ondulée, nervures secondaires env. 10-12(-14) paires; forêts sèches et côtières du Nord, Antsiranana, Vohémar, Daraina ...... 31. A. suarezensis 45a. Inflorescences de 4-9 cm de long; feuille à marge foliaire plane ou faiblement ondulée, nervures secondaires 14 paires ou plus; forêts humides, Ankazobe, E de Fianarantsoa, Ranomafana, Andohahela (Parcelle n° 1), Ivohibe...... 33. A. turkii

46 (36). Nervures secondaires semi-craspédodromes, au moins quelques nervures ramifiées et n’atteignant pas la marge; fruit à indument visible...... 17. A. lenticellata 46a. Nervures secondaires craspédodromes, atteignent toutes la marge; fruit glabre ou à indument évident...... 47

47. Pétiole des feuilles les plus grandes d’au moins de 10 mm long...... 48 47a. Pétiole des feuilles les plus grandes de moins de 10 mm long...... 53

48. Inflorescences exclusivement axillaires; forêts littorales orientales, de Tampolo au nord jusqu’à Mahabo au sud...... 18. A. littoralis 48a. Inflorescences terminales et parfois aussi axillaires...... 49 Taxonomic treatment of Abrahamia (Anacardiaceae) 29

49 (48). Nervures tertiaires à peine visibles, presque absentes; fleurs femelles à corolle glabre sur la surface externe...... 20. A. longipetiolata 49a. Nervures tertiaires visibles au moins sur une surface; corolle de la fleur femelle avec un indument sur la surface externe...... 50

50. Feuilles pas plus de 2.5 fois plus longues que larges; fruits avec un indument évident ...... 51 50a. Feuilles au moins 3 fois plus longues que larges; fruits glabres...... 52

51. Feuilles chartacées; forêts littorales, aux environs de Tolagnaro...... 7. A. delphinensis 51a. Feuilles coriaces; forêts humides orientales...... 8. A. ditimena

52 (50). Feuilles obovales à elliptiques; à 18-33 paires de nervures secondaires, espacées de 1-4 mm; forêts humides et sub-humides de transition du nord-est (Daraina, Vohémar)...... 4. A. capuronii 52a. Feuilles oblancéolées; de 35-40-(47) paires de nervures secondaires, espacées de 0.5-5 mm; forêts sèches et sub-humides de l’ouest et su sud-ouest...... 11. A. grandidieri

53 (47). Nervures secondaires faiblement saillantes à la face supérieure des feuilles; fruits glabres ...... 54 53a. Nervures secondaires distinctement saillantes à la face supérieure des feuilles; fruits à indument évident...... 60

54. Fruits de 3-3.5 cm de long, ellipsoïdes; axes des infructescence robustes, de 1.5-2 mm de diam.; feuilles les plus grandes mesurant 8-9 × 2.5 cm, apex aigu; forêts littorales, aux environs de Tolagnaro...... 9. A. ellipticarpa 54a. Fruits de 1.5-2.6 cm de long, ovoïdes ou ellipsoïdes; axes des infructescence grêles, env. 1(-1.2) mm de diam.; feuilles les plus grandes soit de moins de 8 × 2 cm, soit avec un apex arrondi à émarginé...... 55

55. Feuilles obovales à oblancéolées...... 56 55a. Feuilles elliptiques à oblongues, rarement quelques-unes légèrement ovées ou obovales...... 59

56. Feuilles les plus grandes avec au moins 35 paires de nervures secondaires; forêts sèches ...... 11. A. grandidieri 56a. Feuilles les plus grandes avec pas plus de 25(-28) paires de nervures secondaires; forêts humides...... 57

57. Nervures secondaires largement espacées, 3-4 paires par cm au niveau de la partie médiane du limbe le plus grand; forêts humides, Ankazobe, à l’Est de Fianarantsoa, Ranomafana, Andohahela (Parcelle n° 1), Ivohibe...... 33. A. turkii 57a. Nervures secondaires étroitement espacées, 5-6 paires par cm au niveau de la partie médiane du limbe le plus grand...... 58 30 Boissiera 71

58 (57). Limbe foliaire de la feuille la plus grande de 4(-4.2) cm de long; forêts humides, Analamazaotra, Ambalabe...... 21. A. louvelii 58a. Limbe foliaire de la feuille la plus grande de (4.8-)5 cm de long; forêts semidécidues et h humides, Daraina...... 25. A. patrickii

59 (55). Pétiole grêle, de moins de 1 mm de diam.; cotylédons fusionnés; arbres d’env. 3-12 m; forêts sèches, Isalo to Andohahela...... 27. A. phillipsonii 59a. Pétiole robuste, de 1.2-1.5 mm de diam.; cotylédons séparables; arbuste ou petits arbres de moins de 2 m de haut; forêts sclérophylles, Ibity...... 13. A. ibityensis

60 (53). Feuilles glauques à la face supérieure, presque toujours fortement discolores sur le sec, souvent brun-chocolaté sur la face supérieure (parfois caché par la glaucescence); forêts sèches, partie centrale du haut plateau, Itremo, Ibity, Ambatofinandrahana...... 3. A. buxifolia 60a. Feuilles non glauques à la face supérieure, concolores ou faiblement discolores...... 61

61. Feuilles pubérulentes à la surface supérieure; Rég. d’Itremo...... 14. A. itremoensis 61a. Feuilles glabres à la face supérieure...... 62

62. Feuilles chartacées; forêts littorales, aux environs de Tolagnaro...... 7. A. delphinensis 62a. Feuilles coriaces; forêts humides orientales...... 8. A. ditimena Taxonomic treatment of Abrahamia (Anacardiaceae) 31

1. Abrahamia antongilensis Randrian. & Lowry, spec. nova (Fig. 1).

Typus: Madagascar. Prov. Toamasina: Rég. Analanjirofo, Anjanaharibe forest, 15°10’48’’S 49°37’30’’E, 860 m, 4.II.2003, fr., Antilahimena 1862 (holo-: MO-6671431!; iso-: G [G00341711]!, K!, NY!, P [P00580355]!, TAN [TAN002003]!).

Diagnosis Abrahamia antongilensis Randrian. & Lowry can be distinguished from the other members of the genus by the combination of its leaves with semicraspedodromous venation and a distinctly acuminate apex (10-15 mm long), and inflorescences that are very short (c. 1.8 cm) and terminal.

Description Trees 8-10 m tall; young twigs covered with dense, whitish sericeous indument. Leaves alternate, clustered at the end of branches; blade oblanceolate to obovate, 7-17 3 3-6 cm, chartaceous, apex abruptly acuminate, the acumen 10-15 mm long, margin undulate and very slightly revolute, base cuneate, adaxial surface glabrous or with scattered whitish sericeous indument, abaxial surface covered with dense whitish, appressed sericeous indument when younger, much less dense on older leaves, venation semicraspedodromous, midvein prominent abaxially, secondary veins 25-35 pairs, 2-7 mm apart, forming a wide angle with the midvein (almost perpen- dicular from the base to the leaf mid-section), raised on both surfaces, tertiary veins visible on both surfaces; petiole 10-18 mm long, c. 1.5 mm in diam., canaliculate on adaxial surface, covered with whitish sericeous indument. Inflorescence short, terminal, a panicle, c. 1.8 cm long, axes covered with dense, appressed indument. Male flowers unknown. Female flowers 5-merous, small, c. 3 mm long, subtended by a very small bract with dense indument; pedicel c. 3 mm long, densely sericeous; calyx lobes small, widely ovate, 1 3 1 mm, valvate, adaxial surface glabrous, abaxial surface pubescent; corolla lobes ovate, 2.5 3 1.5 mm, thick, imbricate, adaxial surface glabrous, abaxial surface pubescent; staminodes 5; disk cupuliform, c. 1.5 mm in diam., glabrous; ovary slightly ovoid to spherical, c. 1 3 1 mm, glabrous; style short, thick, c. 1 mm long, glabrous, stigmas capitate. Fruits 2.5-3.5 3 1.7-2 cm, ellipsoid, glabrous, with longitudinal striations.

Distribution, ecology This species is known only from the area around the northern end of the Baie d’Antongil and phenology in northeastern Madagascar (Map 1). Flowering material has been collected in Novem- ber and fruiting specimens in February.

Conservation status With a minimum AOO of 8 km2 and two known localities, one within the newly established Makira natural park and the other based on a historical collection that can be assumed to be have come from within the buffer zone of Masoala national park, 32 Boissiera 71

1 cm

Fig. 1. Abrahamia antongilensis Randrian. & Lowry. Fruiting branch.

[Antilahimena 1862, P] Drawing: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 33

comprising 2 locations, the latter of which is subjected to projected decline in habitat quality, A. antongilensis is assigned a preliminary conservation status of “Endangered” [EN B2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia antongilensis most closely resembles A. lenticellata and A. viguieri but differs by having the abaxial leaf surface covered with whitish appressed indument (vs glabrous in A. lenticellata and with reddish lanate indument in A. viguieri). Moreover, A. antongilensis can be distinguished from A. lenticellata by having glabrous fruits (vs velutinous) and from A. viguieri by its leaves with semicraspedodromous (vs craspedodromous) venation. Abrahamia antongilensis is only known from two collections, both from the area around the Baie d’Antongil in northeastern Madagascar. The type collection (Antilahi- mena 1862), from northwestern of Maroantsetra, has older leaves with less indument on the abaxial surface than on the much younger leaves of the second collection (Service Forestier 8770) from the northern part of the Masoala Peninsula, approximately half way between Antalaha and Maroantsetra.

Paratypi Madagascar. Prov. Toamasina: Rég. SAVA, S du Col d’Ambatondradama, [15°16’S 50°04’E], c. 750 m, 17.XI.1953, fl.,Service Forestier 8770 (MO, P [2 sheets], TEF). 34 Boissiera 71

2. Abrahamia betamponensis Randrian. & Lowry, spec. nova (Fig. 2).

Typus: Madagascar. Prov. Toamasina: Rég. Atsinanana, Betampona RN, [17°55’S 49°13’E], 24.X.1955, bud, Réserves Naturelles 7701 (holo-: P [P06774885]!; iso-: TEF [TEF000880]!).

Diagnosis Abrahamia betamponensis Randrian. & Lowry can easily be distinguished from other members of the genus by its large leaves (9-24 3 4.3-8.5 cm) with a remarkably prominent midvein on the abaxial surface, a long, thick petiole (15-18 mm in length), numerous secondary veins (50-60 pairs), and fruit surface covered with dense, short indument.

Description Trees c. 7 m tall; bark with lenticels; young twigs whitish-gray to light tan, pubescent at the very tips. Leaves opposite or subopposite; blade oblanceolate, 9-24 3 4.3-8.5 cm, subcoriaceous to coriaceous, apex shortly acuminate, retuse or emarginate, margin slightly revolute, base cuneate, adaxial surface glabrous, abaxial surface sparsely strigose along midvein, venation craspedodromous, midvein strikingly prominent abaxially, secondary veins 50-60 pairs, 2-5 mm apart, raised on both surfaces, tertiary veins visible on both surfaces, petiole 15-18 mm long, canaliculate on adaxial surface, c. 2.5-3 mm in diam., glabrous. Inflorescence terminal and axillary, a panicle, 2-7 cm long, axes thick, covered with dense, short, appressed indument; bracts triangular, c. 4 3 2 mm, adaxial surface glabrous, abaxially surface with short, appressed indument. Male flowers 5-merous, small, c. 3 mm long, subtended by a small, deltoid bract; pedicel short, c. 1 mm long, strigose; calyx lobes triangular, 2.5 3 1 mm, adaxial surface glabrous, abaxial surface pubescent; corolla lobes very widely ovate, 2 3 2 mm, adaxial surface glabrous, abaxial surface pubescent; stamens 5, filaments 1.2-1.3 mm long, anthers c. 1 mm long, ovoid; disk thin, dish-shaped and glabrous. Female flowers unknown. Fruits 2.4-3.2 3 1.4-1.8 cm, ellipsoid, covered with dense, short indument, with longitudinal striations.

Vernacular names “Hazombarorana”, “Hazombarorana à grandes feuilles”, “Vatodinga’’, “Voatodinga”.

Distribution, ecology Abrahamia betamponensis is known only from Betampona reserve and its vicinity, with and phenology a single collection from Zahamena national park (Map 1). It occurs in mid-elevation humid forest. Material in bud has been collected in October and fruiting specimens have been made in January.

Conservation status Abrahamia betamponensis has a minimum EOO of 8 km2, but its two known localities are both situated within protected areas (Betampona reserve and Zahamena national park) so in the absence of any data on population size, no future decline is expected Taxonomic treatment of Abrahamia (Anacardiaceae) 35

in terms of AOO, EOO, or quality of habitat. It has therefore assigned a preliminary status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia betamponensis belongs to a group of species with large leaves (up to 37 cm long) that have numerous (40-70) secondary veins. Within this group, A. betamponensis most closely resembles A. sambiranensis, but differs by the dense, short indument on the surface of its mature fruit (vs glabrescent or glabrous).

Paratypi Madagascar. Prov. Toamasina: Rég. Antisnana, Fotsimavo, [Betampona RN], [17°55’S 49°13’E], 300-400 m, 20.I.1950, fr., Réserves Naturelles 2366 (P, TAN); ibid. loco, Réserves Naturelles 2489 (P, TAN); Anjiro, Ambodiriana, [Betampona RN], [17°55’S 49°13’E], 26.X.1957, bud, Réserves Naturelles 9148 (P, TEF); Mangabe, [Zahamena PN], [17°42’S 48°54’E], 20.I.1963, old fr., Réserves Naturelles 12446 (MO, P, TEF); Betampona RN, 17°54’29”S 49°12’16”E, 530 m, 13.I.2017, ster., Schatz et al. 4472 (MO, P, TAN).

1 cm

2 cm

Fig. 2. Abrahamia betamponensis Randrian. & Lowry. A. Fruiting branch; B. Fruit showing indument.

[Réserves Naturelles 2366, P] Drawings: R.L. Andriamiarisoa 36 Boissiera 71

45 E 50 E

Map 1. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. antongilensis Randrian. & Lowry A. betamponensis Randrian. & Lowry A. buxifolia (H. Perrier) Randrian. & Lowry A. capuronii Randrian. & Lowry A. deflexa (H. Perrier) Randrian. & Lowry A. ellipticarpa Randrian. & Lowry 15 S

20 S

25 S Taxonomic treatment of Abrahamia (Anacardiaceae) 37

45 E 50 E

Map 2. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. darainensis Randrian. & Lowry A. delphinensis Randrian. & Lowry A. elongata Randrian. & Lowry A. humbertii (H. Perrier) Randrian. & Lowry A. ibityensis (H. Perrier) Randrian. & Lowry

1515 S S

2020 S S

2525 S S 38 Boissiera 71

45 E 50 E

Map 3. Geographic distribution of Abrahamia ditimena (H. Perrier) Randrian. & Lowry

15 S

20 S

25 S Taxonomic treatment of Abrahamia (Anacardiaceae) 39

45 E 50 E

Map 4. Geographic distribution of Abrahamia Randrian. & Lowry

A. grandidieri (Engl.) Randrian. & Lowry A. lecomtei (H. Perrier) Randrian. & Lowry

15 S

20 S

25 S 40 Boissiera 71

3. Abrahamia buxifolia (H. Perrier) Randrian. & Lowry, comb. nova (Fig. 3).

≡ Protorhus buxifolia H. Perrier in Mém. Mus. Natl. Hist. Nat. Paris 18: 258. 1944.

Lectotypus (designated here): Madagascar. Prov. Toliara: Mt. Belambana, bassin du Mananara (Sud), [22°08’S 46°21’E], 1000 m, VII.1910, Perrier de la Bâthie 5852 (P [P06775177]!; isolecto-: P [P00722637]!). Syntypi: Madagascar. Prov. Fianarant- soa: massif de l’Andringitra, [22°13’S 47°01’E], c. 1800 m, IV.1921, fr., Perrier de la Bâthie 13696 (P [P00722638]!). Prov. Toliara: Mont Vohipolaka, N Betroka, [22°08’S 46°21’E], 1100-1300 m, XI.1933, ster., Humbert 11637bis (P [P06775327]!); ibid. loco, 1100-1200 m, Humbert 11667bis (P [P00722639]!).

Description Trees 2-7 m tall, bark with white milky latex; young twigs pubescent or glabrescent, lenticellate or waxy. Leaves subopposite or opposite; blades obovate to elliptic, 1.5- 6.8 3 1-2.7 cm, subcoriaceous, apex retuse or rounded, margin entire, base cuneate, adaxial surface glabrous, glaucous, often chocolate brown when dry, abaxial surface glabrous (except for sericeous indument on the midvein, and sometimes scattered indument elsewhere on the blade), venation craspedodromous, midvein prominent abaxially, secondary veins 15-25 pairs (rarely to 36 pairs), 1-3 mm apart, prominent on both surfaces, tertiary veins raised on both surfaces; petiole 5-7 mm long, c. 1-1.1 mm in diam., shallowly canaliculate on adaxial surface, glabrescent or pubescent. Inflorescences axillary and terminal, a thyrse, 1.5-4.5 cm long, pubescent. Male and female flowers 5-merous; pedicel c. 0.2-0.3 mm long, pubescent; calyx lobes deltate to widely depressed ovate, 1-1.2 3 1-1.5 mm, valvate, persistent on fruit, adaxial surface glabrous, abaxial surface pubescent; corolla lobes ovate, 2-3 3 1.5-2 mm, pale yellow, imbricate, adaxial surface glabrous, abaxial surface pubescent. Male flowers with 5 or 6 stamens, filaments c. 1-1.5 mm long, glabrous or with sparse indument, white, anthers c. 0.8 mm long, obloid to slightly ovoid, yellow, glabrous; disk c. 2 mm in diam., glabrous; ovary rudimentary. Female flowers with 5 or 6 staminodes, filaments c. 1 mm long, glabrous, white, anthers c. 0.5 mm long, ovoid, sterile, yellowish, glabrous; disk c. 1.5 mm in diam., glabrous; ovary widely ovoid, 1-2 3 1-1.5 mm, pubescent; style shortly trifid, inserted apically, c. 1 mm long, pubescent, stigmas capitate. Fruits c. 1.5 3 1.1 cm, ellipsoid to slightly obovoid, surface smooth, velutinous when young, glabrescent.

Vernacular names “Sohihy’’.

Distribution, ecology Abrahamia buxifolia occurs in the southern and southwestern parts of central Madagas- and phenology car, with populations extending toward the western slopes and the eastern escarpment Taxonomic treatment of Abrahamia (Anacardiaceae) 41

(Map 1). It grows at elevation between 900-1800 m in areas where ecological conditions are harsh, including low rainfall and poor soils. It flowers in September and October, and sets fruits between November and February.

Conservation status With an EOO of 37,259 km², a minimum AOO of 60 km², and 18 recorded localities, most of which are subjected to frequent burning and only 3 of which are within protected areas (Andringitra, Ibity, and Itremo), representing 11 locations, A. buxifolia is assigned a preliminary conservation status of “Near Threatened’’ [NT] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia buxifolia can easily be distinguished from its most similar congener, A. itremoensis, by its glaucous and subcoriaceous leaves (vs non-glaucous and coriaceous) that are somewhat larger in size (1.5-6.8 3 1-2.7 vs 2-6 3 0.6-2 cm), and by its flowers with a glabrous disk (vs with evident indument). Of the four collections cited in the protologue that we examined, we have selected Perrier de la Bâthie 5852 as the lectotype because it is more complete than the other syntypes and is represented by two specimens, whereas each of the three other syntypes comprises only a single sheet, two of which (Humbert 11637bis and Humbert 11667bis) are sterile and one (Perrier de la Bâthie 13696) has just a few fruits. Two other collections were also mentioned in the protologue (Decary 13041 and Humbert 14048) but we have not been able to find these specimens (although a sheet of the latter is reported to be at US) and we have not listed them below in the material examined. The label data for Service Forestier 23500 and 23501 indicate a trunk diameter of 0.5 m, a value that far exceeds any observation we have made in the field. We therefore regard this as a label error and have not included this information in the description.

Additional material Madagascar. Prov. Antananarivo: Antsirabe, Betafo, Antakoboka, 20°03’46’’S examined 46°49’30’’E, 1807 m, 17.IX.2004, y. fl.,Andriamihajarivo 371 (G, MO, P, TAN, TEF); forêt d’Analandraisoa (SW Tsiroanimandidy), [19°09’S 45°49’E], 1200-1300 m, 17-20. XI.1952, fr., Leandri 1852 (G, MO, P [2 sheets], TAN); Mont Ibity, E slope of Kibory, 20°06’S 46°59’E, 1600 m, 2.X.1996, ster., A. Randrianasolo 488 (MO, P, TAN); Tsiro- anomandidy, forêt d’Analandraisoa, [18°59’30’’S 45°41’30’’E], c. 1100 m, XI.1952, fr., Service Forestier 6726 (G, MO, P, TEF). Prov. Fianarantsoa: Ambatofinandrahana, Itremo, forêt d’Antsirakambiaty, 20°35’42’’S 46°33’55’’E, 1605 m, 11.XII.2003, fr., Andriamihajarivo 258 (G, MO, P, TAN); Lovokaloha, 20°29’58’’S 46°51’37’’E, 1411 m, 2.XI.2004, fr., Andriamihajarivo 408 (G, MO, P, TAN); Ambatofinandrahana, 10 km N de RN 35, Antanimena, 20°31’05’’S 46°34’02’’E, 1541 m, 24.VIII.2010, fr., Andrianaivoravelona et al. 363 (K, MO, TAN); entre Ambatofinandrahana et Itremo, [20°33’S 46°45’E], IX.1956, bud, Bosser 10037 (MO, P [3 sheets], TAN); Itremo, [20°32’S 46°33’E], 1600 m, IX.1956, bud, Bosser 10075 (P, TAN); Inselbergs, PK 475, W of Ambalavao, 950-1050 m, 1.II.1975, fr., Croat 30326 (MO); N of Sendrisoa, c. 1200 m, 23.II.1975, fr., Croat 32165 (MO, P, TAN); Itremo massif, along Ambalo- ranogolana creek, 20°34’31’’S 46°34’50’’E, 1500 m, 9.XI.2002, y. fr., Lowry et al. 5861 42 Boissiera 71

A B

Fig. 3. Abrahamia buxifolia Randrian. & Lowry. A. Fruits and upper leaf surfaces, from above; B. Inflorescences and flowers, from above [Lowry et al. 5861].

Photos: G. Schatz

(G, MO, P, TAN); Itremo massif, montane forest on quartz, 20°34’40’’S 46°34’53’’E, 1550 m, 29.X.1997, fl.,Malcomber et al. 2849 (G, MO, P, TAN); Anja Parc, Inselberg Ambohilady, 21°51’11’’S 46°51’14’’E, 1224 m, 10.XII.2010, fr., Ramahefaharivelo 272 (G, MO, P, TAN); Itremo chain, W of Itremo village, 20°35’00’’S 46°36’10’’E, 1500 m, 4.X.1996, fl.,A. Randrianasolo 489 (G, MO, P, TAN); ibid. loco, 1450 m, 6.X.1996, bud, A. Randrianasolo 493 (G, MO, P, TAN); Ambalavao, Sendrisoa, [22°07’S 46°49’E], 15.XI.1953, fr., Réserves Naturelles 5858 (P, TEF); Ivohibe, [Ivohibe RN], Antambohobe, 12.IX.1962, bud, Réserves Naturelles 12154 (P); forêt Ambondrozana, Ihosy, [21°57’S 46°36’E], 17.II.1955, bud, Service Forestier 13771 (P, TEF); massif granitique d’Ifan- dana, entre Ankaramena et Ihosy, [22°01’00’’S 46°22’30’’E], 1300-1400 m, 7.X.1964, fl., Service Forestier 23500 (G, MO, P [2 sheets], TEF); ibid. loco, Service Forestier 23501 (G, MO, NY, P, TEF); massif d’Iandrambaky, SW d’Iarintsena (Ambalavao), [21°51’S 46°51’E], 4.XI.1967, y. fr., Service Forestier 27830 (G, MO, P, TEF); rocher de l’Ifandana, N de Zazafotsy, [22°01’00’’S 46°22’30’’E], 2.XII.1968, fr., Service Forestier 28473 (G, K, MO, NY, P [2 sheets], TEF); Ambatofinandrahanana, Andranomivoaka, 18.III.1952, ster., Service Forestier 45-R-207 (P); Ambalavao, forêt d’Amboandrozana, 9.VI.1954, ster., Service Forestier 63-R-141 (P). Taxonomic treatment of Abrahamia (Anacardiaceae) 43

4. Abrahamia capuronii Randrian. & Lowry, spec. nova (Fig. 4).

Typus: Madagascar. Prov. Antsiranana: Rég. SAVA, Dist. Sambava, S de l’An- jehabe, [14°18’30’’S 49°44’40’’E], c. 600 m, 7.XI.1950, fl.,Service Forestier 767 (holo-: P [P06774827]!; iso-: G [G00341710]!, MO-6684132!, NY!, P [P06774828, P06774829]!, TEF [TEF000881]!).

Diagnosis Abrahamia capuronii Randrian. & Lowry can be distinguished from all other species of Abra- hamia by a combination of its elliptic leaves, narrow corolla lobes (3 3 1.2 mm), and slender inflorescence axes.

Description Trees 10-15 m tall, 20 cm DBH, bark with white or translucid latex; young twigs only pubescent at the tips. Leaves alternate or sub-opposite; blade obovate to elliptic, 4-7.8 3 2-3.6 cm, chartaceous to subcoriaceous, apex rounded or retuse, margin undulate, base cuneate, adaxial surface glabrous, abaxial surface with short appressed indument, mainly along the midvein and at the base, venation craspedodromous, midvein prominent abaxially, secondary veins 18-33 pairs, 1-4 mm apart, prominent abaxially, slightly raised adaxially, tertiary veins slightly visible on adaxial surface; petiole 10-25 mm long, c. 0.5-1 mm in diam., canaliculate on adaxial surface, glabrescent. Inflorescence axillary or terminal, a panicle, 2-8 cm long, axes slender, strigose. Male flowers 5-merous, small, c. 2-2.5 mm long, subtended by a very small, pubescent bract; pedicel short, c. 0.3-0.5 mm long, pubescent; calyx lobes small, widely ovate, 1 3 1 mm, valvate, connate at the base, adaxial surface glabrous, abaxial surface sparsely covered with short indument, glabrescent; corolla lobes 3 3 1.2 mm, lanceolate, imbricate, adaxial surface glabrous, abaxial surface pubescent along its longitudinal axis or glabrescent to glabrous, white (according to field notes); stamens 5, filament c. 1.5 mm, glabrous, anthers c. 1 mm long, obloid, glabrous; disk glabrous, c. 1.2-1.5 mm in diam.; ovary rudimentary. Mature and opened female flowers unknown, developing buds with calyx lobes triangular to deltate, c. 1 3 0.5- 0.8 mm, adaxial surface glabrous, abaxial surface with some indument; corolla lobes ovate, c. 1.5 3 1 mm, mostly glabrous but abaxial surface with a few appressed short trichomes; stamens vestigial; disk glabrous; ovary depressed, ovoid, 0.3-0.5 3 0.5 mm, glabrous; style c. 1 mm, glabrous, stigmas capitate. Young fruits ellipsoid, very slightly asymmetrical, glabrous, with longitudinal striations.

Etymology The species epithet honors René Capuron, whose extensive field work contributed so much to our knowledge of Madagascar’s woody flora, and who personally collected material of 24 of the 34 species of Abrahamia. 44 Boissiera 71

Distribution, ecology Abrahamia capuronii is restricted to far northeastern Madagascar (Map 1), where it and phenology occurs in mid-elevation humid and transitional forests on rocky substrate up to an elevation of 700 m. Material with buds and flowers has been collected in October and early November, and specimens with young fruits have been gathered in late November.

Conservation status With an EOO of 2,585 km² and a minimum AOO of 24 km², A. capuronii would qualify as “Endangered” under Criteria B1 and B2 of the IUCN Red List (IUCN, 2012) if other qualifying criteria were met. However, it is known from a total of 6 locations, one of which is situated within a protected area (Loky Manambato) while the others are at sites potentially subjected to clearing and habitat degradation, and should thus be assigned a preliminary status of “Vulnerable’’ [VU B1ab(iii)+2ab(iii)].

Notes Abrahamia capuronii resembles both A. grandidieri and A. sericea by its relatively closely spaced, parallel and craspedodromous secondary veins, but can be distinguished from these two species by its smaller number of secondary veins (10-33 in A. capuronii vs 35-40(-47) in A. grandidieri and 40-70 in A. sericea).

Paratypi Madagascar. Prov. Antsiranana: Mont Anjenabe, [14°17’S 49°46’E], 650 m, 3-7. XI.1950, fl.,Humbert 24055 (G, K, MO, P [2 sheets], TAN); Vohémar, forêt de Binara, 13°15’S 49°37’E, 700 m, 8.XI.2001, fl.,Gautier et al. 4125 (G, MO, TAN); Vohémar, Analamazava forest, 13°15’43’’S 49°35’22’’E, 25.XI.2004, y. fr., Ranarivelo 446 (MO); Daraina, Ambilobe, 13°44’10’’S 49°23’07’’E, 1299 m, 27.X.2007, bud, Randriambololo- mamonjy et al. 166 (MO, P, TAN); Vohémar, Daraina, forêt d’Analamazava, [13°15’58’’S 49°36’14’’E], 700 m, 29.X.2005, fl.,Randrianaivo 1259 (CNARP, MO, P, TAN); Vohémar, Daraina, forêt d’Antsahabe, 13°12’37’’S 49°33’29’’E, 545 m, 31.X.2005, fl.,Razafit - salama 741 (CNARP, MO, P, TAN); Vohémar, Andrafainkona, Ampisarahana, 13°38’28’’S 49°31’51’’E, 1009 m, 9.XI.2007, bud, Razakamalala et al. 3716 (G, MO, P, TAN). Taxonomic treatment of Abrahamia (Anacardiaceae) 45

5 mm

1 cm

1 mm C 1 cm D

A B

Fig. 4. Abrahamia capuronii Randrian. & Lowry. A. Flowering branch; B. Flowers (one with a petal removed); C. Detail of leaf; D. Inflorescence. [Service Forestier 767, P] Drawings: R.L. Andriamiarisoa 46 Boissiera 71

B A

1 cm 1 cm

Fig. 5. Abrahamia darainensis Randrian. & Lowry. A. Fruiting branch; B. Fruit.

[Ranirison & Nusbaumer 1118, MO] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 47

5. Abrahamia darainensis Randrian. & Lowry, spec. nova (Fig. 5).

Typus: Madagascar. Prov. Antsiranana: Rég. SAVA, Daraina, forêt d’Antsahabe, 13°12’53’’S 49°31’53’’E, 460 m, 25.I.2006, fr., Ranirison & Nusbaumer 1118 (holo-: G [G00090546]!; iso-: MO-6671432!, P [P00580353]!, TEF [TEF000882]!).

Diagnosis Abrahamia darainensis Randrian. & Lowry can be distinguished from other members of the genus by its small, chartaceous, glabrous, elliptic to obovate leaves with a short petiole (5-7 mm long), and its glabrous, ellipsoid fruits that measure 2.6 3 1.5 cm.

Description Trees 6 m tall, 9 cm DBH. Leaves alternate; blade elliptic to obovate, 4-7.5 3 1.2-2.2 cm, chartaceous, apex shortly acuminate, margin undulate, base cuneate, both surfaces glabrous, venation craspedodromous, midvein visible but not prominent on both surfaces, secondary veins 14-18 pairs, 3-5 mm apart, almost impressed adaxially, slightly raised baxially, tertiary veins few and only weakly visible adaxially; petiole 5-7 mm long, c. 0.5-1 mm in diam., canaliculate on adaxial surface, glabrous. Inflorescence and flowers unknown. Fruits 2.6 3 1.5 cm, ellipsoid, light red, glabrous, with longitudinal striations.

Distribution, ecology Abrahamia darainensis is known only from humid forest on the eastern slope of Antsa- and phenology habe in the Daraina region (Map 2). It was collected in fruit in January.

Conservation status While Abrahamia darainensis is only known from a single locality, it is situated within a large block of forest in the Loky Manambato protected area (Daraina), and in the absence of data on population size, no future decline is expected in terms of AOO, EOO, or quality of habitat, so we therefore assign a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia darainensis is similar to A. lenticellata in having chartaceous leaves with a short acuminate apex and undulate margins, and fruits of similar size (2.6 3 1.5 cm in A. darainensis and 2-2.6 3 1.3-1.5 cm in A. lenticellata). It differs, however, by having secondary veins that are fewer in number (14-18 vs 20-25) and are craspedodromous (vs semicraspedodromous), a shorter petiole (5-7 mm vs 7-13 mm), and glabrous (vs velutinous) fruits. Abrahamia darainensis is only known from the type gathering. 48 Boissiera 71

6. Abrahamia deflexa (H. Perrier) Randrian. & Lowry, comb. nova.

≡ Protorhus deflexa H. Perrier in Mém. Mus. Natl. Hist. Nat. Paris 18: 264. 1944.

Lectotypus (designated here): Madagascar. Prov. Mahajanga: Ambongo, XI.1903, fl., Perrier de la Bâthie 1627 (P [P00580302]!; isolecto-: MO-6684133!, P [P00580303]!). Syn- typi: Madagascar. Prov. Mahajanga: Ankatsepy, près Majujnga, s.d., fr., Perrier de la Bâthie 1627bis (P [P00580304]!). Prov. Toliara: Mt. Vohipolaka, N de Betroka, s.d., ster., Hum- bert 11694ter (P, not seen); Mt. Apika, au-dessus de Mahamavo, bassin de réception de la Mananara, affluent du Mandrare, s.d., fr.,Humbert 13850 (P, not seen).

Description Trees 8-15 m tall, 15-20 cm DBH, bark with red latex, young twigs with lenticels, waxy, the very tips scabrous. Leaves alternate, subopposite or opposite; blades ovate, oblong, or elliptic, 2.5-6.5 3 1-3.5 cm, apex retuse, rounded or shortly acuminate, sometimes emarginate, margine entire, base cuneate, adaxial surface glabrous, abaxial surface ferruginous to scabrous when young, later glabrous or glabrescent along the midvein, venation craspedodromous, midvein prominent abaxially, secondary veins 15-20 pairs, more or less parallel, 1-4 mm apart, prominent abaxially, slightly prominent adaxially, tertiary veins very widely spaced, slightly raised on both surfaces; petiole 5-10 mm long, canaliculate on adaxial surface, covered with very short, appressed indument when young, glabrescent. Inflorescence terminal, a thyrse, 2-5 cm long, ferruginous strigose. Male flowers 5-merous; pedicel c. 0.3-0.4 mm long, strigose; calyx lobes depressed ovate, 0.8-1 3 0.5-0.8 mm, imbricate, adaxial surface glabrous, abaxial surface glabrous or pubescent, persistent on the fruit; corolla lobes ovate, c. 2 3 1.5 mm, white or cream, adaxial surface glabrous, abaxial surface glabrous or with appressed short indument, imbricate; stamens 5, filaments 1 mm long, glabrous, anthers c. 0.5 mm long, ovoid to subglobose, yellow, glabrous; disk dish-shaped to cupuliform, c. 1 mm in diam., glabrous; ovary rudimentary. Female flowers unknown. Fruits 1.5-3 3 1-1.5 cm, ellipsoid to ovoid, glabrous, with longitudinal striations.

Vernacular names “Hazombora’’, “Sohihy’’, “Sohy’’, “Tarata’’, “Motsalahy’’ (Sakalava).

Distribution, ecology Abrahamia deflexa occurs in dry forests of western and northwestern Madagascar (Map and phenology 1), usually on sandy or calcareous soils. It flowers between September and November, and has been collected in fruit in November and December.

Conservation status With an EOO of 51,339 km2, a minimum AOO of 40 km2, and an estimated maximum AOO of less than 2,000 km2, as well as 10 known locations, only 2 of which are situated within protected areas (Ankarafantsika and Bemaraha) with the remainder Taxonomic treatment of Abrahamia (Anacardiaceae) 49

projected to be subjected to continuing decline in habitat quality due to a combination of fire and land clearing,A. deflexa is assigned a preliminary conservation status of “Vulnerable’’ [VU B2 ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia deflexa can easily be distinguished from all its congeners by the combination the small, glabrous leaves with a long, thin petiole and glabrous fruits. Of the five collections cited in the protologue, we have selectedPerrier de la Bâthie 1627 as the lectotype because it is represented by three specimens, whereas one of the other syntypes (Perrier de la Bâthie 1627bis) comprises only a single sheet, another (Humbert 5063 = Humbert & Swingle 5063 = Service Forestier 50) is a collection of A. phillipsonii, and the remaining two (Humbert 11694ter and 13850) could not be located.

Additional material Madagascar. Prov. Mahajanga: Katsepy, station d’Antrema, [15°42’37’’S 46°10’30’’E], examined 10.XI.2008, fr., Hladick 7200 (P); Bemaraha, S of the Manambolo River, 19°09’S 44°49’E, 50 m, 11.XII.1996, fr., Jongkind 3505 (MO, P, TAN, WAG); Antsalova, Bemaraha, 18°38’S 44°42’E, 100-200 m, 5.XII.1992, fr., Labat et al. 2308 (MO, P, TAN); Mitsinjo, Katsepy, forêt classée d’Antrema, 15°42’37’’S 46°10’25’’E, 15 m, 12.XI.2005, fr., Labat 3560 (G, MO, P, TAN); 33 km Ankirika-Analalava, 14°49’S 47°41’E, 100 m, 16.XI.1996, fl., Leeuwenberg et al. 14728 (MO, P, TAN, WAG); NW Madagascar, 1841, Pervillé 93 (P [5 sheets]); Ambongo, 1841, Pervillé 590 (P [3 sheets]); Katsepy, station d’Antrema, [15°42’S 46°10’E], 11.XI.2009, fl.,Ramanamisata 157 (P); Port-Berger, Tsingia, Marosely, 15°38’58’’S 47°35’03’’E, 217 m, 16.XI.2004, fl.,Ramananjanahary et al. 49 (MO, P, TAN); Marovoay, W of Ampijoroa reserve, garden A, [16°18’S 46°48’E], 11.XI.1996, ster., A. Randrianasolo 528 (G, MO, P, TAN); forêt sèche de Marosely, 17°38’58’’S 47°35’03’’E, 217 m, 17.XI.2004, y. fr., Razakamalala et al. 1711 (G, K, MO, P, TEF); Port-Bergé, forêt d’Ambohimanga, 6.XII.2004, 15°57’05’’S 47°26’09’’E, 232 m, fr., Razakamalala et al. 1831 (G, K, MO, NY, P, TEF); forêt d’Ankarafantsika, aux env. d’Ampijoroa, [16°18’S 46°48’E], 19-21.XI.1957, fl.,Service Forestier 18470 (MO, P [2 sheets], TEF); Analalava, env. d’Antsangabe, [15°01’S 47°16’E], 4.XI.1958, fr., Service Forestier 18872 (G, MO, P [2 sheets], TEF); Ambongo, près de Benetsy, [15°04’S 47°14’E], 21.XI.1965, Ser- vice Forestier 24266 (P, TEF); Ampijoroa station forestière, fonkontany Ampombolava, [16°18’S 46°49’E], 15.XII.1991, fr., Service Forestier 34333 (MO, TEF); ibid. loco, 200 m, 23.I.1992, fr., Service Forestier 34358 (MO, TEF). 50 Boissiera 71

1 cm

A B

Fig. 6. Abrahamia delphinensis Randrian. & Lowry. A. Branch with flower buds; B. Detail of leaf; C. Inflorescence with flower buds.

[Randriamampionona 631, P] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 51

7. Abrahamia delphinensis Randrian. & Lowry, spec. nova (Fig. 6).

Typus: Madagascar. Prov. Toliara: Rég. Anosy, Vinanibe, W de Fort-Dauphin, [25°03’S 46°56’E], 7.XII.1968, fr., Service Forestier 28606 (holo-: P [P06774807]!; iso-: G [G00341712]!, K!, MO-5056712!, NY!, P [P00580305, P00580306]!, TEF [TEF000883]!).

Diagnosis Abrahamia delphinensis Randrian. & Lowry is easily distinguished from its congeners by its semicraspedodromous secondary veins, its thin petiole, and its fruits densely covered with short indument.

Description Trees 5-12 m tall, 20-21 cm DBH, bark with white latex; tips of young twigs with short, appressed indument. Leaves opposite or sub-opposite; blade obovate, 2.8-5.8-(6) 3 1-3 cm, chartaceous, apex retuse, margins entire, base broadly cuneate, adaxial surface glabrous, abaxial surface sparsely strigose on midvein, venation semicraspedodromous, midvein prominent abaxially, secondary veins 18-25 pairs, 2-3 mm apart, raised and prominent abaxially, slightly raised adaxially, tertiary veins visible only on abaxial surface; petiole 7-15 mm, c. 0.5-1 mm in diam., usually glabrous but sometimes with very sparse short, appressed indument. Inflorescence axillary or terminal, a panicle, 1.5-5.5 cm long, pubescent. Male and female flowers small, 4-merous, c. 2 mm long; calyx lobes 0.5 3 1 mm, triangular, adaxial surface glabrous, abaxial surface pubescent; corolla lobes c. 2 3 1-1.2 mm, ovate, adaxial surface glabrous, abaxial surface pubescent. Male flowers with filaments 1-1.2 mm long, pubescent or sometimes glabrous, anthers c. 0.5-0.8 mm, subglobose, glabrous; disk slightly cupuliform, c. 1.5 mm in diam., glabrous; ovary completely absent. Female flowers with vestigial stamens, filaments c. 1 mm long, pubescent, anthers very small, c. 0.2 mm, sterile; disk glabrous, cupuliform, c. 1.5 mm in diam.; ovary c. 0.6-0.8 3 1 mm, depressed ovoid, completely covered with indument; style very short, inserted apically, partly covered with indument, stigmas capitate. Fruits 2.3-2.5 3 1.2-1.4 cm, ellipsoid, densely covered with short, appressed and slightly tangled indument, with longitudinal striations.

Distribution, ecology Abrahamia delphinensis occurs in low elevation humid forest, including littoral forest and phenology on sand, and is restricted to the area around Fort Dauphin (Map 2). It flowers between September and November, and fruits between October and December.

Conservation status With an EOO of 1442 km2, a minimum AOO of 48 km2, and 12 known localities representing a total of 7 locations, 3 of which are situated within protected areas (Andoha- hela, Mandena, and Sainte Luce) while the others are variously subjected mining as well as fire and land clearing leading to a projected decline in all 5 sub-criteria,A. delphinensis is assigned a preliminary conservation status of “Vulnerable” [VU B1ab(i,ii,iii,iv,v)+2ab (i,ii,iii,iv,v)] using the IUCN Red List Categories and Criteria (IUCN, 2012). 52 Boissiera 71

Notes Abrahamia delphinensis most closely resembles A. lenticellata but can be distinguished by its retuse (vs short acuminate to acuminate) leaf apex and the complete absence (vs presence) of a rudimentary ovary in male flowers.

Paratypi Madagascar. Prov. Toliara: Andohahela PN, NW of Tolagnaro, 24°45’S 46°51’E, 250-500 m, 17-20.X.1992, fl.,Malcomber et al. 1678 (BM, CAS, G, K, MO [2 sheets], NY, P, TAN, US, WAG); N of Fort-Dauphin, near Ste. Luce, 24°47’S 47°10’E, 10 m, 21.X.1989, y. fr., McPherson 14196 (MO, P, TAN, TEF); Mandena, botanical garden, 24°57’S 47°00’E, 25 m, 25.X.1989, fl.,McPherson et al. 14244 (MO, P, TAN, TEF); Ste. Luce, N de Mahatalaky, 0-10 m, 28.IV.1989, ster., Rabevohitra 1959 (MO, P, TAN); Andohahela PN, Isaka-Ivondro, 24°47’S 46°50’E, 0-500 m, 10-27.IX.1993, fl.,Ran - driamampionona 631 (A, BM, BR, BRIT, CAS, F, G, K, MO [2 sheets], NY, P [2 sheets], PRE, S, TAN, US, W, WAG, Z); Andohahela PN, Andranolava, 24°41’S 46°45’E, 28.VIII.1996, ster., Randriamampionona 1338 (MO, P, TAN); Mandena forestry station, 24°57’S 47°00’E, 0-10 m, 18.XI.1996, ster., A. Randrianasolo 546 (MO, P, TAN); N of Tolagnaro, Ste. Luce area, [24°47’S 49°09’E], 0-10 m, 19.XI.1996, ster., A. Ran- drianasolo 569 (MO, P, TAN); Bevoay, Androranga, [24°29’39’’S 47°05’39’’E], 171 m, 18.XI.2009, fr., Razakamalala 4680 (G, MO, P, TAN); Ifarantsa, Ivorona, 24°29’09’’S 47°04’43’’E, 35 m, 10.XII.2010, fr., Razakamalala 5033 (G, K, MO, P, TAN); Man- dena, [24°57’S 47°00’E], 16.XI.1948, fl.,Réserves Naturelles 1348 (G, MO, P); ibid. loco, 18.IX.1950, Service Forestier 1545 (P, TEF); Marofanenitra, 25.IX.1951, Service Forestier 4043 (P, TEF); Mandena, [17°30’S 48°38’E], 29.X.1953, fl.,Service Forestier 7826 (P, TEF); ibid. loco, 3.IX.1953, fl.,Service Forestier 8170 (MO, P, TEF); Col du Maningotry, 300-400 m, 27.IX.1953, fl.,Service Forestier 8510 (G, MO, P, TEF); Man- dena, 15.XII.1954, fr., Service Forestier 15622 (MO, P, TEF). Taxonomic treatment of Abrahamia (Anacardiaceae) 53

8. Abrahamia ditimena (H. Perrier) Randrian. & Lowry, comb. nova (Fig. 7).

≡ Protorhus ditimena H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 263. 1944.

Lectotypus (designated here): Madagascar. Prov. Mahajanga: forêt entre Andilamena et Mandritsara, c. 900 m, IX.1922, fl.,Perrier de la Bâthie 15051 (P [P00722640]!; isolecto-: P [P06775371]!). Syntypi: Madagascar. Prov. Toamasina: Périnet [Analamazaotra], [18°56’S 48°26’E], X.1925, fl.,Louvel 4 (P [P06168986]!); Analamazaotra, [18°56’S 48°26’E], 800 m, XII.1932, fl.,Perrier de la Bâthie 6403 (P [P06775208, P06775370]!, TAN, US-2494672 image seen); ibid. loco, [18°56’S 48°26’E], 800 m, s.d., fl.,Perrier de la Bâthie 10180 (P [P06775219, P06775375]!).

= Protorhus fulva Engl. in Bot. Jahrb. Syst. 1: 421. 1881. Typus: Madagascar: sine loco, s.d., Chapelier s.n. (holo-: P, not found).

= Protorhus sericea var. opaca Lecomte, Madagascar Bois Analamazaotra: 74. 1922. Lectotypus (designated here): Madagascar. Prov. Toamasina: forêt d’Analamazaotra, [18°56’S 48°26’E], s.d., fr., Thouvenot 146 (P [P06169004]!; isolecto-: P [P06169003, P06169005]!, TAN [TAN002001]!).

Description Trees 3-30 m tall, 8-35 cm DBH, bark with white milky latex turning pinkish; young twigs glabrous, with lenticels, waxy. Leaves subopposite; blades obovate to oblanceolate or narrowly elliptic, 2.5-12 3 1.5-5 cm, coriaceous, apex emarginate or retuse, rarely rounded, margin entire, base acute to cuneate, adaxial surface glabrous, abaxial surface very minutely and sparsely pubescent when young, glabrescent, venation craspedodromous, midvein prominent abaxially, secondary veins 20-33 pairs, 1-3 mm apart, prominent on both surfaces, tertiary veins raised on both surface; petiole (5-)7-20 mm long, canaliculate on adaxial surface, glabrous or very lightly pubescent. Inflorescence axillary or terminal, a panicle, 2-10 cm long, axes terete or flattened, tomentose or glabrescent; bracts triangular, deltate or narrowly deltate to narrowly triangular, 1-2.5 3 1-1.5 mm, adaxial surface glabrous, abaxial surface pubescent. Male and female flowers 4- or 5-merous; pedicel c. 0.3 -0.4 mm long, scabrous; calyx lobes deltate, c. 1 3 1-1.2 mm, valvate, adaxial surface glabrous, abaxial surface strigose; corolla lobes ovate, 2.5-3 3 1.5-2 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent; stamens 4 or 5. Male flowers with filaments 1-2 mm long, glabrous, white, anthers 0.5-0.8 mm long, obloid to slightly ovoid, yellow, glabrous; disk 1-1.2 mm in diam., glabrous; ovary rudimentary. Female flowers with vestigial stamens, filaments c. 1 mm long, glabrous, white, anthers 0.2-0.3 mm long, ovoid to sub-globose, sterile, yellow, glabrous; disk c. 1.5 mm in diam., glabrous; ovary widely to very widely ovoid, 1 3 1 mm, pubescent; 54 Boissiera 71

style shortly trifid, subapically inserted, 0.2-0.3 mm long, glabrous, stigmas capitate. Fruits 2-3 3 1.4-1.6 cm, usually globose or ellipsoid to obovoid, slightly asymmetrical, surface velutinous, with longitudinal striations.

Vernacular names “Ditimena’’, “Hazombarorana’’, “Sandramivavy’’.

Distribution, ecology Abrahamia ditimena is the most widespread member of the genus (Map 3). It has been and phenology found in diverse habitats, from the very humid (eastern rainforests or in the Sambirano region) to dry forests (central forests and part of the western slopes), and on a range of substrates, from sand to laterite or on soils derived from gneiss or granite. It flowers between October and December, and its fruits mature in April and May.

Conservation status With an EOO of 376,143 km2, a minimum AOO of 336 km2, and more than 100 recorded localities, A. ditimena is assigned a preliminary conservation assessment of “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Among the species of Abrahamia with velutinous mature fruits, A. ditimena most closely resembles A. sericea, but differs strikingly by its very coriaceous (vs subcoriaceous) leaves and smaller number of secondary veins (20-33 vs 40-70). Among the syntypes of Protorhus ditimena, we have selected Perrier de la Bâthie 15051 as the lectotype because it has more fully developed inflorescences than the three other collections assigned to Abrahamia ditimena (viz. Perrier de la Bâthie 6403 and 10180, and Louvel 4). Two additional syntypes (Louvel 112 and Ursch 195) are referable to A. thouvenotii. The Paris herbarium contains three syntypes of Protorhus sericea var. opaca, from which we have chosen the sheet bearing the original label as the lectotype.

Additional material Madagascar. Prov. Antananarivo: Anjozorobe, forêt NNE d’Andranomay (13 km examined SE d’Anjozorobe), 18°28’38’’S 47°57’18’’E, 1340-1400 m, 18-20.XII.1996, fl.,Rali - manana et al. 44 (MO, P, TAN); Manjakandriana, Mandraka, [18°53’S 47°55’E], 28.V.1950, fr., Service Forestier 1897 (P); Ankazobe, Ambohitantely, [18°09’S 47°18’E], 1600 m, VI.1957, fr., Service Forestier 18012 (P, TEF); forêt relique, PK 169, Maridaza-Ankazobe, [18°19’S 47°06’E], 1475 m, 30.III.1961, fr., Service Forestier 19881 (G, MO, P, TEF). Prov. Antsiranana: Bemanevika, Bandrakorony, 13°45’04’’S 47°58’51’’E, 99 m, 24.I.2009, fr., Bernard et al. 1341 (MO, P, TAN); Andapa, descente de l’Anjanaharibe, 1100 m, 26.XII.1950, bud, Cours 3877 ( P, TAN); Anjanaharibe, W d’Andapa, [14°40’S 49°31’E], 1200-1400 m, 10.XII.1950- 3.I.1951, bud, Humbert 24600 (MO, P); Vohémar, Daraina, forêt de Binara, 13°15’S 49°37’E, 1010 m, 28.XI.2006, bud, Gautier et al. 4936 (G, MO, P, TAN); Sambava, Marojejy NP, 14°26’S 49°45’E, 700-1000 m, 2.X.1994, fr., Lewis et al. 1248 (G, MO, P, TAN); SW Andapa, SW edge of Anjanaharibe-Sud reserve, 14°48’15’’S 49°26’45’’E, 950 m, 8.VIII.1997, y. fr., McPherson 17302 (G, K, MO, NY, P, TAN, WAG); PN Marojejy, 11 km NW Manantenina, 14°16’00’’S 49°44’30’’E, 1075 m, 28.X.1996, Taxonomic treatment of Abrahamia (Anacardiaceae) 55

fr., Rakotomalaza et al. 806 (G, MO, P, TAN); Ambanja, Bandrakorony, 13°45’36’’S 47°59’58’’E, 140 m, 22.I.2009, fr., Rakotovao et al. 4263 (MO, P, TAN); SW d’An- dapa, Anjanaharibe-Sud RS, Mandritsarakely, 14°43’10’’S 49°27’12’’E, 1700 m, 14.II.1995, fr., Ravelonarivo et al. 625 (MO, P, TAN); Andapa, Anjanaharibe-Sud RS, env. d’Ampiferantany, 14°47’45’’S 49°27’54’’E, 1161 m, 22.V.1995, fr., Ravelonarivo et al. 807 (MO, P, TAN); Doany, Anjialavahely, Ankijandambo, 14°14’58’’S 49°26’19’’E, 1546 m, 7.III.2006, fl.,Ravelonarivo et al. 1737 (G, K, MO, P, TAN); Anjialavabe, Ankiakabe, 14°10’47’’S 49°22’01’’E, 1567 m, 20.II.2007, fr., Ravelona- rivo et al. 2161 (G, MO, P, TAN); Anjanaharibe-Sud RS, au S du campement à Man- drintsarahely, 14°46’S 49°30’E, 985 m, 10.VII.1996, y. fr., Razafimandimbison et al. 236 (MO, P, TAN); Ambanja, Ankaraha, canton Marovato, [13°53’S 48°15’E], 24.X.1950, y. fr., Réserves Naturelles 2134 (P, TAN); Anjanaharibe RS, [14°36’S 49°25’E], 1100 m, 26.XII.1950, fl.,Service Forestier 940 (G, P, TEF). Prov. Fian- arantsoa: Andringitra RN, camp II, 22°13’40’’S 47°00’13’’E, 810 m, 22-29.XII.1993, fr., Lewis et al. 881 (MO, P, TAN); ibid. loco, 22°13’40’’S 47°00’13’’E, 810 m, 22-29. XII.1993, fl.,Lewis et al. 912 (G, K, MO, NY, P, TAN); Ranomafana PN, Vatohara- nana, 21°16’S 47°26’E, 100-1100 m, 18-31.I.1993, fl.,Malcomber et al. 2002 (G, K, MO, NY, P, TAN); ibid. loco, 21°16’S 47°26’E, 1000-1100 m, 18-31.I.1993, ster., Malcomber 2051 (MO, P, TAN); Andringitra RN, Ivohibe, de part et d’autre du fleuve Iatara, 22°13’20’’S 47°01’29’’E, 720 m, 12-14.XII.1994, fl.,Rakotovao 303 (G, MO, P, TAN); Befotaka, Forêt d’Andranomangatsy, Parc National de Befotaka, 23°48’54’’ 49°59’56’’E, 2.VI.2012, ster., Rapanarivo et al. 665 (BR, K, MO, P, TAN); Farafan- gana, Ivongo, [22°07’S 46°49’E], 23.IV.1955, fr., Réserves Naturelles 7178 (P, TEF); Ampamaherana, [21°29’15’’S 47°19’20’’E], 15.XII.1949, fl.,Service Forestier 1254 (P, TAN, TEF); ibid. loco, [21°29’15’’S 47°19’20’’E], 5.IV.1949, fr., Service Forestier 2018 (G, MO, P, TAN); Levotsibe, Ft. Carnot, [21°51’S 47°26’E], 23.V.1952, fr., Service Forestier 5453 (MO, P); Midongy du Sud, [23°35’S 47°01’E], 30.VII.1955, fr., Service Forestier 13931 (P, TEF); Ifanadiana, Ranomafana, [21°02’S 47°18’E], 13.VI.1955, y. fr., Service Forestier 14243 (P, TEF); Ft. Carnot, Andrambovato-To- longoina, [21°31’S 47°25’E], 1951, ster., Service Forestier 36B-R-230 (P, TEF). Prov. Mahajanga: Befandriana Nord, Antsakabary, Andapanomby, Makira NW, 15°21’17’’S 49°06’25’’E, 25.IX.2007, fr., Bernard et al. 455 (G, K, MO, P, TAN); Mandritsara, Makira, Ambolotsangana, 15°31’S 49°05’E, 1169 m, 26.IV.2007, fr., Lehavana et al. 451 (MO, P, TAN); ibid. loco, 15°30’S 49°05’E, 1301 m, 28.IV.2007, fr., Lehavana et al. 512 (G, MO, P, TAN); Mangindrano, forêt d’Antsahivo, 14°14’54’’S 49°00’45’’E, 1365 m, 1.XII.2005, fl.,Randrianarivelo et al. 480 (MO, P, TAN); massif de l’Am- batomirahavavy, E de Mangindrano, [14°28’30’’S 48°46’00’’E], 1700 m, 4.II.1951, fr., Service Forestier 977 (MO, P, TEF). Prov. Toamasina: Moramanga, Ambatovy, 18°51’40’’S 48°19’42’’E, 1180 m, 21.XII.2004, fl.,Andriamahefarivo et al. 89 (MO, P, TAN); ibid. loco, 18°49’21’’S 48°20’07’’, fr., Andriatsiferana et al. 2528 (G, MO, P, TAN); ibid. loco, 18°49’27’’S 48°20’13’’E, fr., Andriatsiferana 2551 (MO, TAN); Rant- abe, Beanana, 15°43’47’’S 49°27’16’’E, 175 m, 27.II.2002, fr., Antilahimena et al. 949 (MO, P, TAN); Mahavelona, Masoala PN, Ambohitsitondroina, 15°26’14’’S 56 Boissiera 71

49°57’28’’E, 1158 m, 6.X.2003, fr., Antilahimena 2121 (G, K, MO, NY, P, TAN); Antsirabe-Sahatany, Anjiahely, 15°24’19’’S 49°30’38’’E, 360 m, 28.VI.2004, fr., Antilahimena 2594 (MO, P, TAN); Ambinanitelo, Andampy forest, 15°19’03’’S 49°24’06’’E, 800 m, 1.IX.2004, fr., Antilahimena 2712 (G, MO, P, TAN); Maroantse- tra, Maimborondro forest, 15°19’23’’S 49°23’52’’E, 1106 m, 6.IX.2004, fr., Antila- himena 2797 (MO, P, TAN); Andasibe, Ambatovy forest, 18°51’00’’S 48°19’16’’E, 1097 m, 14.I.2005, y. fr., Antilahimena et al. 3145 (G, K, MO, P, TAN); ibid. loco, 18°49’19’’S 48°20’08’’E, 1094 m, 3.II.2005, fr., Antilahimena et al. 3300 (MO, P, TAN); ibid. loco, 18°51’06’’S 48°18’32’’E, 1105 m, 4.III.2005, fr., Antilahimena 3594 (G, K, MO, P, TAN); ibid. loco, 18°52’08’’S 48°19’09’’E, 1080 m, 9.III.2005, fr., Anti- lahimena et al. 3649 (G, K, MO, P, TAN); ibid. loco, 18°50’53’’S 48°17’47’’E, 1020 m, 10.III.2005, fr., Antilahimena et al. 3678 (G, K, MO, P, TAN); ibid. loco, 18°52’06’’S 48°16’35’’E, 1053 m, 11.XI.2005, y. fr., Antilahimena et al. 4205 (G, K, MO, P, TAN); ibid. loco, 18°51’34’’S 48°18’05’’E, 1137 m, 1.XII.2005, fl.,Antilahimena & Razafin- drabe 4297 (G, P); ibid. loco, 18°50’22’’S 48°18’32’’E, 1183 m, 19.XII.2005, fl.,Anti - lahimena 4485 (G, K, MO, P, TAN); ibid. loco, 18°48’29’’S 48°20’20’’E, 1080 m, 21.XII.2005, fl.,Antilahimena 4505 (G, K, MO, P, TAN); ibid. loco, 18°50’22’’S 48°17’52’’E, 1136 m, 11.XII.2006, fl.,Antilahimena 5121 (G, MO, P, TAN); ibid. loco, 18°51’40’’S 48°18’01’’E, 1116 m, 17.V.2008, fr., Antilahimena 6180 (G, MO, P, TAN); ibid. loco, 18°50’09’’S 48°19’24’’E, 1106 m, 12.III.2009, fr., Antilahimena 7017 (G, K, MO, P, TAN); ibid. loco, 18°50’21’’S 48°18’15’’E, 1090 m, 17.V.2010, fr., Bernard 1572 (MO, P, TAN); Ambatondrazaka, Ankaraoka, N de la route de Nickel- ville, 1200 m, 29.XII.1944, bud, Cours 2114 (P [2 sheets], TAN); sommet de l’Analavorikely, c. 1200 m, 29.XII.1944, bud, Cours 2135 [= Herb. Station Agric. Alaotra 2135] (MO, P [2 sheets], TAN); Périnet [Analamazaotra], [18°56’S 48°26’E], s.d., fr., Descoings s.n. (MO); Masoala PN, E slope of Ambohitsitondroina Mahave- lona, 15°26’00’’S 49°57’34’’E, 1125 m, 24.II.2003, fl.,Lowry et al. 6166 (G, K, MO, NY, P, TAN); Mantadia, beyond the graphite mine, 18°55’S 48°25’E, 1200 m, 8. XI.1994, fl.,McPherson et al. 16546 (G, MO, P, TAN); Ambatovy, 18°51’25’’S 48°17’50’’E, 1050 m, 27.II.1998, fr., McPherson 17481 (G, K, MO, NY, P, TAN, WAG); ibid. loco, 18°50’59’’S 48°18’32’’E, 1115 m, s.d., fr., Miandrimanana 278 (MO, TAN); Masoala, Ambanizana, 15°41’S 49°58’E, 50-200 m, 21-22.VII.1994, fr., Rabe 110 (G, MO, P, TAN); Ambatovy, 18°51’07’’S 48°18’47’’E, 1100 m, 30.I.1997, fr., Rakotomalaza et al. 1033 (MO [2 sheets], P, TAN); ibid. loco, 18°51’34’’S 48°18’25’’E, 3.III.1997, fr., Rakotomalaza et al. 1213 (MO, P, TAN); ibid. loco, 18°51’S 48°18’E, 1121 m, 7.III.1997, fr., Rakotomalaza et al. 1266 (MO, P, TAN); ibid. loco, 18°49’47’’S 48°18’48’’E, 1160 m, 16.V.1997, fr., Rakotomalaza 1330 (MO, P, TAN); ibid. loco, Rakotomalaza 1331 (MO, P, TAN); Zahamena PN, Antanandava, 17°29’30’’S 48°44’48’’E, 900 m, 13.VII.2000, fr., Rakotondrajaona et al. 130 (MO, P, TAN); Zahamena PN, Ambodimangavalo, Moango, 17°33’32’’S 48°53’55’’E, 800 m, 4.V.2003, fr., Rakotondrajaona et al. 276 (CNARP, G, K, MO, P, TEF); Ambohibary, Ampitambe, Ambatovy, 18°51’08’’S 48°17’43’’E, 1106 m, 15.II.2005, fr., Rakotovao et al. 1174 (G, MO, P, TAN); Andasibe, Menalamba, Ambatovy, 18°49’33’’S Taxonomic treatment of Abrahamia (Anacardiaceae) 57

Fig. 7. Abrahamia ditimena (H. Perrier) Randrian. & Lowry. A. Upper and lower leaf surfaces, young fruits, from side [Plunkett et al. 2387]; B. Fruits, from above [Rakotovao et al. 4263].

Photos: A: P. Lowry; B: C. Rakotovao 58 Boissiera 71

48°20’03’’E, 1065 m, 1.IV.2005, fr., Rakotovao et al. 1727 (G, MO, P, TAN); Mor- arano Gara, forêt de Bevetraka, 18°47’10’’S 48°22’09’’E, 974 m, 23.IV.2012, fr., Rakotovao 5758 (G, MO, P, TAN); Ambohibary, Ambatovy, Behontsa, 18°52’35’’S 48°19’54’’E, 987 m, 20.XII.2011, fl.,Ramahenina et al. 71 (G, K, MO, P, TAN); ibid. loco, 18°49’15’’S 48°20’00’’E, 1100 m, 15.V.1999, fr., Ranaivojaona 236 (G, MO, P, TAN); Vatomandry, Ambalabe, Ambinanindrano II, 19°10’09’’S 48°34’40’’E, 598 m, 11.III.2005, fr., Ranaivojaona et al. 1157 (G, K, MO, P, TAN); Analamazaotra RS, SE of entry of the Reserve, [18°56’S 48°26’E], 960 m, 15.VII.1996, fr., A. Randrianasolo 402 (MO, P, TAN); ibid. loco, ster., A. Randrianasolo 403 (MO, TAN); ibid. loco, A. Randrianasolo 409 (MO, TAN); ibid. loco, 950 m, 16.VII.1996, fr., A. Randrianasolo 410 (MO, P, TAN); ibid. loco, A. Randrianasolo 417 (MO, P, TAN); ibid. loco, 950 m, 18.VII.1996, ster., A. Randrianasolo 436 (MO, P, TAN); ibid. loco, A. Randrianasolo 437 (MO, P, TAN); Malompy, W Analamazaotra RS, 900 m, 28.X.1996, bud, A. Randrianasolo 513 (MO, P, TAN); Analamazaotra RS, toward “carrière de sable”, 960 m, 28.X.1996, fr., A. Randrianasolo 514 (MO, P, TAN); Ambalabe, W of Foara Toby, 19°10’02’’S 48°34’24’’E, 650 m, 15.III.2005, fr., A. Randrianasolo et al. 1017 (G, MO, P, TAN); Zahamena PN, Ambodimangavolo, Ambatoharanana, 17°33’57’’S 48°53’58’’E, 740 m, 27.X.2000, fr., Randrianjanaka et al. 585 (MO, P, TAN); Didy, Antsevabe, forêt de Sahananto, 17°59’18’’S 48°35’31’’E, 1236 m, 13.XII.2005, bud, Rasoafaranaivo 92 (G, MO, P, TAN); Andasibe, Andasifahatelo, Ambatoaranana, 18°47’59’’S 48°24’42’’E, 1081 m, 25.IV.2012, fr., Rasoazanany et al. 128 (G, MO, P, TAN); Ambanizana, Anjahana, 15°38’48’’S 49°57’57’’E, 230 m, 21-25.V.1999, fr., Ratovoson 91 (G, MO, P, TAN, US); Zahamena PN, Ambodimangavolo, 17°39’07’’S 48°58’14’’E, 11.VIII.2000, fr., Ratovoson et al. 268 (MO, P, TAN); ibid. loco, 17°30’S 48°44’E, 975 m, 25.IX.2001, fr., Ratovoson et al. 514 (G, CNARP, MO, P, TEF); Ambatovy, Analamay, 18°50’50’’S 48°18’28’’E, 1113 m, 10.II.2005, fr., Razafindra- ibe et al. 94 (G, K, MO, P, TAN); Anosibean’ala, relique forestière d’Anosibeambo, 19°26’12’’S 48°12’15’’E, 747 m, 8.IV.2001, fr., Razafitsalama et al. 52 (G, MO, P, TAN, TEF, US); Lakato, Ambodigavo, 19°07’50’’S 48°24’03’’E, 829 m, 31.V.2007, fr., Razanatsima et al. 253 (G, K, MO, NY, P, TAN); Sahamaloto, Périnet, [18°56’S 48°22’E], 3.VI.1950, y. fr., Service Forestier 1407 (P, TAN, TEF); ibid. loco, [18°56’S 48°26’E], 3.IV.1951, fr., Service Forestier 3313 (P, TAN, TEF); Andramarolasy, Mora- manga, [18°36’S 48°01’E], 30.IV.1952, fr., Service Forestier 5069 (G, K, MO, NY, P); massif de l’Ambohitsitondroina de Mahalevona, [15°25’S 49°58’E], 1100 m, 5.XII.1953, fr., Service Forestier 8722 (MO, P, TEF); massif de Beajanda, [15°16’S 50°04’E], 1000 m, 31.XII.1953, fr., Service Forestier 8832 (MO, P, TEF); ibid. loco, 1100 m, 1.I.1954, fl.,Service Forestier 8837 (P, TEF); Fénérive-Est, Tampolo, [17°17’00’’S 49°23’30’’E], 22.IV.1954, fr., Service Forestier 10302 (P, TEF); ibid. loco, 3.I.1955, fl.,Service Forestier 12496 (P, TEF); Sahavalanina-Antenina, Fénérive, [17°02’S 49°24’E], 25.IV.1955, fr., Service Forestier 15635 (G, MO, P, TEF); Amboat- avy, Manjakanahiana, [18°47’S 47°47’E], 7.V.1955, fr., Service Forestier 15865 ( P, TEF); Ambohimiarina, [19°18’S 48°21’E], 18.IV.1957, fr., Service Forestier 16955 (P); Analamazaotra, Périnet, [18°56’S 48°26’E], 800 m, 9.IV.1958, fr., Service Taxonomic treatment of Abrahamia (Anacardiaceae) 59

Forestier 17957 (P); ibid. loco, [18°56’S 48°26’E], 15.IV.1958, fr., Service Forestier 19026 (P, TEF); ibid. loco, [18°49’00’’S 48°21’30’’E], 8.VIII.1962, ster., Service Forestier 21240 (MO, P); Koubaville, entre Moramanga et Anosibe, [19°16’00’’S 48°13’30’’E], 23.XII.1965, fl.,Service Forestier 24402 (MO, P, TEF); Analamazaotra, Périnet, [18°56’S 48°26’E], II.1966, fl.,Service Forestier 25718 (P); ibid. loco, 25.VI.1966, ster., Service Forestier 25936 (P); Nangaranana, Canton de Marovoay, [18°47’S 48°15’E], II.1970, ster., Service Forestier 26944 (P, TEF); Périnet, [18°56’S 48°26’E], V.1971, ster., Service Forestier 30096 (P, TEF); ibid. loco, [18°56’S 48°26’E], 13.II.1951, ster., Service Forestier 5B-R-172 (P, TEF); ibid. loco, 29.IX.1952, ster., Service Forestier 167B-R-172 (P, TEF); Masoala Peninsula, S of Ambanizana, 15°40’24’’S 49°57’51’’E, 110-260 m, 18.XII.1994, ster., Vasey & Behasy 202 (MO); ibid. loco, Vasey & Behasy 228 (MO); 10 km N of Fénérive-Est, Tampolo STF, 17°17’S 49°25’E, 10 m, 12.V.1991, fr., Zarucchi et al. 7388 (MO, P, TAN). Prov. Toliara: Amboasary, entre Andohahela et l’Elakelaka, 600-800 m, I-II.1934, y. fr., Humbert 14048 (P); Andohahela PN, plot d’Itrotro, N d’Eminiminy, 24°41’S 46°48’E, 400 m, 28.III.1996, ster., Laha et al. 19 (MO, P, TAN); ibid. loco, Laha et al. 59 (MO, P, TAN); Andohahela PN, Andranomitily, plot permanent n° 2, 24°36’18’’S 46°44’25’’E, 800 m, 15.V.2001, fr., Randriampampionona 961 (G, K, MO, NY, P, TAN); Ivorona, ridge above right bank of Andriambe Creek above village, 24°49’11’’S 46°55’31’’E, 628 m, 9.XII.2009, y. fr., Plunkett et al. 2387 (K, MO, NY, P, TAN); Andohahela PN, Emalio, Andranolava, 24°41’S 46°45’E, 28.VIII.1996, ster., Randriamampionona 1319 (MO, P, TAN); Manja, Vondrove, forêt galerie bordant la rivière Mangoky, 21°48’03’’S 44°10’43’’E, 86 m, 20.IX.2007, ster., Rasoafaranaivo et al. 282 (MO, P, TAN); Ranomafana Sud, forêt de Tsitongambarika, 24°35’44’’S 47°02’43’’E, 846 m, 23.XI.2009, fl. & bud,Razakamalala 4854 (G, K, MO, P, TAN). 60 Boissiera 71

9. Abrahamia ellipticarpa Randrian. & Lowry, spec. nova (Fig. 8).

Typus: Madagascar. Prov. Toliara: Rég. Anosy, Vinanibe, à l’W de Fort-Dauphin, [25°03’S 46°56’E], 7.XII.1968, fr., Service Forestier 28611 (P [P00722641]!; iso-: G [G00341714]!, K!, MO-6684134!, P [P00722642]!, TEF [TEF000884]!).

Diagnosis Abrahamia ellipticarpa Randrian. & Lowry can be distinguished from other member of the genus by its large, glabrous, ellipsoid fruits measuring 3-3.5 3 1.5-1.8 cm.

Description Trees c. 10-12 tall, bark with white latex; young twigs with appressed indument. Leaves alternate, subopposite or opposite; blade oblanceolate to narrowly elliptic, 3-9.5 3 1.3-3 cm, subcoriaceous, apex acute or sometimes retuse or rarely emarginate, margin entire, base cuneate, adaxial surface glabrous, abaxial surface with appressed indument, more so along the midvein and margin, venation craspedodromous, midvein prominent abaxially, secondary veins 18-30 pairs, more or less parallel, 2-3 mm apart, slightly raised adaxially and prominent abaxially, tertiary veins very few, impressed and barely visible; petiole 5-10 mm long, c. 1 mm in diam., canaliculate on adaxial surface, pubescent. Inflorescencesand flowers unknown. Fruits 3-3.5 3 1.5-1.8 cm, ellipsoid, glabrous, with longitudinal striations.

Distribution, ecology Abrahamia ellipticarpa has been recorded from sublittoral forest on sand at several sites and phenology in the Fort-Dauphin area, including Mandena, Vinanibe, and Petriky (Map 1). It has been collected in flower in September and in fruit in December.

Conservation status With an EOO of 28 km2, an AOO of 12 km2, and three known locations, 2 of which are situated within protected areas (Mandena and Petriky) whereas the third one is highly degraded and subjected to continued degradation from uncontrolled harvesting and possibly also fire leading to a projected decline in all 5 sub-criteria,A. ellipticarpa is assigned a preliminary conservation status of “Endangered’’ [EN B1ab(i,ii,iii,iv,v)+2ab (i,ii,iii,iv,v)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia ellipticarpa is vegetatively similar to A. oblongifolia but can be distinguished by its fewer secondary veins (18-30 vs 35-40), impressed and barely visible (vs slightly raised and very visible) tertiary venation, and large fruits (3-3.5 3 1.5-1.8 vs 2 3 1 cm) with evident (vs obscure) longitudinal striations.

Paratypi Madagascar. Prov. Toliara: Petriky, N shore of Lake Andranany, 25°03’S 46°53’E, 0-10 m, 14.IV.1989, ster., Gereau 3376 (MO, P, TAN); env. de Fort-Dauphin, entre le Pic St. Louis et la mer, 1-25 m, 20.IX.-6.X.1928, fl.,Humbert 5984 (G, P [2 sheets]); Man- Taxonomic treatment of Abrahamia (Anacardiaceae) 61

dena forestry station, 24°57’S 47°00’E, 0-10 m, 20.XI.1996, ster., A. Randrianasolo 578 (MO, P, TAN); ibid. loco, 3.IX.1953, bud, Service Forestier 8171 (MO, P, TEF); ibid. loco, 12.XII.1952, ster., Service Forestier 207-R-16 (P).

B C

1 cm 2 cm 2 cm

Fig. 8. Abrahamia ellipticarpa Randrian. & Lowry. A. Branch with leaves; B. Infrustescence showing detail of pedicel; C. Fruit.

[A: A. Randrianasolo 578, MO; B-C: Service Forestier 28611, MO] Drawings: R.L. Andriamiarisoa 62 Boissiera 71

10. Abrahamia elongata Randrian. & Lowry, spec. nova (Fig. 9, 10A).

Typus: Madagascar. Prov. Toamasina: Rég. Analanjirofo, Maroantsetra, Nosy Mang- abe, 15°30’S 49°46’E, 0-330 m, 28.IX-5.X.1989, fl.,Schatz 2770 (holo-: MO-3759750!; iso-: K, P [P06169108]!, TAN [TAN001988]!, WAG [WAG1435428] image seen).

Diagnosis Abrahamia elongata Randrian. & Lowry can be distinguished from its congeners by its oblanceolate or narrowly elliptic leaves that are 8.5-26.5 3 2.5-6.5 cm, combined with very pronounced secondary and tertiary veins, long petiole (1.5-3 cm), and a relatively large and glabrous fruits (2-4 3 1-1.5 cm).

Description Trees 4-25 m tall, 10-15 cm DBH, bark with white milky latex; young twigs waxy or lanate to pubescent at the tips. Leaves alternate; blades oblanceolate to narrowly elliptic, 8.5-26.5 3 2.5-6.5 cm, coriaceous, apex emarginate, rounded, shortly acuminate or sometimes acuminate, margin entire, base acute to cuneate, adaxial surface glabrous or with short appressed indument, abaxial surface sericeous to glabrescent, venation craspedodromous, midvein prominent abaxially, secondary veins 25-50 pairs, more or less parallel, 1.5-10 mm apart, very prominent abaxially, tertiary veins raised on both surfaces; petiole 15-30 mm long, canaliculate on adaxial surface, with short, appressed indument when young, later glabrescent or glabrous. Inflorescence axillary, a panicle, 1.5-6 cm long, the branches regular, pubescent or tomentose. Male and female flowers 5-merous; pedicel 0.1-0.3 mm long, pubescent; calyx lobes shallowly deltate to shallowly triangular, c. 1 3 1.2-1.5 mm, adaxial surface glabrous, abaxial surface pubescent, slightly imbricate in male flowers, valvate in female flowers, persistent in fruit; corolla lobes ovate, 2.5-3 3 1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent, white or pale yellow; stamens 5. Male flowers with filaments 1.5-2 mm long, glabrous, white, anthers 1 mm long, obloid to slightly ovoid, yellow, glabrous; disk c. 1 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments c. 1 mm long, glabrous, white, anthers 0.3-0.5 mm long, ovoid, sterile, yellow, glabrous; disk annular, cupuliform, c. 1.5-2 mm in diam., glabrous; ovary very widely ovoid or obovoid, slightly asymmetrical, 1.5-2 3 1.5-1.8 mm, glabrous, placentation subapical; style unbranched, latero-subapically inserted, c. 1 mm long, stigmas capitate, glabrous. Fruits 2-4 3 1-1.5 cm, ellipsoid or slightly asymmetrical, glabrous, with longitudinal striations.

Vernacular names “Ditimena’’, “Hazombarorana’’, “Tarantana’’.

Distribution, ecology Abrahamia elongata occurs from eastern coastal littoral forest on sand to premontane and phenology forests on laterite up to elevations of 900 m, and extends from Marojejy and near Ambi- lobe in the north to Manombo in the south (Map 2). It flowers between September and November, and fruits in November and December. Taxonomic treatment of Abrahamia (Anacardiaceae) 63

Conservation status With an EOO of 147,171 km2, an AOO of 120 km2, and 32 recorded localities, many of which occur within protected areas (Ambalabe, Anjanaharibe-Sud, Betampona, Manombo, Marojejy, Masoala, Nosy Mangabe, and Zahamena), A. elongata is assigned a preliminary conservation assessment of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia elongata most closely resembles A. latifolia, with which it shares large leaves with many secondary veins and exclusively axillary inflorescences. However,A. elongata differs by its coriaceous (vs subcoriaceous) leaves with craspedodromous (vs semicraspedodromous) venation, and its longer petiole (15-30 vs 10 mm). It also resembles A. sambiranensis vegetatively, but differs by having leaves with evident, raised tertiary veins on both surfaces (vs lacking in A. sambiranensis) and whose apex is flat in pressed material (vs folded), and by its fruits that are glabrous (vs pubescent when young and glabrescent at maturity in A. sambiranensis).

Paratypi Madagascar. Prov. Antsiranana: Sambava, forêt de Tsihomanaomby, 14°06’08’’S 50°02’50’’E, 103 m, 15.II.2006, ster., Andrianjafy 1582 (MO, P, TAN); Marojejy PN, campement 1, 14°26’13’’S 49°46’32’’E, 460 m, 12.X.1996, y. fr., Rakotomalaza et al. 752 (MO, P, TAN); Andapa, Anjanaharibe-Sud RS, [14°36’S 49°27’E], 3-7. XII.1996, fr., Ravelonarivo et al. 1052 (MO, P, TAN); Antalaha, forêt Ampohafana, [14°48’30’’S 49°51’30’’E], 9.IX.1955, fl.,Service Forestier 15064 (P [2 sheets], TEF); Andapa, Andilandrano, [14°45’S 49°34’E], 12.XII.1964, fr., Service Forestier 25019 (G, MO, NY, P, TEF); S de Sambava, [14°18’S 50°11’E], 30.XI-1.XII.1966, fr., Service Forestier 27110 (P, TEF); Vohémar, Fanambana, 22.XII.1954, ster., Service Forestier 54-R-151 (P). Prov. Fianarantsoa: forêt de Manombo, S de Farafangana, [23°02’10’’S 47°44’50’’E], 17.X.1964, fl.,Service Forestier 23654 (P, TEF); forêt entre Manakara et Vohipeno (PK 30), [22°22’S 47°51’E], 21.X.1964, y. fr., Service Forestier 23696 (P, TEF). Prov. Toamasina: Manompana, forêt d’Antanambao- Ambodimanga, 16°45’38’’S 49°42’22’’E, 7 m, 5.X.2003, fr., Andrianarivelo 30 (G, MO, P, TEF); Maroantsetra, Ambatoledama forest, 15°17’S 50°00’E, VI.2003, y. fr., Antilahimena 1996 (G, K, MO, NY, P, TAN); Rantabe, Behovana, 15°47’48’’S 49°32’10’’E, 815 m, 31.X.2003, fr., Antilahimena 2251 (G, K, MO, P, TAN); Antsir- abe, Sahatany, Anjiakely, 15°24’19’’S 49°30’28’’E, 360 m, 27.VI.2004, fr., Antila- himena 2579 (G, K, MO, P, TAN); Nosy Mangabe, 15°29’S 49°45’E, 10 m, 18-26. IV.1990, ster., Carlson 126 (MO); ibid. loco, Carlson 150 (MO); ibid. loco, Carlson 188 (MO); Sahalampy (Onibe), [17°50’S 48°25’E], 700 m, XI.1938, fl.,Cours 1160 (MO, P); Mananara, 1.IX.1920, fl.,Decary 59 (P, TAN); Soanierana-Ivongo, forêt d’Antanambao-Ambodimanga, 16°45’40’’S 49°42’35’’E, s.d., fr., Rabehivohitra 4354 (MO, P, TAN, TEF); Sainte Marie, forêt d’Ambohidena, 16°51’11’’S 49°57’10’’E, 10 m, 14.V.2003, ster., Rabenantoandro 1480J (MO); Fénérive-Est, Tampolo station for- estière, 17°17’S 49°23’E, 10 m, 11.X.1999, ster., A. Randrianasolo 596 (MO); Vato- mandry, Ambalabe, Ambinanindrano II, 19°09’38’’S 48°35’02’’E, 600 m, 9.X.2005, fl.,A. Randrianasolo et al. 1045 (MO, NY, P, TAN); ibid. loco, 19°09’01’’S 48°34’55’’E, 64 Boissiera 71

604 m, 6.X.2009, fl.,A. Randrianasolo 1182 (G, K, MO, NY, P, TAN); ibid. loco, 19°09’39’’S 48°36’’E, 568 m, 7.X.2005, ster., Razanatsima et al. 38 (MO, P, TAN); Farankaraina, s.d., fl.,Réserves Naturelles 65 (P); Zahamena NP, Imerimandroso, [17°30’S 48°38’E], 16.IX.1951, fl.,Réserves Naturelles 3171 (P, TAN); Marojejy PN, [14°18’S 49°33’E], 15.IX.1956, fl.,Réserves Naturelles 8265 (G, MO, P); Nosy Mang- abe, 15°30’S 49°46’E, 0-330 m, 28.IX-5.X.1989, fl.,Schatz 2757 (MO, P, TAN); Soanierana-Ivongo, [16°54’30’’S 49°35’00’’E], 27.XII.1949, y. fr., Service Forestier 2361 (P, TAN, TEF); Farankaraina station forestière, [15°25’S 49°52’E], 26.IX.1952, fl., Service Forestier 6154 (G, MO, P [3 sheets], TEF); Betampona RNI, près d’Am- bodiriana, [17°55’S 49°13’E], 6.XI.1953, fl.,Service Forestier 8600 (MO, P, TEF); massif de l’Ambohitsitondroina de Mahalevona (NW de la presqu’île Masoala), [15°25’S 49°58’E], c. 500 m, 30.XI.1953, fr., Service Forestier 8678 (G, MO, NY, P, TEF); Brickaville, Sahamamy, Anivorano, [18°33’S 48°58’E], s.d., ster., Service Forestier 10884 (P, TEF); Farankaraina station forestière, [15°25’S 49°52’E], 11.X.1955, fl. & y. fr.,Service Forestier 15361 (MO, P [3 sheets], TEF); PK 154, quelques km S de Soanierana-Ivongo, [17°00’S 49°32’E], 2.IX.1957, fl.,Service Forestier 18182 (P [2 sheets], TEF); Baie d’Antongil, NW de l’embouchure de la Rantabe, [15°40’30’’S 49°38’30’’E], 50-150 m, 12.IX.1957, fl.,Service Forestier 18262 (MO, TEF); Farankaraina, [15°25’S 49°52’E], 28.IX.1963, fl.,Service Forestier 21392 (G, MO, P [2 sheets], TEF); Tampolo, [17°17’00’’S 49°23’30’’E], 26.XI.1962, fr., Service Forestier 22121 (G, K, MO, NY, P [3 sheets], TEF, WAG); Tamatave, Tanambao-Tamboro, 17.XII.1954, ster., Service Forestier 9-R-195 (P); Lohan’i Sahat- enany, N de Beanana, 2.II.1952, ster., Service Forestier 29-R-139 (P); Maroantsetra, Farankaraina, [15°25’S 49°52’E], 13.VIII.1952, ster., Service Forestier 44-R-139 (P); ibid. loco, 27.XII.1952, ster., Service Forestier 127-R-199 (P); Tampolo, Fénérive Est, 13.I.1951, ster., Service Forestier 16-R-7 (P, TEF); ibid. loco, Service Forestier 323-R- 107 (P, TEF); Masoala Peninsula, S of Ambanizana, 15°40’24’’S 49°57’51’’E, 110- 260 m, 25.X.1994, ster., Vasey et al. 55 (MO, P, TAN). Taxonomic treatment of Abrahamia (Anacardiaceae) 65

4 cm

1 mm

1 mm B

1 cm G

1 cm

E 5 cm A 5 mm

Fig. 9. Abrahamia elongata Randrian. & Lowry. A. Flowering branch; B. Detail of male flower; C. Detail of female flower; D. Leaf; E. Fruit; F. Cross section of fruit; G. Portion of inflorescence in bud.

[A-D, G: Schatz 2770, MO; E-F: Service Forestier 27110, MO] Drawings: B. Alongi 66 Boissiera 71

A C

Fig. 10. Abrahamia elongata Randrian. & Lowry (A) and A. grandidieri (Engl.) Randrian. & Lowry (B-C). A. Inflorescences with open flowers [Schatz 2770]; B. Detail of secondary venation seen from leaf lower surface and fruits, from side [Randrianaivo 1716]; C. Detail of fruits [Randriatsivery 295].

Photos: A: G. Schatz; B: F. Ratovoson; C: M. Randriatsivery Taxonomic treatment of Abrahamia (Anacardiaceae) 67

11. Abrahamia grandidieri (Engl.) Randrian. & Lowry, comb. nova (Fig. 10B-C).

≡ Protorhus grandidieri Engl. in Bot. Jahrb. Syst. 1: 422. 1881. ≡ Rhus grandidieri (Engl.) Baill. in Grandid., Hist. Phys. Madagascar Atlas: tab. 233. 1892.

Typus: Madagascar. Prov. Toliara: Morondava, s.d., fl.,Grevé s.n. (holo-: P [P00722643]!; iso-: P [P00722644]!).

= Rhus venulosa Baker in J. Linn. Soc., Bot. 25: 309. 1890. Typus: Madagascar: N Androna, s.d., fl.,Baron 5756 (holo-: K [K000423274] image seen).

= Protorhus heckelii Dubard & Dop in Bull. Soc. Bot. France 54: 155. 1907. Typus: Madagascar. Prov. Mahajanga: bords des ruisseaux, Ambongo et Boeny, IX.1904, bud & y. fr., Perrier de la Bâthie 859 (holo-: P [P06775232]!).

Description Trees 4-20 m tall, 20-45 cm DBH, bark with milky white latex; young twigs pubescent, with lenticels. Leaves alternate, rarely subopposite; blades oblanceolate or narrowly oblong, 3-16 3 1.5-4 cm, subcoriaceous, apex emarginate, retuse or rounded, margin entire, base acute or cuneate, adaxial surface glaucous, glabrous or sometimes with sericeous indument, especially on and alongside the midvein, abaxial surface with appressed sericeous indument, more so on and alongside the midvein, venation craspedodromous, midvein prominent abaxially, secondary veins 35-40(-47) pairs, parallel, 0.5-5 mm apart, tertiary veins barely visible on both surfaces; petiole 5-15 mm long, canaliculate on adaxial surface, pubescent or glabrescent. Inflorescence axillary or terminal, a panicle, 1.5-16 cm long, pubescent or villous. Male and female flowers 5-merous; pedicel 0-0.5 mm long, pubescent; calyx lobes deltate, 1-1.2 3 c. 1 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent, persistent on fruit; corolla lobes ovate, 2-2.5 3 1-1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent, white to light yellow or yellow green; stamens 5. Male flowers with filaments 1-1.2 mm long, glabrous, white, anthers 0.5 mm long, ovoid to subglobose, yellow, glabrous; disk c. 1 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments 1-1.3 mm long, glabrous, white, anthers 0.5 mm long, subglobose or ovoid, sterile, yellow, glabrous; disk 1-1.5 mm in diam., glabrous; ovary depressed ovoid, slightly asymmetrical, c. 1 3 1-1.2 mm, glabrous, placentation subapical; style shortly trifid, inserted apically, c. 1 mm long, glabrous. Fruits 2-2.8 3 1.1-1.7 cm, ellipsoid, slightly asymmetrical, glabrous, with longitudinal striations.

Vernacular names “Ditimena’’, “Lakalaka’’, “Manavodrevo’’, “Sandramiramy’’, “Soahihy’’, “Sohihy’’, “Sohy’’. 68 Boissiera 71

Distribution, ecology Abrahamia grandidieri occurs in western and southern Madagascar (Map 4), at eleva- and phenology tions ranging from sea level to 1,500 m, and is very often found near rivers or streams. It flowers from August to November and fruits from November through January.

Conservation status With an EOO of 412,300 km2, a minimum AOO of 272 km2, and 70 known localities, including subpopulations in 6 protected areas (Analavelona, Andohahela, Andringi- tra, Bemarivo, Namoroka, and Pic d’Ivohibe), A. grandidieri is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia grandidieri most closely resembles A. sericea, with which it shares leaves with parallel and very closely spaced secondary veins, but can easily be distinguished by its narrower leaves that are glaucous (vs broader and not glaucous), its flowers with a glabrous (vs pubescent) disk and ovary, and its glabrous (vs velutinous) fruits. While the protologue of Protorhus heckelii indicates that it is based on “Perrier de la Bâthie, no. 3, in Herb. Mus”, it is clear that the authors prepared their description using the specimen of Perrier de la Bâthie 859 that bears the original label written in Perrier’s hand [P06775232], along with another label added later that indicates “No. 3 = 859 Perrier de la Bâthie”. Curiously, this specimen also bears a note added some time later by Per- rier that reads “Ces spécimens proviennent d’une autre localité que le no. 859 (les bords de l’Iubohago près de l’Ankirihitra (Boina)”. A second specimen in the Paris herbarium [P06775233], comprising a single leaf and a small part of an inflorescence in a fragment packet on which someone (clearly not Perrier) has indicated “Protorhus (au sp. nov.). Mad- agascar. M. Perrier de la Bâthie, envoyé par Mr. Heckel au Lab. Colonial, fin 1906”, which appears to have been the basis for the choice of the epithet used by Dubard and Dop. This second specimen does not have an original label, but it does bear a note in Perrier’s hand, dated October 1947, that reads “Perrier no. 859! Type (?) de Protorhus Heckelei Dub. & Dop, récolté sur les bords de la Boina Kely, petit affluent de droite de la Betsiboka, à l’est de Mevatenana (Boina)”. It is not clear whether Dubard and Dop examined this specimen, and because according to Perrier de la Bâthie it is not part of the same gathering as the holotype, we have refrained from regarding it as either an isotype or a syntype.

Additional material Madagascar. Prov. Antsiranana: Antsiranana, bord de la rivière Betsieka, I.1938, ster., examined Humbert 19093ter (P); Ambanja, Beampongy, [13°40’30’’S 48°27’00’’E], 4.X.1953, Service Forestier 7715 (P, TEF); Befandriana-Nord, Marosakana, le long de la rivière Tsinjomolava, [14°56’S 47°40’E], 11.IX.1956, fl.,Service Forestier 16094 (P, TEF). Prov. Fianarantsoa: Ihosy, entre Ankaramena et Zazafotsy, [22°12’S 46°22’E], 9.X.1966, fl.,Boiteau 339 (G, MO, P); ibid. loco, Boiteau 341A-C (P); Ambalavao, entre Vohitromby et Vohitrosy, [21°50’S 46°56’E], 8.XII.1974, fr., Cremers 3547 (P [2 sheets], TAN); entre Ihosy et Fianarantsoa, 9. VIII.1938, ster., Grandidier s.n. (P); env. d’Ihosy, [22°23’00’’S 46°07’30’’E], 800-900 m, III.1934, ster., Humbert 14452ter (P); Ambohimahasoa, Ikalamavony, [21°10’00’’S 46°35’30’’E], 1955, ster., Humbert 30255bis (P); RN Andringitra, Vohitsaoka, 17.IX.1951, fl. & bud, Réserves Naturelles 3112 (P [2 sheets], TAN); ibid. loco, 8.XI.1952, bud, Réserves Taxonomic treatment of Abrahamia (Anacardiaceae) 69

Naturelles 4802 (P [2 sheets], TAN); ibid. loco, 27.IX.1957, fr., Réserves Naturelles 9300 ( P, TEF); Ivohibe, [22°31’30’’S 46°59’00’’E], 8.XI.1950, Service Forestier 1449 (P, TAN, TEF); forêt de Mangidy, [21°15’30’’S 46°47’30’’E], 15.X.1951, fl.,Service Forestier 4726 (P, TAN, TEF); Fianarantsoa, forêt Isahafa, [21°27’00’’S 47°04’30’’E], 27.IX.1951, fl.,Service Forestier 4762 (P, TEF); Maintinandrina, [21°15’30’’S 46°47’30’’E], 9.IX.1952, fl.,Service Forestier 5958 (P, TEF); Ihosy, [22°23’00’’S 46°07’30’’E], 30.X.1953, bud, Service Forestier 7615 (P, TEF). Prov. Mahajanga: bord de ruisseau, env. de Bekodoka, [16°58’E 45°06’E], 10.IX.1930, fl., Decary 8303 (G, P, US); Bekodoka, [16°58’S 45°06’E], 19.IX.1930, fl., Decary 8096 (P); Ambanja, Forêt d’Andranomatavy, forêt dense humide, bord de rivière, 13°40’29’’S 47°59’12’’E, 123 m, 28.IX.2013, fl.,Tahinarivony 746 (G, MO, P, TAN); Befandriana Nord, 14.IX.1942, fl.,Herb. Jard. Bot. Tananarive 5253 (P); Ambongo et Boeny, IX.1904, Perrier de la Bâthie 859 (P); Antajiandava, Soalala, [16°33’S 45°49’E], 2.VIII.1951, bud, Service Forestier 3990 (P, TEF); Soalala, bassin moyen du Bemarivo, [17°34’S 45°43’E], IX.1907, fl.,Perrier de la Bâthie 4519 (P [2 sheets], TAN); Bassin du Bemarivo, [16°56’S 44°24’E], c. 80 m, IX.1907, fl.,Perrier de la Bâthie 5863 (P [3 sheets]); ibid. loco, Perrier de la Bâthie 5865 (P); plateau d’An- kara, [17°13’S 46°16’E], VIII.1922, bud, Perrier de la Bâthie 14825 (P [2 sheets]); env. de Mandritsara, [15°50’00’’S 48°49’30’’E], 600 m, IX.1922, fr., Perrier de la Bâthie 15047 (P); Tsaratanana, forêt de Koakala, 16°48’09’’S 47°28’58’’E, 385 m, 7.XII.2003, fr., A. Randriana- ivo et al. 994 (CNARP, MO, P, TAN); Namoroka RN, Andranomavo, [16°36’S 45°34’E], 10. IV.1952, ster., Réserves Naturelles 3888 (P [2 sheets]); ibid. loco, 8.IX.1956, fl.,Réserves Naturelles 8139 (= Service Forestier 17172) (P, TEF); ibid. loco, 17.XI.1960, fr., Réserves Naturelles 11437 (P); Maevatanana, Ankaboka, Ambalajanakomby, [17°30’50’’S 46°04’30’’E], 8.I.1955, fr., Service Forestier 13034 (P, TEF); Morafenobe, Beravina, [18°10’30’’S 45°13’30’’E], 19.VIII.1956, fl.,Service Forestier 16351 (P, TEF); Befandriana Nord, Canton Ambararata, [16°36’S 46°25’E], 16.I.1959, fr., Service Forestier 19403 (P, TEF). Prov. Toliara: PN Andoha- hela, 24°56’23’’S 46°38’40’’E, 120 m, 9-12.V.1997, fr., Birkinshaw et al. 421 (MO, P, TAN); Rivière de l’Onilahy, E de Maromala, s.d., y. fr., Geay 6053 (P); Betroka, Ampandrandava, 8. IV.1942, y. fr., Herb. Jard. Bot. Tananarive 4971 (P); Bekily, Ampandrandava, IX-X.1943, bud & fl.,Herb. Jard. Bot. Tananarive 5793 (P); Andohahela PN, Talakimangery, 24°51’47’’S 46°42’34’’E, 250-500 m, 10.X.2000, fl.,Hoffmann et al. 172 (G, K, P); Betioky, vallée de l’Onilahy, env. de Tongobory, [23°32’S 44°20’E], 100-200 m, 1-8.X.1924, Humbert 2699 (G, P); Amboasary, vallée de la Menarahaka, [22°33’S 46°30’E], 700-800 m, 31.X.1924, fl. & fr., Humbert 3046 (P); près de Bevilany, [25°00’30’’S 46°36’E], 200-300 m, 14.IX.1928, fl.,Hum - bert & Swingle 5712 (P [2 sheets], TAN); haute vallée du Mandrare, [24°39’S 46°26’E], 8-15. XI.1928, ster., Humbert 6487bis (P); Tsivory à Anadabolava, [24°12’40’’S 46°19’00’’E], 300- 400 m, XII.1933, fr., Humbert 12287 (P [2 sheets]); ibid. loco, 200 m, Humbert 12367 (P [2 sheets]); vallée de la Manambolo, env. d’Esomony, [24°32’S 46°41’E], 400-900 m, XII.1933, fr., Humbert 12882 (G, P); vallée de l’Onilahy, en aval de Tongobory, [23°31’30’’S 44°19’30’’E], 0-300 m, 8.XI.1960, ster., Leandri 3759 (P); Beroroha, Makay massif, 21°13’02’’S 45°19’08’’E, 480 m, 22.XI.2010, fr., Phillipson et al. 6188 (G, K, MO, NY, P, TAN); PN Andohahela, 22. IX.1994, fl.,Rakotomalaza 273 (G, K, MO, NY, P, TAN); PN Andohahela, 25°01’S 46°38’E, 15.V.2001, fr., Randriamampionona 1013 (G, K, MO, NY, P, TAN, WAG); ibid. loco, 24°50’20’’S 46°32’17’’E, 6.IV.1996, ster., Randriamampionona 1241 (MO, TAN); PN Andohahela, Eso- 70 Boissiera 71

mony, coté W de la réserve, 24°31’S 46°37E, 20.IX.1991, fl.,Randriamanantena & Durbin 58 (MO, P, TAN); Ambatoabo, forêt d’Ankoba, 24°47’28’’S 46°42’01’’E, 205 m, 24. XI.2008, fr., Randrianaivo 1716 (G, K, MO, P, TAN); Sakaraha, Mahaboboka, Marotsiraka Betsileo, 22°44’20’’S 44°13’24’’E, 594 m, 21.III.2011, fr., Randrianarivony et al. 294 (G, MO, P, TAN); Ranobe, Behalomboro, 24°18’28’’S 46°11’18’’E, 210 m, 1.XI.2007, fr., Ran- driatsivery 295 (MO, P, TAN, TEF); Ampasinampoana, Enato, 24°53’05’’S 46°58’52’’E, 425 m, 16.XII.2009, fr., Razakamalala et al. 5102 (G, MO, P, TAN); Betorabato, forêt d’Androtsy, 21°05’17’’S 45°05’39’’E, 235 m, 11.I.2011, fr., Razakamalala 5920 (MO, P, TAN); PN Andohahela, Behara, [24°48’S 46°32’E], 4.X.1951, fl.,Réserves Naturelles 3407 (MO, P [2 sheets], TAN); Andohahela PN, Canton Enanihila, [24°31’S 46°37’E], 2. VIII.1952, y. fr., Réserves Naturelles 3958 (P [2 sheets], TAN); PN Andohahela, Behara, [24°53’S 46°35’E], 25.XI.1953, fr., Réserves Naturelles 5956 (G, P [2 sheets], TEF); Imanombo, Behara, [24°26’00’’S 45°49’30’’E], 16.IX.1950, fl. & y. fr.,Service Forestier 1518 (P, TAN, TEF); Anapaby, [23°00’S 44°36’E], 1.X.1951, fl.,Service Forestier 3862 (P, TEF); Morondava, Dabara, [20°24’S 44°47’E], 12.VIII.1951, bud, Service Forestier 3915 (P); Ranopiso, [25°03’00’’S 46°41’30’’E], 10.IX.1951, y. fr., Service Forestier 3967 (P, TEF); Betroka, Merarano, [20°30’30’’S 45°00’30’’E], 16.IX.1952, bud, Service Forestier 6337 (P [2 sheets], TEF); Belo sur Tsiribihina, Tsaramandroso, [19°54’30’’S 45°08’00’’E], 1. VII.1953, fr., Service Forestier 7797 (MO, P, TEF); Bekily, [24°13’20’’S 45°19’E], 14.I.1954, fr., Service Forestier 8392 (P, TEF); vallée de l’Onilahy, en aval de Tongobory, [23°31’30’’S 44°19’30’’E], 17.IX.1953, bud & fl., Service Forestier 8462 (P, TEF); Betroka, Benonoka-I, [23°33’30’’S 46°16’30’’E], 10.IV.1954, y. fr., Service Forestier 10199 (P, TEF); Sakaraha, Sohara, [24°08’S 45°28’E], 26.V.1954, y. fr., Service Forestier 10293 (MO, P, TEF); forêt de Befitroa, [20°00’30’’S 45°32’E], 20.IX.1954, fl.,Service Forestier 10876 (P [2 sheets], TEF); km 455, route Ft. Dauphin, [25°02’00’’S 46°39’30’’E], 9.IX.1954, y. fr., Service Forestier 10907 (MO, P, TEF); Sakaraha, Bekily, [24°08’S 45°28’E], 25.XI.1954, fr., Service Forestier 11158 (P, TEF); Miandrivazo, Ankilimadinika, [19°31’S 45°28’E], 23.X.1954, ster., Service Forestier 11185 (P, TEF); Sakatà, Betroka, [22°53’30’’S 45°57’30’’E], 17. IX.1954, fl.,Service Forestier 12124 (P, TEF); Manambolo, [24°34’S 46°36’E], 5.XI.1954, y. fr., Service Forestier 12199 (P, TEF); Sakamarakely, [22°11’S 45°10’E], 14.XI.1954, fr., Service Forestier 12363 (P); ibid. loco, Service Forestier 12364 (P, TEF); Vineta, [23°03’30’’S 44°15’30’’E], 20.XII.1955, fr., Service Forestier 15594 (P, TEF); Morondava, Canton Befasy, Tananadava, [20°28’S 44°08’E], 4.X.1956, fl.,Service Forestier 16556 (P, TEF); Behara, Amboasary, [24°53’S 46°35’E], 3.X.1956, bud, Service Forestier 17038 [= Réserves Naturelles 7920] (MO, P, TEF); Mahabo, Canton Ankilizato, Migodo, [20°24’S 45°03’E], 4.XII.1957, fr., Service Forestier 17869 (P, TEF); Bekily, Ankaranabo-Sud, [24°13’S 45°19’E], 9.IX.1963, bud, Service Forestier 21370 (P, TEF); Bords des ruisseaux, versant W du massif de l’Analavelona, W de Mitia, [22°37’S 44°07’E], 15.XII.1962, fr., Service Forestier 22203 (G, MO, P [2 sheets], TEF); Tsihombe, Manindro, bord rivière Manam- bovo, [25°19’S 45°29’E], 27.IV.1967, y. fr., Service Forestier 26502 (MO, P, TEF); Ampanihy, Bekitro, Tranoroa, [24°42’S 45°04’E], 13.IX.1968, bud & fl.,Service Forestier 26745 (P, TEF); Tanambao Befasy, [20°40’31’’S 44°25’E], 1.IX.1954, ster., Service Forestier 64-R-19 (P, TEF); Ampandrandava, [24°05’S 45°42’E], 700-800 m, IX.1942, fl.,Seyrig 1 (P [2 sheets]). Sine loco: s.d., Baron 9796 (P). Taxonomic treatment of Abrahamia (Anacardiaceae) 71

12. Abrahamia humbertii (H. Perrier) Randrian. & Lowry, comb. nova (Fig. 11A-B).

≡ Protorhus humbertii H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 259. 1944.

Lectotypus (designated here): Madagascar. Prov. Antsiranana: Rég. Diana, collines et plateaux calcaires de l’Ankarana, forêt tropophile, [12°54’S 49°08’E], XII.1937-I.1938, fl., Humbert 18793 (P [P00722645]!; isolecto-: P [P00722646]!). Syntypi: Madagas- car. Prov. Antsiranana: Ankarana, [12°54’S 49°08’E], 300-350 m, XII.1937-I.1938, fr., Humbert 18963 (P [P00722648]!); ibid. loco, Humbert 18967 (P [P00722647]!).

Description Trees 3-20 m tall, 5-13 cm DBH, bark with translucid latex; young twigs glabrous to pubescent towards the tips, with lenticels, waxy. Leaves opposite, deciduous; blades widely elliptic to elliptic or circular to obovate, 3-9 3 1.7-6 cm, subcoriaceous, apex emarginate, retuse, margin entire, base obtuse and oblique, adaxial surface glabrous or waxy, abaxial surface glabrous or sometimes with very scarce short indument, venation craspedodromous, midvein prominent abaxially, secondary veins 10-18 pairs, 2-6 mm apart, prominent abaxially, tertiary veins visible on both surfaces; petiole 7-15 mm long, canaliculate on adaxial surface, pubescent and waxy. Inflorescence axillary and terminal, a thyrse, 1.5-3 cm long, branches flattened, pubescent.Male and female flowers 5-merous; pedicel c. 0.2- 0.3 mm long, glabrous; calyx lobes widely depressed ovate, 0.5-0.7 3 1 mm, imbricate, glabrous on both surfaces, persistent on fruit; corolla lobes ovate, 2.5 3 1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent; stamens 5 or 6. Male flowers with filaments 1-1.5 mm long, glabrous, anthers 0.5-1 mm long, obloid to slightly ovoid, yellow, glabrous; disk cupuliform and crenulate, c. 1 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments 1.5 mm long, glabrous, anthers 0.7-1 mm long, ovoid, sterile, yellow, glabrous; disk c. 1-1.5 mm in diam., glabrous; ovary widely to very widely ovoid, 1.5-2.5 3 1-2 mm, glabrous, placentation parietal or subapical; style trifid, apically inserted, c. 0.5 mm long, glabrous, stigmas capitate. Fruits 2.5-3 3 1.5-2 cm, ellipsoid, symmetrical or very slightly asymmetrical, glabrous, with longitudinal striations.

Vernacular names “Taimbarika’’, “Mampisaraka’’.

Distribution, ecology Abrahamia humbertii occurs in deciduous and dry forests in northwestern and western and phenology Madagascar (Map 2), particularly in several areas with karstic limestone (locally known as ‘tsingy’), viz. Ankarana, Bemaraha, and Kasinga (Tamponketsa). Flowers have been collected in November and December, and fruits are known from November through March.

Conservation status With an EOO of 22,183 km2, a minimum AOO of 32 km2, and 7 known locations, only 1 of which occurs within a protected area (Ankarana) whereas the others are very likely subjected to ongoing pressure from fire and forest degradation and clearing that is pro- 72 Boissiera 71

jected to lead to a reduction in quality of habitat, A. humbertii is assigned a preliminary conservation status of “Vulnerable” [VU B2(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes The most distinctive character of A. humbertii that sets it apart from the other members of the genus is its widely elliptic to circular leaf shape. Among the syntypes of Protorhus humbertii, we have selected Humbert 18793 as the lectotype because it, along with the isolectotype, comprise more complete specimens (albeit in flower) than the other collections cited in the protologue(Humbert 18963 and 18967), both of which are in fruit.

Additional material Madagascar. Prov. Antsiranana: Ankarana, env. du Campement des Anglais, examined 12°55’57’’S 49°04’20’’E, 180-225 m, 19.XI.1996, fr., Andrianantoanina et al. 1018 (G, MO, P, TAN); Ankarana, zone du lac Vert, 2.XI.1990, fl.,Bardot-Vaucoulon 197 (P); Ankarana, camp des Anglais, 10.XI.1990, fl.,Bardot-Vaucoulon 270 (P); ibid. loco, 25.XI.1990, fr., Bardot-Vaucoulon 338 (P); ibid. loco, 12°54’42’’S 49°6’42’’E, 180 m, 15.X.1997, fr., Bardot-Vaucoulon 762 (P); ibid. loco, Bardot-Vaucoulon 763 (MO, P, TAN); Ankarana, XII.1991, fl.,Foury s.n. (P); Sakaramy, Montagne des Français, [12°26’S 49°16’E], 8.XI.1944, y. fr., Homolle 307 (P); Ankarana, near campement des Anglais, 12°54’S 49°08’E, 150 m, 29.I.1994, fr., Leeuwenberg 14345 (MO, P, TAN, WAG); Andranovondronina, Anjiabe, forêt de Belamoty, 12°06’09’’S 49°19’34’’E, 85 m, fr., Ratovoson 1154 (CNARP, MO, P, TAN); Ankara (= Ankarana), Jardin Botanique n° 8, [12°49’S 49°01’E], 28.XI.1952, fr., Service Forestier 6185 (P [2 sheets], TEF); S de Marotaolana (Anivorano-Nord), [12°49’30’’S 49°15’30’’E], 8.III.1964, fr., Service Forestier 23366 (P). Prov. Mahajanga: Antsalova, forêt de Tsiampihy et forêts cotières près de Besaraha, de Bemiha et de Soahanina, 0-20 m, 23.XII.1952, fl.,Leandri 2240 (MO, P); Antsalova, env. de Besara, [18°40’S 44°15’E], 23.XII.1952, fl.,Service Forestier 6863 (G, MO, P). Taxonomic treatment of Abrahamia (Anacardiaceae) 73

A B

Fig. 11. Abrahamia humbertii (H. Perrier) Randrian. & Lowry (A-B) and A. ibityensis (H. Perrier) Randrian. & Lowry (C). A. Detail of leaf base lower surface venation and fruits [Ratovoson 1154]; B. Fruits, from side [Ratovoson 1154]; C. Fruits, from side [Andriambololonera 314].

Photos: A-B: F. Ratovoson; C: F. Rakotoarivony 74 Boissiera 71

13. Abrahamia ibityensis (H. Perrier) Randrian. & Lowry, comb. nova (Fig. 11C).

≡ Protorhus ibityensis H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 263. 1944.

Lectotypus (designated here): Madagascar. Prov. Antananarivo: Mont Ibity, quartzites, [20°07’S 47°01’E], c. 2000 m, IX.1921, fl.,Perrier de la Bâthie 13976 (P [P00722649]!; isolecto-: P [P00722650, P00722651]!). Syntypi: Madagascar. Prov. Antananarivo: Ibity, [20°07’S 47°01’E], 1900-2000 m, s.d., ster., Perrier de la Bâthie 5840 (P [P06169079]!); ibid. loco, c. 2000 m, III.1921, fr., Perrier de la Bâthie 13571 (P [P00722653, P00722654]!); ibid. loco, Perrier de la Bâthie 13571bis (P [P00722652]!). Sine loco: s.d., Decary 13085 “A” (P [P06169081]!).

Description Shrubs 0.25-2 m tall, bark with white or translucid latex; young twigs glabrescent, with lenticels, waxy. Leaves alternate or subopposite; blades oblong to elliptic, 3-7 3 1.2-3 cm, subcoriaceous, apex retuse or emarginate, margin entire and slightly revolute, base cuneate to obtuse, adaxial surface glabrous, sometimes glossy, abaxial surface with scattered, appressed indument, more so along the midvein, venation craspedodromous, midvein prominent abaxially, secondary veins 15-30 pairs, visible, parallel, 0.5-2 mm apart, very slightly raised on both surface, tertiary veins impressed and very rarely visible on both surfaces; petiole 5-7 mm long, shallowly canaliculate on adaxial surface, pubescent when young, glabrescent. Inflorescence axillary or terminal, a thyrse, 1.5-3 cm long, pubescent. Male and female flowers 5-merous; pedicel c. 0.3-0.5 mm long, pubescent; calyx lobes ovate to deltate, 1.5-1.8 3 1.5-2 mm, imbricate, glabrous on both surfaces, margin sparesly ciliate; corolla lobes ovate, 2.5-3 3 1.5-2 mm, imbricate, glabrous, white; stamens 5. Male flowers with filaments 1-1.3 mm long, glabrous on both surfaces, white, anthers c. 1 mm long, ovoid or obloid, sterile, yellow, glabrous; disk dish-shaped, c. 2 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments 1-1.5 mm long, glabrous, white, anthers 0.5 mm long, ovoid, yellow, glabrous; disk c. 1.5-2 mm in diam., glabrous; ovary very widely ovoid, 1 3 1 mm, glabrous, placentation parietal; style apical, shortly trifid, 0.5 mm long, glabrous, stigmas capitate.Fruits 1.5-1.8 3 1.5 cm, widely depressed-ovate, glabrous, with longitudinal striations, pyrenes with a few peripheral resiniferous canals, cotyledons separable.

Distribution, ecology Abrahamia ibityensis is only known from Mt. Ibity and its surroundings, to the SW of and phenology the town of Antsirabe (Map 2), at elevations between 1500 and 2000 m, where it grows mostly on rocky soil in areas that are frequently subjected to fire. Flowers have been recorded in September and October, and fruiting material has been collected in February and March. Taxonomic treatment of Abrahamia (Anacardiaceae) 75

Conservation status With an EOO of 34 km2, a minimum AOO of 28 km2, and 9 known localities representing 7 locations, most of which are situated within a newly established protected area (Ibity) but are still subjected to fire and possible degradation leading to a projected decline in all quality of habitat, A. ibityensis is assigned a provisional conservation status of “Vulnerable’’ [VU B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia ibityensis can be distinguished from all of its congeners by its oblong to elliptic leaves with inconspicuous tertiary venation and the presence of a few peripheral resiniferous canals in the seeds, which results in the two cotyledons being distinct and easily separable. Among the syntypes of Protorhus ibityensis, we have selected Perrier de la Bâthie 13976 as the lectotype because it, along with the two isolectotypes, comprise more complete specimens than the other collections cited in the protologue (Decary 13085 “A”, Perrier de la Bâthie 5840, 13571, and 13571bis [incorrectly cited in the protologue as 1357bis]). We have added the letter “A” to the Decary specimen because this same number has been applied to a specimen of Beilschmiedia microphylla (Kosterm.) Kosterm. (Lauraceae) that bears an original label, whereas the label accompanying the syntype of Protorhus ibityensis was clearly prepared by someone else.

Additional material Madagascar. Prov. Antananarivo: Antsirabe, Ibity (S), W of Ambohimanjaka, [20°15’S examined 47°03’E], 2000 m, 22.XII.1985, y. fr., Dorr & Rakotozafy 4528 (MO, NY, P, TAN); Ibity, 13.X.1970, fl.,Keraudren-Aymonin & Aymonin 24565 (P); Ibity, Ambero, S end of Ambo- ropotsy, 20°10’17’’S 47°02’07’’E, 1700-1800 m, 23.II.2003, fr. & fl.,Koopman 170 (MO, P, TAN); Ibity, 27 km S of Antsirabe, 20°05’S 47°10’E, 1500-2240 m, 13.X.1993, fl., Lewis et al. 741 (G, K, MO, NY, P, TAN); Ibity, 23 km E de la cimenterie, 20°04’40’’S 47°00’10’’E, 1830 m, 3.X.1996, fl.,Randrianaivo et al. 6 (MO, P, TAN); ibid. loco, Randri- anaivo et al. 9 (MO, P, TAN); Ibity, E of Alatsinainy Ibity village and the cement factory, 20°06’S 46°59’E, 1600 m, 2.X.1996, fl.,A. Randrianasolo 485 (G, K, MO, P, TAN); ibid. loco, A. Randrianasolo 486 (MO, P, TAN); Ibity, Falirano, 20°04’24’’S 47°00’10’’E, 1470 m, 20.I.2003, fr., Rogers et al. 82 (G, MO, P, TAN); abords du sommet de l’Analabe, dans la chaine d’Ampandrianombilapa, S de l’Ibity, [20°05’43’’S 47°00’14’’E], 1900 m, 19.XI.1967, fr., Service Forestier 27394 (P, TEF). Prov. Fianarantsoa: Ambatofinanadra- hana, Itremo, E. du Mt. Ambatoandrano, 20°05’25’’S 47°00’07’’E, 1693 m, 1.XI.2001, fl.,Andriamihajarivo 49 (MO, P, TAN). 76 Boissiera 71

14. Abrahamia itremoensis Randrian. & Lowry, spec. nova (Fig. 12).

Typus: Madagascar. Prov. Fianarantsoa: Itremo, W of Itremo village, slope NE of the waterfall, 20°35’S 46°36’E, 1450 m, 6.X.1996, fl.,A. Randrianasolo 491 (holo-: MO-4895266!; iso-: P [P06169070]!, TAN [TAN001990]!).

Diagnosis Abrahamia itremoensis Randrian. & Lowry differs from the species it most closely resembles, A. buxifolia, by its leaves that are pubescent (vs glabrous) and the presence of indument on the disk in male flowers (vs glabrous).

Description Shrubs or small trees 0.3- 4 m tall. Leaves opposite or subopposite; blades oblanceolate, 2-6 3 0.6-2 cm, coriaceous, apex rounded, sometimes retuse or slightly emarginate, margin entire, base cuneate to slightly obtuse, adaxial surface puberulous, abaxial surface pubescent, venation craspedodromous, midvein prominent abaxially, secondary veins 18-26 pairs, parallel, 0.5-1 mm apart, prominent on both surfaces, tertiary veins weakly visible on upper surface, completely invisible on lower surface, petiole short, 1-3 mm long, canaliculate on adaxial surface, pubescent. Inflorescence axillary or terminal, a panicle, short, c. 1-2.5 cm, pubescent; bracts triangular to deltate, c. 1-2 3 0.5-0.8 mm. Male flowers small, 5-merous, 2-4 mm long; pedicel very short, c. 0.1-0.2 mm, pubescent; calyx lobes triangular to deltate, c. 1-1.5 3 1 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent; corolla lobes ovate to oblong, c. 2-4 3 1-2 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent; stamens 5, filaments c. 1 mm long, pubescent, anthers c. 1 3 0.3-0.4 mm, oblong to slightly ovoid, glabrous; disk c. 1 mm in diam., cupuliform, pubescent; ovary rudimentary. Female flowersunknown. Fruits 1.5-2.3 3 1.2-1.5 cm, obovoid to ellipsoid, velutinous.

Distribution, ecology Abrahamia itremoensis occurs mainly at Itremo and in the surrounding area (Map 5), at and phenology elevations between 1000 and 1800 m. It grows on rocky soil in areas that are regularly subjected to fire. Flowering material has been collected in October and specimens with fruit have been gathered between January and April.

Conservation status With an EOO of 154 km2, a minimum AOO of 32 km2, and 6 known locations, most of which are situated within a newly established protected area (Itremo) but are still subjected to fire and possible degradation leading to a projected decline in quality of habitat, A. itremoensis is assigned a preliminary conservation status of “Vulnerable’’ [VU B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia itremoensis can be confused with A. minutifolia due to its small, coriaceous leaves with craspedodromous venation, and it velutinous fruits, but it clearly differs by its oblanceolate (vs obovate) leaf blades, its paniculate (vs thyrsiform) inflorescence, and its larger fruits (1.5-2.3 3 1.2-1.5 vs 1-1.5 3 0.5-0.8 cm). Taxonomic treatment of Abrahamia (Anacardiaceae) 77

One of the collections assigned to this species, Decary 13041, was cited in the protologue of Protorhus buxifolia H. Perrier and therefore represents a syntype of that name.

Paratypi Madagascar. Prov. Fianarantsoa: Ambatofinandrahana, Itremo, SW d’Antsirakambiaty, 20°36’34’’S 46°33’30’’E, 1688 m, 10.XII.2003, fr., Andriamihajarivo 251 (G, MO, P, TAN); Itremo, Andaobatofotsivava, 20°37’54’’S 46°35’53’’E, 1387 m, 21.XII.2003, fr., Antilahimena et al. 2465 (MO, P, TAN); col d’Itremo, 2 km from Vodiharana, 3.VIII.1984, ster., Baum 47 (MO, TAN); Itremo, [20°32’S 46°33’E], 1700 m, IX.1956, bud, Bosser 10074 (TAN [2 sheets]); ibid. loco, X.1963, bud, Bosser 18107 (MO, P, TAN); env. d’Ambatofinandrahana, [20°33’S 46°48’E], 1600-1800 m, 17.II.1938, fr., Decary 13041 (P); W d’Itremo, [20°36’S 46°39’E], 1500-1700 m, 17-22.I. & 18-22.IV.1955, fr., Humbert 28151 (MO, P); Itremo, 20°34’40’’S 46°34’53’’E, 1550 m, 29.X.1997, fl.,Malcomber et al. 2841 (G, MO, P, TAN); forêt d’Ianasana, 7 km W d’Itremo, 20°36’S 46°34’E, 1630 m, 5.II.1999, fr., Messmer et al. 794 (G, MO, P, TAN); Itremo, [20°32’S 46°33’E], X.1964, fl.,Morat 905 (P, TAN); Ambatofinandrah- ana, [20°33’S 46°48’E], 15.X [no year given], bud, Rauh 813 (TAN); Mt. Ambatomenaloha, W d’Itremo, [20°32’S 46°33’E], 20.I.1955, fr., Service Forestier 11576 (P, TEF); env. du col d’Itremo (W d’Ambatofinandrahana), [20°32’S 46°33’E], 1600-1700 m, 27.XI.1969, y. fr., Service Forestier 28897 (P, TEF); Ambatofinandrahana, massif d’Itremo, 20°37’S 46°35’E, 1500 m, 27.X.1994, fl.,van der Werff 13563 (G, MO, P, TAN).

Fig. 12. Abrahamia itremoensis Randrian. & Lowry. A. Flowering branch; B. Leaf (adaxial surface); C. Leaf (abaxial surface); D. Ultimate inflorescence unit with 3 flowers E. Inflorescence.

[A. Randrianasolo 491, P] Drawings: R.L. Andriamiarisoa

1 cm

D E

B 3 mm 5 mm 1 cm 1 cm 78 Boissiera 71

15. Abrahamia latifolia (Engl.) Randrian. & Lowry, comb. nova.

≡ Protorhus latifolia Engl. in Bot. Jahrb. Syst. 1: 421. 1881. ≡ Rhus latifolia (Engl.) Baill. in Grandid., Hist. Phys. Madagascar Atlas: tab. 231. 1892.

Lectotypus (designated here): Madagascar: sine loco, s.d., fl.,Thouars s.n. (P [P00722655]!; isolecto-: P [P00722656, P00722657]!).

Description Trees 6-16 m tall, 30 cm DBH, bark with translucid, white or reddish milky latex; young twigs usually with white corky exoderm, glabrous, with lenticels. Leaves alternate; blades oblanceolate, 11-37 3 3.5-9 cm, subcoriaceous, apex shortly acuminate, retuse or rounded, margin entire, base acute or cuneate, adaxial surface glabrous, abaxial surface glabrous, venation craspedodromous, midvein prominent abaxially, secondary veins 40-60 pairs, parallel, 1-4 mm apart, prominent abaxially, tertiary veins visible on abaxial surface, slightly less so on adaxial surface; petiole 10 mm long, shallowly canaliculate on adaxial surface, glabrous or glabrescent. Inflorescence axillary, a panicle, 2.5-8 cm long, branches flattened or slightly triangular, pubescent or glabrescent. Male and female flowers 5-merous; pedicel c. 0.5 mm long, pubescent; calyx lobes deltate, c. 0.8 3 1 mm, valvate, adaxial surface glabrous, abaxial surface pubescent, persistent on fruit; corolla lobes ovate, 3 3 1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent, pale yellow; stamens 5. Male flowerswith filaments 1.5 mm long, glabrous, white, anthers 0.5-0.7 mm long, subglobose, yellow, glabrous; disk, c. 1-1.5 mm in diam., cupuliform, glabrous; ovary rudimentary. Female flowers with filaments 0.7-0.8 mm long, anthers 0.5 mm long, ovoid, sterile, yellow, glabrous; disk c. 1.5 mm in diam., glabrous; ovary depressed ovoid, c. 1 3 1- 1.5 mm, glabrous, placentation subapical; style shortly trifid, apicaly inserted, c. 1 mm long, glabrous, stigmas capitate. Fruits 2.5 3 1.7 cm, ellipsoid, surface smooth, glabrous.

Vernacular names “Hazombarora’’, “Hazombarora beravina’’, “Hazombarorana’’.

Distribution, ecology Abrahamia latifolia occurs at low- to mid-altitude sites (at elevations from sea level to and phenology 780 m) on sandy or lateritic soils in eastern coastal and sublittoral humid forests (Map 5). It has been recorded in flower in October and November and in fruit in December.

Conservation status With an EOO of 15,645 km2, a minimum AOO of 44 km2, but a maximum AOO that is estimated to exceed 2,000 km2, and 11 known localities representing 9 locations, only 2 of which are in protected areas (Makira and Mananara) while the remainder are subjected to land clearing leading to a projected decline in quality of habitat, A. latifolia is assigned a preliminary conservation status of “Vulnerable’’ [VU B1ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012). Taxonomic treatment of Abrahamia (Anacardiaceae) 79

Notes Abrahamia latifolia can be distinguished from all other members of the genus by its particularly large leaves (11-37 3 3.5-9 cm), which are glabrous on both surfaces. Among the three syntypes collected by du Petit-Thouars, we have selected as the lectotype the sheet that is accompanied by a hand-written description and a line drawing of a flower.

Additional material Madagascar. Prov. Mahajanga: Mandritsara, village le plus proche Lambafotsy, [16°10’S examined 48°50’E], 26.X.1955, fl.,Service Forestier 15129 (MO, P [3 sheets], TEF). Prov. Toama- sina: Antsirabe, Anjiahely, Ankahaminivola forest, 15°23’57’’S 49°26’08’’E, 780 m, 15.XII.2002, fr., Antilahimena 1493 (G, K, MO, P, TAN); Foulpointe, [17°41’S 49°31’E], s.d., Humblot s.n. (P [4 sheets]); Saharamy, [16°24’30’’S 49°41’00’’E], 14.X.1990, fl., Raharimalala 2320 (P); Tampolo forest station, 0-3 m, 31.VII.1996, ster., A. Randriana- solo 470 (G, MO, P, TAN); Mahatsara forest station, 17°38’S 49°29’E, 0-5 m, 26.X.1996, fl., A. Randrianasolo 511 (MO, P, TAN); Sainte Marie, Lonkintsy, forêt d’Ambohidena, 16°46’23’’S 49°42’47’’E, 15.V.2003, ster., Razakamalala et al. 451B (MO, P, TAN); Maroantsetra, massif de l’Androrona, [15°51’S 49°31’E], 700 m, II.1954, fr., Service Forestier 8999 (G, MO, P [2 sheets]); Tampolo, Jardin Botanique n° 21, [17°17’S 49°23’E], 29.XI.1956, fl.,Service Forestier 16469 (MO [2 sheets], P [3 sheets], TEF); ibid. loco, [17°17’00’’S 49°23’30’’E], 21.XI.1957, bud & fl.,Service Forestier 17807 (G, MO, P [2 sheets], TEF); ibid. loco, [17°17’S 49°23’E], 16.X.1958, fl.,Service Forestier 19229 (MO, P [3 sheets], TEF); Vavatenina, forêt de Sahavolo, [17°40’50’’S 49°06’30’’E], 25.X.1964, fl.,Service Forestier 21919 (P, TEF); forêt de Mangalimaso, W de Foulpointe, [17°41’30’’S 49°29’30’’E], 23.XI.1962, fl.,Service Forestier 22103 (MO, P, TEF); Nosy Sainte Marie, forêt de Kalalao, 5.VIII.1964, ster., Service Forestier 8-R-64 (P). 80 Boissiera 71

45 E 50 E

Map 5. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. itremoensis Randrian. & Lowry A. latifolia (Engl.) Randrian. & Lowry A. lokobensis Randrian. & Lowry A. louvelii (H. Perrier) Randrian. & Lowry

15 S

20 S

25 S Taxonomic treatment of Abrahamia (Anacardiaceae) 81

45 E 50 E

Map 6. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. lenticellata Randrian. & Lowry A. nitida (Engl.) Randrian. & Lowry A. oblongifolia (Engl.) Randrian. & Lowry A. patrickii Randrian. & Lowry A. pauciflora( Engl.) Randrian. & Lowry

15 S

20 S

25 S 82 Boissiera 71

45 E 50 E

Map 7. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. littoralis Randrian. & Lowry A. longipetiolata Randrian. & Lowry A. minutifolia Randrian. & Lowry

15 S

20 S

25 S Taxonomic treatment of Abrahamia (Anacardiaceae) 83

45 E 50 E

Map 8. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. phillipsonii Randrian. & Lowry A. sericea (Engl.) Randrian. & Lowry A. suarezensis Randrian. & Lowry

15 S

20 S

25 S 84 Boissiera 71

16. Abrahamia lecomtei (H. Perrier) Randrian. & Lowry, comb. nova (Fig. 13).

≡ Protorhus lecomtei H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 262. 1944.

Lectotypus (designated here): Madagascar. Prov. Toamasina: Rég. Atsinanana, Vato- mandry, bois littoraux en arrière des lagunes, fl., IX.1921,Perrier de la Bâthie 14301 (P [P00722658]!; isolecto-: MO-6684135!, P [P00722659]!).

Description Trees 5-15 m tall, 5-20 cm DBH, bark with white latex; young twigs glabrous, waxy, with lenticels, white or pale brown, spongy, sometimes the very tip with sericeous red brown indument. Leaves alternate, closely spaced, more so toward the branch ends; blades, oblanceolate, rarely widely elliptic, 5.3-17.5 3 1.5-5 cm, subcoriaceous, apex shortly acuminate, acuminate or rarely rounded, margin entire, base acute or cuneate to obtuse, adaxial surface glabrous, abaxial surface sericeous on midvein, sometimes sparsely so, venation semicraspedodromous, midvein prominent abaxially, secondary veins 20-30 pairs, more or less parallel, 2-8 mm apart, rather prominent abaxially, tertiary veins often obscure on adaxial surface, fairly visible on abaxial surface; petiole 5-8-(10) mm long, shallowly canaliculate on adaxial surface, red-brown pubescent when young, glabrescent. Inflorescence axillary, a panicle, 1-5 cm long, axes regular, red-brown pubescent; bracts triangular, 1-2 3 0.5-1 mm, abaxial surface red-brown pubescent outside. Male and female flowers 5-(or 6)-merous; pedicel c. 0.3-0.5 mm long, red-brown pubescent; calyx lobes ovate to deltate, 1-1.2 3 0.8-1.5 mm, imbricate (valvate in female flowers), adaxial surface glabrous, abaxial surface glabrescent or with red brown indument; corolla lobes ovate or oblong, 2-3 3 1-1.5 mm, imbricate, adaxial surface glabrous, abaxial surface glabrous or with sparse reddish indument, yellow green; stamens 5. Male flowers with filaments 0.5-0.7 mm long, glabrous, anthers 0.5-0.6 mm long, obloid to slightly ovoid, white, glabrous; disk cupuliform, c. 1 mm in diam., glabrous; ovary completely absent. Female flowerswith filaments c. 0.8 mm long, glabrous, anthers c. 0.5 mm long, ovoid to obloid, sterile, yellow, glabrous and pubescent; disk c. 1-1.5 mm in diam., glabrous; ovary very widely ovoid, c. 1 3 1 mm, glabrous, placentation subapical (nearly apical); style simple, subapically to apically inserted, 1 mm long, glabrous, stigmas capitate. Fruits c. 1.7 3 1 cm, ellipsoid, symmetrical, glabrous, with weakly visible longitudinal striations.

Vernacular names “Ditimena’’, “Hazombarorana’’, “Tavolo malamatra’’, “Tsiramiramy mena’’.

Distribution, ecology Abrahamia lecomtei occurs in eastern rainforest from coastal areas to elevations as high as and phenology 950 m (Map 4). It has been recorded in flower between September and December, and fruits have been collected in December and January. Taxonomic treatment of Abrahamia (Anacardiaceae) 85

Conservation status With an EOO of 7,413 km2, a minimum AOO of 32 km2 but an estimated AOO > 500 km2, and 7 recorded localities, only some of which are in protected areas (Ambalabe, Betampona, and Zahamena), A. lecomtei is assigned a preliminary conservation status of “Vulnerable” [VU B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia lecomtei can be distinguished from all other members of the genus by the presence of spongy, pale or white bark on the twigs and small branches, its leaves with semicraspedodromous venation and a short petiole (5-8 mm long), and its male flowers with short filaments (not exceeding 0.8 mm) and a disk that lacks a rudimentary ovary. Perrier de la Bâthie 5283 (P) was cited in the protologue of Protorhus lecomtei and thus represents a syntype, but it is clearly referable to Abrahamia thouvenotii.

Additional material Madagascar. Prov. Toamasina: Zahamena PN, W d’Ampasimaventy, 17°44’42’’S examined 48°58’26’’E, 300-350 m, 9.VII.2003, fl.,Andrianjafy 396 (CNARP, MO, P); Ambaton- drazaka, Sahalampy, Onibe, [17°45’S 48°51’E], 700 m, XI.1938, bud, Cours 1171 (P [2 sheets]); RNI Betampona, 17°51’55’’S 49°12’15’’E, 275-650 m, 19-25.VIII.1990, ster., Carlson 364 (MO, P, TAN); Ambodimangavolo, 3 km W d’Ambatoharanana, 17°34’17’’S 48°53’49’’E, 950 m, 30.IV.2003, fl.,Rakotondrajaona 266 (CNARP, G, K, MO, P, TEF); Vatomandry, Ambalabe, forêt de Vohibe, 19°09’07’’S 48°34’33’’E, 704 m, 20.I.2012, fr., Rakotoarivelo 608 (G, K, MO, P, TAN); ibid. loco, 19°09’42’’S 48°34’41’’E, 667 m, 20.XI.2013, y. fr., A. Randrianasolo 1456 (G, K, MO, P, TAN); Zahamena PN, Ihofika, 17°39’13’’S 48°57’50’’E, 550 m, 22.X.2002, fl.,S. Randrianasolo 335 (CNARP, MO, P); Ambalabe, forêt de Vohibe, 19°09’32’’S 48°34’39’’E, 617 m, 16.X.2008, fl.,Razanatsima 629 (G, K, MO, P, TAN); ibid. loco, 624 m, 1.XII.2008, y. fr. & fl.,Razanatsima & Celestin 635 (MO, P, TAN); Betampona RNI, [17°55’S 49°13’E], 22.XII.1950, fl.,Réserves Naturelles 2656 (MO, P [2 sheets]); Betampona RNI, Betavoana, [17°56’S 49°17’E], 12.XI.1957, fl., Réserves Naturelles 9156 (P, TEF). Sine loco: 19.XII.1938, y. fr., Lam & Meeuse 6022 (P).

Fig. 13. Abrahamia lecomtei (H. Perrier) Randrian. & Lowry. Fruits, from side [Razanatsima & Celestin 635].

Photo: A. Razanatsima 86 Boissiera 71

17. Abrahamia lenticellata Randrian. & Lowry, spec. nova (Fig. 14).

– Protorhus ditimena var. lanceolata H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 261. 1944 [nom. inval.].

Typus: Madagascar. Prov. Antsiranana: Rég. Analanjorofo, N de Nosiarina (basse Bemarivo), [14°14’30’’S 50°03’30’’E], 50 m, 7.XII.1966, fr., Service Forestier 27167 (holo-: P [P06775145]; iso-: G [G00341713]!, MO-6684136!, P [P00580350]!, TEF [TEF000885]!).

Diagnosis Abrahamia lenticellata Randrian. & Lowry most closely resembles A. ditimena but differs by having chartaceous (vs coriaceous) leaves with an acuminate or shortly acuminate apex (vs emarginate or retuse) and semicraspedodromous (vs craspedodromous) secondary veins.

Description Trees 5-20 m tall, 10-35 cm DBH, bark with white milky latex; young twigs glabrous or with ferruginous pubescence, with lenticels, waxy. Leaves alternate or subopposite; blades obovate or oblanceolate, 2.5-8(-14.5) 3 1.2-4 cm, chartaceous, apex shortly acuminate or acuminate, rarely rounded, margin sometimes undulate, base cuneate, adaxial surface glabrous, abaxial surface with sparse short, appressed indument, sometimes glabrous, venation semicraspedodromous, midvein prominent abaxially, secondary veins 20-25 pairs, 1-5 mm apart, slightly prominent abaxially, tertiary veins raised on abaxial surface; petiole 7-13 mm long, 0.5-1mm in diam., canaliculate on adaxial surface, glabrous. Inflorescence axillary or terminal, a panicle or sometimes a thyrse, 1-4 cm long, axes terete, pubescent. Male and female flowers 4-merous (rarely 5-merous), small, c. 2 mm long; pedicel c. 0.5-0.8 mm, scabrous; calyx lobes deltate to triangular, c. 1 3 1-1.5 mm, valvate, adaxial surface glabrous, abaxial surface pubescent, persistent on fruit; corolla lobes ovate, 2-2.5 3 1-2 mm, imbricate, adaxial surface glabrous, light yellow or cream, abaxial surface pubescent; stamens 4. Male flowers with filaments 1-1.2 mm long, white, glabrous, sometimes with sparse trichomes toward the base, anthers 0.5 mm long, ovoid to subglobose, yellow, glabrous; disk 1-1.5 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments c. 0.5 mm long, pubescent, anthers 0.2-0.3 mm long, sub-spherical, sterile, brown, glabrous; disk c. 2 mm in diam., slightly cupuliform, glabrous; ovary depressed ovoid, 1 3 1.1 mm, pubescent, slightly asymmetrical, style short, thick, pubescent, c. 0.3-0.4 mm long, stigmas capitate. Fruits 2-2.6 3 1.3-1.5 cm, ellipsoid to ovoid, slightly asymmetrical, surface velutinous, with longitudinal striations.

Vernacular names “Hazombarorana’’, “Hazombarorana à petite feuille’’, “Sandramy mena’’.

Distribution, ecology Abrahamia lenticellata occurs in eastern rainforest (Map 6) at elevations from sea level to and phenology 1200 m. Material in flower has been collected between August and October, and fruits have been gathered in November and December. Taxonomic treatment of Abrahamia (Anacardiaceae) 87

Conservation status With an EOO of 69,556 km2, a minimum AOO of 104 km2, and 28 recorded localities, with subpopulations in at least 11 protected areas (Zahamena, Betampona, Ambalabe, Pic d’Ivohibe, Ranomafana, Analamazoatra, Makira, Nosy Mangabe, Masoala, Anjana- haribe-Sud, and Marojejy) that comprise about 22 locations, A. lenticellata is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Cate- gories and Criteria (IUCN, 2012).

Notes Material attributable to this species was assigned by Perrier de la Bâthie (1944, 1946) to an invalidly published variety of Protorhus ditimena (≡ Abrahamia ditimena). Despite an unquestionable similarity to that species, A. lenticellata clearly warrants recognition at the species level, differing from A. ditimena in having leaves with an apex that is almost always acuminate, sometimes shortly so (vs emarginate or retuse) and whose blades are chartaceous (vs coriaceous) in texture.

Paratypi Madagascar. Prov. Antsiranana: Sambava, Marojejy PN, N of Mandena, 14°27’S 49°47’E, 230-550 m, 21.X.1989, fl.,Miller 4327 (MO, P, TAN); Sambava, Marojejy PN, Mandena, près du campement 1, WP-2-4, 14°26’13’’S 49°46’32’’E, 460 m, 10.X.1996, fl., Rakotomalaza et al. 728 (G, K, MO, NY, P, TAN); Andapa, Anjanaharibe Sud RS, 14°43’10’’S 49°27’12’’E, 1700 m, 14.II.1995, fr., Ravelonarivo et al. 669 (MO, P, TAN); Doany, Andrinomilolo, 13 km W d’Andranopositra, 14°20’04’’S 49°17’50’’E, 856 m, 13.XI.2006, fr., Ravelonarivo et al. 2088 (MO, P, TAN); Sambava, [Marojejy PN], 24.IX.1956, fl.,Réserves Naturelles 8266 (P, TAN). Prov. Fianarantsoa: Pic d’Ivohibe RS, forêt de Marovitsika, 22°28’49’’S 46°56’49’’E, 950 m, fl., s.d.,Hoffmann et al. 241 (G, K, P); Ranomafana Park, Parcelle n° 3, Talatakely, 21°15’S 47°27’E, 800-1000 m, 3-8. IX.1993, fl.,Kotozafy et al. 230 (MO, P, TAN); Ranomafana PN, c. 15 km W de Tsarata- nana, 21°09’S 47°32’E, 750 m, 8.X.1994, ster., Kotozafy 703, 724 (MO, P, TAN); Ivohibe, 7.5 km ENE d’Ivohibe, 22°28’12’’S 46°57’36’’E, 1000 m, 11.X.1997, fl.,Rakotovao et al. 830 (G, P, MO, TAN, G); Ranomafana PN, piste T 200, 21°16’S 47°25’E, 800-900 m, 8.X.1996, ster., A. Randrianasolo 501 (G, K, MO, P, TAN); Nosy Varika, Besakoana-Am- pasinambo, 7.VIII.1952, ster., Service Forestier 68-R-116 (P). Prov. Toamasina: Maro- antsetra, Ambinanihely, 15°26’S 49°29’E, 358 m, 20.XI.2003, y. fr. & fl.,Antilahimena 2328 (G, MO, P, TAN); Vinanibe, Andogona, Makira forest, 15°27’40’’S 49°17’46’’E, 1153 m, 28.XI.2003, fr., Antilahimena 2427 (G, K, MO, NY, P, TAN); Toamasina, Betam- pona RN, 17°51’55’’S 49°12’15’’E, 275-650 m, 25.VIII.1990, ster., Carlson 383 (MO, P, TAN); Ambatondrazaka, Manambato à Amboditafonana, [17°27’S 48°46’E], 1200 m, 11.X.1945, fl.,Cours 2826 (MO, P, TAN); Ambatondrazaka, Sahalampy, Onibe, [17°45’S 48°15’E], 700 m, XI.1938, fl.,Cours 1042 (P); Moramanga, Analamazaotra, [18°56’S 48°26’E], 800-900 m, VIII.1925, bud & fl.,Louvel 48 (P); near Andasibe, forest of Mantadia, 18°55’S 48°25’E, 900 m, 5.XI.1994, fl.,McPherson et al. 16505 (G, K, MO, P, TAN); Masoala PN, Antalavia, 15°47’S 50°02’E, 22.XI.1994, ster., Rahajasoa et al. 965 (MO, P, TAN); Amparafaravola, SE d’Ambodisaina, 17°17’25’’S 48°40’30’’E, 850 m, 10.XII.2001, fr., Rakotondrafara et al. 98 (CNARP, MO, P, TAN); Vatomandry, Commune Ambalabe, Tobin’i Foara, 19°09’28’’S 48°34’30’’E, 560 m, 4.X.2005, fl., 88 Boissiera 71

5 mm

4 cm

A 4 cm

1 mm 5 mm

1 cm B C E Fig. 14. Abrahamia lenticellata Randrian. & Lowry. A. Flowering branch; B. Leaf (adaxial surface); C. Detail of male flower; D. Fruiting branch; E. Fruit; F. Inflorescence in bud.

[A-C, F: Miller 4327, MO; D-E: Schatz & Carlson 2848, MO] Drawings: B. Alongi Taxonomic treatment of Abrahamia (Anacardiaceae) 89

Ranaivojaona et al. 1197 (G, K, MO, P, TAN); Commune Ambalabe, N of Tsitondroina waterfall, 19°10’15’’S 48°’33’02’’E, 500 m, s.d., fr., A. Randrianasolo et al. 927 (G, K, MO, NY, P, TAN, WAG); Commune Ambalabe, fokontany Sahanionaka, 19°11’24’’S 48°34’30’’E, 288 m, 13.XI.2010, fr., A. Randrianasolo 1313 (G, K, MO, NY, P, TAN); Commune Ambalabe, Vohibe forest, 19°09’15’’S 48°34’44’’E, 629 m, 21.XI.2013, fr., A. Randrianasolo 1458 (G, K, MO, P, TAN); Périnet, [18°56’S 48°26’E], 800-900 m, 23.XII.1947, y. fr., Réserves Naturelles 1261 (P); Nosy Mangabe, 15°30’S 49°46’E, 0-330 m, 28.IX-5.X.1989, fl.,Schatz et al. 2777 (MO, P, TAN); ibid. loco, 17.XII.1989, fr., Schatz & Carlson 2848 (MO, NY, P, TAN); Soanieran’Ivongo, [16°54’30’’S 49°35’00’’E], 0-50 m, 27.XII.1949, fr., Service Forestier 2370 (P, TAN, TEF); Farankaraina, [15°25’S 49°52’E], 20-25 m, 9.X.1952, bud, Service Forestier 6248 (G, P); massif de l’Ambatotsitondroina de Mahalevona, NW de la presqu’île Masoala, [15°25’S 49°58’E], 600 m, 21.XII.1953, y. fr., Service Forestier 8697 (MO, P, TEF); forêt orientale, entre Ankovana et le Col d’Am- batondradama, 500 m, 13.XII.1953, fr., Service Forestier 8750 (MO, P, TEF); Farankara- ina, Andranofotsy, [15°25’S 49°52’E], 20-25 m, 18.X.1963, fl.,Service Forestier 21483 (G, MO, NY, P, TEF); Vavatenina, sommet Mahatsinjo, [17°27’S 49°00’E], 20.IX.1966, bud, Service Forestier 26139 (P); Mahatsinjo, Sahatavy, 27.IX.1966, fl.,Service Forestier 26246 (P [2 sheets]); Brickaville, Lohariandava, Ankarangina, 14.VI.1954, ster., Service Forestier 70-R-77 (P); Maroantsetra, Jardin Botanique 17, [Farankaraina], 12.XII.1952, ster., Service Forestier 99-R-199 (P). 90 Boissiera 71

18. Abrahamia littoralis Randrian. & Lowry, spec. nova (Fig. 15).

Typus: Madagascar. Prov. Toamasina: Rég. Analanjirofo, Fénérive-Est, Tampolo forestry station, along the “Grand Layon”, just 20 m before the first bridge going toward the Indian Ocean, 0-5 m, 25.X.1996, fl.,A. Randrianasolo 509 (holo-: MO-04895278!; iso-: G!, K!, P [P06775163]!, TAN [TAN001996]!, WAG!).

Diagnosis Abrahamia littoralis Randrian. & Lowry is easily distinguished from its congeners by its medium sized (4-17 3 1.3-5.2 cm), oblanceolate leaves bearing scattered, very often obscure tertiary venation, its long petiole (10-30 mm), and its axillary inflorescences.

Description Trees 4-15 m tall, 12-18 cm DBH, bark with white milky or translucid latex; young twigs pubescent or ferruginously scabrous. Leaves alternate; blades oblanceolate, 4-17 3 1.3-5.2 cm, coriaceous to subcoriaceous, apex very often shortly acuminate, rounded, retuse or emarginate, margin entire, base acute, adaxial surface sericeous when young, later glabrescent, abaxial surface sericeous when young, glabrescent, venation craspedodromous, slightly prominent adaxially, midvein prominent abaxially, secondary veins c. 25 pairs, more or less parallel, 2-10 mm apart, prominent abaxially, forming an acute angle with the midrib, tertiary veins widely spaced, often obscure on adaxial surface; petiole 10-30 mm long, canaliculate on adaxial surface, pubescent when young, later glabrescent. Inflorescence axillary, a panicle, 2-5 cm long, axes, strigose. Male and female flowers 5-merous; pedicel 0.3-0.5 mm long, with short, appressed indument; calyx lobes triangular, 1.5 3 1 mm, imbricate, adaxial surface glabrous, abaxial surface with short appressed indument, persistent on fruit; corolla lobes narrowly ovate, 2-3 3 c. 1 mm, imbricate, adaxial surface glabrous, abaxial surface strigose, pale yellow, whiter inside; stamens 5. Male flowers with filaments 1.2-1.5 mm long, glabrous, white, anthers 0.8-1 mm long, obloid to slightly ovoid, white, glabrous; disk cupuliform, c. 1.5 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments c. 1 mm long, glabrous, white, anthers c. 0.2-0.3 mm long, subglobose, sterile, white, glabrous; disk c. 1.5 mm in diam., glabrous; ovary widely ovoid to obovoid, c. 1.5 3 c. 1 mm, glabrous, placentation subapical; style very shortly trifid, apical, c. 1-1.1 mm long, glabrous, stigmas capitate. Fruits 2-2.5 3 1-1.5 cm, ellipsoid, slightly asymmetrical, glabrous, with longitudinal striations.

Vernacular names “Hazombarorana’’, “Tarantana’’.

Distribution, ecology Abrahamia littoralis is found in eastern littoral or sublittoral forests near sea level and phenology on sandy substrates (Map 7). It flowers in October and very young fruits have been collected in November. Taxonomic treatment of Abrahamia (Anacardiaceae) 91

0.5 mm

4 cm A

2.5 mm

1 mm 1 mm D

Fig. 15. Abrahamia littoralis Randrian. & Lowry. A. Flowering branch; B. Infrutescence. C. Fruit; D. Detail of female flower; E. Detail of male flower. [ A. Randrianasolo 509, MO] Drawings: B. Alongi 92 Boissiera 71

Conservation status With an EOO of 18,722 km2, a minimum AOO of 56 km2, and 10 recorded localities, with a subpopulation in a single protected area (Agnalazaha), comprising 8 locations subjected to pressure from habitat degradation due to forest exploitation and land clearing, A. littoralis is assigned a preliminary conservation assessment of “Vulnerable” [VU B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes With its oblanceolate leaf blade, craspedodromous venation, axillary inflorescence, presence of a rudimentary ovary in male flowers, and glabrous fruit,A. littoralis most closely resembles A. elongata but differs by its much smaller leaves (4-17 3 1.3-5.2 vs 8.5-26.5 3 2.5-6.5 cm), fewer secondary veins (25 vs 25-50), and obscure (vs prominent) tertiary venation on the abaxial surface.

Paratypi Madagascar. Prov. Fianarantsoa: Farafangana, Mahabo, forêt littorale d’Apoakafo, 23°11’02’’S 47°41’38’’E, 34 m, 19.IX.2003, fl.,Ludovic et al. 435 (MO, P, TAN); ibid. loco, 23°11’10’’S 47°43’02’’E, 30 m, 5.VIII.2004, fl.,Ludovic et al. 880 (G, MO, P, TAN); ibid. loco, 23°11’S 47°41’E, 21 m, 29.X.2004, fr., Ludovic et al. 928 ( MO, TAN); Mahabo, Baboaka, forêt d’Andranomainty, 23°10’S 47°44’E, 21 m, s.d., fr., Ludovic & Emilson 1006 (G, K, MO, P, TAN); near Mahabo village, 23°10’51’’S 47°42’29’’E, 30 m, 5.XI.2001, fr., McPher- son & Rabenantoandro 18338 (G, K, MO, NY, P, TEF, WAG); Fokontany Mahabo, Firaisana Mahabo-Mananivo, 23°11’01’’S 47°42’44’’E, 17 m, 6.XI.2001, fr., Rabenantoandro et al. 701 (MO, P, TAN, TEF); ibid. loco, 23°11’01’’S 47°42’44’’E, 17 m, 6.XI.2001, y. fr., Rabenan- toandro et al. 706 (MO, P, TAN,TEF); ibid. loco, 23°10’39’’S 47°43’06’’E, 10 m, 25.IX.2002, fr., Rabenantoandro et al. 1011 (G, MO, P, TEF); ibid. loco, 23°11’26’’S 47°43’17’’E, 11 m, 1.IX.2007, bud, Rakotoarivelo 198 (MO, P, TAN); ibid. loco, 23°11’13’’S 47°42’27’’E, 22 m, 26.VIII.2003, fl.,Razakamalala et al. 708 (G, K, MO, P, TEF); Farafangana, Nosiala, [23°12’00’’S 47°42’30’’E], 19.XI.1955, fr., Service Forestier 16211 (P, TEF). Prov. Toamasina: Fénérive-Est, Station forestière de Tampolo, 17°16’52’’S 49°24’44’’E, 0-15 m, 24.I.1995, ster., Raholivelo et al. 150 (MO); ibid. loco, 0-5 m, 1.VIII.1996, ster., A. Randrianasolo 467, 467a (MO, P, TAN); ibid. loco, 25.X.1996, y. fr., A. Randrianasolo 510 (MO, P, TAN); Brick- aville, Vohibola forest, 18°33’39’’S 49°14’33’’E, 27 m, 28.XI.2002, fl.,A. Randrianasolo et al. 737 (G, K, MO, P); Ambila Lemaitso, [18°49’S 49°08’E], 26.X.1950, y. fr., Service Forestier 1583 (G, MO, P, TAN, TEF); Tampolo, [17°17’00’’S 49°23’30’’E], 15.X.1954, fl., Service Forestier 11071 (MO, P [2 sheets]); ibid. loco, [17°17’00’’S 49°23’30’’E], 19.X.1955, fl.,Service Forestier 15187 (MO, P, TEF); ibid. loco, [17°17’S 49°23’E], 28.IX.1957, fl.,Ser - vice Forestier 17797 (G, MO, NY, P, TEF); forêt sublittorale de Vohibola, W de Tampina, [18°31’S 49°17’E], 12-14.VIII.1957, y. fr., Service Forestier 18086 (G, MO, P [2 sheets]); forêt sublittorale, entre Foulpointe et l’embouchure de l’Onibe, [17°40’30’’S 49°29’30’’E], 29.IX.1963, fl.,Service Forestier 22764 (P). Prov. Toliara: Tolagnaro (Fort-Dauphin), Man- dena Forest, along road SW of QIT Fer et Titane work camp, 24°57’S 47°00’E, 0-10 m, 19.IV.1989, ster., Gereau et al. 3393 (MO, P, TAN); Tolagnaro, Ambinanibe, port d’Ehoala, 25°03’S 46°56’E, 2 m, 5.III.2007, ster., Rabenantoandro et al. 1777 (MO). Sine loco: s.d., fl.,Humblot 358 (P); s.d., fl.,Louvel 147 (P). Taxonomic treatment of Abrahamia (Anacardiaceae) 93

19. Abrahamia lokobensis Randrian. & Lowry, spec. nova (Fig. 16).

Typus: Madagascar. Prov. Antsiranana: Rég. Diana, RNI Lokobe, 13°24’40’’S 48°18’50’’E, 40-80 m, 24.XI.1994, fr., Antilahimena 198 (holo-: MO-6671433!; iso-: G!, K!, NY!, P [P06775168]!, TAN [TAN001991]!).

Diagnosis Abrahamia lokobensis Randrian. & Lowry can be distinguished from its congeners by its large (14.6-23.9 3 4.8-7.2 cm), chartaceous to subcoriaceous, oblanceolate leaves and its obovoid fruit borne in a terminal infructescence.

Description Trees c. 8-25 m tall, 10-80 cm DBH, bark with white latex; young twigs lenticellate, glabrous. Leaves opposite to subopposite or alternate; blade oblanceolate, 14.6-23.9 3 4.8-7.2 cm, chartaceous to subcoriaceous, apex short acuminate, retuse or sometimes emarginate, margin entire, base cuneate, sometimes asymmetric, adaxial surface glabrous, abaxial surface nearly glabrous except sparsely scabrous indument on midvein or around it, venation craspedodromous, midvein prominent abaxially, secondary veins c. 30-45 pairs, 2-10 mm apart, forming wide angle with the midvein, raised on both surfaces, tertiary veins visible on both surfaces; petiole 13-30.5 mm long, 1.5-3 mm in diam., canaliculate on adaxial surface, very often pubescent, glabrescent or sometimes glabrous. Inflorescence terminal or pseudo- terminal, a panicle, 2.5 3 7 cm, with buds at very early stage of development, dark brown pubescent. Flowers unknown. Fruits 2-3.1 3 1-1.5 cm, obovoid, pale green, glabrous, with longitudinal striations.

Distribution, ecology Abrahamia lokobensis is only known from northwestern Madagascar; it occurs at eleva- and phenology tions between 10 and 100 m at the Lokobe reserve on the island of Nosy-be, and up to 1200-1400 m in Tsaratanana national park (Map 5). Flowers have been collected in August and fruits in November.

Conservation status Abrahamia lokobensis has an EOO of 3,124 km2 and a minimum AOO of 16 km2, with two subpopulations occurring within a protected area (Lokobe and Tsaratanana) whereas the other subpopulation was collected in the area of Maromandia where future decline is expected in terms of quality of habitat, and thus exists at three locations with respect to the threat of habitat degradation. It has therefore been assigned a preliminary status of “Endangered” [EN B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia lokobensis most closely resembles A. sambiranensis but can be distinguished by its smaller number of secondary veins (35-45 vs 50-60), its tertiary venation that is visible on both leaf surfaces (vs not visible), its shorter petiole (13-19 vs (20-)25-65 mm), and exclusively terminal (vs axillary) infructescence. 94 Boissiera 71

Paratypi Madagascar. Prov. Antsiranana: Rég. Diana, Lokobe RNI, 13°25’S 48°18’E, 10-100 m, 14.XI.1996, fr., Antilahimena et al. 315 (G, K, MO, P, TAN); ibid. loco, [13°25’S 48°19’E], 13-15.III.1964, fr., Service Forestier 23459 (MO, P); Maromandia, Ambanja, [14°12’S 48°05’E], 15.VIII.1952, ster., Service Forestier 64-R-152 (P [2 sheets]); Tsaratanana RNI, Ambinany Beangona, 14°01’S 48°47’E, 1200-1400 m, 1.VIII.2000, y. bud, Antilahimena et al. 533 (G, K, MO, NY, P, TAN).

1 cm

1 cm 1 cm Fig. 16. C Abrahamia lokobensis Randrian. & Lowry. A. Fruiting branch; B. Fruit; C. Inflorescence with buds.

[A-B: Antilahimena 198, P; C: Antilahimena et al. 533, P] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 95

20. Abrahamia longipetiolata Randrian. & Lowry, spec. nova (Fig. 17-18).

Typus: Madagascar. Prov. Antsiranana: Rég. Diana, commune Mahavonona, Montagne des Français, campement Ampitiantsambo, 12°23’13’’S 49°22’53’’E, 239 m, 11.IX.2004, bud & fl.,Randriambololomamonjy 7 (holo-: MO-6278643!; iso-: P [P04800624]!, TAN [TAN001993]!).

Diagnosis Abrahamia longipetiolata Randrian. & Lowry can be distinguished from other members of the genus by the combination of its strikingly long petiole (15-30 mm) and small leaves (6-11 3 1.5-3 cm) with impressed secondary veins on the adaxial surface.

Description Trees c. 7-25 m tall, 19-60 cm DBH, bark with white latex. Leaves alternate, sometimes sub-opposite; blade narrowly elliptic to narrowly oblong, 6-11 3 1.5-3 cm, chartaceous to subcoriaceous, discolorous (glaucous above), apex acute and retuse, margin slightly revolute, base broadly cuneate, sometimes slightly asymmetric, adaxial surface glabrous, abaxial surface with appressed sericeous indument along the midvein, especially towards the base, venation craspedodromous, midvein prominent below, secondary veins 25-35 pairs, more or less parallel, 2-4 mm apart, impressed above, barely raised below, forming a wide angle with the midvein, tertiary veins very few, barely visible; petiole 15-30 mm long, 1-2 mm in diam., canaliculate on adaxial surface, strigose. Inflorescence terminal, a panicle, branched, 9 cm long, axes pubescent; bracts triangular, 1-2.5 3 0.5-1 mm, adaxial surface glabrous, abaxial surface pubescent. Male and female flowers5-merous , small, 3-4 mm long; pedicel c. 1 mm long, pubescent; calyx lobes 1-1.1 mm, ovate, valvate, adaxial surface glabrous, abaxial surface pubescent; corolla lobes ovate, 3-4 3 1.5-2 mm, imbricate, adaxial surface glabrous, abaxial surface sparsely pubescent (much less so than calyx); stamens 5 (rarely 6). Male flowers with filaments c. 1-1.2 mm long, glabrous, anthers 1.1-1.2 3 1 mm, ovoid, yellow, glabrous; disk c. 1.5-2 mm in diam., cupuliform, glabrous; ovary rudimentary. Female flowers with 5 staminodes, sometimes 6, filaments c. 1 mm long, glabrous, anthers 1 3 0.5 mm, ovoid, sterile, yellow, glabrous; disk c. 2-2.5 mm in diam., glabrous; ovary widely ovoid, 1-1.1 3 1.1.5 mm, glabrous; style short, c. 0.5 mm, stigmas capitate. Fruits 2.4-2.6 3 1.5-1.8 cm, asymmetrical, ellipsoid, glabrous, with weak longitudinal striations.

Distribution, ecology Abrahamia longipetiolata occurs in forests in far northeastern Madagascar (Map 7). and phenology Flowering material has been collected in August and September, and fruits have been recorded from October to December.

Conservation status With an EOO of 799 km2, a minimum AOO of 20 km2, and 6 localities, with subpopulations in only 2 protected areas (Loky Manambato and Montagne des Français), representing 5 locations, 3 of which are subjected to pressure from forest exploitation and 96 Boissiera 71

Fig. 17. Abrahamia longipetiolata Randrian. & Lowry. A. Flowering branch; B. Fruiting branch; C. Fruit.

[A: Randriambololomamonjy 7, P; B-C: Randriambololomamonjy 2, P] Drawings: R.L. Andriamiarisoa

A B C

1 cm 1 cm 4 mm

land clearing, A. longipetiolata is assigned a provisional conservation status of “Endan- gered’’ [EN B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia longipetiolata resembles A. ibityensis by the impressed venation of its leaves, but differs by its larger leaves (6-11 3 1.5-3 vs 3-7 3 1.2-3 cm), longer petiole (15-30 vs 5-7 mm), terminal paniculate (vs terminal or axillary thyrsiform) inflorescences, and fruit with united (vs separate) cotyledons.

Paratypi Madagascar. Prov. Antsiranana: Vohémar, forêt d’Ambilondava, N du village de Daraina, 13°10’S 49°39’E, 400 m, 25.X.2004, fr., Callmander & Anselme Be 268 (G, MO, P, TAN); Sadjoavato, forêt de Sahafary, 12°35’17’’S 49°26’45’’E, 207 m, 15.X.2007, fl.,Rakotoar - ivelo 172 (CNARP, G, MO, P, TAN); Mahavonona, Montagne des Français, campement Ampitiantsambo, 12°23’13’’S 49°22’53’’E, 239 m, 11.IX.2004, y. fr., Randriambololoma- monjy 2 (G, MO, P [2 sheets], TAN); Andrafiabe, Sahafary, 12°35’43’’E 49°26’54’’E, 233 m, 11.IX.2004, fl. & y. fr.,Razafitsalama et al. 638 (CNARP, G, MO, P, TAN, TEF); ibid. loco, [12°34’S 49°26’E], 10.XII.1952, fr., Service Forestier 6287 (MO, P, TEF); ibid. loco, 9.IV.1953, ster., Service Forestier 216-R-6 (P); N de Vohémar, versant du Massif de l’Ankerana, [13°17’S 49°51’E], 50-450 m, 17.XII.1966, fr., Service Forestier 27359 (G, MO, P [2 sheets], TEF). Taxonomic treatment of Abrahamia (Anacardiaceae) 97

Fig. 18. Abrahamia longipetiolata Randrian. & Lowry. Inflorescence with buds and opened flowers, from side [Razafitsalama et al. 638].

Photo: L.J. Razafitsalama 98 Boissiera 71

Fig. 19. Abrahamia louvelii (H. Perrier) Randrian. & Lowry. A. Branch with fruits; B. Fruits, from side. [Rakotoarivelo et al. 643].

Photos: N. Rakotoarivelo B Taxonomic treatment of Abrahamia (Anacardiaceae) 99

21. Abrahamia louvelii (H. Perrier) Randrian. & Lowry, comb. nova (Fig. 19).

≡ Protorhus louvelii H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 263. 1944 [as louveli].

Lectotypus (designated here): Madagascar. Prov. Toamasina: Rég. Alaotra-Man- goro, Analamazaotra, c. 1000 m, fl., 1919,Thouvenot 36 (P [P00722660]!; isolecto-: P [P00722661]!, TAN [TAN000075]!, US-2494483 image seen).

Description Trees 14-15 m tall, bark with white-yellow milky latex; young twigs glabrous but with scabrous indument at the tips, with a few lenticels. Leaves subopposite to opposite; blades obovate to oblanceolate, 2-4 3 1-2 cm, coriaceous to subcoriaceous, apex emarginate, retuse or rounded, margin entire, base cuneate, adaxial surface glabrous, abaxial surface with very sparse, appressed, short indument, venation craspedodromous, midvein prominent abaxially, secondary veins 15-28 pairs, 1-2 mm apart, parallel or nearly so, prominent abaxially, tertiary veins obscure on both surfaces; petiole 3-7 mm long, canaliculate on adaxial surface, glabrous. Inflorescence axillary and terminal, a panicle, 2.5-4.5 cm long, axes flattened or somewhat triangular, pubescent to strigose. Male flowers 5-merous; pedicel 0.2-0.3 mm long, very sparsely strigose; calyx lobes deltate, 1 3 1 mm, valvate, glabrous; corolla lobes ovate, 2.5 3 1.5 mm, imbricate, glabrous; stamens 5, filaments 0.5-0.7 mm long, glabrous, anthers 0.5-0.6 mm long, obloid to slightly ovoid, glabrous; disk dish-shaped or slightly cupiliform, 1.5 mm in diam., glabrous; ovary rudimentary. Female flowers unknown. Fruits 1.5- 1.7 3 1-1.2 cm, green when fresh, black when dry, ellipsoid to slightly obovoid, glabrous, with longitudinal striations.

Vernacular names “Ditimena’’, “Soretry’’.

Distribution, ecology Abrahamia louvelii is known only from the Analamazaotra and Vohibe (Ambalabe) for- and phenology ests (Map 5) at elevations between 600 and 1000 m, in areas with eastern humid forest. Material has been collected in flower in November and in fruit in January.

Conservation status With a minimum AOO of 8 km2 and 3 localities, all of which are situated within protected areas (Ambalabe and Analamazaotra), A. louvelii is assigned a preliminary conservation status of “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia louvelii most closely resembles A. minutifolia but differs by having flowers borne in panicles (vs thyrses), a glabrous (vs abaxially pubescent) corolla, and glabrous (vs velutinous) fruits. 100 Boissiera 71

We have selected one of the two sheets of Thouvenot 36 in the Paris herbarium as the lectotype of Protorhus louvelii because it comprises more complete material.

Additional material Madagascar. Prov. Toamasina: Réserve spéciale au S d’Andasibe Mahaveriky, [18°56’S examined 48°26’E], 4.XI.1967, ster., Jacquemin 687 (P); Vatomandry, Ambalabe, forêt de Vohibe, 19°09’39’’S 48°34’43’’E, 657 m, 22.I.2012, fr., Rakotoarivelo et al. 643 (G, K, MO, P, TAN); ibid. loco, 19°09’31’’S 48°34’40’’E, 536 m, 19.II.2009, ster., Razanatsima 667 (MO, P, TAN); Moramanga, Analamazaotra, Périnet, [18°56’S 48°26’E], II.1966, ster., Service Forestier 25719 (P). Taxonomic treatment of Abrahamia (Anacardiaceae) 101

22. Abrahamia minutifolia Randrian. & Lowry, spec. nova (Fig. 20-21).

– Protorhus ditimena var. minutifolia H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 261. 1944 [nom. inval.].

Typus: Madagascar. Prov. Toamasina: Rég. Alaotra-Mangoro, Moramanga, Analamazaotra RS, “carrière de sable”, [18°56’S 48°26’E], 900-950 m, 29.X.1996, fl.,A. Randrianasolo 516 (holo-: MO-04895273!; iso-: G!, K!, P [P06169142]!, TAN [TAN002000]!).

Diagnosis Abrahamia minutifolia Randrian. & Lowry can be distinguished by having leaves that are smaller (1-3 3 0.9-1.8 cm) than any other member of the genus and fruits that are among the smallest (1-1.5 3 0.5-0.8 cm).

Description Trees 3.5-12 m tall, 11.9-40 cm DBH, bark with translucid latex; young twigs waxy, lenticellate, with short, red, appressed indument at the tip. Leaves subopposite; blades, obovate, 1.8-4 3 0.9-1.8 cm, subcoriaceous to coriaceous, apex emarginate, retuse, or rounded, margin entire, base acute, adaxial surface glabrous, abaxial surface with very sparse appressed reddish indument when young, glabrescent, venation craspedodromous, midvein prominent below, secondary veins 15-20 pairs, 0.5-2 mm apart, prominent on both surfaces, tertiary veins raised on both surfaces; petiole 3-8 mm long, canaliculate on adaxial surface, reddish pubescent when young, later glabrescent. Inflorescence axillary or terminal, a thyrse, 1-3 cm long, axes somehow flattened, with ferruginous indument. Male and female flowers 4- or 5-merous; pedicel 0.2-0.3 mm long, pubescent; calyx lobes deltate, 1 3 1 mm, imbricate, glabrous, persistent on fruit; corolla lobes ovate, 2-2.2 3 1-1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent; stamens 4 or 5. Male flowers with filaments 1-1.5 mm long, glabrous, white, anthers c. 0.8 mm long, obloid to slightly ovoid, yellow, glabrous; disk cupuliform, c. 1 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments 0.3-0.5 mm long, glabrous to slightly pubescent, white, anthers c. 0.5 mm long, subglobose to ovoid, sterile, yellow, glabrous; disk c. 1.5 mm in diam., glabrous; ovary widely to very widely ovoid, c. 1.5 3 1 mm, pubescent, placentation subapical; style shortly trifid, subapically to apically inserted, 0.5 mm long, partly pubescent, stigmas capitate. Fruits 1-1.5 3 0.5-0.8 cm, ellipsoid, slightly asymmetrical, surface velutinous, with longitudinal striations.

Distribution, ecology Abrahamia minutifolia has been collected primarily in premontane forest along Madagas- and phenology car’s eastern escarpment at an elevation of c. 900 m and in central high elevation forest at c. 1500 m, but it also occurs in humid forests in both the eastern and northwestern parts of the island (Map 7). It flowers between September and November, and fruits have been collected from November to April. 102 Boissiera 71

5 mm

5 mm B C

5 mm

D E 4 cm 2 mm

5 mm 5 mm

F G Fig. 20. Abrahamia minutifolia Randrian. & Lowry. A. Flowering branch; B. Detail of male flower; C. Detail of female flower; D. Fruit showing indument; E. Cross section of fruit; F. Leaf (abaxial surface); G. Young infructescence.

[A, C, F: A. Randrianasolo 516, MO; B: Service Forestier 18400, MO; D-E, G: Service Forestier 7361 and 8417, P] Drawings: B. Alongi Taxonomic treatment of Abrahamia (Anacardiaceae) 103

Conservation status With an EOO of 117,262 km2, a minimum AOO of 84 km2, and 21 known localities, with subpopulations in at least 5 protected areas (Ambohitantely, Kalabenono, Maso- ala, Manongarivo, and Analamazaotra-Perinet), A. minutifolia is assigned a preliminary conservation status of “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia minutifolia is part of a group of species, along with A. delphinensis, A. ditimena, A. itremoensis, A. lenticellata and A. sericea, that has velutinous fruits, but is distinguished by its thyrsiform (vs paniculate) inflorescence and smaller fruits (1-1.5 3 0.5-0.8 vs 2-3-2.5 3 1.2-1.4 cm in A. delphinensis, 2-3 3 1.4-1.6 cm in A. ditimena, 1.5-2.3 3 1.2-1.5 cm in A. itremoensis, 2-2.6 3 1.3-1.5 cm in A. lenticellata and 2-3 3 0.8-1.3 cm in A. sericea).

The name Protorhus ditimena var. minutifolia, published by Perrier de la Bâthie (1944) on the basis of a fragmentary specimen (Louvel 21), is invalid because it lacked a Latin diagnosis, as required for all names published between 1 January 1935 and 31 December 2011 under Article 39.1 of the International Code of Nomenclature (McNeill et al., 2012). We have chosen to retain Perrier de la Bâthie’s epithet, which aptly characterizes this taxon, by formally describing it as a new species.

Paratypi Madagascar. Prov. Antananarivo: Ankazobe, Ambohitantely RS, 18°11’S 47°17’E, 1620 m, 14.I.1997, fl.,Randrianaivo et al. 53 (G, K, MO, NY, P, TAN, WAG); forest E of Anjozorobe, E Antsahabe village, [18°22’S 47°47’E], 1400 m, 22.X.1996, fl.,A. Randrianasolo 508 (MO, P, TAN); Ambohitantely RS, 18°10’42’’S 47°17’00’’E, 1530 m, 23-24.XI.1993, fl.,Schatz et al. 3565 (G, K, MO, P, TAN); ibid. loco, [18°09’S 47°14’E], V.1953, fr., Service Forestier 7361 (P, TEF); ibid. loco, [18°08’S 47°16’E], 1600 m, IV.1953, fr., Service Forestier 8417 (P, TEF); ibid. loco, 26.VII.1954, ster., Service Forestier 11-R-282 (P, TEF); Poste forestier Manankazo, Ankazobe, [18°09’S 47°14’E], 22.VI.1955, fr., Service Forestier 16814 (P, TEF). Prov. Antsiranana: Ambanja, Bekolosy, 14°02’S 48°18’E, 1200 m, 28.III.1996, fr., Gautier & Be 2909 (G, MO, P, TAN, TEF); ibid. loco, 14°02’S 48°18’E, 1070 m, 23.IX.1996, fl.,Gautier et al. 3123 (G, MO, P, TAN, TEF); Ambilobe, forêt de Kalabenono, 13°38’45’’S 48°40’45’’E, 934 m, 27.XI.2006, fr., Callmander et al. 631 (G, MO); Andapa, Doany, forêt d’Andramanalana, 14°23’02’’S 49°21’36’’E, 1280 m, IV.2006, fr., Rakotovao et al. 3074 (G, MO, P, TAN); Andramilolo, SE Anjanaha- ribe, 14°19’43’’S 49°18’08’’E, 1412 m, 12.XI.2006, fl.,Razakamalala et al. 2939 (G, MO, P, TAN). Prov. Fianarantsoa: Tolongoina, Andrambavato, 2 km S Tatamaly, 21°30’S 47°24’E, 1075 m, 13.X.2000, fl.,Randriatafika 142 (G, K, MO, P, TAN). Prov. Toamasina: Alaotra-Mangoro, Ambatovy, Analamay, 18°48’28’’S 48°20’10’’E, 1083 m, 2.XII.2005, fl.,Antilahimena et al. 4316 (G, MO, P, TAN); ibid. loco, fl.,Anti - lahimena et al. 4318 (G, MO, P, TAN); Moramanga, Ambatovy/Andranovery forest, 18°52’09’’S 48°18’15’’E, 1013 m, 26.X.2007, fl.,Antilahimena 5937 (G, MO, P, TAN); Analamazaotra, IX.1925, bud, Louvel 21 (P); Maroantsetra, Masoala PN, E slope of Ambohitsitondroinan’Mahalevona summit, 15°25’34’’S 49°57’22’’E, 1160 m, 104 Boissiera 71

A B

Fig. 21. 25.II.2003, fr., Lowry et al. 6190 (G, K, MO, P, TAN, WAG); along road to Lakato, Abrahamia minutifolia 18 km S of RN2, 19°04’06’’S 48°22’01’’E, 1050 m, 17.X.2004, fl.,Lowry et al. 6547 Randrian. & Lowry. A. Branches showing (G, MO, P, TAN); Ambatovy, Analamay, 18°49’26’’S 48°20’16’’E, 1000 m, 26.I.1997, inflorescences and fr., Rakotomalaza et al. 1005 (MO, P, TAN); Analamazaotra, 950 m, 29.X.1996, fl.,A. flower buds Randrianasolo 515 (G, MO, P, TAN); Mantadia PN, c. 11 km N of village of Andasibe, [Antilaimena 5937]; 18°49’47’’S 48°25’56’’E, 939 m, 10.I.2003, ster., A. Randrianasolo 781 (MO, P, TEF); B. Branch with young fruits Ambatovy, 18°49’28’’S 48°19’23’’E, 1134 m, 26.XI.2005, bud, Razafindraibe 135 [Callmander et al. 631]. (MO, P, TAN); Analamazaotra-Périnet, [18°56’S 48°26’E], 6.XI.1957, bud, Service Forestier 18400 (G, K, MO, NY, P [2 sheets]); Ankazomanitra, PK 45, route Anosi- Photos: A: P.. Antilahimena; be-Moramanga, [19°58’S 48°08’E], 6.XI.1968, bud, Service Forestier 26800 (G, MO, B: M. Callmander NY, P, TEF); W d’Antanandava (PK 45 Moramanga-Anosibe), [19°08’S 48°15’E], XI.1968, fl.,Service Forestier 28453 (G, MO, P, TEF); ibid. loco, XI.1968, bud & fl., Service Forestier 28454 (G, MO, NY, P, TEF). Taxonomic treatment of Abrahamia (Anacardiaceae) 105

23. Abrahamia nitida (Engl.) Randrian. & Lowry, comb. nova.

≡ Protorhus nitida Engl. in Bot. Jarhb. Syst. 1: 421. 1881.

Typus: Madagascar: sine loco, s.d., fl.,Thouars s.n. (holo-: P [P00722662]!; iso-: P [P00722664]!).

Description Trees 4-25 m m tall, 10-22 cm DBH, bark with red or white milky latex; young twigs glabrous or glabrescent, with lenticels, waxy. Leaves usually whorled, sometimes opposite; blades oblanceolate, 3.5-10.5 3 1.5-3.8 cm, coriaceous, apex emarginate or retuse, margin revolute, base cuneate, glabrous on both surfaces, venation craspedodromous, midvein prominent abaxially, secondary veins 20-30 pairs, parallel, 3 mm apart, prominent abaxially, tertiary veins very slightly raised on abaxial surface; petiole 5-15 mm long, canaliculate on adaxial surface, glabrous. Inflorescence axillary or terminal, a panicle, 2-4 cm long, axes flattened, glabrous or glabrescent. Male and female flowers 5-merous; pedicel 0.5 mm long, pubescent; calyx lobes deltoid to triangular, 0.5-0.7 3 1-1.2 mm, valavate, glabrous, persistent on fruit; corolla lobes ovate, 2-2.2 3 1-1.5 mm, imbricate, adaxial surface glabrous, abaxial surface with short reddish indument; stamens 5. Male flowers with filaments c. 1 mm long, glabrous, anthers 0.5 mm long, ovoid to subglobose, glabrous; disk c. 1 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments c. 1 mm long, anthers 0.5 3 0.5 mm, obloid to ovoid, sterile, glabrous; disk c. 1 mm in diam., cupuliform, glabrous; ovary c. 0.5-0.6 3 1 mm, broadly ovoid, glabrous, placenation subapical, style very short, c. 0.2-0.3 mm long, apical, stigmas capitate. Fruits 2-2.5 3 1-1.5 cm, ellipsoid, glabrous, surface smooth.

Vernacular names “Ditimenafotsy’’, “Hazombarorana’’, “Tahaka’’, “Tsiramiramy’’.

Distribution, ecology Abrahamia nitida occurs in humid forest in coastal eastern Madagascar (Map 6). It has and phenology been recorded in flower in December and in fruit in late January.

Conservation status With an EOO of 45,042 km2, a minimum AOO of 76 km2, and 14 known locations, with subpopulations in 4 protected areas (Agnalazaha, Makira, Nosy Mangabe, and Pointe à Lareée) and the remainder subjected to pressure from forest exploitation and land clearing, A. nitida is assigned a preliminary conservation status of “Near Threat- ened” [NT] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes The leaves of A. nitida are usually whorled, a feature found in only a few other members of the genus, the most similar being A. sericea, from which A. nitida can be distinguished by its leaves with widely (vs very closely) spaced secondary veins, glabrous (vs pubescent) disk, and glabrous (vs velutinous) fruit. 106 Boissiera 71

We have followed René Capuron (note in herb.) in considering the second collection made by Thouars listed above as an isotype despite the fact that we cannot be certain it represents part of a single gathering.

Additional material Madagascar. Prov. Antsiranana: Antalaha, PN Masoala, 15°35’S 50°25’E, 22.IX.1994, examined ster., Rahajasoa et al. 510 (MO, TAN); ibid. loco, 22.IX.1994, ster., Rahajasoa et al. 523 (MO, P, TAN). Prov. Fianarantsoa: Farafangana, Mahabo, forêt de Marovahatry, 23°10’41’’ 47°43’49’’E, 23 m, 21.IV.2004, y. fr., Ludovic 738 (MO, P, TAN); Mahabo Mananivo, Nosiala, bloc de Betaindambo, 23°08’42’’S 47°42’18’’E, 31 m, 22.IV.2004, fr., Ludovic 766 (MO, P, TAN); ibid. loco, 23°11’S 47°41’E, 21 m, 29.X.2004, fl.,Ludovic et al. 927 (G, K, MO, P, TAN); Mahabo Mananivo, Baboaka, [23°10’S 47°44’E], 21 m, 20.XI.2004, fl.,Ludovic et al. 998 (G, K, MO, P, TAN); near village of Mahabo, 23°10’51’’S 47°42’29’’E, 30 m, 5.XI.2001, fl.,McPherson et al. 18334 (MO, TAN, TEF); ibid. loco, McPherson et al. 18347 (MO, P, TAN, TEF); ibid. loco, 23°10’42’’S 47°42’24’’E, 20 m, 7.XI.2001, fl.,McPherson et al. 18373 (MO, P, TAN, TEF); ibid. loco, 23°08’53’’S 47°43’04’’E, 22.IX.2002, fr., Rabevohitra 4077 (MO, NY, P, TAN, TEF); ibid. loco, 10 m, 19.VI.2004, ster., A. Randrianasolo et al. 867 (MO); ibid. loco, 23°12’05’’S 47°42’48’’E, 24 m, 26.VIII.2003, fr., Razakamalala et al. 744 (G, MO, P, TEF); Mananjary, forêt d’Alimamba, 21°28’04’’S 48°17’43’’E, 13.X.2004, fl.,Razakamalala et al. 1655 (MO, NY, P, TAN, TEF); Amporoforo, [22°29’00’’S 47°47’30’’E], 26.VII.1955, ster., Service Forestier 13912 (MO, P, TEF); Ambalavotaka, [21°19’30’’S 48°21’30’’E], 7.I.1955, fr., Service Forestier 15459 (P, TEF). Prov. Toamasina: Maroantsetra, commune d’Ambi- nanitelo, Ankirindro, 15°28’29’’S 49°33’10’’E, 200 m, 21.XI.2002, fl.,Antilahimena et al. 1430 (MO, P, TAN); Fénérive-Est, Mahatsara forest station, [17°38’S 49°29’E], 23.I.1986, fr., Dorr 4617 (MO, P, TAN); Soanieran’Ivongo, forêt d’Antanambao-Am- bodimanga, 16°46’24’’S 49°42’51’’E, 30.I.2003, fr., Rabevohitra et al. 4305 (G, K, MO, P, TEF); Mahanoro, Ambodibonara, forêt Nakinà, 20°21’34’’S 48°36’21’’E, 13.XI.2003, fl.,Rabevohitra et al. 4753 (G, K, MO, NY, P, TAN, TEF); Soanieran’Ivongo, NW Tanam- bao, 16°45’12’’S 49°42’56’’E, 31.I.2004, fr., Rabevohitra et al. 4980 (MO, NY, P, TEF); Brickaville, Andranokoditra, Vohibola, 18°33’23’’S 49°14’58’’E, 5 m, 10.XII.2001, fr., Rabenantoandro et al. 871 (MO, P, TAN, TEF); Tampolo forest station, 0-5 m, 31.VII.1996, ster., A. Randrianasolo 457 (MO, P, TAN); Ambila Lemaitso, 18°49’S 49°08’E, 0-3 m, 28.X.1996, fr., A. Randrianasolo 512 (G, K, MO, P, TAN); ibid. loco, 18°51’38’’S 49°08’34’’E, 0-5 m, 9.X.1999, ster., A. Randrianasolo et al. 586 (G, MO, P, TAN); Maroantsetra, Nosy Mangabe, 15°30’S 49°46’E, 0-330 m, 13-23.IV.1988, ster., Schatz 2131 (MO, TAN); Mahanoro, [19°44’00’’S 48°48’30’’E], 29.I.1951, fr., Service Forestier 2998 (P, TAN, TEF); Ambila Lemaitso, [18°49’S 49°08’E], 11.XII.1952, fl., Service Forestier 6448 (P [2 sheets], TEF); ibid. loco, 12.XII.1952, fl.,Service Forestier 6486 (P, TEF); ibid. loco, 24.I.1953, fr., Service Forestier 7253 (MO, P, TEF). Prov. Toliara: Fort-Dauphin, Mahatalaky St Luce S7, 2 km S d’Ambadrika, 24°47’47’’S 47°08’37’’E, 20 m, 25.XI.2011, fl.,Ratovoson 1793 (MO, P, TAN). Taxonomic treatment of Abrahamia (Anacardiaceae) 107

24. Abrahamia oblongifolia (Engl.) Randrian. & Lowry, comb. nova.

≡ Protorhus oblongifolia Engl. in Bot. Jahrb. Syst. 1: 422. 1881.

Typus: Madagascar. Prov. Antsiranana: Rég. Diana, Nossi-Bé, 1840, fl.,Richard 366 (holo-: P [P00722663]!).

Description Trees 10-20 m tall, 23-25 cm DBH; young twigs very often triangular, lenticellate, with ferruginous indument. Leaves subopposite to opposite; blades obovate to oblanceolate or narrowly oblong to elliptic, 4-17 3 1.5-5.5 cm, chartaceous to subcoriaceous, apex retuse, rounded, or emarginate, margin entire, minutely thickened and pale in color on abaxial surface, base cuneate or cuneate-obtuse, adaxial surface glabrous, somewhat waxy, abaxial surface pubescent when young, later nearly glabrous, venation craspedodromous (rarely a few secondary veins not quite reaching the margin), midvein prominent abaxially, secondary veins 35-40 pairs, 2-6 mm apart, more or less parallel, prominent on both surfaces but more so abaxially, tertiary veins slightly raised on abaxial surface; petiole 5-15 mm long, canaliculate on adaxial surface, glabrous, waxy. Inflorescence axillary or terminal, a panicle, 3-5 cm long, the axes terete or flattened, with ferruginous indument.Male and female flowers 5-merous (rarely 6-merous); pedicel 0-0.5 mm long, ferruginous pubescent or strigose; calyx lobes ovate, c. 1 3 1 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent or ferruginous pubescent; corolla lobes ovate, 2 3 1 mm, imbricate, adaxial surface glabrous, abaxial surface ferruginous pubescent or scabrous; stamens 5 or 6. Male flowers with filaments 1-1.5 mm long, glabrous, anthers c. 0.5 mm long, subglobose, glabrous; disk cupuliform, sometimes shallowly so, 1-1.5 mm in diam., glabrous; ovary rudimentary. Female flowers with filaments 0.5-0.7 mm long, glabrous, anthers c. 0.3 mm long, subglobose to slightly ovoid, glabrous; disk annular, cupuliform, sometimes shallowly so, 1.5 mm in diam., sterile, glabrous; ovary very widely ovoid, slightly asymmetric, 1 3 1 mm, glabrous, placentation apical; style simple, very short, inserted more sub-apically than laterally, 1 mm long, glabrous, stigma capitate, 3-4-lobed. Fruits c. 2 3 1 cm, ellipsoid, slightly asymmetrical, glabrous, with obscure longitudinal striations.

Vernacular names “Hazombarorana’’, “Manavodrevo’’, “Tarata’’, “Tsiramiramy’’.

Distribution, ecology Abrahamia oblongifolia occurs in far northern and northwestern Madagascar (Map 6), and phenology from sea level to an elevation of c. 300 m, usually along rivers or at the base of hills or mountains. Flowering material has been recorded in October and November, and fruits have been collected in December and January. 108 Boissiera 71

Conservation status With an EOO of 3,220 km2, a minimum AOO of 32 km2, and estimated maximum AOO of < 500 km2, and 7 recorded localities representing 7 locations with respect to the threat of forest exploitation, and with subpopulations in 2 protected areas (Analamera and Montagne des Français) and projected decline outside of protected areas, A. oblongifolia is assigned a preliminary conservation status of “Vulnerable’’ [VU B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia oblongifolia can be distinguished from all other members of the genus by the combination of its numerous secondary veins (35-40), both terminal and axillary inflorescences, and glabrous fruits.

Additional material Madagascar. Prov. Antsiranana: Analamera RS, Betsieka, [12°42’S 49°20’E], 10-20 examined m, I.1938, ster., Humbert 19091ter (P); Mahavanona, Montagne des Français, Ambato- sarolo, 12°22’27’’S 49°21’26’’E, 204 m, 8.X.2004, y. fr., Randriambololomamonjy et al. 42 (MO, P, TAN); Sakaramy, près de Diego, [12°27’S 49°13’E], bud, Perrier de la Bâthie 17672 (P); Ramena, Montagne des Français, Andavakoera, 12°20’48’’S 49°21’28’’E, 88 m, 7.IX.2004, fl.,Randrianaivo 1081 (CNARP, G, MO, P, TAN, TEF); Mahava- nona, Montagne des Français, campement Antafiankoroka, 12°21’39’’S 49°21’34’’E, 281 m, 24.IX.2004, y. fr., Randrianarivelo et al. 139 (G, K, MO, P, TAN); Nosy Be, [13°19’S 48°16’E], 1840, fr., Richard 315 (P); Montagnes des Français, [12°22’S 49°21’E], 31.XI.1952, bud, Service Forestier 5885 (MO, P [2 sheets], TEF); ibid. loco, 23.XI.1955, bud, Service Forestier 15355 (G, MO, P [2 sheets], TEF) ; Forêt de Sah- ankazo, [12°29’S 49°26’E], 11.XII.1955, fr., Service Forestier 15507 (G, MO, P [2 sheets], TEF); Montagnes des Français, [12°22’S 49°21’E], 21.IV.1954, ster., Service Forestier 28-R-61 (P); ibid. loco, 21.V.1951, ster., Service Forestier 51-R-150 (P); ibid. loco, 24.III.1955, ster., Service Forestier 442-R-6 (P). Sine loco: 17.X.1927, fl.,Ursch 186 (P). Taxonomic treatment of Abrahamia (Anacardiaceae) 109

25. Abrahamia patrickii Randrian. & Lowry, spec. nova (Fig. 22).

Typus: Madagascar. Prov. Antsiranana: Rég. SAVA, Vohémar, commune rurale de Daraina, forêt de Bobankora, partie N, 510 m, 13°13’15’’S 49°45’42’’E, 510 m, 29.I.2005, fr., Ranirison & Nusbaumer 919 (holo-: G [G00019652]!; iso-: MO-6671434!, P [P00580356]!, TEF [TEF000886]!).

Diagnosis Abrahamia patrickii Randrian. & Lowry can easily be distinguished from other members of the genus by its sub-verticillate to subopposite leaves, narrowly oblanceolate leaf blade with very closely spaced secondary veins, and the presence of circular orange-brown scales on the abaxial leaf surface.

Description Trees 13.5 m tall, 20 cm in diam.; young twigs lenticellate. Leaves sub-verticillate or subopposite; blade oblanceolate, 3-5.5 3 1-1.5 cm, subcoriaceous, apex emarginate or round, margin undulate, base cuneate to acute, adaxial surface glabrous, abaxial surface mostly glabrous but with scattered short reddish indument along midvein and margin, as well as circular, orange-brown scales, venation craspedodromous, midvein prominent abaxially, secondary veins 24-28 pairs, 1-3 mm apart, very often closely spaced, raised on both surfaces, tertiary veins not visible on either surface; petiole 6-13 mm long, c. 0.8-1 mm in diam., glabrous, canaliculate on adaxial surface. Inflorescence and flowers unknown. Fruits 2.3-2.6 3 1.1-1.3 cm, slightly asymmetric, ellipsoid, glabrous, with longitudinal striations.

Etymology The species epithet honors Patrick Ranirison for his dedication to the study of the Mal- agasy flora, most notably his work in the Daraina region of northeastern Madagascar.

Distribution, ecology Abrahamia patrickii is known only from the type collection from the Daraina area and a and phenology second gathering made N of Sambava (Map 6), in semi-deciduous and evergreen humid forest. It has been collected with fruits in January and February.

Conservation status With a minimum AOO of 8 km2 and 2 localities, both of which situated are within protected areas (Loky Manambato and Makirovona), A. patrickii is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia patrickii closely resembles A. nitida by its sub-verticillate leaves with an oblanceolate blade, similar number of craspedodromous secondary veins (24-28 in A. patrickii and 20-30 in A. nitida), similar petiole length (6-13 mm in A. patrickii and 5-15 mm in A. nitida), and glabrous fruit. However, the leaves of A. patrickii differ from those of A. nitida by their undulate (vs revolute) margins, orange-brown circular scales (vs glabrous) on the abaxial surface, narrowly spaced secondary veins (1-3 vs 3 mm apart), obscure (vs visible and raised) tertiary veins, and fruit with longitudinal striations (vs smooth) on the surface. 110 Boissiera 71

Paratypus Madagascar. Prov. Antsiranana: Rég. SAVA, Sambava, forêt d’Antsahaha Kasaka, 2 km NW du village Ambavala, 14°05’39’’S 50°02’42’’E, 146 m, 16.II.2006, fr., Andrianjafy et al. 1590 (MO, TAN, P).

1 cm 1 cm B A

Fig. 22. Abrahamia patrickii Randrian. & Lowry. A. Fruiting branch; B. Fruit. [Ranirison & Nusbaumer 919, MO] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 111

26. Abrahamia pauciflora (Engl.) Randrian. & Lowry, comb. nova.

≡ Protorhus pauciflora Engl. in A. DC. & C. DC., Monogr. Phan. 4: 313. 1883.

Lectotypus (designated here): Madagascar. Prov. Antsiranana: Rég. Diana, Nossibe, I.1850, fl. & y. fr.,Boivin 2257 (P [P00580352]!; isolecto-: G!, P [P00364873, P06774888, P06774892]!, MO-6684876!, TAN!).

Description Trees 6-11 m tall, 11-15 cm DBH; young branch tips covered with short and appressed indument. Leaves opposite or subopposite; blade obovate, 3.9-9.2 3 1.8-4 cm, chartaceous, apex rounded to emarginate, margin undulate, base cuneate to attenuate, adaxial surface glabrous, abaxial surface with scattered short appressed indument, venation craspedodromous, midvein prominent abaxially, secondary veins 14-17 pairs, more or less parallel and ascending, 3-9 mm apart, widely spaced at leaf mid-section, raised on both surfaces or some impressed near margin, tertiary veins impressed, not very visible, especially on adaxial surface; petiole 8-15 mm, c. 1 mm in diam., canaliculate on adaxial surface, covered with short, appressed indument when young, glabrescent. Inflorescence always axillary, a panicle, very short, up to 1.5 cm long, with no more than 5 flowers, axes strigose. Male flowers unknown. Female flowers5-merous, small, c. 1 mm long, subtended by a triangular bract with indument; pedicel 1 mm long; calyx lobes depressed ovate, very small, 0.5 3 1 mm, glabrous on both surfaces; corolla lobes ovate, thick, c. 2 x 1 mm, glabrous on both surfaces; staminodes with filaments c. 0.8 mm long, glabrous, anthers c. 0.3 mm long, ovoid, sterile, glabrous; disk cupuliform, glabrous, c. 1.5 mm in diam.; ovary spherical glabrous, style short, thick, 0.5 mm long. Fruits unknown.

Distribution, ecology Abrahamia pauciflora is restricted to NW Madagascar, where it has been collected on the island and phenology of Nosy Be and in the Ambongo-Boina area (Map 6) in humid forest. Material with floral buds and flowers has been collected in November and January (the fruiting period is not known).

Conservation status With a minimum AOO of 4 km2 and a single known extant subpopulation, which is within a protected area (Ampasindava), and considering that the historical locality at Maevarano was documented in 1908, well more than 3 generations ago, A. pauciflora is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia pauciflora can easily be distinguished from other members of the genus by the combination of its leaves with an attenuate base and very widely spaced secondary veins (to 9 mm apart in the median section of the blade), and very short inflorescence (not exceeding 1.5 cm long) usually bearing only 1-5 flowers. 112 Boissiera 71

Contrary to what is indicated in many online databases, the name Protorhus pauciflora (≡ Abrahamia pauciflora) was not treated in Engler (1881: 421 or 422), but rather was published two years later by the same author in his treatment of Anacardiaceae for the Monographiae Phanerogamarum (Engler, 1883: 313). Curiously, Engler (1883) erroneously cited that name as having been published in his earlier work (Engler, 1881: 421). While one of the specimens of Boivin 2257 in the Paris herbarium has the hand- written word ‘type’ on the label, this was probably added subsequent to the publica- tion of Protorhus pauciflora. We have chosen to designate this specimen as the lectotype because it bears Boivin’s original label. Two additional specimens of Boivin 2257 have labels indicating that they came from Mayotte in the Comoro Islands, but as indicated subsequently by Perrier de la Bâthie (in herb.), they are without question part of the same gathering as the material from Nossi Be.

Additional material Madagascar. Prov. Antsiranana: Ampasindava, forêt d’Andranomatavy, 13°40’40’’S examined 47°58’35’’E, 215 m, 2.XII.2009, bud, Ammann et al. 467 (G, MO, P, TEF); ibid. loco, 13°40’04’’S 47°59’21’’E, 295 m, 25.XI.2009, fl.,Madiomanana et al. 272 (G, MO, P, TEF). Prov. Mahajanga: Morarivo, Maevarano, près de Majunga, [15°56’S 46°31’E], VII.1908, ster., Perrier de la Bâthie 2341 (P). Taxonomic treatment of Abrahamia (Anacardiaceae) 113

27. Abrahamia phillipsonii Randrian. & Lowry, spec. nova (Fig. 23-24).

Typus: Madagascar. Prov. Toliara: Rég. Androy, Andohahela PN, parcelle 2, 24°50’S 46°32’E, 100 m, 7.XII.1988, fr., Phillipson 2853 (holo-: MO-3767006!; iso-: K, P [P06775412]!, TAN [TAN001997]!, US-3661611 image seen, WAG [WAG1523550] image seen).

Diagnosis Abrahamia phillipsonii Randrian. & Lowry most closely resembles A. lenticellata, with which its range overlaps, but can be easily distinguished by its craspedodromous (vs semicraspedodromous) venation, flowers borne in thyrses (vs panicles), and glabrous (vs velutinous) fruits.

Description Trees 3-12(-20) m tall, 12-30 cm DBH, bark with white latex; young twigs glabrous or pubescent (sometimes ferruginous). Leaves alternate or subopposite; blades oblanceolate to obovate or narrowly elliptic to narrowly oblong, 2.6-7.8 3 0.6-2(-2.2) cm, chartaceous to subcoriaceous, apex emarginate, retuse or rounded, sometimes acute, margin entire, base acute or cuneate, adaxial surface glabrous, very often waxy, abaxial surface with very sparse indument, slightly more appressed on the midvein, nearly glabrous oth- erwise, venation craspedodromous, midvein prominent abaxially, secondary veins 11-19 (-23) pairs, prominent abaxially, 1-5 mm apart, tertiary veins rarely visible on both surfaces; petiole c. 0.5-10 mm long, glabrous, canaliculate on adaxial surface. Inflorescence axillary or terminal, a thyrse, 1.5-5.5 cm long, the axes flattened, with short, appressed, ferruginous or reddish indument. Male flowers 5-merous; calyx lobes widely depressed ovate to depressed ovate or triangular, 0.8-1 3 1-1.5 mm, valvate, glabrous, persistent on fruits; corolla lobes ovate, 2-2.2 3 1-1.5 mm, imbricate, glabrous, stamens 5, filaments c. 1.5 mm long, white, glabrous, anthers 0.8-1 mm long, ovoid to subglobose, yellow, glabrous; disk 1-1.2 mm in diam., glabrous; ovary rudimentary. Female flowers unknown. Fruits 1.5-2 3 1-1.3 cm, obovoid to ellipsoid, glabrous, with longitudinal striations.

Etymology The species epithet honors our friend and colleague Peter. B. Phillipson, whose work has contributed so much to our knowledge of the Malagasy flora, including that of the drier parts of the island, where he has conducted most of his field work over the last three decades.

Vernacular names “Manantsadrano’’, “Tarantantily’’, “Tsitaké’’.

Distribution, ecology Abrahamia phillipsonii is restricted to sub-arid southern Madagascar (Map 8), where it and phenology occurs at elevations between 100 and 1100 m. Flowering material has been collected between September and November, and fruits have been gathered from December through February. However, in drought years flowering can be delayed, which accounts for the limited number of collection with flowers gathered in March and April. 114 Boissiera 71

A B C D

1 cm 2 mm 1 mm 1 cm

Fig. 23. Abrahamia phillipsonii Randrian. & Lowry. A. Flowering branch; B. Inflorescence; C. Detail of male flower; D. Fruit.

[A-C: Service Forestier 8523bis, MO; D: Phillipson 2853, MO] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 115

Conservation status With an EOO of 64,857 km2, a minimum AOO of 112 km2, and 29 recorded localities, with subpopulations in 4 protected areas (Analavelona, Andohahela, Isalo, and Zom- bitsy), A. phillipsonii is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia phillipsonii vegetatively most closely resembles A. deflexa, but it differs by its oblanceolate to obovate, narrowly elliptic to narrowly oblong (vs ovate, oblong or elliptic) leaf blades and its axillary or terminal (vs only terminal) inflorescence.

Paratypi Madagascar. Prov. Fianarantsoa: Ihosy, Analalava forest, 2-3 km W of western bound- ary of Isalo PN, [22°35’38’’S 45°08’20’’E], 770 m, 18.V.2004, fr., Rogers et al. 456 (G, K, MO, P, TAN); massif de l’Isalo, [22°08’S 45°10’E], 20.XII.1968, fr., Service Forestier 28504 (P, TEF); Ranohira, PK 712, [22°38’S 45°20’E], 23.VII.1955, ster., Service For- estier 13889 (MO, P, TEF); Isalo, 20.VII.1954, ster., Service Forestier 135-R-141 (P); Isalo, Benenitra, 8.VIII.1954, ster., Service Forestier 180-R-18 (P); Isalo, Ranohira, 23.VII.1955, ster., Service Forestier 288-R-239 (P); Ihosy, 30.X.1953, ster., Service Forestier 777-R-1 (P). Prov. Toliara: Tolagnaro, Andohahela PN, N du mont Apiky, [24°47’02’’S 46°43’12’’E], 708 m, 18.I.2007, fr., Andriamihajarivo 1156 (MO, P, TAN, TEF); Amboasary, fourré d’Ambatofotsy Masay, 24°18’16’’S 46°08’00’’E, 350 m, 11.XI.2008, fr., Andriamihajarivo et al. 1574 (G, MO, P, TAN); Sakaraha, forêt de Zombitsy, [22°48’S 44°41’E], XII.1959, fr., Bosser 13402 (MO, P [2 sheets], TAN); Andohahela PN, parcelle 3, [25°00’S 46°40’E], 17.XI.1990, fr., Dumetz 1413 (MO, P, TAN); vallée de Fiherenana, 300-500 m, 1-2.VIII.1928, fr., Humbert & Swingle 5063 (= Service Forestier 50) (P); Betroka, Mont Vohipolaka, 1200-1400 m, XI.1933, ster., Humbert 11694ter (P, TAN); Amboasary, entre Andohahela et l’Elakelaka, [24°45’S 46°43’E], 700 m, I-II.1934, fr., Humbert 13770 (P); entre Andohahela et l’Elakelaka, mont Apiky, [24°45’30’’S 46°43’30’’E], 800-900 m, I-II.1934, fr., Humbert 13850 (G, P [2 sheets]); Beroroha, bassin de la Malio, [21°58’S 45°15’E], 400-450 m, 23-27.XI.1946, bud, Humbert 19403 (P); forêt de Zombitsy, [22°48’S 44°41’E], 600-850 m, 26-29.III.1955, fl. & bud,Humbert 29595 (G, MO [2 sheets], P); Beronono-Makay, 21°18’48’’S 45°08’55’’E, 531 m, 16.I.2010, fr., Rakotovao et al. 5117 (MO, P, TAN); PN Andohahela, Isaka Ivondro, 24°48’S 46°52’E, 100-500 m, 16-24.III.1993, Randriamampionona 202 (G, MO, P, TAN); PN Andohahela, parcel 3, 25°04’S 46°41’E, 100-300 m, 8-10.IV.1993, y. fr., Randriamampionona 285 (G, MO, P, TAN); Andohahela PN, parcel 1, Tsimelahy, 24°50’20’’S 46°32’17’’E, 6.IV.1996, ster., Randriamampionona 1259 (MO, TAN); Sakaraha, Mahaboboka, Ambinanintelo, forêt d’Analavelona, 22°41’14’’S 44°09’32’’E, 961 m, 3.XII.2016, fr., Randrianarivony et al. 745 (MO, P, TAN); PN Andohahela, parcelle n° 3, [25°04’S 46°41 E], 100-300 m, 19.X.1990, fl.,A. Randrianasolo 181 (MO [2 sheets], P, TAN, TEF); Sakaraha, at start of Kinanamboribory valley, 22°39’44’’S 44°11’02’’E, 1208 m, 8.XII.2009, fr., A. Randriana- solo 1246 (G, MO, P, TAN); Ambatoabo, 5 km N d’Imonty, 24°46’44’’S 46°42’56’’E, 500 m, 25.XI.2009, fr., Ratovoson 1534 (G, K, MO, P, TAN); Betorabato, Abotorabatorano, 21°34’04’’S 45°34’34’’E, 296 m, 13.I.2011, fr., Razakamalala 6000 (G, MO, P, TAN); Ankomaka, [22°36’S 44°02’E], 26.I.1952, fr., Service Forestier 4556 (P, TEF); forêt de 116 Boissiera 71

Zombitsy, [22°46’S 44°42’E], 24.IV.1953, fl.,Service Forestier 7757 (P, TEF); env. de Tsimela, S d’Ambatoabo, [24°56’S 46°38’E], 28.IX.1953, fl.,Service Forestier 8523bis (G, NY, MO, P, TEF); forêt de Zombitsy, [22°46’30’’S 44°38’00’’E], III.1931, fl.,Ser - vice Forestier 11931 (G, MO, P); Analavelona, [22°40’S 44°10’E], 25.XI.1954, fr., Ser- vice Forestier 12089 (P, TEF); ibid. loco, 28.XI.1954, fr., Service Forestier 12140 (P, TEF); Betioky, vallée Sakamarakely, [22°11’S 45°10’E], 14.XI.1954, fr., Service Forestier 12362 (P); Zombitsy, [22°46’S 44°42’E], 7.XI.1954, fl.,Service Forestier 12428 (P [2 sheets], TEF); Ambovombe, Beanantara, [25°03’30’’S 46°12’30’’E], 17.IX.1955, fl.,Service Forestier 15656 (P, TEF); Ampolitsy, Beroroha, [21°40’30’’S 45°10’E], 16.IV.1956, y. fr., Service Forestier 15800 (P, TEF); Amboasary, bush entre Bevilany et Ranomainty, [25°00’S 46°34’E], 1.XII.1961, fr., Service Forestier 20481 (P [2 sheets], TEF); crête du massif de l’Angavo, [24°50’30’’S 45°48’00’’E], c. 500 m, 21.I.1963, fr., Service Forestier 22487 (P, TEF); Zombitse, Andranolava, 13.V.1954, ster., Service Forestier 27-R-224 (P); Kitranga, Ihosy, 25.VIII.1955, ster., Service Forestier 43-R-230 (P); Analavelona, 7.IV.1954, ster., Service Forestier 76-R-18 (P); forêt de Zombitsy, 26.II.1955, ster., Service Forestier 86-R-161 (P); forêt d’Ihera, 25.IV.1955, ster., Service Forestier 104-R-161 (P); ibid. loco, 9.IV.1954, ster., Service Forestier 108-R-18 (P); Bejangoa, Ankazoabo, 25.V.1954, ster., Service Forestier 112-R-13 (P); env. d’Ampandrandava (entre Bekily et Tsivory), [24°05’S 45°42’E], 1100 m, XI.1942, fl.,Seyrig 265 (P [2 sheets]); ibid. loco, Seyrig 265B (MO, P).

Fig. 24. Abrahamia phillipsonii Randrian. & Lowry. Fruits [Ratovoson et al. 1534]. Photo: F. Ratovoson Taxonomic treatment of Abrahamia (Anacardiaceae) 117

28. Abrahamia revoluta Randrian. & Lowry, spec. nova (Fig. 25).

Typus: Madagascar. Prov. Antsiranana: Rég. SAVA, Vohémar, forêt littoral au S de Vohémar, [13°21’S 50°00’E], 1-19.XII.1966, fr., Service Forestier 24318A (holo: P [P06168913]!; iso-: MO-05056713!).

Diagnosis Abrahamia revoluta Randrian. & Lowry can be distinguished from its congeners by its glabrous leaves with undulate and revolute margins and semicraspedodromous secondary veins.

Description Trees 15-20 m tall, 21.6 cm DBH; young twigs glabrous, lenticellate. Leaves opposite or subopposite; blade obovate, 3.2-9 3 1.4-3.8 cm, subcoriaceous, apex retuse, sometimes emarginate, margin revolute, undulate, base cuneate, both surfaces glabrous, venation semicraspedodromous, midvein prominent abaxially, secondary veins 15-30 pairs, more or less parallel, 2-4(-5) mm apart, raised on both leaf surfaces, tertiary veins not visible on both surfaces; petiole 5-13 mm long, 0.5-1 mm in diam., canaliculate on adaxial surface, usually glabrous. Inflorescenceand flowers unknown. Infrutescence terminal. Fruits 2.5-2.7 3 1.3-1.5 cm, glabrous, with longitudinal striations.

Distribution, ecology Abrahamia revoluta occurs in NE Madagascar from S of Vohémar to the southern part of and phenology the Masoala peninsula (Map 9). Fruiting material has been gathered between October and December.

Conservation status Abrahamia revoluta has a minimum AOO of 12 km2 and only one of its three extant subpopulation occurs within a protected area (Masoala) whereas the other two were collected in areas where future decline is expected in terms of quality of habitat. It has therefore been assigned a preliminary status of “Endangered’’ [EN B2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia revoluta most closely resembles A. ditimena, with which it shares obovate leaves with a retuse apex that are similar in size and the number of secondary veins. However, A. revoluta differs by its subcoriaceous (vs coriaceous) leaves, semicraspedodromous (vs craspedodromous) venation, revolute and undulate (vs entire and flat) leaf margins, obscure (vs raised and visible) tertiary venation on the lower leaf surface, terminal (vs terminal or axillary) infrutescences, and glabrous (vs velutinous) fruits.

Paratypi Madagascar. Prov. Antsiranana: versant NW de la Montagne d’Ambre, forêt d’An- karefo (Ankazobe), 19-21.X.1954, y. fr., Service Forestier 11357bis (P, MO). Prov. Toama- sina: Maroantsetra, Masoala Peninsula, S of Ambanizana, 15°40’24’’S 49°57’51’’E, 110- 260 m, 29.XI.1994, ster., Vasey et al. 176 (MO, TAN). 118 Boissiera 71

1 cm

1 cm Fig. 25. Abrahamia revoluta Randrian. & Lowry. A. Fruiting branch; B. Fruit. [Service Forestier 24318A, MO] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 119

29. Abrahamia sambiranensis Randrian. & Lowry, spec. nova (Fig. 26).

Typus: Madagascar. Prov. Mahajanga: Rég. Diana, Ambato forêt classée, c. 40 km N d’Ambanja, 13°26’S 48°33’E, 90-459 m, 23.XI.1996, fr., Randrianaivo et al. 29 (holo-: MO-6671436!; iso-: G!, MO-6671435!, P [P06775160]!, TAN [TAN001998]!).

Diagnosis Abrahamia sambiranensis Randrian. & Lowry can be distinguished from other members of the genus by the combination of large (10-19 3 2.5-5.6 cm), coriaceous leaves with numerous (50- 60) distinctly raised secondary veins and a folded apex in dry material, its long petiole (30-65 mm), and its large, glabrous fruits (2.5-3 cm long).

Description Trees 10-15 m tall, 15-18 cm DBH, bark with white latex; young twigs pubescent towards the tips. Leaves alternate, clustered toward the branch ends; blade oblanceolate or narrowly elliptic to lanceolate, 10-19.4 3 3.2-5.5 cm, coriaceous, apex acuminate, folded in dry material, margin undulate, base cuneate to broadly cuneate, adaxial surface glabrous, often glaucous, abaxial surface sparsely sericeous, more so on midvein, venation craspedodromous, midvein prominent abaxially, secondary veins 50-60 pairs, more or less parallel, 2-4 mm apart, raised on both surfaces, tertiary veins not visible on both surfaces; petiole (20)-25-58 mm long, 2-3.5 mm in diam., canaliculate on adaxial surface, tomentose. Inflorescence axillary, a panicle, 2-9.5 cm long, very few flowers reaching maturity, axes pubescent. Male flowers 5-merous; pedicel c. 0.5-1 mm long, pubescent; calyx lobes triangular to ovate, 1 3 1 mm, connate at base, adaxial surface glabrous, abaxial surface pubescent; corolla lobes ovate, 2.5 3 1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent; stamens 5, filaments 1 mm long, anthers c. 1 3 1 mm, obloid to ovoid; disk cupuliform, glabrous; rudimentary ovary absent. Female flowersunknown. Fruits 2.5-3 3 1.5-1.8 cm, ellipsoid, pubescent when young, glabrescent, with fine longitudinal striations.

Distribution, ecology Abrahamia sambiranensis is restricted to humid forests in the Sambirano region of NW and phenology Madagascar and extends into the northern part of the island (Map 9). It flowers in July and August, and fruits have been collected in October and November.

Conservation status With an EOO of 1,717 km2, a minimum AOO of 28 km2, and seven localities, with only a single subpopulation within a protected area (Galoko), A. sambiranensis exists at 7 locations, six of which are subjected to pressure from forest exploitation and land clearing; it is therefore assigned a preliminary conservation status of “Vulnerable” [VU B1ab(iii)+2ab(iii)] using the IUCN Red List Categories and Criteria (IUCN, 2012). 120 Boissiera 71

Notes Abrahamia sambiranensis vegetatively resembles A. elongata, but differs by having leaves that lack tertiary venation (vs present and raised on both surfaces in A. elongata) and whose apex is usually folded in pressed material (vs flat), and by its fruits that are glabrescent when mature (vs glabrous in A. elongata).

Paratypi Madagascar. Prov. Antsiranana: Rég. Diana, Ambilomangodra, 13°02’42’’S 49°09’11’’E, 50 m, 14.VII.2005, fl.,Randrianaivo et al. 1201 (CNARP, MO, P, TAN); forêt de Kalabenono, 13°37’27’’S 48°40’05’’E, 369 m, 10.XI.2006, fr., Razafitsalama et al. 1064 (G, MO, P, TAN); Galoko, [13°33’45’’S 48°45’00’’E], 29.VII.1955, y. fr., Service Forestier 15748 (P, TEF); forêts à l’E d’Ampondrabe, 28.X.1954, fr., Service Forestier 18931 (MO, P [2 sheets]); massif de l’Anda- vakoera, N d’Angodromena, [13°06’30’’S 49°18’30’’E], 1.X.1958, fr., Service Forestier 18947 (G, MO, P [2 sheets], TEF); Ampandrikoera, 12.X.1951, ster., Service Forestier 181-R-64 (P).

1 cm

Fig. 26. Abrahamia sambiranensis Randrian. & Lowry. A A. Fruiting branch; B. Fruit.

[Randrianaivo et al. 29, P] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 121

30. Abrahamia sericea (Engl.) Randrian. & Lowry, comb. nova (Fig. 27).

≡ Protorhus sericea Engl. in Bot. Jahrb. Syst. 1: 421. 1881.

Typus: Madagascar: sine loco, s.d., y. fr., Thouars s.n. (holo-: P [P00580351]!).

Description Trees 5-25 m tall, 11-50 cm DBH, bark with white or translucid latex; young twigs ferruginous-pubescent or with short, yellow indument, lenticellate, waxy. Leaves alternate or subopposite, sometimes verticellate; blades oblanceolate to obovate, sometimes obovate, 5-15.8 3 2.7-4.5 cm, subcoriaceous, apex emarginate, retuse or shortly acuminate, sometimes truncate, margin entire, base acute or cuneate, adaxial surface glabrous, abaxial surface pubescent when young, later glabrescent, venation craspedodromous, midvein prominent abaxially, secondary veins 40-70 pairs, more or less parallel, 0.5-3 mm apart, prominent on both surfaces, tertiary veins raised on adaxial surface; petiole 8-20 mm long, canaliculate on adaxial surface and very often flattened at base, pubescent when young, glabrescent. Inflorescence terminal, sometimes axillary, a panicle or thyrse, 5-10 cm long, the axes flattened, covered with yellowish, ferruginous strigose indument.Male and female flowers 5- or 6-merous; pedicel 0-0.3 mm long, strigose or pubescent; calyx lobes deltate or narrowly deltate, 1-1.2 3 1-1.5 mm, imbricate, rarely valvate, adaxial surface glabrous, abaxial surface pubescent or ferruginous, persistent on fruit; corolla lobes ovate, 2-3 3 1-1.8 mm, imbricate or quincuncial, adaxial surface glabrous, abaxial surface pubescent; stamens 5 or 6. Male flowers with filaments 1-1.5 mm long, glabrous, anthers 0.8-1 mm long, ovoid, glabrous; disk 1-2 mm in diam., pubescent; ovary rudimentary. Female flowers with filaments c. 1 mm long, glabrous, anthers 0.2-0.5 mm long, ovoid, sterile, glabrous; disk 1.5-2 mm in diam., pubescent; ovary very widely ovoid, 1-1.5 3 1 -1.5 mm, covered with sericeous indument, placentation subapical; style very shortly trifid (apparently simple), apical, 0.5-1 mm long, pubescent, stigmas capitate. Fruits 2-3 3 0.8-1.5 cm, ellipsoid, slightly asymmetrical, velutinous, with longitudinal striations.

Vernacular names “Ditimena’’, “Hazombarorana’’, “Rodrano’’ (Tanala), “Sandramy’’, “Sosoka’’, “Tsimalazo’’, “Varoa’’, “Vodihazo’’ (Tanala).

Distribution, ecology Abrahamia sericea occurs in eastern humid domain forest, extending from coastal areas and phenology to the escarpment (Map 8), where it can reach 800-1200 m elevation. It flowers from September through December, and sets fruit from December until May.

Conservation status With an EOO of 178,062 km2, a minimum AOO of 236 km2, and 64 known localities, with subpopulations in numerous protected areas, A. sericea is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012). 122 Boissiera 71

45 E 50 E

Map 9. Geographic distribution of species of Abrahamia Randrian. & Lowry

A. revoluta Randrian. & Lowry A. sambiranensis Randrian. & Lowry A. viguieri (H. Perrier) Randrian. & Lowry

15 S

20 S

25 S Taxonomic treatment of Abrahamia (Anacardiaceae) 123

45 E 50 E

Map 10. Geographic distribution of Abrahamia Randrian. & Lowry

A. thouvenotii (Lecomte) Randrian. & Lowry A. turkii Randrian. & Lowry

15 S

20 S

25 S 124 Boissiera 71

Notes Abrahamia sericea and A. grandidieri are distinctive within the genus in having very closely spaced, parallel secondary veins, but A. sericea differs by the presence of indument on its disk and ovary, a character absent in A. grandidieri but shared with another species, A. thouvenotii, which, however, has very widely spaced, arcuate secondary veins.

Additional material Madagascar. Prov. Antsiranana: Rég. Diana, Tsaratanana RNI, Mandrizavona village, examined 13°52’33’’S 48°48’44’’E, 700 m, 27.III.2000, fr., Antilahimena et al. 429 (G, K, MO, NY, P, TAN); Ambodimanga Ramena, Tsaratanana Reserve, 13°52’37’’S 48°51’02’’E, 547 m, 15.XI.2001, y. fr., Antilahimena et al. 817 (G, K, MO, P, TAN); Antalaha, forêt littorale Ambodipont, Masoala PN, 15°44’45’’S 50°19’25’’E, 3.X.1996, bud, Bernard 360 (MO, P, TAN); Ambodimanga, Antsahabe, 13°49’30’’S 48°46’59’’E, 500 m, 7.X.1998, fl.,Birkin - shaw et al. 527 (MO, P, TAN); Manongarivo RS, bras gauche de l’Ambatra, 13°59’S 48°26’E, 470 m, 5.VI.2000, old fr., Gautier et al. 3751 (G, MO, P, TAN); vallée de la Lokoho, [14°26’S 49°49’E], 75 m, 13.XII.1948, bud, Humbert 22383 (G, MO, P); Sam- bava, vallée de l’Andalangy, [14°18’S 49°43’E], 200-800 m, 12-14.XI.1950, bud & fl., Humbert 24164 (MO, P); Ambilobe, massif de Marivorahona au SW de Manambato, [13°46’40’’S 48°59’30’’E], 1750-2000 m, 18-26.III.1951, fr., Humbert 25713 (MO, P); E of Ankaramy, Manongarivo RS, Antsatrotro, 14°05’S 48°23’E, 600-900 m, 26-30. III.1993, fr., Malcomber et al. 2291 (G, K, MO, NY, P, TAN); Sambirano, forêt de l’Ankaizina, [14°30’S 48°55’E], 1000 m, VII.1908, bud, Perrier de la Bâthie 5859 (P [2 sheets]); Antalaha, Tampolo, 2 km S of Befotaka, 14°44’S 50°13’E, 27.I.2007, fr., Rana- rivelo 1255 (CAS, G, MO); Ambodimanga-Ramena, le long de la piste vers Antsaharatsy, 13°48’33’’S 48°45’07’’E, 460 m, 18.IV.2000, y. fr. & fr., Ratovoson 236 (P); Masoala PN, Ambohitralanana, [15°33’S 50°07’E], 24.I.1953, fl.,Réserves Naturelles 4942 (P, TEF); Masoala PN, Ampanavoana, [15°33’S 50°07’E], 6.XII.1954, fl.,Réserves Naturelles 6808 (P [2 sheets], TEF); bord d’un affluent de l’Andalangy, Sambava, [14°18’S 49°43’E], 350 m, 12.XII.1950, bud, Service Forestier 810 (MO, P [3 sheets], TEF); Masoala PN, Ambo- hitralanana, 15°33’S 50°07’E, 10.II.1954, fl.,Réserves Naturelles 6618 (P, TEF); massif du Marivorahona, Ankijabe, [13°46’40’’S 49°00’00’’E], 1050 m, 26.III.1951, fr., Service Forestier 3070 (P [2 sheets], TEF); Tsiroanimandidy-Ambanja, [13°52’30’’S 48°21’00’’E], 200 m, 26.IX.1952, fl.,Service Forestier 6113 (P [3 sheets], TEF); Andrambavato, [14°15’00’’S 50°01’30’’E], 100 m, 26.XI.1954, fl.,Service Forestier 12232 (P [2 sheets], TEF); forêt de Betafononana, Maromandia, [14°53’S 50°16’E], 0-50m, 16.XI.1954, fl., Service Forestier 12449 (P, TEF); Ambohitralanana, Antalaha, [former Masoala RN], [15°33’S 50°07’E], 0-50 m, 23.X.1956, fl.,Service Forestier 17438 [= Réserves Naturelles 8592] (P, TEF). Prov. Fianarantsoa: Fort-Carnot, [21°51’30’’S 47°26’30’’E], 1986, bud, Beaujard 12 (P); Rég. Tanala, [21°51’S 47°26’E], 1994, ster., Beaujard 441 (P); Mananjary, entre les bassins du Matilana et du Mananjary, [22°26’S 47°55’E], 800 m, X.1911, fl., Perrier de la Bâthie 5851 (P [2 sheets]); Vohipeno, Ampasina-Maroala, [22°21’S 48°11’E], 0-100 m, 20.X.1952, bud & fl.,Service Forestier 6431 (P [2 sheets], TEF); Sahavana, Ft. Carnot, [21°51’30’’S 47°26’30’’E], 23.XII.1952, fr., Service Forestier 6664 (P, TEF); Van- gaindrano, Vohipaho, [23°33’30’’S 47°30’30’’E], 10 m, 15.XI.1952, fl.,Service Forestier 7107 (MO, P, TEF); Tolongoina, Mandriandry, [21°34’30’’S 47°31’00’’E], 500 m, Taxonomic treatment of Abrahamia (Anacardiaceae) 125

Fig. 27. Abrahamia sericea (Engl.) Randrian. & Lowry. Branch with fruits, from below [Razanatsima et al. 1442].

Photo: A. Razanatsima 126 Boissiera 71

28.IX.1952, bud, Service Forestier 7113 (P, TEF); Maromiandra, Manampatroana, [21°41’S 47°31’E], 300 m, 29.XI.1952, bud, Service Forestier 7150 (P, TEF); Nosy Varika, Nosiarivo, [20°30’30’’S 48°00’00’’E], 500 m, 28.I.1953, fr., Service Forestier 7302 (P, TEF); rive droite de Sahanana Ampasinambo, [20°31’S 48°00’E], 530 m, 14.XI.1953, bud & fl.,Service Forestier 9955 (P, TEF); Sakahana, Tolongoina, Ft. Carnot, [21°31’30’’S 47°45’30’’E], 878-1000 m, 17.XI.1954, fl.,Service Forestier 13653 (P, TEF); Sahavana, Ft. Carnot, [21°51’30’’S 47°26’30’’E], 31.XI.1954, fl.,Service Forestier 14511 (P, TEF); Ambalangavo, Vohipeno, [22°21’S 47°40’E], 0-100 m, 15.X.1954, fr., Service Forestier 14929 (MO, P, TEF). Prov. Toamasina: Moramanga, Analamazaotra, 18°55’11’’S 48°25’17’’E, 100-1200 m, 28.II.2000, fr., Andrianjafy 27 (MO, P, TAN); Vavatenina, Manakambahiny I, 17°39’07’’S 48°58’14’’E, 550-600 m, 21.X.2002, bud, Andrianjafy et al. 293 (CNARP, MO, P, TAN); Maroantsetra, Rantabe, Mafaipoza, 15°39’24’’S 49°34’53’’E, 80 m, 21.II.2002, fr., Antilahimena 872 (G, MO, P, TAN); Maroantsetra, along Andramo- fotsy river, 15°07’30’’S 49°49’13’’E, 99 m, 6.XI.2003, bud, Antilahimena et al. 2264 (G, K, MO, P, TAN); Toamasina II, Fotsimavo, Ambodiriana, [17°55’S 49°13’E], 300-400 m, XII.1962, fr., Bosser 16913 (MO, P, TAN); Nosy Mangabe, 15°29’S 49°45’E, 15 m, IV.1990, ster., Carlson 149 (MO); ibid loco, Carlson 195 (MO); Ambatondrazaka, Ampita- nonoka à Fotsialanana, [17°46’S 48°54’E], 1200 m, 19.I.1945, fr., Cours 2465 (MO, P); Brickaville, itinéraire de Didy, [18°07’00’’S 48°32’40’’E], 100-1100 m, bud, Cours 4907 (P [4 sheets]); Moramanga, PK 163-164, W of Andasibe (Périnet), [18°56’S 47°57’E], 1105 m, 19.I.1986, fr., Dorr et al. 4582 (MO, P, TAN); forêt d’Analamazaotra, [18°55’S 48°25’E], 1000 m, fl., Louvel 3 (P); Andasibe, Mantadia NP, [18°49’30’’S 48°28’00’’E], 930-1040 m, 10.I.2003, y. fr., Pell et al. 621 (MO, NY, P, TAN); forêt d’Analamazotra, [18°55’48’’S 48°25’48’’E], s.d., fl. & fr.,Perrier de la Bâthie 3 (P); bords des rivières, Ambatovola, [18°53’S 48°34’E], I.1928, fr., Perrier de la Bâthie 18365 (P [2 sheets]); Zahamena PN, entre rivières Laindrotra et Saharahara, 17°45’34’’S 48°42’04’’E, 900 m, 9.III.1990, fr., Rakotondrajaona 168 (MO, P, TAN); Analamazaotra RS, 18°56’S 48°26’E, 960 m, 15.VII.1996, ster., A. Randrianasolo 404 (MO, P, TAN); Tampolo forestry station, 17°17’S 49°23’E, 5 m, 1.VIII.1996, ster., A. Randrianasolo 468 (MO, P); Analamazaotra RS, 18°56’S 48°26’E, c. 900 m, 28.X.1996, bud, A. Randrianasolo 513 (G, K, MO, NY, P, TAN); Mantadia PN, c. 11 km N of village of Andasibe, 18°49’47’’S 48°25’56’’E, 939 m, 10.I.2003, ster., A. Randrianasolo 783 (MO, P, TEF); Vatomandry, Ambalabe, N of Maro- toho, 19°10’19’’S 48°37’26’’E, 367 m, 26.XI.2004, y. fr., A. Randrianasolo et al. 969 (G, K, MO, NY, P, TAN, WAG); Ambalabe, Antsofotra forest, 1 km NW of Mahatsara village, 19°13’48’’S 48°37’57’’E, 226 m, 3.II.2011, ster., A. Randrianasolo 1341 (G, K, MO, P, TAN); Manakambahiny-Est, Amparafarabe, Zahamena PN, 17°43’42’’S 48°40’27’’E, 875 m, 2.XI.2001, fl.,S. Randrianasolo et al. 251 (CNARP, MO, P, TAN, TEF); Ambatovy, Andranovery, 18°51’51’’S 48°18’49’’E, 1005 m, 14.II.2008, ster., S. Randrianasolo et al. 670 (MO, P, TAN); Manakambahiny-Est, Zahamena PN, forêt Alaindrotra, 17°45’07’’S 48°43’01’’E, 2.III.2001, fr., Randrianjanaka et al. 595 (CNARP, MO, P, TAN, TEF); Zahamena PN, W de Mitanonoka, 17°45’53’’S 48°54’01’’E, 606 m, 26.XI.2001, fl.,Ran - drianjanaka et al. 654 (CNARP, G, MO, NY, P, TAN, TEF); Maroseranana, bord d’Iam- panga, 18°23’53’’S 48°49’50’’E, 554 m, 3.II.2012, fr., Ravelonarivo et al. 4323 (MO, P, Taxonomic treatment of Abrahamia (Anacardiaceae) 127

TAN); Ambalabe, forêt de Vohibe, 19°09’31’’S 48°34’40’’E, 536 m, 19.II.2009, ster., Razanatsima et al. 660 (G, MO, P, TAN); Ambendrantsara, Manakambahiny-Est, [17°46’S 48°39’E], 750-1512 m, 29.III.1949, fr., Réserves Naturelles 1923 (P, TAN); Zahamena PN, Mangabe, 15.II.1952, fr., Réserves Naturelles 3728 (P); Toamasina I, Zahamena PN, Ampasimalaza, [17°42’S 48°54’E], 22.V.1952, fr., Réserves Naturelles 3732 (P [2 sheets]); Ambodiriana, Tamatave, [Betampona RN], [17°56’S 49°17’E], 20.XI.1960, fr., Réserves Naturelles 11656 (P [2 sheets], TEF); Nosy Mangabe, 15°30’S 49°46’E, 0-330 m, 12-23. IV.1988, ster., Schatz et al. 2131 (MO, TAN); ibid. loco, fl.,Schatz et al. 2492 (MO, P, TAN); ibid. loco, 23-28.II.1989, fr., Schatz 2639 (MO, P, TAN); Analamazaotra, [18°56’S 48°26’E], 1000 m, 21.III.1949, fr., Service Forestier 561 (P, TEF); ibid. loco, 3.IV.1957, fr., Service Forestier 3723 (P, TEF); bords d’un cours d’eau entre Amboditavolo et le col d’Antandrokolaka, [15°46’S 49°32’E], 350 m, 13.II.1954, fr., Service Forestier 9015 (MO, P, TEF); Andriantantely, [18°44’S 48°38’E], 200-300 m, 19.X.1954, bud, Service Forestier 12620 (P, TEF); Farankaraina, Maroantsetra, [15°25’S 49°52’E], 20-25 m, 2.IV.1955, fr., Service Forestier 14361 (P [2 sheets], TEF); Fénérive-Est, Ambodiampana, [17°03’S 49°21’E], 100 m, 16.X.1955, bud, Service Forestier 15645 (P, TEF); forêt d’Analamazaotra, [18°56’S 48°26’E], 6.XI.1957, fl.,Service Forestier 18397 (MO, P, TEF); ibid. loco, 1000 m, 22.IX.1966, ster., Service Forestier 26018 (P); canton Manakamba- hiny-Est, Mandrivo Morafeno, [17°46’S 48°39’E], 1000 m, 9.II.1967, fr., Service For- estier 26286 (P, TEF). Prov. Toliara: Commune d’Enaniliha, forêt entre Eminiminy et le pic Trafonaomby, Andohahela PN, parcelle 1, 24°38’03’’S 46°45’56’’E, 445 m, 25.X.2009, fl., Bernard et al. 1474 (MO, P, TAN); Ampitanonoka à Fotsialana, [17°46’S 48°54’E], 1200 m, 19.I.1945, fr., Homolle 2465 (P); Andohahela PN, parcelle 1, NW of Emin- iminy, 24°38’S 46°46’E, 500-1000 m, 6-13.II.1993, fr., Malcomber et al. 2174 (G, K, MO, NY, P, TAN, WAG); Andohahela PN, parcelle 1, 24°37’33’’S 46°45’55’’E, 440 m, 25.X.1995, fl.,Rakotomalaza et al. 471 (G, K, MO, NY, P, TAN); PN Andohahela, parcelle 1, à proximité d’Eminiminy, 24°40’S 46°48’, 200-700 m, 13-25.I.1993, fr., Randriama- mpionona 37 (MO, P, TAN); Commune Emagnobo, Malamamay, [24°36’27’’S 47°00’52’’E], 20.X.2016, fl.,A. Randrianasolo 1665 (G, K, MO, NY, P, TAN); Commune Bevoay, forêt Mahabo, 24°30’00’’S 47°05’14’’E, 144 m, 18.XI.2009, fl. & bud,Razakamalala 4706 (G, MO, P, TAN); Commune Bevoay, Androangabe, forêt d’Ampitanitanony, [24°30’39’’S 47°05’08’’E], 238 m, 12.I.2016, fr., Razanatsima et al. 1442 (MO, P, TAN). 128 Boissiera 71

31. Abrahamia suarezensis Randrian. & Lowry, spec. nova (Fig. 28-29).

Typus: Madagascar. Prov. Antsiranana: Rég. Diana, NW, près de Ramena, 12°14’S 49°21’E, 9.XI.1961, Service Forestier 20083 (holo-: P [P06169101]!; iso-: G!, MO-6684137!, NY!, P [P00722667]!, TEF [TEF000887]!).

Diagnosis Abrahamia suarezensis Randrian. & Lowry can easily be distinguished from other members of the genus by its small, narrow oblanceolate leaves (2.5-6.5 3 0.7-2.5 cm) with ascending secondary veins that form an acute angle with the midvein at the leaf base.

Description Shrubs or trees 1.5-20 m tall, 6-40 cm DBH; young twigs waxy, lenticellate and pubescent at the very tip. Leaves opposite to subopposite; blades oblanceolate to obovate, 2.5-7.5 3 0.7-3.5 cm, subcoriaceous to chartaceous, apex rounded, retuse, or emarginate, margin entire, undulate, base acute to cuneate, adaxial surface glabrous, abaxial surface with sparse sericeous indument when young, glabrescent, venation craspedodromous, midvein slightly prominent abaxially, secondary veins 9-12 pairs, 1-5 mm apart, not very prominent abaxially but well defined, tertiary veins obscure on both surfaces; petiole 4-10 mm long, canaliculate on adaxial surface, sericeous when young, glabrescent. Inflorescence axillary or terminal, a panicle (sometimes depauper- ate), 1.5-3.5 cm long, the axes flattened, sericeous.Male and female flowers with a pedicel 0.5-1 mm long, pubescent; calyx lobes very widely ovate, c. 1 3 1 mm, glabrous on both surface; corolla lobes ovate, 2.5-3 3 2 mm, imbricate, glabrous on both surfaces, yellow green or light yellow; stamens 4 or 5. Male flowers 5-merous, filaments 1.5-2 mm long, glabrous, white, anthers 1 mm long, obloid to slightly ovoid, yellow, glabrous; disk 1 mm in diam., glabrous; ovary rudimentary. Female flowers 4-merous, filaments c. 1.5 mm long, glabrous, white, anthers c. 0.8 mm long, ovoid, sterile, yellow, glabrous; disk annular, cupuliform, c. 1.5 mm in diam., glabrous; ovary very widely ovoid, 1 3 1 mm, glabrous, placentation apical or slightly subapical; style shortly trifid, latero-subapical or apical, 0.7-0.8 mm long, glabrous, stigmas capitate. Fruits 1.8-2.7 3 1-1.5 cm, ellipsoid, slightly asymmetrical, glabrous, with longitudinal striations.

Distribution, ecology Abrahamia suarezensis is known only from far northern and northeastern Mada- and phenology gascar, from the vicinity of Antsiranana to the area around Daraina and extending along the east coast to S of Vohémar (Map 8). It appears to occur primarily in coastal sandy dune forests although several collections have been made in dry forest on other substrates in the Daraina area (i.e., to the W of Vohémar). Flowering material has been collected in October and November, and fruits are known from November and December. Taxonomic treatment of Abrahamia (Anacardiaceae) 129

4 cm

D 1 cm

C 1 mm

1 mm

A B

Fig. 28. Abrahamia suarezensis Randrian. & Lowry. A. Detail of male flower; B. Detail of female flower; C. Flowering branch; D. Leaf (adaxial surface).

[Service Forestier 20083, P] Drawings: R.L. Andriamiarisoa 130 Boissiera 71

Fig. 29. Abrahamia suarezensis Randrian. & Lowry. A. Flowers, from above [Rakotonandrasana et al. 1026]; B. Group of fruits at the end of a branch [Randrianaivo et al. 1131].

Photos: R. Randrianaivo Taxonomic treatment of Abrahamia (Anacardiaceae) 131

Conservation status With an EOO of 9,014 km2, a minimum AOO of 116 km2, and 35 recorded localities, with subpopulations in 3 protected areas (Loky Manambato, Oronjia, and Montagne des Français), representing c. 23 locations, A. suarezensis is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Crite- ria (IUCN, 2012).

Notes Abrahamia suarezensis most closely resembles A. patrickii, from which it differs by having leaves with fewer secondary veins (9-12 vs 24-28) that are widely (vs closely) spaced, and shorter petioles (4-10 vs 6-13 mm). Typical material of this species from dunes in the area around Antsiranana has narrowly obovate to oblanceolate leaves up to 5 cm long, but collections from farther South near Daraina and Mahavanona have more broadly obovate leaves that can reach 5-7 cm in length, as well as slightly larger fruits. While these differences appear to show some geographic structuring, we have chosen to refrain from recognizing the larger-leaved material from the South as a distinct taxon because no clear limits can be seen between the forms and certain specimens bear leaves spanning the range of variation in both shape and size.

Paratypi Madagascar. Prov. Antsiranana: Vohémar, Ampisikina, Misoribe, 12°58’10’’S 49°34’58’’E, 179 m, 28.IX.2013, fl.,Bernard 2170 (MO, P, TAN); Antsiranana II, Ramena, Orangea, Mamelon vert, 12°14’38’’S 49°22’04’’E, 39 m, 17.XII.2008, fr., Claude Chris- tian et al. 14 (MO, P, TAN); Antsiranana, Ramena, forêt d’Orangea, 12°15’S 49°23’30’’E, 24.I.2002, fr., De Block et al. 1378 (BR, G, K, MO, P, TAN); Vohémar, Daraina, forêt d’Ambohitsitondroina, 13°08’S 49°27’E, 180 m, 20.III.2004, fr., Gautier et al. 4655 (G, MO, TAN); Vohémar, Ampasiramena, 12°50’34’’S 49°49’44’’E, 7 m, 12.XI.2005, fl.,Jul - let Be et al. 260 (CNARP, MO, P, TAN); Daraina, Ambilondambo, 13°09’55’’S 49°38’52’’E, 292 m, 17.XI.2005, fl.,Jullet Be et al. 294 (CNARP, MO, P, TAN); Nosibe, forêt d’Anal- abe, Andavakabe, 13°03’49’’S 49°54’02’’E, 25 m, 4.XII.2004, y. fr., Manjakahery et al. 58 (MO, P, TAN); ibid. loco, 13°04’06’’S 49°54’24’’E, 29 m, 8.XII.2004, y. fr., Manjakahery et al. 76 (MO, P, TAN); ibid. loco, 13°04’20’’S 49°54’04’’E, 18 m, 15.XII.2004, y. fr., Man- jakahery et al. 94 (MO, P, TAN); Sakalava Bay, 12°14’S 49°22’E, 20 m, 26.XI.1989, fl., McPherson 14563 (MO, P, TAN); Antsahalalina, 12 km E of Daraina, 14°07’S 49°46’E, 205-607 m, 24.II.1990, fr., Meyers 45 (MO, P, TAN); Analamazava, SW of Daraina, [14°73’S 49°38’E], 200-1180 m, 23.III.1990, ster., Meyers 73 (MO, TAN); ibid. loco, 26.IV.1990, ster., Meyers 93 (MO, P, TAN); ibid. loco, 24.XI.1990, fl.,Meyers 204 (MO, P, TAN); ibid. loco, 19.II.1991, fr., Meyers 273 (P, MO, TAN); Manatamena, 7 km NE of Daraina, 14°60’S 49°42’E, 112-330 m, IV.1990, ster., Meyers et al. 106 (MO, P, TAN); Daraina, forêt d’Ampondrabe, 12°58’S 49°42’E, 199 m, 15.II.2005, fr., Nusbaumer et al. 1474 (G, MO, TAN); Daraina, forêt d’Ambohitsitondroina, 13°08’S 49°27’E, 272 m, 20.I.2006, fr., Nusbaumer et al. 1966 (G, MO, TAN); Ramena, forêt de l’Orangea, 12°15’13’’S 49°21’44’’E, 13-25 m, 3.IX.2001, fl.,Rabenantoandro et al. 627 (MO, P, TAN); forêt d’Anaborano, près du Lac Sahaka, 13°04’42’’S 49°54’13’’E, 25 m, 2.XI.2002, fl., Rabenantoandro et al. 1095 (G, K, MO, P, TAN, TEF); Tsarabaria, forêt d’Ambondrobe, 132 Boissiera 71

13°43’05’’S 50°05’’52’’E, 13 m, 3.III.2003, fr., Rabenantoandro et al. 1412 (G, MO, P, TEF); Daraina, E du village d’Ambondrombe, 13°41’48’’S 50°05’18’’E, 14.III.2004, fr., Rabevohitra et al. 5149 (G, K, MO, P, TEF [2 sheets]); forêt d’Orangea, 12°15’S 49°20’E, 3.X.2006, fl.,Rakotonandrasana et al. 1026 (CNARP, MO, P, TAN); Ambolobozobe, Ambovomavo, 12°31’13’’S 49°31’21’’E, 24.XI.2007, fr., Rakotonandrasana et al. 1208 (CNARP, G, MO, P, TAN); Mahavanona, forêt d’ Analabe, 12°24’46’’S 49°22’25’’E, 219 m, 15.II.2005, fr., Rakotondrafara 295 (CNARP, G, MO, P, TAN); Andrafiabe, Anjialava, 12°26’57’’S 49°30’48’’E, 2 m, 13.XII.2007, fr., Rakotondrafara et al. 519 (CNARP, G, MO, P, TAN); forêt E de Diego-Suarez, [12°16’55’’S 49°17’40’’E], X.1975, fl.,Rakotozafy 1529 (MO, TAN); Baie de Sakalava, 12°16’39’’S 49°22’49’’E, 21 m, 5.XI.2006, fl.,Rana - ivojaona 1480 (G, MO, P, TAN); Orangea, 12°14’12’’S 49°22’08’’E, 33 m, 7.XI.2006, fr., Ranaivojaona 1571 (G, MO, P, TAN); Ankorangana, forêt d’Andramadana, 12°39’18’’S 49°32’11’’E, 15 m, 22.XI.2001, fr., Randrianaivo et al. 803 (MO, P, TAN); forêt d’Oran- gea, 12°15’01’’S 49°21’40’’E, 76 m, 8.XII.2004, fr., Randrianaivo et al. 1131 (G, MO, P, TAN); Daraina, forêt de Bakaraoka Sud, 13°11’13’’S 49°42’40’’E, 191 m, 15.XI.2005, fl., S. Randrianasolo et al. 551 (CNARP, MO, P, TAN); Daraina, forêt d’Ambilondomba, [13°10’S 49°39’E], 471 m, 27.I.2004, fr., Ranirison et al. 326 (G, MO, TAN); Daraina, Befarafara, Solanampilaka, 13°05’56’’S 49°35’29’’E, 110 m, 16.XI.2005, fl.,Ratovoson et al. 1109 (CNARP, G, MO, P, TAN); fourrée d’Orangea, 12°16’59’’S 49°23’03’’E, 46 m, 30.XI.2005, fl.,Razafitsalama et al. 868 (CNARP, G, MO, P, TAN); Vohémar, Tsarabaria, forêt E du village Ambondrombe, 13°43’08’’S 50°05’50’’E, 13.III.2004, fl.,Razakamalala et al. 1006 (G, K, MO, P, TAN); Near village of Ankorikakely, 2-3 km NE Sakalava bay, 12°16’40’’S 49°23’05’’E, 48 m, 4.XI.2006, fr., Rogers et al. 1089 (MO, P, TAN); Oran- gea, Ramena, 12°14’13’’S 49°22’08’’E, 10 m, 12.II.2005, fr., Schatz et al. 4190 (MO, P, TAN); Vohémar, 27.XII.1951, ster., Service Forestier 4348 (P); forêt d’Andranotsimaka, [12°23’00’’S 49°07’40’’E], 4.X.1952, fl. & fr.,Service Forestier 5788 (MO, P, TEF); forêt d’Orangea, [12°16’55’’S 49°17’40’’E], 18.X.1954, fl.,Service Forestier 11337 (G, MO, NY, P [2 sheets], TEF); Analabe, Vohémar, [14°03’S 49°43’E], 13.IX.1955, fl.,Service Forestier 15498 (G, MO, NY, P, TEF); dunes littorales, près de Ramena, [12°14’S 49°21’E], 9.XI.1961, fl.,Service Forestier 20368 (G, K, MO, MY, P [2 sheets], TEF); ibid. loco, Service Forestier 20369 (MO, P, TEF); forêt littorale, S de Vohémar, [13°21’S 50°00’E], 11-19.XII.1966, bud & fl.,Service Forestier 27279 (MO, P, TEF); ibid. loco, fr., Service Forestier 27280 (P); versant E du massif de l’Ankerana, N de Vohémar, [13°17’S 49°51’E], 50-450 m, 17.XII.1966, fl.,Service Forestier 27352 (P, TEF); forêt d’Orangea area, 12°14’S 49°21’E, 55 m, 23.X.1993, fl.,van Nek 1978 (TAN); ibid. loco, 12°14’S 49°22’E, 65 m, 23.X.1993, fl.,van Nek 1984 (G, MO, P, TAN). Taxonomic treatment of Abrahamia (Anacardiaceae) 133

32. Abrahamia thouvenotii (Lecomte) Randrian. & Lowry, comb. nova (Fig. 30).

≡ Protorhus thouvenotii Lecomte, Madagascar Bois Analamazaotra: 73. 1922.

Lectotypus (designated here): Madagascar. Prov. Toamasina: Rég. Alaotra-Mangoro, Moramanga, Forêt d’Analamazaotra, [18°56’S 48°26’E], 1000 m, s.d., fr., Thouvenot 80 (P [P00580346]!; isolecto-: MO-6684138!, P [P00580347]!, TAN [TAN000076]!).

Description Trees 5-20 m tall, 5-54 cm DBH, bark with white milky latex; young twigs ferruginous pubescent. Leaves alternate; blades oblanceolate to obovate or elliptic to narrowly elliptic or narrowly oblong, 3-15 3 1.5-5 cm, coriaceous, apex emarginate, retuse, or shortly acuminate, sometimes truncate, margin entire, base acute to cuneate or obtuse, adaxial surface ferruginous pubescent when young, later glabrescent or pubescent only on the midvein, abaxial surface pubescent, venation craspedodromous, midvein prominent abaxially, secondary veins 10-20 pairs, arcuate, 2-10(-15) mm apart, prominent on both surfaces, tertiary veins raised on both surfaces; petiole 5-15 mm long, canaliculate on adaxial surface, pubescent, sometimes glabrescent. Inflorescence axillary or terminal, a panicle, 3-8 cm long, axes terete, tomentose. Male and female flowers5-merous; pedicel 0-0.3 mm long, pubescent; calyx lobes deltate to ovate, 1.2-1.5 3 1-1.5 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent, persistent on fruit; corolla lobes ovate, 2-3 3 1-1.2 mm, imbricate, adaxial surface pale yellow, glabrous, abaxial surface brown, pubescent; stamens 5. Male flowers with filaments 1 mm long, glabrous, white, anthers 0.6-0.8 mm long, ovoid, yellow, glabrous; disk 1-1.2 mm in diam., pubescent; ovary rudimentary. Female flowers with filaments c. 0.8-1 mm long, glabrous, white, anthers c. 0.5-0.6 mm long, subglobose or ovoid, sterile, yellow, glabrous; disk 1.5-2 mm in diam., pubescent; ovary widely to very widely ovoid, c. 1-1.2 3 1 mm, pubescent, placentation subapical; style very shortly trifid, apical, 0.3-0.4 mm long, covered with indument.Fruits 1.5-2.5 3 0.8-1.5 cm, ellipsoid, very often glabrate or glabrescent, rarely completely glabrous, with longitudinal striations.

Vernacular names “Ditimena’’, “Menavahatra’’, “Tandraviravy’’ (North East).

Distribution, ecology Abrahamia thouvenotii occurs mainly in eastern Madagascar and the Sambirano and phenology region (Map 10). It is found principally in mid-elevation rainforests at an elevation between 800 and 900 m, although a few subpopulations have been recorded at much higher elevations. Flowers have generally been collected from August to October, although some individuals may flower as early as July. Fruits have been recorded between November and January. 134 Boissiera 71

Conservation status With an EOO of 116,472 km2, a minimum AOO of 180 km2, and 46 recorded localities, with subpopulations in at least 10 protected areas (Anjanaharibe-Sud, Makira, Manon- garivo, Mantadia, Marojejy, Masoala, Montagne d’Ambre, Analamazaotra, Ranomafana, and Zahamena), A. thouvenotii is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia thouvenotii can be distinguished from other members of the genus by the presence of widely spaced secondary leaf venation and indument on the ovary and disk. Three collections from northern and northeastern Madagascar (Miller 3625, Service Forestier 15067 and Ursch 195) differ from typical material by having flowers with a glabrous disk. Additional collections from this area, especially from female individuals, will be needed, however, to determine whether they may represent a distinct taxon. Until such time, we have chosen to include them in our circumscription of A. thouvenotii.

Perrier de la Bâthie (1944) cited two collections in the protologue of Protorhus thouvenotii, one in bud (Thouvenot 7) and the other in fruit (Thouvenot 80). We have designated a sheet of Thouvenot 80 as the lectotype because it is represented by two additional specimens, whereas there is only a single sheet of Thouvenot 7. Protorhus lecomtei (≡ Abrahamia lecomtei) was based on two collections that belong to different taxa, one of which (Perrier de la Bâthie 5283) we have placed in A. thouvenotii. As indicated above under A. lecomtei, we have designated a specimen of the other syntype gathering, Perrier de la Bâthie 14301, as the lectotype in order to retain a circumscription that includes the additional material assigned by Perrier de la Bâthie (1944) to Protorhus lecomtei.

Additional material Madagascar. Prov. Antananarivo: Anjozorobe, 2 km NNE d’Andranomay, 18°28’08’’S examined 47°57’03’’E, 1340-1400 m, 20.XII.1996, fr., Birkinshaw et al. 373 (G, K, MO, P, TAN). Prov. Antsiranana: Ambilobe, forêt de Kalabenono, 13°38’43’’S 48°40’41’’E, 971 m, 27.XI.2006, fr., Callmander et al. 643 (G, NY, MO, P, TAN); Marojejy PN, versant S de Beondroka, 14°25’S 49°50’E, 600 m, 14.XI.1989, bud, Deroin & Badré 96 (MO, P, TAN); Ambanja, Antsatrotro (Manongarivo), [14°05’S 48°23’E], 800 m, XI.1954, fr., Service Forestier 11505 (P [3 sheets], TEF); Andapa, SW edge of Anjanaharibe-Sud Reserve, 14°48’15’’S 49°26’45’’E, 1000-1100 m, 6.VIII.1997, bud, McPherson 17266 (G, K, MO, P, TAN); Marojejy PN, NW Manantenina, Campement n° 3, 14°26.2’S 49°44.5’E, 1200 m, 1.XI.1996, fl.,Messmer et al. 305 (G, MO, P, TAN); Montagne d’Ambre, 12°32’S 49°10’E, 1050 m, 21.X.1988, fl.,Miller 3625 (MO [2 sheets], P, TAN); Ambilobe, Anta- nambao Belinta village, Kalabinono, 13°39’22’’S 48°40’11’’E, 1004 m, 3.XI.2007, fr., Rakotovao 3897 (G, MO, P, TAN); Marojejy PN, env. d’Andohan’Antsahamberokely et Marojejybe, 14°25’45’’S 49°40’20’’E, 1672 m, 7.III.1994, fr., Rasoavimbahoaka 176 (MO, P, TAN); Andapa-Bealanana, Befingotra, 14°12’S 49°27’E, 1100 m, 13-18.IX.1996, fl., Ravelonarivo et al. 1029 (G, K, MO, P, TAN); Marojejy PN, between Campsite II and Campsite III, 14°25’09’’-14°25’24’’S 49°44’46’’-49°45’29’’E, 1280-1350 m, 23.XI.2003, fl.,Schmidt et al. 4308 (G, K, MO, P, TAN); Antalaha, Ambodigavo-Maromandia, [14°51’S 50°07’E], 120 m, 11.IX.1955, bud, Service Forestier 15067 (P [2 sheets], TEF); forêt Taxonomic treatment of Abrahamia (Anacardiaceae) 135

Fig. 30. Abrahamia thouvenotii (Lecomte) Randrian. & Lowry. Tip of a branch showing lower surface of leaves and fruits [Callmander et al. 643].

Photo: M. Callmander 136 Boissiera 71

d’Ambre, [12°36’S 49°09’E], 16.VI.1954, ster., Service Forestier 160-R-152 (P); Diego Suarez, [12°16’S 49°17’E], 17.X.1927, bud, Ursch 195 (P). Prov. Fianarantsoa: Ifanadi- ana, 7 km W of Ranomafana, 21°16’S 47°25’E, 1000 m, 19.X.1987, ster., Daniels 95 (MO, P, UMO); Ranomafana PN, parcelle 3, Talatakely, Piste C, [21°15’S 47°27’E], 1645 m, 3-8.IX.1993, fl.,Kotozafy et al. 230 (G, K, MO, NY, P, TAN, WAG); Ranomafana PN, Vatoharanana, Parcelle 3, 21°16’S 47°26’E, 900-1100 m, ster., Kotozafy et al. 544 (MO, P, TAN); Ranomafana PN, parcelle 3, 21°16’S 47°26’E, 980 m, 16-19.II.1994, ster., Rakoto et al. 514 (MO, P, TAN); Ranomafana PN, Talatakely area, 21°16’S 47°25’E, 800- 900 m, 8.X.1996, ster., A. Randrianasolo 497 (MO, P, TAN); Ranomafana NP, parcelle 3, 21°15’30’’S 47°25’E, 900-1100 m, 23.IX.1994, fl.,J. Randrianasolo et al. 154 (MO, P, TAN); ibid. loco, J. Randrianasolo et al. 155 (MO, P, TAN); ibid. loco, J. Randrianasolo et al. 157 (MO, P, TAN); Andrambovato, E de Fianarantsoa, [21°31’S 47°25’E], 878-1000 m, 24-25.I.1955, ster., Service Forestier 11587 (P, TEF); ibid. loco, Service Forestier 13260 (P); Ranomafana PN, Talatakely trail system, 21°15’30’’S 47°25’00’’E, 950-1150 m, 23. IX.1993, fl.,Solo & J. Randrianasolo 8 (MO, P, TAN); ibid. loco, 16.VIII.1993, fl.,Turk et al. 527 (G, K, MO, NY, P, TAN, WAG); Ranomafana PN, parcelle I, [21°14’S 14°27’E], 850 m, 29.IX.1994, fl.,Turk et al. 637 (MO, P, TAN). Prov. Mahajanga: Anjanahari- be-Sud RS, [14°42’S 49°27’E], 1100 m, 14.XII.1994, fr., Ravelonarivo et al. 552 (MO, P, TAN). Prov. Toamasina: Vavatenina, Zahamena PN, 17°38’27’’S 48°52’32’’E, 700-1040 m, 4.X.2001, fl.,Andrianjafy et al. 194 (CNARP, MO, P, TAN); Masoala PN, Ambohitsi- tondroina, 15°25’37’’S 45°57’24’’E, 1158 m, 10.X.2003, bud, Antilahimena et al. 2169 (G, MO, P, TAN); Commune Vinanibe, Makira forest, 15°27’40’’S 49°17’46’’E, 1153 m, 28.XI.2003, fl.,Antilahimena 2421 (G, K, MO, NY, P, TAN); Maroantsetra, Ambatofotsy, Andampy forest, 15°19’03’’S 49°24’06’’E, 800 m, 4.IX.2004, fr., Antilahimena 2746 (G, MO, P, TAN); Ambatovy, Sahavarina forest, 18°51’37’’S 48°20’25’’E, 1020 m, 24. XI.2005, fr., Antilahimena et al. 4230 (G, MO, P, TAN); Ambatovy, Ampangadiantran- draka Forest, 18°51’43’’S 48°17’39’’E, 1049 m, 9.XII.2006, fr., Antilahimena & Edmond 5087 (G, MO, P, TAN); forêt d’Analamay, 18°49’37’’S 48°19’11’’E, 1153 m, 6.VI.2007, ster., Bernard et al. 529 (MO, P, TAN); Mantadia PN, 700 m N from point PK 13, 18°48’09’’S 48°25’39’’E, 870-930 m, 23.VI.1996, y. fl.,Birkinshaw et al. 313 (MO, P, TAN); moyenne vallée du Mangoro, [18°53’S 48°07’E], 10.IX.1942, fl.,Decary 18516 (MO, P); Analamazaotra, [18°56’S 48°26’E], X.1925, bud, Louvel 44 (P); ibid. loco, 1925, bud, Louvel 112 (P); Mantadia PN, 10.I.2003, ster., Pell et al. 633 (MO, NY, P, TAN); forêt d’Analamazaotra, [18°56’S 48°26’E], 800 m, s.d., fr., Perrier de la Bâthie 5283 (P [4 sheets]); Ambatondrazaka, Commune Didy, Fokontany Antsevabe, 17°59’S 48°36’E, 1165 m, 28.IX.2005, fl.,Rakotonasolo 1066 (G, K, MO, P, TAN); Amparafaravola, E d’Ambodisaina, 17°16’49’’S 48°49’18’’E, 850 m, 11.XII.2001, fr., Rakotondrafara et al. 102 (CNARP, MO, P, TAN); 1.5 km E du campement du PK 14, piste de Mantadia PN, 18°49’S 48°26’E, 940-1012 m, 21.IX.1996, fl.,Ralimanana et al. 10 (G, MO, P, TAN); Lakato, village Agnalatsara, 19°07’29’’S 48°23’32’’E, 815 m, 1.VI.2007, ster., Ranaivoja- ona et al. 1724 (MO, P, TAN); Analamazaotra RS, [18°56’S 48°26’E], 950 m, 15.VII.1996, ster., A. Randrianasolo 413 (MO, P, TAN); Analamazaotra RS, along the Indri trail, about 300 m from observation point, 18°56’12’’S 48°25’10’’E, 1000 m, 10.I.2003, ster., Taxonomic treatment of Abrahamia (Anacardiaceae) 137

A. Randrianasolo 779 (MO, TEF); Analamazaotra forestry station, 18°56’S 48°26’E, 950 m, 19.X.1996, fl.,A. Randrianasolo 506 (MO, P, TAN); ibid. loco, 18.VII.1996, ster., A. Randrianasolo 433 (MO, P, TAN); ibid. loco, 16.VII.1996, ster., A. Randrianasolo 414 (MO, P, TAN); Zahamena PN, forêt communautaire d’Antenina, 17°32’32’’S 48°43’20’’E, 1250 m, 6.XII.2002, fr., Randrianjanaka et al. 762 (CNARP, G, MO, NY, P, TAN, TEF); Ambodi- mangavolo, Ambatoharanana, Moango, 17°33’57’’S 48°53’58’’E, 740-1200 m, 25.X.2005, fl.,Ratovoson et al. 336 (CNARP, G, MO, P, TAN, TEF); Ambohibary, Ambatovy, 18°52’28’’S 48°19’19’’E, 1036 m, 12.XII.2011, fr., Ravelonarivo et al. 3972 (G, K, MO, NY, P, TAN); entre Analamay et Ambatovy, 18°49’52’’S 48°18’45’’E, 1192 m, 22.XI.2005, fl.,Razafindraibe et al. 104 (MO, P, TAN); Ambatovy, forêt de cuirasse de Vatovy, 18°51’21’’S 48°18’29’’E, 1104 m, 1.XII.2005, fr., Razanatsoa et al. 194 (MO, NY, P, TAN); Masoala Peninsula, SE of Ambanizana, 15°38’S 49°59’E, 150-170 m, 1.XI.1992, bud, Schatz et al. 3401 (G, MO, P, TAN); Sahamaloto, Périnet, [18°56’S 48°26’E], 1000 m, 11.X.1952, fl.,Service Forestier 6052 (P [2 sheets], TEF); massif de l’Ambohitsiton- droina de Mahalevona, NW de la presqu’île Masoala, [15°25’S 49°58’E], 900 m, 5. XII.1953, bud, Service Forestier 8714 (G, MO, P, TEF); Sahamaloto, Périnet, [19°03’S 48°12’E], 1054 m, 20.IV.1954, y. fr., Service Forestier 10359 (P, TEF); Amparafaravola, forêt d’Andranobe, W de Manohilahy, [17°46’S 47°58’E], 1000-1200 m, 31.VIII.1958, fl., Service Forestier 18772 (MO, P [2 sheets], TEF); vestiges de forêts près du village d’Ankarahara (PK 100-102 Antananarivo-Moramanga), Mangoro, [18°54’S 48°09’E], 800 m, 8.VIII.1961, bud, Service Forestier 20337 (MO, P [3 sheets], TEF); Analamazaotra, [18°56’S 48°26’E], 1000 m, 21.V.1962, ster., Service Forestier 21142 (MO, P); ibid. loco, 25.V.1962, ster., Service Forestier 21143 (MO, P); Andasitsimanga, Périnet, [18°53’30’’S 48°22’30’’E], 800-1000 m, 2.VIII.1962, bud, Service Forestier 21224 (MO, P); Perinet, [18°56’S 48°26’E], 1000 m, II.1966, ster., Service Forestier 25720 (P); ibid. loco, Service Forestier 25721 (P); ibid. loco, Service Forestier 25722 (P); ibid. loco, 29.VI.1966, ster., Service Forestier 25960 (P); ibid. loco, N. de la pisciculture, III.1966, ster., Service Forestier 25778 (P); Ankazomanitra, PK 45 route Moramanga-Anosibe, [18°56’20’’S 48°13’40’’E], 900 m, 9.XI.1968, y. fr., Service Forestier 26830 (P, TEF); forêt W d’Antanandava (PK 45 de la route Moramanga-Anosibe), [19°17’S 48°12’E], 900 m, XI.1968, fr., Service Forestier 28443 (P); Sahamaloto, Périnet, [19°03’S 48°12’E], 1054 m, 20.VII.1952, bud, Service Forestier 45B-R-172 (P, TEF); ibid. loco, 27.IX.1952, bud, Service Forestier 165B-R- 172 (P, TEF); Ampatsakambe, 28.V.1950, ster., Service Forestier 189B-R-56 (P). 138 Boissiera 71

C B A

2 cm 2 mm 1 cm

Fig. 31. Abrahamia turkii Randrian. & Lowry. A. Flowering branch; B. Ultimate inflorescence unit; C. Leaf (abaxial surface).

[Solo & J. Randrianasolo 35, P] Drawings: R.L. Andriamiarisoa Taxonomic treatment of Abrahamia (Anacardiaceae) 139

33. Abrahamia turkii Randrian. & Lowry, spec. nova (Fig. 31).

Typus: Madagascar. Prov. Fianarantsoa: Rég. Vatovavy-Fitovinany, 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, 20.X.1993, fl.,Solo & J. Randrianasolo 35 (holo-: MO-6671437!; iso-: G, K, MO-6671438!, P [P06168911]!, TAN [TAN002002]!, WAG).

Diagnosis Abrahamia turkii Randrian. & Lowry can be distinguished from other members of the genus by its coriaceous leaves, inflorescence with thick axes and ferruginous indument, and fruits that are small in size (1.8-2 3 1 cm) and glabrous.

Description Trees 11-30 m tall, 18-100 cm DBH, bark with white latex; young twigs sericeous on tips, lenticellate farther from the tip. Leaves usually alternate, sometimes subopposite; blade obovate, 2.5-8 3 1.5-3.5 cm, coriaceous, apex usually emarginate or retuse, sometimes rounded, margin slightly revolute, sometimes undulate, base cuneate, adaxial surface sparsely pubescent, abaxial surface sericeous or strigose, venation craspedodromous, midvein prominent abaxially, secondary veins 14-24 pairs, 1-4 mm apart, raised on both surfaces, tertiary veins visible on both surfaces; petiole 0.5-1 cm long, 1-1.5 mm in diam., pubescent or scabrous, canaliculate on adaxial surface. Inflorescence axillary or terminal, a panicle, erect, axes thick, 1-5 cm long, reddish brown pubescent. Male flowers 5-merous, c. 3-4 mm long; pedicel c. 0.3-0.5 mm long, reddish brown pubescent; calyx lobes triangular, 1.5-2 3 1-2 mm, adaxial surface glabrous, abaxial surface, pubescent; corolla lobes ovate, 2-4 3 2 mm, adaxial surface glabrous, abaxial surface pubescent; stamens 5, filament 1-2 mm long, glabrous, anthers 1 mm long, ovoid to obloid, glabrous; disk slightly cupuliform, c. 2 mm in diam., glabrous; ovary very rudimentary or absent. Female flowers unknown. Fruits 1.8-2 3 1 cm, ovoid, glabrous, wrinkled when dry (longitudinal striations not as evident as in other species).

Distribution, ecology Abrahamia turkii is distributed mainly in south-central and southern Madagascar, and phenology although one collection has been made at the Ambohitantely forest, far to the north of the other recorded subpopulations (Map 10). Flowering material has been recorded in October and November, and fruits have been collected in January.

Conservation status With an EOO of 85,992 km2, a minimum AOO of 40 km2 but an estimated value well exceeding 2,000 km2, and 9 localities, with subpopulations in 6 protected areas (Ambo- hitantely, Andohahela, Betampona, Kalambatritra, Pic d’Ivohibe, and Ranomafana), representing 9 locations, only some of which may be subjected to pressure from forest exploitation and land clearing, A. turkii is assigned a preliminary conservation status of “Least Concern” [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012). 140 Boissiera 71

Notes Abrahamia turkii closely resembles A. ditimena by its medium sized, coriaceous leaves with craspedodromous venation, usually emarginate or retuse apex, and tertiary venation that is visible on both surfaces, and its paniculate inflorescence, but it can easily be distinguished by its short (1-5 vs 2-10 cm) inflorescence covered with reddish brown pubescence (vs glabrescent or tomentose), and its glabrous (vs velutinous) fruits.

Paratypi Madagascar. Prov. Antananarivo: forêt d’Ambohitantely, 18 km NW of Ankazobe, [18°10’S 47°18’E], 1400-1646 m, 21.XI.1985, bud, Barnett et al. 533 (MO, P, TAN). Prov. Fianarantsoa: Ivohibe RS, piste II et III, 22°29’00’’S 46°58’06’’E, 1275 m, 20.X.1997, fl., Rakotomalaza et al. 1434 (G, MO, P, TAN); Ivohibe RS, 5.5 km SE d’Angondongodona, camp 4, 22°25’18’’S 46°53’54’’E, 1150 m, 4.XI.1997, fl.,Rakotomalaza et al. 1494 (G, K, MO, P, TAN); Haute Matsiatra, Ialatsara forest, 21°03’07’’S 47°12’42’’E, 12.X.2005, fl., Ranarivelo et al. 719 (CAS, MO); Ranomafana PN, parcelle 3, Vatoharana trail system, 21°17’S 47°26’E, 1100-1250 m, 18.XI.1993, fl.,Turk & J. Randrianasolo 585 (G, MO, P, TAN); Andrambovato, E de Fianarantsoa, [21°31’S 47°25’E], 878-1000 m, 24.I.1955, fr., Service Forestier 11590 (G, MO, P, TEF). Prov. Toamasina: RN1 [Betampona], s.d., fl. & y. fr.,Anonymous s.n. (G, MO, P [2 sheets]). Prov. Toliara: Betroka, Kalambatritra RS, forêt d’Analamaro, 23°28’28’’S 46°25’15’’E, 1423 m, 9.I.2004, fl.,Andrianjafy et al. 601 (MO, P, TAN); Andohahela PN, parcelle 1, 13.5 km NW d’Eminiminy, 24°35’S 46°44’1’’E, 1200 m, 15.XI.1995, fl.,Messmer et al. 105 (G, MO). Taxonomic treatment of Abrahamia (Anacardiaceae) 141

34. Abrahamia viguieri (H. Perrier) Randrian. & Lowry, comb. nova.

≡ Protorhus viguieri H. Perrier in Mém. Mus. Natl. Hist. Nat. 18: 262. 1944.

Lectotypus (designated here): Madagascar. Prov. Toamasina: Rég. Alaotra-Mangoro, Moramanga, forêt d’Analamazaotra, [18°56’S 48°26’E], 1000 m, 19.X.1912, fl.,Hum - bert & Viguier 815 (P [P00580348]!; iso-: K, MO-6684877!, P [P00580349, P02141537, P06242027]!).

Description Trees 2-25 m tall, 10-15 cm DBH; young twigs lanate. Leaves alternate, sometimes verticellate; blades oblanceolate, 8-30 3 2-6 cm, chartaceous, apex acuminate, margin slightly revolute, base acute or cuneate, adaxial surface lanate, abaxial surface pubescent or lanate only on midvein, venation craspedodromous, midvein prominent abaxially, secondary veins 30-40 pairs, 2-15 mm apart, prominent abaxially, tertiary veins visible on both surfaces; petiole 5-20 mm long, canaliculate on adaxial surface, lanate. Inflorescence axillary, a panicle, 3-11 cm long, axes terete, villous or lanate. Male and female flowers5-merous, glabrous; pedicel 0.2-0.3 mm long, villous; calyx lobes triangular, 0.5 3 0.5-0.7 mm, imbricate, adaxial surface glabrous, abaxial surface pubescent to lanate, persistent on fruit; corolla lobes ovate, 2 3 1 mm, imbricate or quincuncial, adaxial surface glabrous, abaxial surface pubescent on the longitudinal main axis; stamens 5; disk c. 1-1.2 mm in diam., pubescent. Male flowers with filaments 1.5 mm long, anthers 0.5 mm long, ovoid to subglobose, yellow, glabrous; ovary completely absent. Female flowers with filaments 0.5-0.7 mm long, anthers 0.3- 0.5 mm long, ovoid, sterile, yellow, glabrous; ovary widely depressed ovate, 1-1.2 3 0.8-1 mm, glabrous, placentation subapical, style simple (or with a minute trace of division), apically inserted, 0.5-0.7 mm long, glabrous; stigma 4-lobed, capitate. Fruits 3 3 1.5 cm, ellipsoidal, symmetrical or very slightly asymmetrical, glabrous, with longitudinal striations.

Etymology The species is dedicated to Dan Turk, a forest botanist who has contributed tremendously to our knowledge of the trees of Ranomafana National Park.

Vernacular names “Lambafohala’’, “Natoboka’’, “Sefana’’, “Tsitsebona’’.

Distribution, ecology Abrahamia viguieri ranges from east coast humid littoral forests on sand and laterite and phenology at low elevation to premontane forests along the eastern escarpment at an elevation of up to c. 900 m (Map 9). Material has been collected with flowers in October and November; very young fruits have been recorded in December and mature fruits in late February. 142 Boissiera 71

Conservation status With an EOO of 28,641 km2, a minimum AOO of 60 km2, and 17 localities, with subpopulations in six protected areas (Ambalabe, Analalava, Betampona, Makira, Analamazaotra, and Zahamena), A. viguieri is assigned a preliminary conservation status of “Least Concern’’ [LC] using the IUCN Red List Categories and Criteria (IUCN, 2012).

Notes Abrahamia viguieri is easily distinguished from other members of the genus by the lanate upper surface of its leaves and its male flowers with a pubescent disk and completely lacking even a rudimentary or vestigial ovary.

Perrier de la Bâthie (1944) cited two collections in the protologue of Protorhus viguieri, one with male flowers(Humbert & Viguier 815) and the other with fruits (Perrier de la Bâthie 4387, erroneously cited as 4397). We have selected a specimen of Humbert & Viguier 815 as the lectotype because it is the best preserved among the syntypes and because an isolectotype was available for distribution to MO.

Additional material Madagascar. Prov. Fianarantsoa: Fort Carnot, exploitation Charlier, [21°51’30’’S examined 47°26’30’’E], 23.II.1954, fr., Service Forestier 14652 (MO, P, TEF). Prov. Toamasina: Vato- mandry, Ambalabe, Ambinanindrano II, forêt de Vohibe, Sahamanahirana, 19°09’32’’S 48°34’39’’E, 635 m, 28.X.2012, bud & fl.,Andriamiarinoro 304 (MO, P, TAN); Bricka- ville, Sahatsara, 18°26’23’’S 48°49’40’’E, 698 m, 28.X.2005, fl.,Andriamihajarivo et al. 620 (MO, P, TAN, TEF); Brickaville, Anivorano-Sud, Ankerana, 18°26’03’’S 48°49’53’’E, 575 m, 31.X.2005, bud, Andriantiana 297 (P, MO, TAN); Maroantsetra, Rantabe, Beho- vana village, 15°47’34’’S 49°32’45’’E, 521 m, 31.X.2003, fl.,Antilahimena 2237 (G, K, MO, NY, P, TAN, WAG); Brickaville, Didy à Brickaville, [18°07’S 48°32’E], s.d., fl.,Cours 4823 (MO, P [3 sheets]); Moramanga, station forestière d’Andasibe, [18°56’S 48°26’E], 900 m, 8.XII.1989, bud & fl.,E vrard 11232 (P, TAN); Betampona RN, 17°31’S 49°07’E, 275-650 m, 26.IX.1993, bud, Lewis et al. 619 (G, K, MO [2 sheets], NY, P, TAN, WAG); forêt d’Analamazaotra, [18°55’48’’S 48°25’48’’E], 800 m, s.d., fr., Perrier de la Bâthie 4387 (P); Zahamena PN, NE à partir de la rivière de Moango, 17°33’49’’S 48°54’01’’E, 794 m, 20.X.2000, fl.,Rakotonandrasana et al. 441 (CNARP, MO, P, TAN); Vavatenina, Zahamena PN, Besakay, 17°39’20’’S 48°54’22’’E, 780 m, 7.X.2001, bud, Rakotonandrasana et al. 573 (CNARP, MO, P, TAN); Analamazaotra RS, along the Indri trail, about 300 m from observation point, 18°56’12’’S 48°25’10’’E, 1000 m, 10.I.2003, ster., A. Randrianasolo 776 (MO, TEF); Fotsimavo, canton Tamatave, [17°56’S 49°13’E], 10.XI.1955, bud & fl., Réserves Naturelles 7712 (MO, P, TEF); Ambodiriana, [17°55’S 49°13’E], 100-200 m, 12.IV.1959, bud, Réserves Naturelles 10847 (MO, P); Sahatavy, [Zahamena PN], [17°38’S 48°50’E], 750-1912 m, 18.X.1960, bud, Réserves Naturelles 11360 (MO, P [2 sheets], TEF); Fénérive-Est, forêt d’Analalava, Foulpointe, [17°42’S 49°29’E], 0-50 m, 30.X.1963, bud & fl.,Service Forestier 22781 (P, TEF); Ambatonandrohina, Sahatavy, [17°27’S 49°00’E], 13.X.1966, bud & fl.,Service Forestier 26261 (P, TEF). Taxonomic treatment of Abrahamia (Anacardiaceae) 143

Acknowledgments e wish to thank the botanists of the Missouri Botanical Garden’s Madagascar W Program for their assistance in collecting specimens of Abrahamia, in particular Fortunat Rakotoarivony, Nivo Rakotoarivelo, Aina Razanatsima, Tefy Andriamihajarivo, Tabita Randrianarivony and Monique Randriatsivery, as well as Ny Lafatra Ravalomana- rivo for specimen and data management. The first author would like to express his special thanks to Zachary Rogers and Peter Phillipson for their valuable assistance and advice on a wide variety of subjects relating to this project. We are grateful to Roger Lala Andriamiarisoa and Barbara Alongi for the beautiful illustrations. We also wish to thank the staff at the Herbier National at the Muséum National d’Histoire Naturelle in Paris for their hospitality and assistance during the first author’s visits, and we are grateful to the staff of the two national herbaria in Madagascar (TAN and TEF) for allowing us to examine their specimens. We express thanks to John Mitchell, Susan Pell, and Martin Callmander for valuable comments and suggestions on the manuscript that resulted in its improvement. We are also grateful to Patrick Perret for his attentive reading of the proofs and Laurent Gautier for his assistance with the French identification key. Finally we thank Matthieu Berthod for the expertly done layout of this volume. Field work was conducted under the collaborative agreements between the Missouri Botan- ical Garden and the Parc Botanique et Zoologique de Tsimbazaza, and the Ministère de l’Environnement, de l’Ecologie, de la Mer et des Forêts. We gratefully acknowledge courtesies extended by the Government of Madagascar and by Madagascar National Parks. This research was conducted with support of the National Geographic Society (grant no. 8818-10 and 9401-13 to AR), the William L. Brown Center of the Missouri Botanical Garden, and the U.S. National Science Foundation (grant no. DEB-0743355 to GES and PPL). 144 Boissiera 71

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Index Abrahamia Randrian. & Lowry...... 13 Abrahamia antongilensis Randrian. & Lowry...... 31

Abrahamia betamponensis Randrian. & Lowry...... 34

Abrahamia buxifolia (H. Perrier) Randrian. & Lowry...... 40

Abrahamia capuronii Randrian. & Lowry...... 43

Abrahamia darainensis Randrian. & Lowry...... 47

Abrahamia deflexa (H. Perrier) Randrian. & Lowry...... 48

Abrahamia delphinensis Randrian. & Lowry...... 51

Abrahamia ditimena (H. Perrier) Randrian. & Lowry...... 53

Abrahamia ellipticarpa Randrian. & Lowry ...... 60

Abrahamia elongata Randrian. & Lowry...... 62

Abrahamia grandidieri (Engl.) Randrian. & Lowry...... 67

Abrahamia humbertii (H. Perrier) Randrian. & Lowry...... 71

Abrahamia ibityensis (H. Perrier) Randrian. & Lowry...... 74

Abrahamia itremoensis Randrian. & Lowry ...... 76

Abrahamia latifolia (Engl.) Randrian. & Lowry...... 78

Abrahamia lecomtei (H. Perrier) Randrian. & Lowry...... 84

Abrahamia lenticellata Randrian. & Lowry...... 86

Abrahamia littoralis Randrian. & Lowry...... 90

Abrahamia lokobensis Randrian. & Lowry...... 93

Abrahamia longipetiolata Randrian. & Lowry...... 95

Abrahamia louvelii (H. Perrier) Randrian. & Lowry...... 99

Abrahamia minutifolia Randrian. & Lowry...... 101

Abrahamia nitida (Engl.) Randrian. & Lowry...... 105

Abrahamia oblongifolia (Engl.) Randrian. & Lowry...... 107

Abrahamia patrickii Randrian. & Lowry...... 109

Abrahamia pauciflora (Engl.) Randrian. & Lowry...... 111

Abrahamia phillipsonii Randrian. & Lowry...... 113

Abrahamia revoluta Randrian. & Lowry ...... 117 Taxonomic treatment of Abrahamia (Anacardiaceae) 149

Abrahamia sambiranensis Randrian. & Lowry...... 119

Abrahamia sericea (Engl.) Randrian. & Lowry...... 121

Abrahamia suarezensis Randrian. & Lowry...... 128

Abrahamia thouvenotii (Lecomte) Randrian. & Lowry...... 133

Abrahamia turkii Randrian. & Lowry...... 139

Abrahamia viguieri (H. Perrier) Randrian. & Lowry...... 141

Protorhus buxifolia H. Perrier...... 40

Protorhus deflexa H. Perrier...... 48

Protorhus ditimena H. Perrier...... 53

Protorhus ditimena var. minutifolia H. Perrier...... 101

Protorhus fulva Engl...... 53

Protorhus grandidieri Engl...... 67

Protorhus heckelii Dubard & Dop ...... 67

Protorhus humbertii H. Perrier...... 71

Protorhus ibityensis H. Perrier...... 74

Protorhus latifolia Engl...... 78

Protorhus lecomtei H. Perrier ...... 84

Protorhus louvelii H. Perrier...... 99

Protorhus nitida Engl...... 105

Protorhus oblongifolia Engl...... 107

Protorhus pauciflora Engl...... 111

Protorhus sericea Engl...... 121

Protorhus sericea var. opaca Lecomte ...... 53

Protorhus thouvenotii Lecomte ...... 133

Protorhus viguieri H. Perrier ...... 141

Rhus grandidieri (Engl.) Baill...... 67

Rhus latifolia (Engl.) Baill...... 78

Rhus venulosa Baker...... 67

Cover image Abrahamia lenticellata Randrian. & Lowry based on drawings by Barbara Alongi BOISSIERA 71 / 1-152 (2017) ISSN: 0373-2975