Taxonomic revision of Phyllanthus (Phyllanthaceae) in Madagascar and the Comoro Islands III: subgenera Swartziani, Afroswartziani and Emblica

Summary

The third part of the revision of the genus Phyllanthus L. (Phyllanthaceae) in Madagascar and the Comoro Islands treats the subgenera Swartziani comb. & stat. nov., Afroswartziani subgen. nov. and Emblica. Taxonomic history of the subgenus Phyllanthus is given and its circumscription is revised. Phyllanthus subgenus Afroswartziani subgen. nov. comprises thirteen species. One species P. ankazobensis is described as new to science. P. vatovaviensis and P. madagascariensis var. kalambatitrensis are validly published by this paper. Identification keys, descriptions and species notes (habitat, distribution and threats) are provided. Distribution maps and IUCN conservation assessments are provided for all species and varieties.

Introduction

This paper is the third part of our series of revisions of the genus Phyllanthus L. in Madagascar and the Comoro Islands. In the first part (Ralimanana & Hoffmann 2011) we gave a general synopsis of the subgenera and sections occurring on these islands and dealt with the subgenera Isocladus G. L. Webster, Betsileani (Jean F. Brunel) Ralim. & Petra Hoffm., Kirganelia (A. Juss.) G. L. Webster and Tenellanthus Jean F. Brunel. In the second part (Ralimanana & Hoffmann, submitted) we treated subegenera Anisonemoides (Jean F. Brunel) Ralim. & Petra Hoffm. and Menarda (Müll. Arg.) Ralim. & Petra Hoffm. (= both treated within Bruneliathus Ralim. & Petra Hoffm., subgen. nov. ined. (Ralimanana & Hoffmann 2011)). In the present paper we review the circumscription of subgenus Phyllanthus sensu Webster (1956 – 1958) which has recently been shown to be non-monophyletic in molecular phylogenetic studies (Kathriarachchi et al. 2006). In the first modern systematic treatment of the genus, his monograph of West Indian Phyllanthus, Webster (1956 − 1958) recognised four sections within subgenus Phyllanthus. They are Callitrichoides G. L. Webster, Cyclanthera G. L. Webster, Phyllanthus and Urinaria G. L. Webster. Webster divided section Phyllanthus into three subsections: Niruri G. L. Webster, Swartziani G. L. Webster and Pentaphylli G. L. Webster. He pointed out that subgenus Phyllanthus included the majority of the various herbaceous and undershrub representatives, it was also a heterogenous group and possibily polyphyletic.

When Brunel (1987) revised the subgenus Phyllanthus in tropical Africa, he proposed wide-ranging changes to the classification. He moved the monospecific Cuban section Callitrichoides into a new subgenus Microglochidion (Müll. Arg.) Jean F. Brunel, and transferred the West Indian section Cyclanthera to subgenus Xylophylla (L.) Pers. In subgenus Phyllanthus, Brunel (1987) retained only two of Webster’s sections which are Phyllanthus (containing the type of the genus, neotropical P. niruri L., and one African species P. benguelensis Müll. Arg.) and Urinaria (containing P. urinaria L.). All other species of Webster’s subgenus Phyllanthus were placed in section Anthophyllus Jean F. Brunel which he divided into four subsections: Swartziani, Pentaphylli, and two new subsections Callidisci Jean F. Brunel and Odontadenii Jean F. Brunel & Jacq. Roux. He proposed several other, pr edominantly African sections to be included in subgenus Phyllanthus.

According to recent molecular data (Kathriarachchi et al. 2006), African, Madagascan and some Asian representatives of section Phyllanthus subsection Swartziani sensu Webster, are grouped in a strongly supported clade that is genetically divergent from the clade containing the type species of subsection Swartziani, P. amarus Schumach. & Thonn. and Reverchonia A. Gray. Both groups can be circumscribed by the morphological characters listed below. Based on both the morphological and genetic divergences, we here raise the neotropical subsection Swartziani (typified by the pantropical weed P. amarus) to subgeneric level, and create a new subgenus Afroswartziani Ralim. & Petra Hoffm. subgen. nov. to accomodate the paleotropical species. More African species need to be examined with molecular methods to establish whether other sections that were proposed by Brunel (1987) should be supported, but the inclusion of Phyllanthus hutchinsonianus S. Moore (a member of section Praephyllanthus Jean F. Brunel) in the Afroswartziani clade makes this at least partially doubtful. The herbaceous subgenus Swartziani has mainly bisexual cymules, membranous cataphylls and cataphyllary stipules. In contrast, Afroswartziani species are mostly shrubs or small trees and very rarely herbs but woody at the base. They have unisexual cymules (except very rarely bisexual in P. ampandrandavae Leandri), cataphylls and cataphyllary stipules that are mostly subcoriaceous to coriaceous. Thirteen endemic Madagascan species of subgenus Afroswartziani are treated in this paper. Of those, Phyllanthus lokohensis Leandri and P. madagascariensis Müll. Arg. were included in the molecular phylogenetic analyses of Kathriarachchi et al. (2006). The other members of clades F and G in Kathriarachchi et al. (2006) included in those analyses (all from continental Africa or Asia) were P. debilis Klein ex Willd., P. gossweileri Hutch., P. hutchinsonianus S. Moore, P. kaessneri Hutch., P. mannianus Müll. Arg., P. rheedii Wight and P. sepialis Müll. Arg. The only member of subgenus Swarziani occurring in Madagascar and the Comoro Islands is P. amarus, an introduced weed of American origin.

With 1269 accepted species (Govaerts et al. 2000 ), Phyllanthus s.l. is the largest genus in the Phyllanthaceae. It is known as a taxonomically “difficult” genus due to its complexity and morphological diversity. Circumscription of Phyllanthus has been widely discussed (Wurdack et al. 2004; Kathriarachchi et al. 2006; Pruesapan et al. 2012). Wurdack et al. (2004) demonstrated that Phyllanthus is polyphyletic using the rbcL region. Katriarachchi et al. (2006) was the first study to present a broad molecular analysis of Phyllanthus and its allies and to evaluate the existing classification of Phyllanthus. They found that Breynia J. R. Forst. & G. Forst, Glochidion J. R. Forst. & G. Forst, Reverchonia A. Gray and Sauropus Blume are nested within Phyllanthus and should be combined with it, thus proposing an expansion of the generic concept. Pruesapan et al. (2012) argued that the number of species per genus analysed by Kathriarachchi et al. (2006) was insufficient to support their findings. Pruesapan et al. (2012) support the maintenance of Breynia and Glochidion at generic level and the splitting of Phyllanthus into smaller genera, backing up their argument with a new analysis of nuclear and chloroplast markers.

Lack of molecular data still remains problematic for Madagascan Phyllanthus. Kathriarachchi et al. (2006) have included only two of the 13 species of the subgenus Afroswartziani presented in this paper, and none of the seven Madagascan Gomphidium. This data is insufficient to support the recognition of the subgenera in this paper at generic rank. Therefore we focus on the classification change at infrageneric level.

The last taxon treated here is Phyllanthus subgenus Emblica section Urinaria. This group was classified in subgenus Phyllanthus by both Webster (1956 – 1958) and Brunel (1987). Although the morphology of these pantropical herbaceous weeds (invasive from Asia) otherwise fits relatively well into subgenus Phyllanthus, some morphological characters support one of the most surprising results of Kathriarachchi et al. (2006, see there for more discussion): that P. urinaria is most closely related to a group of Asian tree species that include the well-known P. emblica L. (clade N). We therefore here transfer section Urinaria from subgenus Phyllanthus to subgenus Emblica.

Materials and methods

The revision presented here is based on studies of herbarium material, spirit collections, and field observations. Field work was conducted in different areas of Madagascar. Apart from our own collections, herbarium material of Phyllanthus was consulted at BM, K, LINN, P and the national herbaria in Madagascar (TAN and TEF). The descriptions and measurements of all vegetative and floral characters are mostly based on dried specimens or softened plant parts, supplemented by field observations. Distribution maps and conservation assessments were obtained using GIS (Arcview 3.3) and applying IUCN criteria (IUCN 2001). The floristic domains here referred to are as defined by Humbert (1965).

Taxonomic treatment

I. Phyllanthus subgen. Swartziani (G. L. Webster) Ralim. & Petra Hoffm., comb. & stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77134143-1

Subgenus Phyllanthus section Phyllanthus subsection Swartziani G. L. Webster (1955: 53). Type species: Phyllanthus amarus Schumach. & Thonn.

Monoecious herbs with phyllanthoid (pinnatiform) or non-phyllanthoid (phyllotaxy spiral) branching. Brachyblasts absent. Cataphylls and cataphyllary stipules thinly membranous. Stipules entire or irregularly toothed. Leaves alternate. Inflorescences on leafy branches, axillary, bisexual, 1 – 7-flowered. Tepals (4 – )5 – 6. Disc free glands in male flowers, annular in female flowers (but described as discrete glands for Phyllanthus abnormis Baill. by Johnston & Warnock 1963; Webster 1970). Stamens 2 – 3; filaments entirely or partially fused into a column; anthers entirely or partially fused, dehiscing obliquely to less commonly horizontally; pollen 3-colporate, reticulate (Webster 1956 – 1958; Brunel 1987). Ovary 3-locular, smooth (sometimes slightly rugose in P. amarus); styles free, bifid; stigmas obtuse (somewhat dilated in Reverchonia according to Webster & Miller (1963)). Fruits dehiscent; tepals persistent, not accrescent in fruit. Seeds smooth or finely striate longitudinally.

This description includes the North American Reverchonia arenaria A. Gray, Phyllanthus abnormis and the pantropical weed P. amarus, which are the only taxa so far known to belong definitely to this subgenus (= clade E in Kathriarachchi et al. 2006).

1. Phyllanthus amarus Schumach. & Thonn. (Schumacher 1827: 421).

Phyllanthus niruri L. var. amarus Leandri (1958: 73). Type: Africa, Guinea, Thonning 4 (holotype fragment, K!; C).

Phyllanthus amarus Schumach. & Thonn. var. baronianus Leandri (1939: 184), nom. invalid. Phyllanthus niruri L. var. baronianus (Leandri) Leandri (1958: 74).

Phyllanthus niruroides Müll. Arg. var. madagascariensis Leandri (1939: 185), nom. invalid.

Monoecious herbs 10 – 45 cm tall. Branching pinnatiform; orthotropic branches terete, 1 – 3 mm in diam., striate, glabrous. Cataphylls triangular, 1 – 2 × 0.3 – 0.4 mm, brown yellowish or reddish, thinly membranous, entire, glabrous. Cataphyllary stipules triangular, 1 – 1.8 × 0.3 – 0.8 mm, yellowish brown or reddish, thinly membranous, entire, glabrous. Plagiotropic branches solitary or in fascicles of two, terete, 4 – 12 cm long, 0.4 – 0.5 mm in diam., striate, glabrous. Stipules persistent, triangular or linear, 1 – 1.2 × 0.2 – 0.3 mm, membranous, entire, glabrous, green or reddish when fresh. Leaves 12 – 27 per branch. Internodes 0.7 – 2.8 mm long. Petioles terete, 0.3 – 0.5 × c. 0.1 mm, glabrous. Leaf blades elliptic to oblong, 3 – 11 × 3 – 4 mm, 1 – 1.7 times longer than wide, obtuse or rounded at the base, rounded or obtuse at the apex, membranous, entire, glabrous on both sides, green or reddish green adaxially, light green or greyish green abaxially when fresh; midvein flattened adaxially, prominent abaxially; secondary veins 4 – 5 pairs, indistinct adaxially, flattened abaxially. Inflorescences bisexual, consisting of 1 – 2 male flowers. Bracts linear or triangular, grouped in tufts of 3 – 6 mainly sterile bracts, 0.3 – 0.4 × 0.2 – 0.3 mm, entire, glabrous. Male flowers 0.4 – 0.7 × 1 – 1.2 mm. Pedicels 0.6 – 1.2 × c. 0.2 mm, glabrous. Tepals 5 (very rarely 6), subequal, elliptic or ovate, 0.4 – 0.7 × 0.2 – 0.4 mm, obtuse to acute, membranous, glabrous on both sides, margin entire, hyaline for ¹/₄ of total tepal width on each side, veins unbranched. Disc glands 5 (very rarely 6), globose, c. 0.1 mm in diam., smooth. Stamens 3 (rarely 2), filaments fused into a column; column terete, 0.2 – 0.3 × 0.1 – 0.2 mm; anthers partially or entirely fused, ovoid or elongate (sometimes reduced to one or two functional anthers), c. 0.1 mm long, dehiscing obliquely to less commonly horizontally. Female flowers 0.7 – 1 × 0.8 – 1 mm. Pedicels 1 – 1.3 × 0.1 – 0.3 mm, glabrous. Tepals 5 (or very rarely 6), equal, elliptic to obovate, 0.8 – 1 × 0.3 – 0.6 mm, acute, membranous, glabrous on both sides, margin entire, hyaline for ¹/₄ of total tepal width on each side, veins unbranched. Disc lobed, smooth. Ovary 3-locular, globose, 0.4 – 0.6 × 0.4 – 0.5 mm, smooth or slightly rugose, glabrous; styles free, bifid for c. ¹/₂ of their length, terete, 0.1 – 0.2 mm long; stigmas obtuse. Fruits globose or depressed globose, 1.5 – 2 × 1.5 – 2 mm; fruiting pedicels 1.5 – 2 × 0.1 – 0.3 mm, glabrous; columella 0.6 – 1 mm long. Seeds 1 – 1.5 × 0.8 – 1 mm, finely striate longitudinally.

distribution. This species originated in America, pantropical by introduction; in Madagascar distributed in eastern, central, western and southern floristic domains. Map 1.

Map 1

Distribution of Phyllanthus amarus.

specimens examined. madagascar . s.loc., Baron 1611 (P); Baron 4886 (K, P); Baron 4081 (K); Baron 6715 (P); Chapelier s.n. (P); alt. 1200 m, 1 March 1945, Cours 2701 (P); 11 Aug. 1932, Decary 10268 (P); Herbier du Jardin Botanique de Tananarive 3987 (P); Humblot 96 (P); vers 1882, Lanz s.n. (P); Leandri 27264 (P); Perrier de la Bâthie 9538 (P); Perrier de la Bâthie 9866 (P); Du Petit Thouars s.n. (P). Antananarivo. Manankazo, 2 Jan. 1942, Decary 17093 (P); Ambatolampy, Station forestière de Manjakatompo, Anosiarivo, 19°19'09"S, 47°19'01"E, alt. 1800 m, 26 Jan. 2004, Ralimanana 434 ( K, TAN); Antsirabe, Jan. 1914, Perrier de la Bâthie 9847 (P). Fianarantsoa. Befotaka, Farafangana, 11 Aug. 1926, Decary s.n. (P); Mananjary, s. dat. 1909, Geay 7029 (P); & Mananjary, s. dat. 1909, Geay 7030 (P); Andringitra, 22°7'20"S, 46°55'22"E, alt. 1440 m, 24 March 2004, Ralimanana 524 (K, MO, P, TAN); Ankatafana, forêt d’ Ambahisosotra, 21°29'12"S, 48°16'20"E, 19 June 2004, Ranaivojaona 820 (TAN); Angalapena Betsileo, Scott Elliott 2157 (P). Antsiranana. Nosy be, May 1847, Boivin 2190 (P); Sambava, Bemanevika, 14°07'36"S, 50°09'26"E, alt. 3 m, 7 June 2013, Rakotonasolo et al. 2225 (K, TAN). Mahajanga. Ste Marie de Marovoay, 29 Aug. 1911, Kaudre s.n. (P); Amborovy, 18 May 1965, Peltier & Peltier 5266 (P); Aux environs de Maevatanana, July 1900, Perrier de la Bâthie 274 (P); Parc National d’Ankarafantsika, 16°19'09"S, 46°48'08"E, alt. 120 m, 22 Oct. 2002, Ralimanana et al. 242 (K, TAN); Station forestière Antsanitia, 15°35'20"S, 46°25'56"E, alt. 19 m, 23 Oct. 2002, Ralimanana 250 (G, K, MO, P, TAN). Toamasina. Vicinity of Brickaville, 18°50'S, 49°04'E, alt. 100 m, 1 May 1983, d’Arcy 15253 (MO); Station Agricole de Lac Alaotra, 24 Aug. 1937, Boiteau 2708 (P); Analamazaotra, Oct. 1939, Boiteau 4246 (P); Toamasina, 15 Jan. 1989, Brunel 12670 (P); Aux environs de Toamasina, May 1989, Brunel 12671 (P); Ambila, 29 April 1928, Decary s.n. (P); Ambila, 29 April 1928, Decary 6407 (K); Ambila, alt. 3 – 9 m, 9 Oct. 1946, Cours 2893 (P); Vatomandry, 31 Jan. 1904, Guillot 85 (P); Aux environs de Toamasina, 25 Sept. 1912, Viguier & Humbert 287 (P); Côte Est, Lantz s.n. (P); Brickaville, forêt de Vohibola, sable blanc, 18°33'35"S, 49°15'05"E, alt. 0 – 10 m, 26 Oct. 2002, Ranaivojaona 456 (K, MO, TAN); Mahanoro, Ambalavontaka, forêt littorale d’Ampamanara, 20°22'22"S, 48°33'56"E, alt. 13 m, 14 April 2004, Ranaivojaona 611 (K, MO, TAN); Ambalavontaka, forêt littorale d’Ampamanara, 20°22'22"S, 48°33'56"E, alt. 13 m, 14 April 2004, Ranaivojaona 612 (K, MO, TAN); Fénérive Est, station forestière de Tampolo, 17°17'S, 49°26'E, alt. 0 – 5 m, 8 April 1997, Randrianaivo 85 (MO, P, TAN); Ankirindro, Maroantsetra, 15°19'20"S, 49°33'26"E, alt. 100 m, 2 Feb. 1999, Schatz et al. 3936 (MO, P); Andovoranto, alt. 300 m, 12 Oct. 1912, Viguier & Humbert 667 (P); Andovoranto, Anivorano, 14 Oct. 1912, Viguier & Humbert 708 (P). Toliara. Sud de Toliara, Feb. 1989, Brunel 12515 (P); Tolagnaro, 25 May 1932, Decary 9810 (K, P); Tolagnaro, 11 Aug. 1932, Decary 10268 (P); Tolagnaro, 25 May 1932, Decary 9810 (P); Tolagnaro, 25°2'S, 46°59'E, 11 March 1989, Gereau & Dumetz 3227 (K, MO, P, TAN, TEF); Miandrivazo, Hure s.n. (P); Morondava, Kirindy Nord, 20°4'21"S, 44°40'37"E, alt. 41 m, 24 March 2003, Ralimanana et al. 328 (K, MO, P, TAN); St Augustin, 23°31'19"S, 43°45'4"E, alt. 7 m, 16 Feb. 2004, Ralimanana et al. 478 (K, TAN); Ampanihy, Besavoa, 24°43'41"S, 44°43'0.7"E, alt. 227 m, 18 Feb. 2004, Ralimanana et al. 500 (K, TAN); Tolagnaro, Scott Elliott 2380 (K, P). comores. Mayotte, Coconi, 12°50'6"S, 45°8'12"E, alt. 95 m, 28 Nov. 2000, Labat & Barthelat 3313 (K, P).

habitat in madagascar. Species growing on different types of vegetation on different types of soil; sea level – 1800 m.

II. Phyllanthus subgen. Afroswartziani Ralim. & Petra Hoffm., subgen. nov. Type: Phyllanthus lokohensis Leandri.

http://www.ipni.org/urn:lsid:ipni.org:names:77134144-1

Subgenus Afroswartziani is similar to subgenera Swartziani and Tenellanthus because of the dehiscing fruits, but differs from Swartziani in the habit (shrub or small tree), very rarely herbaceous but even then woody at the base (vs always herbs), the cataphylls and cataphyllary stitpules subcoriaceous to coriaceous (vs membranous); differs from Tenellanthus in (2 –) 3 stamens partially or entirely fused (5 (– 7), entirely free), in anthers dehiscing obliquely or horizontally (vs longitudinally), the pollen 3-coloporate, exine microperforate or scabrous (vs 4-colporate, reticulate) and the seeds smooth or thinly striated (vs papillose or verrucose).

Monoecious or dioecious shrubs, small trees or rarely herbs with phyllanthoid (pinnatiform or bipinnatiform) branching. Brachyblasts absent. Cataphylls and cataphyllary stipules subcoriaceous to coriaceous. Leaves alternate. Stipules entire. Inflorescences on leafy branches, axillary, unisexual or very rarely with bisexual cymules, 1 – 2-flowered, female flowers generally solitary. Tepals 5 – 6. Disc free glands in male flowers, annular lobed or sub-entire in female flowers. Stamens 2 – 3; filaments entirely or partially fused into a column (free in aberrant P. kaessneri (Brunel 1987)); anthers entirely or partially fused or entirely free, dehiscing obliquely or horizontally; pollen 3-colporate, microperforate or scabrous. Ovary 3-locular, smooth to rugose, sometimes on short gynophore; styles free or fused at the base, bifid; stigmas obtuse or acute. Fruits dehiscent; tepals persistent, sometimes accrescent in fruit. Seeds 2 per locule, smooth or finely striate longitudinally.

This description is provisional because it is restricted to the Madagascan representatives and the few continental African and Asian species (Phyllanthus debilis Klein ex Willd., P. gossweileri Hutch., P. hutchinsonianus S. Moore, P. mannianus Müll. Arg., P. rheedii Wight and P. sepialis Müll. Arg.; and maybe P. kaessneri Hutch.) that were shown to belong to this subgenus by Kathriarachchi et al. (2006: clades G and F). Wider sampling for phylogenetic molecular analysis is likely to show that more species belong to subgenus Afroswartziani.

Key to species of Phyllanthus subgenus Afroswartziani

• 1. Annual or perennial herbs, up to 30 cm high.…...........2

• 1’. Shrubs or small trees, more than 30 cm high.…...........4

• 2. Plants sprawling on ground; leaf blades ovate, obovate or suborbicular 1.7 – 3 × 1 – 2 mm; bracts in a tuft of 4 – 6 sterile bracts in male flowers; disc glands rugose to verrucose in male inflorescence; filaments entirely fused in column; anthers entirely fused.…...........2. P. venustulus

• 2’. Plants erect; leaf blades elliptic, lanceolate or oblong, 5 – 8 × 2 – 4 mm; bracts not tufted; disc glands smooth; filaments fused in column for ¹/₄ – ²/₃ of their total length; anthers free.…...........3

• 3. Midvein prominent on both sides; secondary veins indistinct on both sides. Tepals 6 in male flowers; anthers dehiscing obliquely; disc 3-lobed in female flowers; fruits 1 – 1.3 × 1.2 – 2 mm; seeds 0.6 – 1 × 0.4 – 0.6 mm.…...........3. P. ampandrandavae

• 3’. Midvein flattened on both sides; secondary veins 4 – 6 pairs, flattened abaxially, indistinct or flattened adaxially. Tepals 5 in male flowers; anthers dehiscing horizontally; disc 5-lobed in female flowers; fruits 2 – 2.5 × 2.3 – 2.7 mm; seeds 1.6 – 2.3 × 1 – 1.4 mm.…...........4. P. andranovatensis

• 4. Bracts not tufted in male inflorescences; filaments partially united into a column for ¹/₂ – ³/₄ of their total length, anthers ovoid, free and dehiscing obliquely; female flower tepals 4 mm long or more.…...........5

• 4’. Bracts in a tuft in male inflorescences; filaments entirely fused in column, anthers partially or entirely fused in a circumscissile synandrium dehiscing horizontally; female flower tepals 1 – 4 mm long.…...........8

• 5. Branches trailing and flexible, plagiotropic branches glabrous; gynophore 0.3 – 0.4 mm; styles fused for ²/₃ of their length.…...........5 P. ankazobensis

• 5’. Branches erect, plagiotropic branches pubescent or scabridulous; gynophore absent, styles free or only fused at the base.…...........6

• 6. Leaves subcoriaceous to coriaceous; plant of littoral forest.…...........6a. P. madagascariensis var. madagascariensis

• 6’. Leaves membranous or chartaceous; not a littoral forest plant.…...........7

• 7. Tepals 5 in male flower, veins unbranched; disc glands globose; female disc annular lobed; stigmas obtuse.…...........6b. P. madagascariensis var. kalambatitrensis

• 7’. Tepals 5 – 6 in male flower, veins branched; disc glands elliptic transversally; female disc annular subentire; stigmas acute.…...........7. P. melleri

• 8(4). Plagiotropic branches grouped at the distal part of the orthotropic branches; rheophyte. 8. P. rangoloakesis

• 8’. Plagiotropic branches spread regularly along orthotropic branches; not a rheophyte.…...........9

• 9. Plagiotropic branches slightly flattenned; anthers partially fused.…...........9. P. coursii

• 9’. Plagiotropic branches terete; anthers entirely fused into a disciform circumscissile synandrium, unlobed or 2 – 3 lobed.…...........10

• 10. Internodes 0.5 – 1 mm; leaf blades 1.5 – 3 mm wide, asymmetric at the base.…...........10. P. ivohibeus

• 10’. Internodes 2 – 10 mm; leaf blades 5 – 12 mm wide, symmetric at the base.…...........11

• 11. Cataphylls and cataphyllary stipules c. 0.4 mm long; petioles c. 0.4 mm long; leaf apex round; male flower tepals 1 – 1.3 mm long; fruiting pedicel 2.5 – 3 mm long.…...........11. P. vatovaviensis

• 11’. Cataphylls and cataphyllary stipules 0.8 – 1.5 mm long; petioles 0.8 – 1.5 mm long; leaf apex obtuse, acute or mucronulate; male flower tepals 2 – 2.5 mm long; fruiting pedicel 5 – 10 mm long.…...........12

• 12. Leaves obtuse at the apex, margin non revolute. Flowering and fruiting pedicels terete; gynophore 0.2 – 0.5 mm; ovary 0.5 – 1 mm diam.; disc glands smooth in male flowers; anthers fused into a disciform trilobed synandrium…...........12. P. lokohensis

• 12’. Leaves acute to mucronulate at the apex, margin revolute. Flowering and fruiting pedicels terete and becoming flattened and widened approaching the receptacle; gynophore absent; ovary 1.3 – 1.5 mm diam.; disc glands rugose or verrucose in male flowers; anthers entirely fused into a disciform bilobed (very rarely tri-lobed) synandrium…...........13. P. moramangicus

2. Phyllanthus venustulus Leandri (1938: 198). Type: Madagascar, Antananarivo, Ambatolampy, Tsiafajavona, 22 Nov. 1912, Humbert & Viguier 1739 (lectotype P(P000782930)!, selected here; isolectotype P(P000482887)!). This collection was chosen as lectotype because it contains male and female plant parts on the same sheet.

Dioecious herbs, sprawling on the ground. Branching pinnatiform; orthotropic branches terete, 0.5 – 1 mm in diam., striate, glabrous. Cataphylls triangular, 1.2 – 1.8 × 0.5 – 0.7 mm, coriaceous, entire, glabrous. Cataphyllary stipules triangular, 1.2 – 1.7 × 0.5 – 0.7 mm, coriaceous, entire, glabrous. Plagiotropic branches solitary, terete or slightly compressed, 0.8 – 2.5 cm long, 0.4 – 0.6 mm in diam., glabrous, rugose. Stipules persistent, triangular or subulate, 1 – 2 × 0.2 – 0.3 mm, membranous, entire, glabrous, yellowish, with white hyaline margin. Internodes 0.7 – 2 mm long. Leaves 8 – 17 per branch. Petioles terete or flattened, 0.2 – 0.4 × 0.2 – 0.3 mm, glabrous. Leaf blades ovate, obovate or suborbicular, 1.7 – 3 × 1 – 2 mm, 1.1 – 1.7 times longer than wide, rounded at the base, slightly mucronulate or obtuse at the apex, chartaceous, entire, sometimes slightly revolute, glabrous on both sides; midvein prominent or not visible adaxially, prominent abaxially; secondary veins 3 – 4 pairs, not visible adaxially, prominent abaxially. Inflorescences unisexual, consisting of 1 flower, male inflorescence on proximal part of plagiotropic branches, female inflorescence on distal part. Bracts triangular or elliptic, 0.5 – 0.7 × 0.2 – 0.3 mm, entire, glabrous, in a tuft of 4 – 6 mainly sterile in male flowers. Male flowers 0.8 – 1.2 × 0.7 – 1 mm, whitish when fresh. Pedicels 0.5 – 0.8 × c. 0.3 mm, glabrous. Tepals 5, subequal, elliptic or obovate, 0.8 – 1 × 0.4 – 0.6 mm, round or obtuse, membranous, glabrous on both sides, margin entire or slightly erose, hyaline for ¹/₄ of total tepal width on each side, veins unbranched. Disc glands 5, lenticular, peltate, c. 0.2 × 0.2 – 0.3 mm, rugose to verrucose. Stamens 2 – 3, filaments entirely fused into a column; column terete, 0.4 – 0.6 × 0.2 – 0.3 mm; anthers entirely fused into a disciform circumscissile synandrium, 0.2 – 0.3 mm in diam., dehiscing horizontally. Female flowers 1.9 – 3 × 1.9 – 4 mm. Pedicels 0.8 – 1 × 0.2 – 0.4 mm, glabrous. Tepals 5, inequal, ovate or obovate, 1.5 – 3 × 0.8 – 1.5 mm, obtuse, membranous, glabrous on both sides, margin entire, hyaline for ¹/₄ of total tepal width on each side, veins unbranched. Disc lobed, smooth. Ovary 3-locular, globose, 0.5 – 0.7 × 0.5 – 0.7 mm, smooth or slightly rugose, glabrous; styles entirely free, bifid for c. ¹/₂ of their length, terete, 0.4 – 0.5 mm long; stigmas obtuse and papillose. Fruits globose, 1 – 1.4 mm in diam., glabrous; fruiting pedicels 1 – 1.3 × 0.3 – 0.5 mm, glabrous; tepals 1.8 – 2.3 × 0.9 – 1.3 mm; styles persistent; columella 0.4 – 0.8 × 0.2 – 0.3 mm. Seeds 1 – 1.2 × 0.8 – 0.9 mm, striate longitudinally.

distribution. Endemic to Madagascar. Central floristic domain; 1200 – 1800 m. Map 2.

Map 2

Distribution of Phyllanthus venustulus.

specimens examined. madagascar. Antananarivo. Ambatolampy, Tsiafajavona, 22 Nov. 1912, Humbert & Viguier 1739 (lectotype P); Ambatolampy, Tsiafajavona, 22 Nov. 1912, Humbert & Viguier 1739 (isolectotype P) Ambatolampy, Tsiafajavona, s. dat., Perrier de la Bâthie 13371 (lectoparatype P); Ambatolampy, Tsiafajavona, s. dat., Perrier de la Bâthie 13530 (lectoparatype P); Manjakatompo, 21 Dec. 1950, Benoist 545 (P); Manjakatompo, 19 Dec. 1951, Benoist 1675 (P); Ambohimandroso, bord de l’Onive, alt. 1500 m, Dec. 1955, Bosser 8836 (P, TAN); Marécage, Ankaratra, entre Antsampandrahana et Namokely, alt. 1200 m, Feb. 1957, Bosser 10994 (TAN); Marécage, route d’Anjozorobe, P. K. 34, Dec. 1958, Bosser 12392 (TAN); Bords des rizières, aux environs de Manjakandriana, Feb. 1960, Bosser 13733 (TAN); Bords des rizières, Manjakatompo, March 1961, Bosser 15275 (P, TAN).

habitat. Peat and rocky areas; subhumid bioclimate; 1500 – 2500 m.

conservation status. Endangered (EN (B1abi-iv + B2abi-iv)). Phyllanthus venustulus has an area of occupancy (AOO) of 312.5 km² and an extent of occurence (EOO) of 3582 km². This species is found outside of protected areas and it is threatened by habitat loss due to agriculture and annual bush fire which lead to a decline of the EOO, the AOO, and the number of locations or subpopulations.

3. Phyllanthus ampandrandavae Leandri (1957: 224). Type: Madagascar, Toliara, aux environs d’Ampandrandava, entre Bekily et Tsivory, Nov. 1942, Seyrig 72 B (holotype P(P00535932)!; isotype P(P00535933)!).

Monoecious herbs, 20 – 30 cm high. Branching pinnatiform; orthotropic branches woody, terete, 0.8 – 1 mm in diam., striate, glabrous. Cataphylls triangular or lanceolate, 0.8 – 1 × 0.3 – 0.5 mm, subcoriaceous, entire, glabrous. Cataphyllary stipules triangular or lanceolate, 0.8 – 1 × 0.3 – 0.5 mm, subcoriaceous, entire, glabrous. Plagiotropic branches solitary, terete or subangular, 3.5 – 9.5 cm long, c. 1 mm in diam., glabrous, striate. Stipules persistent, triangular, 0.8 – 1 × 0.3 – 0.5 mm, membranous, entire, glabrous. Internodes 1.5 – 4 mm long. Leaves 10 – 27 per branch. Petioles terete, c. 0.5 × 0.2 mm, glabrous. Leaf blades elliptic or sometimes lanceolate, 5 – 6 × 2 – 3 mm, 2 – 2.5 times longer than wide, attenuate or rounded at the base, acute at the apex, sub-coriaceous or chartaceous, entire, glabrous on both sides; midvein prominent on both sides; secondary veins not visible on either side. Inflorescences unisexual or bisexual, consisting of 1 male and 1 female flowers, or 1 female flower, female inflorescences on proximal part of plagiotropic branches, bisexual inflorescences on distal part of plagiotropic branches. Bracts triangular, 0.4 – 0.8 × 0.3 – 0.5 mm, entire, glabrous. Male flowers c. 0.6 × 1 mm. Pedicels c. 0.6 × 0.1 mm, glabrous. Tepals 6, inequal, obovate or suborbicular, c. 0.6 × 0.6 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for ³/₄ of total tepal width on each side, veins unbranched. Disc glands 5 or 6, globose, c. 0.1 mm in diam., smooth. Stamens 3, filaments fused into a column for c. ¹/₄ of their length; filaments terete, c. 0.5 × 0.1 mm; column c. 0.1 × 0.3; anthers ovoid to globose, free, c. 0.2 mm in diam., dehiscing obliquely. Female flowers c. 0.9 × 1 mm. Pedicels c. 1 × 0.2 mm, glabrous. Tepals 5 – 6, equal, obovate or suborbicular, c. 0.8 × 0.5 mm, rounded or obtuse, membranous, glabrous on both sides, margin entire, hyaline for ¹/₃ of total tepal width on each side, veins branched. Disc annular 3-lobed, smooth. Ovary 3-locular, globose, c. 1 × 1 mm, smooth, glabrous; styles entirely free, bifid for c. ¹/₂ of their length, terete, c. 0.15 mm long; stigmas obtuse and papillose. Fruits depressed globose, 1 – 1.3 × 1.2 – 2 mm, glabrous, smooth; fruiting pedicels c. 1 × 0.1 mm, glabrous; tepals c. 1 × 0.6 mm; styles persistent; columella c. 0.7 × 0.2 mm. Seeds 0.6 – 1 × 0.4 – 0.6 mm, striate longitudinally.

distribution. Endemic to Madagascar: central and southern floristic domains. Map 3.

Map 3

Distribution of Phyllanthus ampandrandavae.

specimen examined. madagascar. Toliara. Ankilibe, Feb. 1989, Brunel 12515 (P); aux environs d’Ampandrandava, entre Bekily et Tsivory, Nov. 1942, Seyrig 72 B (holotype P); aux environs d’Ampandrandava, entre Bekily et Tsivory, Nov. 1942, Seyrig 72 B (isotype P).

habitat. Wooded grassland or degraded dry spiny forest; 600 – 800 m.

conservation status. Endangered (EN (B2abii-iv)). Phyllanthus ampandrandavae has an AOO of 46.219 km². This species is found in southern areas of Madagascar and localities are outside of protected areas. The species is threatened by habitat loss due to annual bush fire and pasture which lead to a decline in the AOO, the quality of habitat, and the number of locations or subpopulations. We assume that this species is very rare because it is only known from two collections from Ankilibe and Ampandrandava.

4. Phyllanthus andranovatensis Jean F. Brunel & Jacq. Roux (1981: 400). Type: Madagascar, Tolanaro, Andranovato, près de Vinanibe, 8 Oct. 1975, Brunel 2130 (holotype STR; isotypes TOGO, P(P00535927)!, TAN(TAN000574)!)

Monoecious herbs 10 – 20 cm high. Branching pinnatiform; orthotropic branches terete, 0.5 – 1.1 mm in diam., slightly rugose or striate, glabrous. Cataphylls triangular, 0.4 – 0.9 × 0.2 – 0.3 mm, coriaceous, entire, glabrous. Cataphyllary stipules 0.3 – 0.9 × 0.2 – 0.3 mm, coriaceous, entire, glabrous. Plagiotropic branches solitary, terete or slightly compressed, 1 – 4 cm long, 0.3 – 0.4 mm in diam., glabrous, rugose. Stipules persistent, linear, 1 – 1.5 × 0.1 – 0.3 mm, membranous, entire, glabrous. Leaves 7 – 14 per branch. Internodes 0.4 – 2 mm long. Petioles terete, 0.4 – 0.5 × 0.2 – 0.3 mm, glabrous. Leaf blades elliptic or oblong, 5 – 8 × 2 – 4 , 2 – 2.5 times longer than wide, obtuse at the base, obtuse or rounded at the apex, membranous, entire, glabrous on both sides, light green adaxially and reddish abaxially when fresh; midvein flattened on both sides; secondary veins 4 – 6 pairs, flattened abaxially, indistinct or flattened adaxially. Inflorescences unisexual, consisting of 1 – 2 male flowers or only 1 female flower, male inflorescence on proximal part of plagiotropic branches and female inflorescence on distal part of plagiotropic branches. Bracts linear, 0.3 – 0.5 × c. 0.1 mm, entire, glabrous. Male flowers 1.3 – 1.5 × 1.8 – 2.5 mm. Pedicels 0.8 – 1 × 0.2 – 0.4 mm, glabrous. Tepals 5, equal, obovate, 1 – 1.3 × 0.6 – 0.9 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₄ of total tepal width on each side, veins unbranched. Disc glands 5, obovate, thin, c. 0.2 × 0.2 mm, smooth. Stamens 2 – 3, filaments fused into a column for ²/₃ their length; filaments terete, 0.4 – 0.6 × 0.1 – 0.2 mm; anthers free, c. 0.3 mm long, dehiscing horizontally. Female flowers 1.3 – 1.9 × 2 – 2.5 mm. Pedicels 1 – 1.4 × 0.3 – 0.4 mm, glabrous. Tepals 5, equal, elliptic, 1 – 1.9 × 0.8 – 1.2 mm, rounded, membranous, glabrous, margin entire, hyaline for c. ¹/₄ of total tepal width on each side, veins unbranched. Disc annular 5-lobed, smooth. Ovary 3-locular, globose or depressed globose, 0.7 – 1 × 0.7 – 1.3 mm, smooth, glabrous; styles entirely free, terete, bifid for c. ¹/₂ of their length, 0.4 – 0.5 mm long; stigmas obtuse. Fruits globose or depressed globose, 2 – 2.5 × 2.3 – 2.7 mm, glabrous; fruiting pedicels 1 – 2 × 0.3 – 0.5 mm, glabrous, smooth at base and slightly papillose at apex; tepals 1 – 2 × 0.8 – 1.3 mm; styles caducous; columella c. 0.3 × 0.2 mm. Seeds 1.6 – 2.3 × 1–1.4 mm, with 9 – 11 longitudinal striae.

distribution. Endemic to Madagascar: eastern floristic domain. Map 4.

Map 4

Distribution of Phyllanthus andranovatensis.

specimen examined. madagascar. Toliara. Tolanaro, Andranovato, près de Vinanibe, 8 Oct. 1975, Brunel 2130 (isotypes P, TAN).

habitat. Littoral forest and swamp; alt. sea level – 10 m.

conservation status. Data deficient (DD). Phyllanthus andranovatensis is only known from the type and has not been collected in recent years.

5. Phyllanthus ankazobensis Ralim. & Petra Hoffm. sp. nov. Type: Madagascar, Antananarivo, Anjozorobe, Antsahabe, 18°23'47"S, 47°56'9"E, alt. 1330 m, 12 Dec. 2002, Ralimanana & Ranaivojaona 306 (holotype K!; isotypes G!, MO!, P!, TAN!).

http://www.ipni.org/urn:lsid:ipni.org:names:77134145-1

Monoecious shrubs with trailing and flexible orthotropic and plagiotropic branches, 50 – 80 cm high. Branching pinnatiform; orthotropic branches terete, 0.5 – 2 mm in diam., rugose, glabrous. Cataphylls triangular, 0.6 – 1.5 × 0.3 – 0.5 mm, subcoriaceous or coriaceous, light brown with white hyaline margin, entire, glabrous. Cataphyllary stipules triangular, 0.4 – 1.5 × 0.3 – 0.5 mm, subcoriaceous or coriaceous, glabrous. Plagiotropic branches solitary, terete, 2 – 5.5 cm long, 0.3 – 0.5 mm in diam., glabrous, striate or smooth. Stipules persistent, triangular, 0.7 – 1.2 × 0.3 – 0.5 mm, membranous, entire, glabrous. Internodes 1 – 7 mm long. Leaves 5 – 14 per branch. Petioles terete, 0.5 – 1 × 0.3 – 0.6 mm, glabrous. Leaf blades ovate, 3 – 10 × 2 – 5.5 mm, obtuse or rounded, asymmetric at the base, obtuse or acute, often mucronulate at the apex, chartaceous, revolute, glabrous on both sides, green adaxially and light green abaxially when fresh; midvein flattened adaxially, prominent abaxially; secondary veins 3 – 5 pairs, indistinct adaxially, flattened abaxially. Inflorescences unisexual, consisting of 2 or 3 male flowers (one open and the others in bud) or only 1 female flower; male inflorescences on proximal or middle parts of plagiotropic branches and female inflorescence on distal part. Bracts linear or triangular, 0.3 – 0.5 × 0.2 – 0.3 mm, entire, glabrous. Male flowers 2 – 3 × 1.5 – 2 mm, cream when fresh. Pedicels 2 – 3 × 0.1 – 0.2 mm, glabrous. Tepals 5, equal, obovate, elliptic to obovate, 2 – 3 × 1 – 1.5 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of total tepal width on each side, veins branched. Disc glands 5, reniform or lenticular, 0.2 – 0.3 × c. 0.2 mm, verrucose. Stamens 3, filaments fused into a column for ¹/₂ – ²/₃ of their length; filaments terete, free part 0.3 – 0.5 mm long; column terete, 0.3 – 0.6 mm in diam.; anthers free, ovoid, 0.2 – 0.4 mm long, dehiscing obliquely. Female flowers 4 – 5 × 4 – 6 mm, cream when fresh. Pedicels 1.4 – 2 × 0.2 – 0.3 mm (1 mm wide near the receptacle), glabrous. Tepals 5, equal or inequal, elliptic to obovate, 5 – 6 × 3 – 4 mm, rounded, membranous or slightly coriaceous, glabrous on both sides, margin entire, hyaline for ¹/₁₀ – ¹/₈ of total tepal width on each side, veins branched. Disc lobed, smooth. Ovary 3-locular, ovoid or globose, 0.8 – 1 × 0.8 – 1 mm, on gynophore 0.3 – 0.4 mm, smooth, glabrous; styles fused for ²/₃ of their length, bifid for ¹/₃ their length, terete, 0.6 – 0.8 mm long; stigmas obtuse. Fruits only immature ones seen, globose, 1 – 2 × 1 – 2 mm, glabrous; fruiting pedicels terete, 3 – 5 × 0.3 – 0.5 mm glabrous, tepals 5 – 7 × 3 – 4 mm. Fig. 1.

Fig. 1

Phyllanthus ankazobensis. A habit; B leaves; C part of plagiotropic branch; D male flower; E female flower. From H. Ralimanana 306 (TAN). drawn by r. l. andriamiarisoa.

recognition. Phyllanthus ankazobensis is similar to P. madagascariensis in leaf shape, stamen number, filament fusion, and the capsular fruits but differs in the trailing and flexible branches (vs erect), gynophore 0.3 – 0.4 mm (vs gynophore absent) and the styles fused into a column for ²/₃ of their length (vs styles free or only fused at the base).

distribution. Endemic to Madagascar: central floristic domain. Map 5.

Map 5

Distribution of Phyllanthus ankazobensis.

specimens examined. madagascar. Central Madagascar, Baron 3739 (K, P); Antananarivo. Manankazo, près d’Ankazobe, 1913, Perrier de la Bâthie 9881 (P); Anjozorobe, Antsahabe, 18°23'47"S, 47°56'9"E, alt. 1330 m, 12 Dec. 2002, Ralimanana & Ranaivojaona 306 (holotype K, isotypes G, MO, P, TAN). Toamasina. Alaotra Mangoro, Ambatovy, 18°51'12"S, 48°19'04"E, alt. 1104 m, 11 June 2008, Antilahimena 6298 (MO, P, TAN); Ambatovy, 1 July 1966, Peltier & Peltier 5999 (P); Moramanga, Andasibe, Andasifahatelo, Marinjo, corridor forestier Analamay Mantadia, 18°47'05"S, 48°24'12"E, alt. 1044 m, 29 May 2012, Rasoazanany & T. Ratolojanahary 222 (MO, TAN); Moramanga, Ambatovy, Ampitambe, 18°51'03"S, 48°43'E, alt. 1144 m, 15 Feb. 2005, Razanatsoa et al. 160 (MO, P, TAN).

habitat. Open humid forest; alt. 1000 – 1200 m.

conservation status. Vulnerable (VU (B1abi-iv + B2abi-iv)). Phyllanthus ankazobensis has an EOO of 7292 km² and an AOO of 1016 km². Habitat is threatened by annual fire, agriculture and mining which lead to a decline in the AOO, and the EOO, the quality of habitat, and the number of locations or subpopulations.

6. Phyllanthus madagascariensis Müll. Arg. (Müller 1863: 35). Type: Madagascar, aux environs de Tamatave, Chapelier s.n. (holotype P(P00078221)!; isotype P(P00078222)!).

Phyllanthus tampinensis Leandri (1957: 233). Type: Madagascar, Tampina, Sud de Tamatave, Decary 17701 (holotype P(P00535931)!).

Monoecious shrubs 50 – 100 cm high. Branching pinnatiform; orthotropic branches terete, 2 – 6 mm in diam., striate, glabrous. Cataphylls triangular, 0.3 – 1 × 0.2 – 0.4 mm, coriaceous entire, glabrous. Cataphyllary stipules 0.3 – 1 × 0.2 – 0.4 mm, coriaceous entire, glabrous. Plagiotropic branches solitary or fascicled in twos, terete, 3 – 15 cm long, 0.3 – 0.4 mm in diam., pubescent or scabridulous, striate. Stipules persistent or caducous, triangular, 0.4 – 1.8 × 0.3 – 0.4 mm, membranous or subcoriaceous, entire, glabrous. Internodes 1.5 – 4 mm long. Leaves 4 – 16 per branch. Petioles terete or terete, 0.3 – 1 × 0.2 – 0.3 mm, glabrous. Leaf blades elliptic, ovate, obovate, 5 – 15 × 4 – 10 mm, 1.3 – 1.5 times longer than wide, attenuate and asymmetric at the base, acute or obtuse at the apex, subcoriaceous to coriaceous or chartaceous, entire, sometimes slightly revolute, glabrous adaxially, thinly papillose or glabrous abaxially, green adaxially and light green abaxially when fresh; midvein flattened on both sides; secondary veins distinct or indistinct on both sides. Inflorescences unisexual; consisting of 1 or 2 male flowers or only 1 female flower; male inflorescences and female inflorescences on different branches or male inflorescences on proximal of plagiotropic branches and female inflorescence on distal part. Bracts triangular or elliptic, 0.5 – 0.8 × c. 0.3 mm, entire or slightly erose, glabrous. Male flowers 1.7 – 3 × 2 – 4 mm, whitish green when fresh. Pedicels 1 – 3.5 × 0.2 – 0.3 mm, glabrous. Tepals 5, equal, elliptic or obovate, 1.7 – 3 × 1.5 – 2 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of total tepal width on each side, veins unbranched. Disc glands 5, globose, c. 0.4 × 0.4 mm, smooth. Stamens 3, filaments fused for ²/₃ – ³/₄ of their length; column terete, 1.5 – 1.7 mm in diam.; free parts terete, 0.3 – 0.5 mm in diam.; anthers free, ovoid, c. 0.3 mm long, dehiscing obliquely. Female flowers 3.2 – 7 × 6 – 10 mm, whitish green when fresh. Pedicels 3 – 3.5 × 0.3 – 0.5 mm, glabrous. Tepals 5, equal, elliptic or obovate, 5 – 7 × 3 – 5 mm, rounded, coriaceous, glabrous on both sides, margin entire, hyaline for ¹/₁₂ of total tepal width on each side, veins branched. Disc annular lobed, smooth. Ovary 3-locular, globose, 1 – 2 × 1 – 2 mm, smooth, glabrous; styles free or fused at the base, bifid for c. ¹/₂ of their length, terete, 1.2 – 2 mm long; stigmas obtuse. Fruits depressed globose, 2 – 3 × 3 – 4.5 mm, glabrous; fruiting pedicels terete, 3 – 6 × 0.3 – 0.5 mm, glabrous; tepals 5 – 8 × 4 – 5.2 mm; styles persistent; columella 0.8 – 1.2 × 0.2 – 0.4 mm. Seeds 1.7 – 2 × 1 – 1.5 mm, smooth.

6a. Phyllanthus madagascariensis var. madagascariensis Leaf blades subcoriaceous to coriaceous; secondary veins indistinct in both sides. Female flower tepals 6 – 7 × 3 – 5 mm. Species growing in littoral forest.

distribution. Endemic to Madagascar: Eastern floristic domain (Antsiranana, Fianarantsoa and Toamasina province); sea level – 20 m. Map 6.

Map 6

Distribution of Phyllanthus madagascariensis var. madagascariensis (circles) and P. madagascariensis var. kalambantitrensis (squares).

specimens examined. madagascar. s. loc., Oct. 1883, Humblot 145 (K, P). Antsiranana. Antalaha, Ambohitralanana, Ambodirafia, littoral forest, 15°16'31"E, 50°28'60"E, 8 April 2003, Ranaivojaona 562 (TAN). Fianarantsoa. Mananjary, March – May 1909, Geay 7090 (P); Mananjary, March – May 1909, Geay 7230 (P); Mananjary, March – May 1909, Geay 7259 (P); Mananjary, March – May 1909, Geay 7947 (P); Farafangana, Mahabo, 23°11'9"S, 47°42'22"E, alt. 25 m, 9 Nov. 2001, McPherson & Rabenantoandro 18390 (K, MO); Mahanoro, Ambodibonara, forêt littorale Antaimby, 20°22'22"S, 48°33'16"E, alt. 10 m, 16 May 2004, Razakamalala et al. 1126 (K, MO, TAN); Ampasimaneva, forêt littorale d’Ambolo, 20°43'59"S, 48°27'09"E, alt. 13 m, 15 June 2004, Razakamalala et al. 1449 (MO, P, TAN). Toamasina. Ste Marie, Lokintsy, forêt littorale d’Ambohidena, 16°51'9"S, 49°57'13"E, alt. 19 m, 9 Oct. 2003, Andrianarivelo et al. 87 (K, MO, P, TEF); Ste Marie, Boivin 1886² (P); aux environs de Tamatave Chapelier s.n. (holotype P); aux environs de Tamatave Chapelier s.n. (isotype P); Ambila, 4 May 1928, Decary 6445 (K, P, TAN); Tampina, sud de Toamasina, 5 March 1942, Decary 17701 (P); Ste Marie, Lokintsy, forest of Ambohidena, 16°51'S, 49°57'E, alt. 10 m, 13 May 2003, McPherson et al. 18925 (K, MO, P); Ambila, 18°48'S, 49°09'E, alt. 10 m, 23 Sept. 2000, Miller et al. 10640 (K, MO, P); Tampina, Nov. 1921, Perrier de la Bâthie 13323 (P); Brickaville, S of Ambila, 18°54'S, 49°7'E, alt. 0 – 10 m, 4 Aug. 1996, Randrianasolo 479 (K, MO, P); Ste Marie, Lokintsy, forêt d’Ambohidena, 16°50'25"S, 49°57'9"E, alt. 10 m, 19 Feb. 2004, Razakamalala et al. 897 (K, MO); Ste Marie, Lokintsy, forest of Ambohidena, 16°51'11"S, 49°57'10"E, 1 June 2004, Rabehevitra 1182 (MO, P, TEF); Ambila, road to Andevoranto, 18°49'S, 49°08'E, alt. 10 m, 3 July 1995, Razafimandimbison 182 (K, MO).

habitat. Littoral forest.

conservation status . Vulnerable (VU (B1abi-iv + B2abi-iv)]. Phyllanthus madagascariensis var. madagascariensis has an EOO of 14968.4 km² and an AOO of 972 km². The habitat is found in littoral forest which is fragmented and is severely threatened by slash and burn agriculture.

6b. Phyllanthus madagascariensis var. kalambatitrensis Leandri ex Ralim. & Petra Hoffm., var. nov. Type: Madagascar, Kalambatritra massif, Humbert 12118 (holotype P!).

http://www.ipni.org/urn:lsid:ipni.org:names:77134147-1

Phyllanthus madagascariensis var. kalambatitrensis Leandri (1958: 82), nom. invalid.

recognition. Phyllanthus madagascariensis var. kalambatitrensis differs from var. madagascariensis in the leaf blades being chartaceous (vs subcoriaceous to coriaceous), the secondary veins distinct (vs indistinct) and the female flower with tepals 4 – 4.5 × 1.8 – 2 mm (vs 6 – 7 × 3 – 5 mm). Plant growing in mid-elevation humid forest (vs littoral forest).

distribution. Endemic to Madagascar: Central floristic domain. Map 6.

specimens examined. madagascar. Fianarantsoa. Iakora, Begogo, Bekora, forêt de Sahalava au sud du village d’Androizaha, forêt humide, 23°31'59"S, 46°32'04"E, alt. 1180 m, 28 Jan. 2005, Andrianjafy 751 (MO, TAN). Toliara. Betroka, Ivahona, Befarafara, forêt humide de moyenne altitude, 23°24'48"S, 46°26'55"E, alt. 1561 m, 27 May 2005, Andrianjafy 1136 (MO, TAN).

habitat. Mid-altitude humid forest; alt. 1100 – 1600 m.

conservation status. Least Concern (LC). Phyllanthus madagascariensis var. kalambatritrensis has an AOO of 5 km² with two locations inside a protected area. The location is in remote area and very rarely visited by people so this species is not under human pressure.

7. Phyllanthus melleri Müll. Arg . (Müller 1864: 514). Type: Madagascar, between Toamasina and Antananarivo, Meller s.n. (holotype K!).

Phyllanthus melleri var. campenoni Leandri (1939: 185), nom. invalid.

Monoecious shrubs 50 – 100 cm high. Branching pinnatiform; orthotropic branches terete, 2 – 4 mm in diam., glabrous. Cataphylls triangular, 0.5 – 1.7 × 0.2 – 0.4 mm, coriaceous, entire, glabrous. Cataphyllary stipules triangular, 0.4 – 1.7 × 0.2 – 0.4 mm, coriaceous, entire, glabrous. Plagiotropic branches fascicled in twos or solitary, terete, 3 – 16 cm long, 0.3 – 0.8 mm in diam., scabridulous to pubescent, striate. Stipules persistent or caducous, triangular or subulate, 0.4 – 1.8 × 0.3 – 0.4 mm, slightly coriaceous, entire, glabrous. Internodes 3 – 5 mm long. Leaves 4 – 10 per branch. Petioles terete, 0.5 – 1 × 0.2 – 0.3 mm, glabrous. Leaf blades ovate or elliptic, 4 – 15 × 3 – 10 mm, 1.3 – 1.5 times longer than wide, attenuate or obtuse, sometimes suboblique or asymmetric at the base, acute or obtuse at the apex, chartaceous, entire, glabrous on both sides; midvein flattened on both sides; secondary veins 3 – 5 pairs, flattened on both sides. Inflorescences unisexual, consisting of 1 or 2 male flowers, or only 1 female flower; male inflorescences on proximal part of plagiotropic branches and female inflorescences on distal part, or inflorescences on different branches. Bracts triangular or elliptic, 0.4 – 0.6 × c. 0.3 mm, entire, glabrous. Male flowers 1.8 – 4 × 2 – 4 mm, whitish green when fresh. Pedicels 1 – 4 × 0.2 mm, glabrous. Tepals 5 – 6, equal, elliptic or obovate, 2 – 4 × 1.5 – 2 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of total tepal width on each side, veins branched. Disc glands 5 or 6, elliptic transversally, c. 0.4 mm in diam., smooth. Stamens 3, filaments fused into a column for ²/₃ – ³/₄ of their length; filaments 1.5 – 2 mm long; column terete, 1.5 – 1.7 mm in diam.; free parts terete, 0.3 – 0.6 mm in diam.; anthers free, ovoid, c. 0.3 mm long, dehiscing obliquely. Female flowers 3 – 5 × 6 – 10 mm, whitish green when fresh. Pedicels c. 3 × 0.5 mm, glabrous. Tepals 5, equal, elliptic or obovate, c. 5 × 2 – 3 mm, rounded, chartaceous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of tepal width, veins branched. Disc annular subentire, smooth. Ovary 3-locular, globose, 1 – 1.5 mm in diam., smooth, glabrous; styles fused at the base, bifid for c. ¹/₂ of their length, terete, 1.2 – 1.5 mm long; stigmas acute. Fruits globose, only immature ones seen, glabrous; fruiting pedicels terete, c. 4 × 0.5 mm, glabrous, tepals c. 5 × 2 – 3 mm.

distribution. Endemic to Madagascar: Central floristic domain. Map 7.

Map 7

Distribution of Phyllanthus melleri.

specimens examined. madagascar. Central Madagascar, Baron 411 (K); Central Madagascar, Baron 3741 (K); Central Madagascar, Baron 4080 (K); Campenon s.n. (P). Antananarivo. Between Toamasina and Antananarivo, Meller s.n. (holotype K); station forestière d’Angavokely, 18°55'35"S, 47°45'1"E, alt. 1715 m, 2 Nov. 2002, Lowry II et al. 5814 (K, MO); Andramasina, Antakafatra, Andranomena forest; 19°27'10"S, 47°58'52"E, alt. 1594 m, 16 Dec. 2009, Ralimanana et al. 1594 (TAN, TEF).

habitat. Open humid forest or humid forest boundaries; alt. 1590 – 1720 m.

conservation status. Endangered (EN B2ab(ii,iii,iv)). Phyllanthus melleri is known from six collections but four of them are without precision of localities. It has an AOO of 130.162 km² and it is found outside of protected areas. Therefore this species is threatened by habitat loss due to human activities such as fire and forest logging which lead to a decline of area of occupancy, quality of habitat and number of locations.

8. Phyllanthus rangoloakensis Leandri (1938: 198). Type: Madagascar, Central Madagascar, Andrangoloaka, Nov. 1880, Hildebrandt 3707 (holotype P(P00078271)!; isotypes P(P00078272, P00078273)!, G, JE, US).

Monoecious shrubs 50 – 70 cm high. Branching pinnatiform; orthotropic branches terete, 2 – 5 mm in diam., rugose (scars of old plagiotropic branches visible), glabrous. Cataphylls triangular, 1 – 1.5 × 0.5 – 1 mm, coriaceous, entire or slightly erose, glabrous. Cataphyllary stipules 0.9 – 1.5 × 0.5 – 1 mm, coriaceous, entire or slightly erose, glabrous. Plagiotropic branches solitary, grouped at the distal part of the orthotropic branch, terete, slightly dilated at the base, 3.5 – 9.5 cm long, 0.6 – 1 mm in diam., glabrous, striate. Stipules persistent, triangular, 1.5 – 1.8 × 0.5 – 0.6 mm, coriaceous, entire, glabrous. Leaves 20 – 36 per branch. Internodes 1 – 2 mm long. Petioles terete or flattened, 0.5 – 1 × 0.4 – 0.6 mm, glabrous. Leaf blades oblong sub-falcate, 6 – 10 × 2.5 – 3 mm, 2.4 – 3.3 times longer than wide, obtuse at the base, acute at the apex, chartaceous, entire, slightly revolute, glabrous on both sides, green adaxially when fresh, light green abaxially when fresh; midvein prominent adaxially, flattened abaxially; secondary veins indistinct on both sides. Inflorescences unisexual, male inflorescences with 1 – 2 flowers (one open and one still in bud), female inflorescences with one female flower, male inflorescence on proximal part and female inflorescence on distal part of plagiotropic branches. Bracts linear or triangular, grouped in a tuft of 4 – 6 mainly sterile bracts in male inflorescences, 0.4 – 0.8 × 0.1 – 0.2 mm, entire, glabrous. Male flowers 1.2 – 1.5 × 1.8 – 1.9 mm, whitish green when fresh. Pedicels 0.4 – 0.6 × c. 0.3 mm, glabrous. Tepals 5, equal, obovate, 1 – 1.5 × 0.8 – 1 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for ¹/₈ – ¹/₆ of total tepal width on each side, veins unbranched. Disc glands 5, lenticular, c. 0.3 mm in diam., smooth. Stamens with filaments entirely fused in a column; column terete, 0.4 – 0.6 × c. 0.3 mm; anthers completely fused into a disciform circumscissile synandrium dehiscing horizontally. Female flowers 1.7 – 2.2 × 2.3 – 3 mm, whitish green when fresh. Pedicels 0.5 – 0.7 × 0.3 – 0.4 mm, glabrous. Tepals 5, equal, elliptic, 1.5 – 2 × 0.7 – 1 mm, slightly coriaceous, obtuse, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of total tepal width on each side, veins branched near the apex. Disc lobed, smooth. Ovary 3-locular, globose, 1 – 1.2 mm, smooth, glabrous; styles fused at the base, bifid for ¹/₃ – ¹/₂ of their length, terete, 0.5 – 0.8 mm long; stigmas obtuse. Fruits globose, 1.5 – 2 mm in diam., glabrous; fruiting pedicels 1 – 1.4 × 0.3 – 0.5 mm, glabrous; tepals 1.5 – 2 × 1 – 1.3 mm; styles persistent; columella c. 0.7 × 0.3 mm. Seeds 1 – 1.5 × 0.7 – 1 mm, smooth. Rheophyte.

distribution. Endemic to Madagascar: Central and eastern floristic domains. Map 8.

Map 8

Distribution of Phyllanthus rangoloakensis.

specimens examined. madagascar. Central Madagascar, Andrangoloaka, Nov. 1880, Hildebrandt 3707 (holotype P); Central Madagascar, Andrangoloaka, Nov. 1880, Hildebrandt 3707 (isotypes P). Toamasina. Roches humides, route d’Anosibe An’ala, Aug. 1953, Bosser 6099 (P); Roches humides, route d’Anosibe An’ala, Aug. 1953, Bosser 6100 (P); Moramanga, rocher humide, route d’Anosibe, Aug. 1953, Bosser 6115 (TAN); Entre Sandrangato et Anosibe, sud de Moramanga, alt. 800 – 1000 m, 3 – 7 Nov. 1952, Capuron & Leandri 1696 (P); Above Anevoka, 18°57'S, 48°28'E, alt. 900 m, Gillespie 4027 (K, MO, P); Vallée de Manampanihy, col de Sahindro, Feb. 1934, Humbert 14004 (P); 41 km S of Moramanga, road to Anosibe An’ala, 19°14'52"S, 48°13'54"E, 28 Nov. 1996, Schatz et al. 3709 (K, MO, TEF); Road to Lakato, 19°00'52"S, 48°20'47"E, alt. 1040 m, 10 Nov. 2003, Schatz et al. 4141 (MO, P). Toliara. Haut vallée de Manampanihy, col de Saindro, alt. 1200 m, Feb. 1934, Humbert 14004 (MO, P).

habitat. Species growing on rocks near stream or in river-beds; alt. 800 – 1200 m.

conservation status. Vulnerable (VU (B1ab i-iv + B2ab i-iv)). Phyllanthus rangoloakensis has an EOO of 16548.7 km² and an AOO of 567 km². All locations are outside of protected areas and the habitat is threatened by bush fire and cut and slash.

9. Phyllanthus coursii Leandri (1957: 226). Type: Madagascar, Sud-Est du Lac Alaotra, Cours 191 (holotype P(P00535905)!; isotype US).

Monoecious shrubs 80 – 100 cm high. Branching bipinnatiform; orthotropic branches terete, 0.5 – 2.5 mm in diam., striate, glabrous. Cataphylls triangular, c. 1 × 0.5 mm, coriaceous, entire, glabrous. Cataphyllary stipules triangular, c. 0.8 × 0.5 mm, coriaceous, entire, glabrous. Plagiotropic branches solitary, slightly flattened, 3 – 8.5 cm long, c. 0.3 mm in diam., puberulent, striate. Stipules persistent or caducous, triangular, c. 0.8 × 0.4 mm, membranous, entire, glabrous. Internodes 3 – 6 mm long. Leaves 5 – 12 per branch. Petioles terete, c. 1 × 0.2 mm, glabrous. Leaf blades elliptic or ovate, 4 – 15 × 2.5 – 7 mm, 1.5 – 1.6 times longer than wide, slightly attenuate or obtuse at the base, obtuse or acute, mucronulate at the apex, chartaceous, entire or slightly revolute, glabrous on both sides; midvein flattened adaxially, prominent abaxially; secondary veins 4 – 5 pairs, slightly prominent adaxially, flattened abaxially. Inflorescence unisexual, consisting of 2 – 4 male flowers or only 1 female flower; male inflorescences on proximal part and female inflorescences on distal part of plagiotropic branches. Bracts triangular, grouped in a tuft of 9 – 10 mainly sterile bracts in male inflorescences, c. 0.7 × 0.3 mm, entire, glabrous. Male flowers c. 1.4 × 1 – 1.2 mm, whitish green when fresh. Pedicels c. 3 × 0.2 mm, glabrous. Tepals 5, subequal, broadly obovate, 1 – 1.4 × c. 1.2 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of total tepal width on each side, veins reticulately branched. Disc glands 5, peltate, lenticular, c. 0.3 × 0.3 mm, rugose or verrucose. Stamens 3, filaments entirely fused into a column; column terete, c. 1 × 0.2 mm; anthers partially fused, c. 0.3 mm long, dehiscing horizontally. Female flowers 1.8 – 2 × 2.5 – 3 mm, whitish green when fresh. Pedicels 5 – 7 × c. 0.2 mm, glabrous. Tepals 5, subequal, obovate, c. 2 × 1.4 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for ¹/₁₂ – ¹/₈ of total tepal width on each side, veins branched. Disc annular, lobed, smooth. Ovary 3-locular, globose, c. 0.8 mm in diam., smooth, puberulent or pubescent; styles free or fused only at the base, bifid for c. ¹/₃ of their length, terete, c. 0.6 mm long; stigmas obtuse. Fruits globose, c. 2 mm in diam., puberulent or pubescent, striate; fruiting pedicels terete, 4 – 5 × c. 0.2 mm, glabrous; tepals c. 3 × 1.6 mm; styles persistent; columella c. 1 × 0.4 mm. Seeds 0.8 – 1.2 × 0.3 – 0.4 mm, smooth.

distribution. Endemic to Madagascar: central floristic domain. Map 9.

Map 9

Distribution of Phyllanthus coursii.

specimen examined. madagascar. Sud-Est du Lac Alaotra, Cours 191 (holotype P).

habitat. Mid-altitude humid forest.

conservation status. Data Deficient (DD). Phyllanthus coursii is only known from the type and has not been collected in recent years.

10. Phyllanthus ivohibeus Leandri (1938: 197). Type: Madagascar, Fianarantsoa, Ivohibe, bassin de Manampatra, Perrier de la Bâthie 9760 (lectotype P!, selected here). Because of the quality and the preservation of the specimens, this collection is chosen as lectotype.

Dioecious shrubs 40 – 70 cm high. Branching pinnatiform; orthotropic branches terete, 0.5 – 2 mm in diam., smooth, glabrous. Cataphylls triangular, 0.5 – 0.8 × 0.3 – 0.4 mm, coriaceous, entire, glabrous. Cataphyllary stipules 0.4 – 0.8 × 0.3 – 0.4 mm, coriaceous, entire, glabrous. Plagiotropic branches solitary, terete, 2 – 8 cm long, 0.3 – 0.4 mm in diam., glabrous, striate. Stipules persistent, triangular, 0.5 – 1 × c. 0.3 mm, membranous, entire, glabrous. Internodes 0.5 – 1 mm long. Leaves 9 – 48 per branch. Petioles terete, 0.4 – 0.6 × 0.2 – 0.3 mm, glabrous. Leaf blades elliptic (asymmetric at the base), c. 2.5 – 5 × 1.5 – 3 mm, 1.6 times longer than wide, rounded at the base, acute, obtuse or rounded at the apex, chartaceous, entire, sometimes revolute, glabrous on both sides; midveins prominent adaxially, impressed abaxially; secondary veins 3 – 4 pairs, indistinct adaxially, flattened abaxially. Inflorescences unisexual, consisting of 2 male flowers or only 1 female flower. Bracts triangular, grouped in a tuft of 5 – 8 mainly sterile bracts in male inflorescences, 0.4 – 0.6 × 0.2 – 0.3 mm, entire or slightly erose, glabrous. Male flowers 1.8 – 2 × 2 – 2.2 mm, whitish when fresh. Pedicels 1.5 – 2 × 0.2 – 0.3 mm, glabrous. Tepals 5, equal, elliptic or suborbicular, 1.5 – 1.7 × 1 – 1.5 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₃ of total tepal width on each side, veins unbranched. Disc glands 5, peltate, lenticular, c. 0.4 × 0.3 mm, rugose or verrucose. Stamens 2 – 3, filaments entirely fused into a column; column terete, 0.8 – 1 × 0.2 – 0.3 mm; anthers completely fused into a disciform circumscissile synandrium, unlobed or with two or three lobes, dehiscing horizontally. Female flowers 2 – 2.5 × 3 – 3.5 mm, whitish when fresh. Pedicels 1.5 – 2 × c. 0.2, glabrous. Tepals 5, elliptic or obovate, 1.8 – 2.5 × 1.3 – 1.7 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for ¹/₆ – ¹/₄ of tepal width on each side, veins unbranched. Disc lobed, smooth. Ovary 3-locular, globose, 1 – 1.7 mm in diam., smooth, glabrous; styles entirely free, bifid for c. ¹/₂ of their length, terete, 0.8 – 1 mm long; stigmas obtuse. Fruits depressed globose, 1.5 – 2 × 1.7 – 2.2 mm, glabrous; fruiting pedicels 3 – 5 × 0.2 – 0.3 mm, glabrous; tepals 2 – 3 × 1.5 – 2 mm. Seeds c. 1 × 0.8 mm, smooth. Fig. 2.

Fig. 2

Phyllanthus ivohibeus. A habit; B leaves; C nodes with stipules, petiole, leaves and inflorescence; D part of plagiotropic branch with fruits; E female flower; F fruit. From F. Rakotonasolo 803 (TAN). drawn by r. l. andriamiarisoa.

distribution. Endemic to Madagascar: central floristic domain (Antananarivo and Fianarantsoa province). Map 10.

Map 10

Distribution of Phyllanthus ivohibeus.

specimens examined. madagascar. Antananarivo. Antananarivo, Perrier de la Bâthie 9735 (P); Tsinjoarivo, NE of queen’s palace, 19°37'91"S, 47°41'49"E, alt. 1555 m, 18 Nov. 2004, Ralimanana & Rajaonarison 649 (TAN); Vakinakaratra, N of Pic Vohimalaza near Betafo, alt. 1600 m, 18 Nov. 1912, Viguier & Humbert 1363 (P). Fianarantsoa. Vangaindrano, Midongy Atsimo, Ambangivelo, Telorano, 23°43'33", 47°02'13"E, alt. 705 m, 22 Sept. 2008, Bussmann et al. 15074 (MO, TAN); Manjarivolo, alt. 1650 m, 1 Nov. 1970, Guillaumet 3457 (P); Bassin de Matitanana, Sept. 1934, Hein s.n. (P); Haute vallée de Rienana, alt. 1000 – 1400 m, Humbert 3621 (lectoparatype P); Ivohibe, bassin de Manampatra, Perrier de la Bâthie 9760 (lectotype P); Itomampy, mont Papanga, alt. 1300 – 1700 m, 2 – 3 Dec. 1928, Humbert 6917 (P); Ambalavao, Ambalamanasy, Ambondrombe, 21°25'S, 44°57'E, 26 May 2004, Rakotonasolo 803 (MO, TAN); Ivohibe, Ambarongy, N of Iantara R, 22°13'20"S, 47°01'29"E, alt. 720 m, 12 Oct. 1994, Rakotovao et al. 153 (MO, TAN); limite nord de la reserve, le long de la riviere Ifefitany, 22°28'12"S, 46°57'36"E, alt. 900 m, Rakotovao et al. 838 (MO, TAN); Andrambovato, côte Talatamaty river, mid altitude forest, 21°30'07"S, 47°24'06"E, alt. 1075 m, 13 Oct. 2000, Rakotovao & Randriantafika 998 (K, MO, TAN).

habitat. Open humid forest and secondary vegetation; alt. 700 – 1700 m.

conservation status. Vulnerable (VU (B1ab i, iii, iv)). Phyllanthus ivohibeus has an EOO of 19706 km². This species is threatened by habitat loss due to fire and forest logging which lead to a decline in extent of occurrence, the quality of habitat, and the number of locations.

11. Phyllanthus vatovaviensis Leandri ex Ralim. & Petra Hoffm. sp. nov. Type: Madagascar, Fianarantsoa, Mont Vatovavy, Decary 13730 (holotype P(P00482889)!; isotypes P(P00078287, P00078288)!).

http://www.ipni.org/urn:lsid:ipni.org:names:77134149-1

Phyllanthus vatovaviensis Leandri (1958: 68), nom. invalid.

Monoecious shrubs 40 – 50 cm high with flexible branches. Branching pinnatiform or bipinnatiform; orthotropic branches terete, 0.5 – 2 mm in diam., striate, glabrous. Cataphylls triangular, c. 0.4 × 0.5 mm, coriaceous, entire, glabrous. Cataphyllary stipules triangular, c. 0.4 × 0.5 mm, coriaceous, entire, glabrous. Plagiotropic branches solitary, flexible, terete, 3 – 11 cm long, 0.3 – 0.5 mm in diam., glabrous, striate. Stipules persistent, triangular, 0.8 – 1.5 × 0.3 – 0.5 mm, membranous, entire, glabrous. Internodes 2 – 4 mm long. Leaves 9 – 25 per branch. Petioles terete, c. 0.4 × 0.2 mm, glabrous. Leaf blades elliptic to oblong (symmetric at the base), 8 – 14 × 5 – 6 mm, 1.6 – 2.3 times longer than wide, rounded or obtuse at the base, rounded at the apex, chartaceous, entire, slightly revolute, glabrous on both sides; midvein slightly prominent adaxially, flattened abaxially; secondary veins 3 – 5 pairs, flattened on both sides. Inflorescences unisexual, consisting of 2 or 3 male flowers or only 1 female flower; male and female inflorescences on different branches. Bracts triangular, grouped in a tuft of 9 – 12 mainly sterile bracts in male inflorescences, 0.2 – 0.4 × 0.2 – 0.3 mm, entire, glabrous. Male flowers 1 – 1.3 × 1 – 1.2 mm, whitish green when fresh. Pedicels 1 – 2 × 0.2 – 0.3 mm, glabrous. Tepals 5, inequal (the outer ones larger than the inner ones), obovate or suborbicular, 1 – 1.3 × 0.7 – 1.2 mm, rounded, membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₄ of total tepal width on each side, veins unbranched. Disc glands 5, lenticular, c. 0.3 × 0.2 mm, rugose or verrucose. Stamens 2, filaments entirely fused into a column; column terete, c. 0.6 × 0.3 mm; anthers entirely fused into a disciform circumscissile bilobed synandrium, dehiscing horizontally. Female flowers 1.8 – 2 × 2.2 – 2.8 mm, whitish green when fresh. Pedicels 2 – 3 × 0.2 – 0.3 mm (diam. increasing apically to 0.4 – 0.5 mm), glabrous. Tepals 5, equal, obovate or elliptic, 1.8 – 2 × 1 – 1.2 mm, rounded, membranous, glabrous on both sides, margin entire or slightly erose, hyaline for c. ¹/₆ of total tepal width on each side, veins branched. Disc ± flattened, irregularly lobed, smooth. Ovary 3-locular, sub-globose, 0.7 – 1.2 mm in diam., smooth, glabrous; styles fused at the base, bifid for c. ¹/₂ of their length, terete, 0.6 – 0.7 mm long; stigmas obtuse. Fruits sub-globose, 1.3 – 2 × 1.2 – 1.8 mm, glabrous; fruiting pedicels terete in the basal half, becoming flattened apically, 2.5 – 3 × 0.2 – 0.4 mm, glabrous; tepals c. 3 × 2 mm; styles caducous; columella c. 1 × 0.3 mm. Seeds c. 1.3 × 1 mm, finely longitudinally striate.

recognition. Phyllanthus vatovaviensis is similar to P. moramangicus in the bracts of the male flowers being tufted, the stamen filaments being entirely fused, the anthers fused into a disciform circumscissile synandrium but differs from P. moramangicus by the petiole c. 0.4 mm long (0.8 – 1 mm), the leaf blades oblong (vs ovate to lanceolate), leaf blade length up to 14 mm (vs up to 25 mm) and the apex rounded (vs acute or mucronulate).

distribution. Endemic to Madagascar: Eastern floristic domain. Map 11.

Map 11

Distribution of Phyllanthus vatovaviensis.

specimens examined. madagascar. Central Madagascar, Baron 1887 (P). Fianarantsoa. Mont Vatovavy, Decary 13730 (holotype P); Mont Vatovavy, Decary 13730 (isotypes P). Toamasina. Vatomandry, Ambalabe, 19°11'18"S, 48°33'6"E, alt. 650 m, 13 March 2005, Razanatsima et al. 9 (MO, P, TAN).

habitat. Humid forest and degraded humid forest; alt 100 – 650 m.

conservation status. Endangered (EN (B1abi-iv + B2ab i-iv)). Phyllanthus vatovaviensis has an EOO of 3158 km² and an AOO of 1875 km². All locations are found outside of protected areas. This species is mainly found in the east of Madagascar where the habitat is threatened by cut and slash agriculture and forest logging.

notes. Phyllanthus vatovaviensis is among the taxa cited by Leandri (1958) in Flore de Madagascar et des Comores. He referred to the first description of this species. However, after checking the journal we noted that no diagnosis of this species was included. Investigations into Leandri’s other publications have yielded no results. The species had not been validly published previously.

12. Phyllanthus lokohensis Leandri (1957: 229). Type: Madagascar, Antsiranana, Massif de Marojejy (Nord-Est), bassin de Lokoho Est d’Ambalamanasy II, alt. 600 m, Humbert & Capuron 22206 (holotype, P(P00078219)!; isotype P(P00535937)!).

Monoecious shrubs 40 – 100 cm high. Branching pinnatiform; orthotropic branches terete, 1.5 – 3 mm in diam., striate or smooth, glabrous. Cataphylls triangular, 1.3 – 1.5 × 0.4 – 0.6 mm, subcoriaceous, entire, glabrous. Cataphyllary stipules triangular, 1.3 – 1.5 × 0.4 – 0.6 mm, subcoriaceous, entire, glabrous. Plagiotropic branches solitary, terete, 4 – 15 cm long, 0.4 – 0.9 mm in diam., glabrous or pubescent, striate. Stipules persistent, triangular, 0.8 – 1.3 × 0.4 – 0.6 mm, membranous, entire or slightly erose, glabrous. Internodes 2 – 10 mm long. Leaves 7 – 20 per branch. Petioles terete, 1.2 – 1.5 × 0.3 – 0.5 mm, glabrous or rarely pubescent. Leaf blades elliptic to ovate (symmetric at the base), 5 – 18 × 5 – 12 mm, 1 – 1.5 times longer than wide, obtuse at the base and apex, membranous to chartaceous, glabrous on both sides, green or reddish green adaxially, reddish or greyish green abaxially when fresh; midvein flattened on both sides; secondary veins 3 – 7 pairs, slightly prominent adaxially, flattened abaxially. Inflorescences unisexual, consisting of 1 or 2 male flowers or only 1 female flower; male inflorescences on proximal part and female inflorescences on distal part of plagiotropic branches, or male and female inflorescences on different branches. Bracts triangular or elliptic, grouped in a tuft of 5 – 7 mainly sterile bracts in male inflorescences, 0.9 – 1.5 × 0.4 – 0.7 mm, dentate or erose, glabrous. Male flowers 2 – 2.5 × 2.5 – 4 mm, whitish green when fresh. Pedicels terete, 2.5 – 5 × c. 0.1 mm, glabrous. Tepals 5, equal, elliptic or obovate, 2 – 2.5 × 1.8 – 2 mm, rounded or obtuse, membranous, glabrous on both sides, margin entire, hyaline for ¹/₈ – ¹/₆ of total tepal width on each side, veins branched. Disc glands 5, reniform, 0.4 – 0.5 mm in diam., smooth. Stamens 3, filaments entirely fused into a column; column terete, 0.6 – 0.8 × c. 0.3 mm; anthers completely fused into a disciform circumscissile tri-lobed synandrium, dehiscing horizontally. Female flowers 3.5 – 5 × 4 – 8 mm, whitish green when fresh. Pedicels 3 – 7 × 0.1 – 0.2 mm, glabrous. Tepals 5, equal, elliptic or obovate, 3 – 4 × 2 – 3 mm, rounded, slightly coriaceous or membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₁₀ of total tepal width on each side, veins branched. Disc lobed, rugose or smooth. Ovary 3-locular, globose, 0.5 – 1 mm diam., on short gynophore 0.2 – 0.5 mm, smooth, glabrous; styles fused into a column for c. ¹/₄ of their length, bifid for c. ¹/₃ of their length, terete, 1.5 – 2 mm long; stigmas obtuse. Fruits globose, 1.5 – 2.5 mm in diam., glabrous; fruiting pedicels terete, 6 – 7 × c. 0.3 mm, glabrous; tepals 3 – 7 × 3 – 4 mm; styles persistent; columella 1 – 2 × 0.3 – 0.4 mm. Seeds 1 – 1.8 × 0.7 – 1 mm, smooth or thinly striate longitudinally, light green or yellowish when fresh. Fig. 3.

Fig. 3

Phyllanthus lokohensis. A habit; B leaf; C node; D male flower; E female flower. From G. Cours 4305 (TAN). drawn by r. l. andriamiarisoa.

distribution. Endemic to Madagascar; Central and eastern floristic domains. Map 12.

Map 12

Distribution of Phyllanthus lokohensis.

specimens examined. madagascar. Antsiranana. Massif de Marojejy (Nord-Est), bassin de Lokoho Est d’Ambalamanasy II, alt. 600 m, Humbert & Capuron 22206 (holotype P); Massif de Marojejy (Nord-Est), bassin de Lokoho Est d’Ambalamanasy II, alt. 600 m, Humbert & Capuron 22206 (isotype P); Andapa, Ambalamanasy II, Marojejy National Park, Betsiasisa forest, east of Antanimbaribe, 14°29'10"S, 49°34'02"E, alt. 865 m, 1 Feb. 2013, Ralimanana et al. 1752 (K, MO, P, TAN). Fianarantsoa. Réserve Spéciale d’Ivohibe, forest of Marovitsika, 22°28'49"S, 49°56'49"E, alt. 850 m, Oct. 2000, Hoffmann 224 (K, TAN). Toamasina. Brickaville, Anjahamamy, Anivoranokely, Ankerana forest, 18°24'12"S, 48°48'24"E, alt. 975 m, 25 Jan. 2012, Antilahimena 8041 (MO, TAN); Rahobevava, c. 1250 m, 12 March 1951, Herbier de Station Agricole Alaotra, Cours 4305 (TAN); Soanierana Ivongo, between Ambatovaky and Amberomanitra, 16°51'40"S, 49°17'20"E, alt. 350 m, 17 Dec. 2003, Ralimanana et al. 410 (G, MO, P, TAN); Soanierana Ivongo, between Amberomanitra and Ambatobe, 16°51'S, 49°17'E, alt. 348 m, 18 Dec. 2003, Ralimanana et al. 416 (K, MO, P, TAN); Brickaville, Maroseranana, Ambodilendemy, forêt d’Ankerana, 18°24'48"S, 48°47'32"E, alt. 1180 m, 23 March 2011, Ravelonarivo et al. 3817 (MO, TAN); Brickaville, Maroseranana, Ambatolampy, forêt d’Ankerana, 18°24'11"S, 48°47'04"E, alt. 1131 m, 1 Feb. 2012, Ravelonarivo & Felix 4264 (MO, P, TAN).

habitat. Humid forest; alt. 350 – 1250 m.

conservation status. Vulnerable (VU (B1abi-iv)). Phyllanthus lokohensis has an EOO of 13317 km² and an AOO of 4870.59 km². This species has a disjunct distribution. It is threatened by habitat loss due to human activities such as cut and slash which leads to a decline of area of occupancy, quality of habitat and number of locations or subpopulations.

13. Phyllanthus moramangicus (Leandri) Leandri (1958: 84). Phyllanthus melleri Müll. Arg. subsp. moramangicus Leandri (1938: 197). Type: Madagascar, Toamasina: Moramanga, forêt au sud de Moramanga, Decary 7185 (holotype P!).

Monoecious shrubs 40 – 60 cm tall. Branching pinnatiform; orthotropic branches terete, becoming slightly flattened apically, 2 – 3 mm in diam., striate, glabrous. Cataphylls triangular, 0.8 – 1 × 0.5 – 0.7 mm, coriaceous, entire, glabrous. Cataphyllary stipules triangular, 0.8 – 1 × 0.5 – 0.6 mm, coriaceous, entire, glabrous. Plagiotropic branches solitary or fascicled in twos, terete, 3.5 – 9.5 cm long, 0.5 – 0.7 mm in diam., puberulous or glabrous, striate. Stipules persistent, triangular, 0.8 – 1 × 0.4 – 0.6 mm, membranous, entire, glabrous. Leaves 7 – 13 per branch. Internodes 2 – 6 mm long. Petioles terete, 0.8 – 1 × 0.4 – 0.5 mm, glabrous. Leaf blades ovate to lanceolate (symmetric at the base), 8 – 25 × 5 – 7 mm, 2 – 4.1 times longer than wide, obtuse to attenuate at the base, acute to mucronulate at the apex, chartaceous, revolute, glabrous on both sides, green adaxially, green greyish abaxially when fresh; midvein prominent on both sides; secondary veins 3 – 7 pairs, slightly prominent adaxially, flattened abaxially. Inflorescences unisexual, consisting of 2 male flowers or only 1 female flower; male inflorescences and female inflorescences on different branches. Bracts triangular, grouped in a tuft of 8 – 15 mainly sterile bracts in male inflorescence, 0.5 – 0.8 × 0.3 – 0.4 mm, entire or slightly erose, glabrous. Male flowers 1.8 – 2 × 2 – 2.2 mm, whitish green when fresh. Pedicels 2 – 3 × c. 0.5 mm, glabrous. Tepals 5, subequal, obovate, 2 – 2.2 × 1 – 1.5 mm, rounded, slightly coriaceous or membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of tepal width on each side, veins unbranched. Disc glands 5, lenticular, c. 0.3 × 0.2 mm, rugose or verrucose. Stamens 2 (– 3); filaments entirely fused into a column; column terete, 0.4 – 1 × 0.3 – 0.4 mm; anthers entirely fused into a disciform circumscissile synandrium, bilobed or very rarely tri-lobed, c. 0.2 × 0.3 mm in diam., dehiscing horizontally. Female flowers 2 – 3 × 3 – 4 mm, whitish when fresh. Pedicels 5 – 8 × c. 0.4 mm, apically flattened and slightly thickened (up to 0.8 mm), glabrous. Tepals 5, inequal, ovate or obovate, 2 – 3 × 1.5 – 1.8 mm, rounded or obtuse, slightly coriaceous or membranous, glabrous on both sides, margin entire, hyaline for c. ¹/₈ of total tepal width on each side, veins branched. Disc lobed, verrucose. Ovary 3-locular, globose, 1.3 – 1.5 mm diam., smooth, glabrous; styles fused only at the base, bifid for c. ¹/₃ of their length, terete, 0.6 – 0.8 mm long; stigmas obtuse. Fruits depressed globose, 2 – 2.5 × 2 – 3 mm, glabrous; fruiting pedicels terete, apically slightly thickened, 5 – 10 × 0.4 – 0.8 mm, glabrous; tepals 2.5 – 3 × 1.5 – 1.8 mm; styles caducous; columella 1.2 – 1.4 × 0.3 – 0.4 mm. Seeds 1 – 1.3 × 1 – 1.2 mm, smooth. Fig. 4.

Fig. 4

Phyllanthus moramangicus. A habit; B leaf; C node with stipule, petiole and tuft of sterile bracts; D male flower; E female flower. From R. Ranaivojaona 556 (TAN). drawn by r. l. andriamiarisoa.

distribution. Endemic to Madagascar: Central floristic domain. Map 13.

Map 13

Distribution of Phyllanthus moramangicus.

specimens examined. madagascar. Toamasina. Moramanga, forêt au sud de Moramanga, Decary 7185 (holotype P). Road to Lakato, c. 14 km S of RN2, 19°01'43"S, 48°21'06"E, alt. 1040 m, 13 Nov. 2003, Lowry II et al. 6246 (MO, TAN); Aux environs de la Chutes de la Mort, PK 55 route de Moramanga Anosibe An’ala, 11 Dec. 1965, Services Forestiers 24397 SF (Capuron) (TEF); Andasibe, Morafeno Anevoka, forêt d’Andasihamaka, 18°57'55"S, 48°27'33"E, 29 Dec. 2002, Ranaivojaona 556 (TAN); Ambatovy, forêt de Berano, 18°51'35"S, 48°19'56"E, alt. 1070 m, Ranaivojaona 1046 (MO, TAN); Alaotra Mangoro, Andasibe, Parc d’Analamazaotra, Barika Amboasary, 18°56'46"S, 48°26'17"E, alt. 1071 m, 13 March 2013, Rasoazanany et al. 466 (MO, P, TAN).

habitat. Open humid forest, degraded humid forest and secondary vegetation.

conservation status. Endangered (EN (B1abi-iv + B2ab i-iv)). Phyllanthus moramangicus has an EOO of 187 km² and an AOO of 30 km². All locations are found outside protected areas. This species is threatened by human activities such as cut and slash, mining and annual fire.

III. Phyllanthus subgen. Emblica (Gaertn.) G. L. Webster sect. Urinaria G. L. Webster

Phyllanthus subgenus Phyllanthus section Urinaria G. L. Webster (1955: 51). Type of section: Phyllanthus urinaria L.

Monoecious herbs with pinnatiform phyllanthoid branching. Brachyblasts absent. Leaves alternate. Stipules entire. Inflorescences on leafy branches, axillary, unisexual, 1 to 2-flowered. Tepals 5 or 6. Disc: free glands in male flowers, annular in female flowers. Stamens 3; filaments entirely fused into a column; anthers basifixed, free, dehiscing longitudinally; pollen prolate, 4 – 5-colporate, exine tectate, scabrous and microperforate (Brunel 1987). Ovary 3-locular, tuberculate; styles free, bifid; stigmas obtuse. Fruits dehiscent; tepals persistent, not accrescent; seeds 2 per locule, deeply and coarsely transversally ribbed.

14. Phyllanthus urinaria L., Species Plantarum (Linnaeus 1753: 982). Type: Sri Lanka. Herb. Hermann 1: 15; 2: 7; 3: 55; 4: 41, No. 332 (lectotype BM!, selected by Fawcett & Rendle (1920: 255); Jarvis (2006)).

Monoecious herbs 10 – 40 cm high. Orthotropic branches terete, 0.4 – 0.6 mm in diam., rugose or smooth, glabrous. Cataphylls triangular, acute, 1.5 – 2.5 × 0.3 – 0.4 mm, subcoriaceous or chartaceous, with hyaline margin, entire, glabrous. Cataphyllary stipules triangular, auriculate at the base, 1 – 2 × 0.3 – 0.4 mm, membranous or chartaceous, entire, glabrous. Plagiotropic branches solitary, flattened and winged, 1 – 10 cm long, 0.3 – 0.4 mm in diam., rugose or smooth, glabrous or hirsutulous. Stipules persistent, triangular or lanceolate, 0.4 – 0.6 × 0.2 – 0.3 mm, membranous or chartaceous, entire, glabrous. Leaves 4 – 11 per branch. Internodes 1 – 2.5 mm long. Petioles terete, 0.4 – 0.6 × c. 0.3 mm glabrous. Leaf blades mostly oblong or obovate-oblong, 3.5 – 8 × 1.6 – 3 mm, 1.7 – 2.6 times longer than wide, rounded or obtuse and sometimes slightly asymmetric at the base, obtuse or acute and mucronulate at the apex, membranous, glabrous on both sides, margin slightly denticulate; midvein prominent on both sides; secondary veins 3 – 5 pairs, flattened or indistinct adaxially, prominent abaxially. Inflorescences unisexual, consisting of 1 – 2 male flowers on distal part of plagiotropic branches, and of 1 female flower on proximal part of plagiotropic branches. Bracts triangular or ovate, grouped in a tuft of 4 – 6 mainly sterile bracts in male inflorescences, 0.2 – 0.4 × 0.2 – 0.3 mm, entire, glabrous, whitish green when fresh. Male flowers 0.5 – 0.6 × 0.4 – 0.6 mm, greenish white when fresh. Pedicels 0.5 – 0.7 × c. 0.3 mm, glabrous. Tepals 5 or 6, equal, obovate or oblong, membranous, 0.3 – 0.6 × 0.3 – 0.4 mm, obtuse or acute, glabrous, margin entire, hyaline for ¹/₆ of total tepal width on each side, veins unbranched. Disc glands 5 or 6, globose. Stamens 3; filaments entirely fused into a column; column terete, 0.1 – 0.2 mm long, c. 0.1 mm in diam.; anthers free, ovoid, 0.3 – 0.5 mm long, dehiscing longitudinally. Female flowers 0.5 – 0.8 × 0.4 – 0.6 mm, whitish green when fresh. Pedicels 0.5 – 1 × c. 0.4 mm, glabrous. Tepals 5 or 6, equal or unequal, membranous, oblong or lanceolate, 0.5 – 0.8 × 0.3 – 0.4 mm, obtuse or acute, glabrous on both sides, margin entire, hyaline for ¹/₆ of total tepal width on each side, veins unbranched. Disc lobed, c. 0.1 mm thick. Ovary 3-locular, globose or depressed globose, 0.3 – 0. 5 × 0.3 – 0.5 mm, tuberculate, glabrous; styles 3, fused at the base, bifid for ²/₃ – ³/₄ of their length, flattened, 0.2 – 0.4 mm long; stigmas obtuse. Fruits depressed globose, c. 2 × 3 mm, glabrous; fruiting pedicels 0.6 – 1.3 × c. 0.4 mm; tepals 0.7 – 1 × 0.3 – 0.4 mm; styles persistent or caducous; columella 0.5 – 1 × 0.3 mm. Seeds 0.6 – 1.2 × 0.5 – 0.9 mm, dorsally and laterally with deep and coarse transversal ridges.

distribution. Native to Asia, introduced and becoming common in the tropics worldwide; in Madagascar: Eastern and Sambirano floristic domain. Map 14.

Map 14

Distribution of Phyllanthus urinaria.

specimens examined. sri lanka. Hermann 1: 15; 2: 7; 3: 55; 4: 41, No. 332 (lectotype BM). madagascar. s. loc., Nov. 1819, Perrier de la Bathie s.n. (P); 21 June 1866, Gerrard 72 (K). Antsiranana. Nosy Be, Hildebrandt 3156 (K, P). Fianarantsoa. Fort Carnot, 1986, Beaujard 77 (P). Toamasina. Ambatondrazaka, Herb. Station Agric. Alaotra 27646 (P, TAN); Soanierana Ivongo, Ambatovaky Reserve, 16°49'19"S, 49°10'14"E, 475 m, 10 Dec. 2003, Ralimanana et al. 350 (K, MO, TAN); Mahanoro, Ambodibonara, forêt littorale d’Ampamanara, 20°22'22"S, 48°33'56"E, 13 m, 14 May 2004, Ranaivojaona 611 (K, MO, TAN); Ankirindro massif, Maroantsetra, 15°19'20"S, 49°33'26"E, 100 m, 2 Feb. 1999, Schatz et al. 3936 (K, MO); Andevoranto, Anivorano, 300 m, 12 Feb. 1912, Viguier & Humbert 667 (P). comoro islands . mayotte . Grande Terre, Dembeni, Hauts Tsararano, 2 Aug. 2002, Barthelat 979 (K, Mayotte); Coconi, 12°49'87"S, 45°8'23"E, 100 m, 17 Nov. 2002, Ralimanana & Hoffmann 279 (K, TAN).

habitat in madagascar. Humid forest boundaries, littoral forest, fields; alt. 10 – 500 m.