FIVE nEw spEcIEs oF GuattErIa ( annonacEaE ) From FrEnch GuIana , Guyana and surInamE

Amongst the Neotropical genera of Annonaceae, Guatteria Ruiz & Pav. is one of the largest. Chatrou et al. (2004) reports 265 species, though the actual number may ex­ ceed this. The genus is easily recognised by the combination of axillary flowers/ inflo­ rescences with clearly visible articulation on the pedicel, an apocarpous fruit of stipitate monocarps and an impressed primary vein on the upper side of the leaf. The taxonomy of the genus is still based on the treatments of Fries (1939, 1941, 1943, 1948a, b, c, 1949, 1950a, b, 1952, 1953, 1957a, b, 1960), whose work was based on the relatively few collections available at that time, a situation which in recent years has greatly im­ proved. While accounts covering parts of the genus have appeared in several regional floras and checklists (e.g. Steyermark et al., 1995; Boggan et al., 1997; Hollowell et al., 2001), a new revision of this diverse and taxonomically challenging genus as a whole has yet to be made. First steps towards this goal can best be directed towards amply collected areas, providing a good reflection of the diversity present. The Flora of the Guianas project, coordinated by the Utrecht branch of the National Herbarium of the Netherlands, offered such an opportunity. The large number of specimens of Guatteria collected in the Guianas in the last 250 years and deposited at the Herbarium of the Utrecht branch of the National Herbarium of the Netherlands (U) and in parts on loan at the Herbarium of the Leipzig University (LZ) were the generous base for improved understanding of formerly insufficiently known taxa. Many recent collections of high quality, particularly from more remote places and/or from higher elevations, have until now been difficult or impossible to identify to known species, e.g. using Fries’ keys (1939). Such collections could not even be unambiguously assigned to any particular


INTRODUCTION
Amongst the Neotropical genera of Annonaceae, Guatteria Ruiz & Pav. is one of the largest.Chatrou et al. (2004) reports 265 species, though the actual number may ex ceed this.The genus is easily recognised by the combination of axillary flowers/inflo rescences with clearly visible articulation on the pedicel, an apocarpous fruit of stipitate monocarps and an impressed primary vein on the upper side of the leaf.The taxonomy of the genus is still based on the treatments of Fries (1939Fries ( , 1941Fries ( , 1943Fries ( , 1948aFries ( , b, c, 1949Fries ( , 1950aFries ( , b, 1952Fries ( , 1953Fries ( , 1957aFries ( , b, 1960)), whose work was based on the relatively few collections available at that time, a situation which in recent years has greatly im proved.While accounts covering parts of the genus have appeared in several regional floras and checklists (e.g.Steyermark et al., 1995;Boggan et al., 1997;Hollowell et al., 2001), a new revision of this diverse and taxonomically challenging genus as a whole has yet to be made.First steps towards this goal can best be directed towards amply collected areas, providing a good reflection of the diversity present.The Flora of the Guianas project, coordinated by the Utrecht branch of the National Herbarium of the Netherlands, offered such an opportunity.The large number of specimens of Guatteria collected in the Guianas in the last 250 years and deposited at the Herbarium of the Utrecht branch of the National Herbarium of the Netherlands (U) and in parts on loan at the Herbarium of the Leipzig University (LZ) were the generous base for improved understanding of formerly insufficiently known taxa.Many recent collections of high quality, particularly from more remote places and/or from higher elevations, have until now been difficult or impossible to identify to known species, e.g. using Fries' keys (1939).Such collections could not even be unambiguously assigned to any particular of Fries' sections within Guatteria, which according to recent work appear to include many non-monophyletic groupings (R.H.J. Erkens, pers.com.).Scharf et al. (2005) described five new species from the Pakaraima Mountains in Guyana.In advance of a revision of Guatteria (Annonaceae) for the three Guianas five more new species from the Guianas are described here.Diagnostic characters of the new species are listed below the respective descriptions.Measurements of the pedicel relate to the whole length of the flower stalk, including both the part below the articulation ('peduncle') and the part above the articulation ('pedicel s.str.') 1 .Coordinates, if not mentioned on the label, were inferred from printed books (Hoff &Cremers, 1996, The Lands andSurveys Department, 2001), maps (see list of references) as well as from sources on the internet (Geonames, Falling Rain).Scharf & Maas,; Map 1 Species foliis anthracinis, angustis, costa supra pilis longis ornatis distincta.-Typus: Lindeman, Stoffers et al. 429 (holo U;iso BBS, C, F, NY, S), Suriname, Lely Mts, 550-710 m, N 4° 25', W 54° 39', 26 Nov. 1975.= Guatteria aff.oblonga sensu Maas & Maas-van de Kamer (2002: 62); not R.E.Fr. = Guatteria sp. 2, in Ribeiro et al. (1999: 133).
Phenology -Flowering: June to January; fruiting: July to December.Etymology -This species has been named anthracina ('coal-black') after the very dark appearance of the dried leaves.
2. Guatteria procera R.E.Fr. is another species with dark drying leaves, but that species has a dense indument of erect, long, brownish hairs on all parts, except for the upper side of the lamina.
Phenology -Flowering: May to June; fruiting: June.Etymology -The species epitheton of 'G.elegans' refers to the slender and elegant appearance of its small leaves, which is caused by an extraordinary low number of secondary veins and an unusually sharp angle between primary vein and secondaries.The fruits are also quite small, and with very slender stipes.
Note -A sharp angle of 30-40° between primary vein and secondaries is not oc curring in any other Guatteria species in the Guianas and Brazilian Amazonia.Tree 20-30 m tall, 35-45 cm diam., with steep, simple, round buttresses; young twigs densely covered with appressed, short hairs, very soon glabrous.Leaves: petiole 4-5 by 1-1.5 mm; lamina narrowly elliptic to narrowly obovate, 9-14 by 2-3 cm, charta ceous to subcoriaceous, base attenuate, apex acuminate (acumen 10-15 mm long), margin recurved, dull and greyish green to brown above, brown below, densely covered with appressed, silvery, long hairs above, very soon glabrous, densely covered with appressed, silvery hairs below, secondary veins indistinct, slightly raised above, 8-13 on either side of primary vein, angle of secondary veins with primary vein 60 -70°, indistinctly loopforming.Flowers unknown, but following observations on fruiting specimens: solitary or in pairs on older leafless parts of whip-like branches; pedicels c. 10 mm long, sparsely covered with appressed, whitish hairs.Monocarps 5-20, ma turing dark purple in vivo, shiny black in sicco, ellipsoid to obovoid, 7-9 by 5-7 mm, apex rounded to minutely apiculate (apiculum < 0.5 mm long), sparsely covered with Phenology -Fruiting: September and December.Etymology -Guatteria leucotricha is named after its whitish or silvery long hairs at the lower side of the leaves.
Note -Guatteria leucotricha is highly typical by its narrow, long-attenuate leaves with recurved margins and by its indument of whitish or silvery hairs on the lower side of the lamina, a set of characters not met in any other Guianan species.Tree, 4-15 m tall; young twigs sparsely to densely covered with erect, long, dark brown hairs, soon glabrous.Leaves: petiole 3-7 by 1-1.5 mm; lamina narrowly elliptic to ovate, 5-10 by 2-3.5 cm, chartaceous, base acute, apex acuminate (acumen 5-15 mm long), leaden greyish black to whitish brown above, blackish to dark brown below, glabrous above, except for some incurved, blackish brown hairs along the primary vein, sparsely covered with erect hairs below, secondary veins indistinct, raised above, 7-10 on either side of primary vein (intermingling with intersecondaries), angle of secondary veins with primary vein 65-75°, loop-forming at acute to right angles, smallest distance between loops and margin 2-3 mm.Flowers solitary, rarely 2 per axil; pedicels 7-12(-20) mm long, densely covered with erect hairs; sepals triangular, 3-4 by 2-3 mm, apically reflexed, outer side densely covered with appressed, long, reddish brown hairs, inner side glabrous; petals ovate, c. 5 by 3 mm (probably immature), green in vivo (Henkel et al. 1156: black tipped in green), outer petals rather densely covered with appressed or curly, long hairs on both sides, inner petals on the outer side with a characteristic long-haired triangle in the centre, inner side glabrous or nearly so; stamens yellow in vivo and sicco, 90-110, c. 1 mm long, slightly umbonate, connective shield densely papillate; carpels 30-40, blackish, sparsely covered with very short, whitish hairs.Monocarps 10-30, green in vivo, black to dark reddish brown in sicco, ellipsoid, 9-12 by c. 5 mm, apiculate (apiculum < 1 mm long), glabrous, stipes purple to red when ripe, 5-15 mm long, very slender.Seeds shiny, maroon, slightly grooved, ellipsoid to ovoid, c. 8 by 5 mm.
Distribution -Central Suriname and Western Guyana.
Habitat & Ecology -In high altitude forest, on mountain tops or on slopes, together with typical Guayana Shield highland vegetation, on moist places (seepage bog), on white sand or sandstone with peat and grey sandy clay.Elevation 750-1500 m.
Phenology -Flowering: January to February; fruiting: January to March.Etymology -The specific epithet minutiflora was chosen regarding the small flowers present in the material available.
Notes -1.Guatteria minutiflora is very easily distinguishable from all Guianan species by its small, often blackish drying leaves and by its minute flowers with petals of not more than 5 by 3 mm.
2. The collections BW 5654 and BW 5711 were labelled G. cf.trichostemon R.E.Fr.G. trichostemon is a lowland species occurring further South in Western Amazonian Brazil, near São Paulo de Olivença.Morphologically, G. trichostemon is different by e.g. a glossy upper surface of the leaf, longer, very slender pedicels of 25-30 mm length, and a conspicuous bend in the middle of the pedicel which marks the articula tion.The latter feature was not observed in any collection from the Guianas.Tree, 4-25 m tall; young twigs densely covered with erect, curly, brown hairs, per sistent one growing period or even longer.Leaves: petiole 6-8 by 1-2 mm, distinctly canaliculate above; lamina narrowly oblong-elliptic, rarely elliptic, 10-20 by 2.5-4 cm, coriaceous, base acute, apex shortly acuminate (acumen 5-10 mm long), greyish green (in vivo dark green) above, pale brown (in vivo reddish) below, glabrous above, but primary vein with some wavy hairs, densely covered with appressed, brown hairs below, secondary veins distinct, prominent above, 10-15 on either side of primary vein, angle of secondary veins with primary vein c. 70°, loop-forming at right to obtuse angles, smallest distance between loops and margin 2-3 mm.Flowers solitary or in pairs; flower buds distinctly pointed, pedicels 8-10 by 1 mm, to 15 by 3 mm in fruit, densely covered with a felt-like indument of curly, brown hairs (in fruit sparsely so to glabrous), almost woody; sepals triangular, 8-10 by 5-6 mm, both sides densely covered with a felt-like indument of curly, reddish brown hairs; petals narrowly triangular, 20-25 by 6-8 mm, both sides densely covered with curly, reddish brown hairs, margins reflexed; stamens pale yellow in sicco, 200-250, c. 1.2 mm long, connective shield impressed, densely papillate; carpels arranged in a trigonal pyramid-like structure, c. 60, dark brown, sparsely covered with whitish, short hairs.Monocarps 25-30, maturing black ish green in vivo, dark to rusty brown in sicco, narrowly ellipsoid, 10 -14 by 6-7 mm, apex minutely apiculate (apiculum < 0.3 mm long), young monocarps densely, older ones sparsely covered with curly, brown hairs, stipes 20-25 by 1 mm, densely covered with curly, short, brown hairs.Seeds c. 12 by 5 mm, smooth, shiny, redbrown.Distribution -French Guiana and adjacent Brazil (Amapá).
Habitat and Ecology -In upland.
Phenology -Flowering: February; fruiting: August to October.Vernacular names -French Guiana: Baka pao (Saramaka, Boni), Pandiecou.Etymology -Guatteria pannosa has been so named because of its dense felt-like (pannose) indument of appressed to curly, reddish brown hairs on all parts except the upper surface of the leaf.
Notes -1.Guatteria pannosa is a very distinct species by its narrow and coriaceous leaves, pointed flower buds, narrow petals, and its felt-like indument.The monocarps are distinctively narrowly ellipsoid.To complete the description, two collections from adjacent Brazil, which doubtless belong to G. pannosa, are included.
2. Guatteria pannosa has a number of features in common with an as yet undescribed Panamanian species: coriaceous leaves, pointed flower buds, flowers' and fruits' general appearance, indument of erect and curly, brown hairs on the young twigs.Guatteria pannosa differs from the Panamanian species by its narrowly oblong-elliptic leaves (10-20 by 2.5-4 cm, vs narrowly elliptic to narrowly ovate, 20-23 by 7-8 cm), the indument on the lower side of the leaf (densely covered with long, appressed hairs vs sparsely covered with very short, appressed hairs or glabrous), and by its longer fruiting stipes (20-25 vs 9-14 mm).The Panamanian material (cf.list of specimens below) will be described soon, and further discussion will be given then (R.H.J. Erkens et al., in this volume).