Olivier Lachenaud
email: olivier.lachenaud@meisebotanicgarden.be

Piero G Delprete

Elmar Robbrecht

C Affinis

L Andersson

C Dolichophylla

C M Taylor

C Necopinata

C Panurensis

Müll Arg

C Adinantha

C Cardenasiana

C Verrucosa

C M Taylor Bahia

C Lucida

J G Jardim

Zappi

Revision of Carapichea (Rubiaceae-Psychotrieae) in the Guianas, with two new combinations and transfer of three species to Notopleura

Keywords: Carapichea, Psychotria, Uragoga, Stachyococcus, Guiana Shield, French

Background and aims -The genus Carapichea (Rubiaceae), recently resurrected and separated from Psychotria, currently includes 24 Neotropical species and is morphologically heterogeneous. A revision of the genus in the Guianas is presented here, as part of the authors' work on the Rubiaceae treatment in the Flora of the Guianas series. Material and methods -This paper is based on a study of herbarium specimens from BBS, BM, BR, BRB, CAY, INPA, K, P, P-JJR, and U; type specimens from other herbaria were consulted online. Some of the species were also studied in the field by the authors. Normal practices of herbarium taxonomy have been applied. Results -Three species previously included in Carapichea (C. altsonii, C. nivea, and C. sandwithiana) show aberrant characters for the genus and are transferred to Notopleura; new combinations are published for the former two (N. altsonii and N. nivea). Eight species of Carapichea, one of them still imperfectly known, are recorded from the Guianas; a ninth species, C. araguariensis, is included in this revision as it is expected to occur in the region. New combinations are published for two species: Carapichea galbaoensis and C. squamelligera, based on Psychotria galbaoensis and P. squamelligera, respectively; the former species was previously included in synonymy of C. guianensis, which is here redefined in a narrower sense. The delimitation of C. ligularis is expanded to include C. pacimonica as a synonym. The first records of C. adinantha in French Guiana -and the Guiana Shield -are documented. The taxonomy of the two remaining species, C. tillettii and C. urniformis, is left unchanged. For every species a complete description is presented, as well as data on distribution, ecology, phenology, local names (when known), and a list of collections studied.

INTRODUCTION

The genus Carapichea Aubl. (Rubiaceae) was described by Aublet (1775) with a single species from French Guiana, C. guianensis. For the following two centuries, most authors, with the exception of [START_REF] Candolle | Prodromus systematis naturalis regni vegetabilis[END_REF], did not accept the genus as distinct and included it successively in the synonymy of Cephaelis Sw. [START_REF] Willdenow | Caroli a Linné Species Plantarum[END_REF], Tapogomea Aubl. [START_REF] Poiret | Tapogomea carapichea[END_REF]Baillon 1880[START_REF] Baillon | The Natural History of Plants (English version)[END_REF], Uragoga Baill. [START_REF] Pulle | An enumeration of the vascular plants known from Surinam: together with their distribution and synonymy[END_REF], Cephaelis again [START_REF] Standley | Studies on American plants-II[END_REF][START_REF] Bremekamp | Notes on the Rubiaceae of Surinam[END_REF], and finally Psychotria L. [START_REF] Steyermark | Rubiaceae. In: Maguire B & coll. (Eds) The Botany of the Guayana Highland -Part IX[END_REF]. Based on molecular studies, [START_REF] Andersson | Re-establishment of Carapichea (Rubiaceae, Psychotrieae)[END_REF] resurrected and expanded the genus Carapichea, which, according to him, "differs from Psychotria (sensu [START_REF] Petit | Les espèces africaines du genre Psychotria L. (Rubiaceae) -I[END_REF] and most other members of the Psychotria complex in having the combination of non-caducous stipules, pyrenes with distinct germination slits on abaxial ridges (rather than marginally), and testa without ethanol-soluble pigment". [START_REF] Jardim | Carapichea lucida (Rubiaceae: Psychotrieae), a new species from eastern Bahia, Brazil[END_REF] described a new species from north-eastern Brazil, C. lucida, which is unusual in its deeply bifid stipules and pyrenes with a basal-ventral germination slit. [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] further expanded the delimitation of Carapichea and published a synopsis of the genus. They described the generic diagnostic characters as follows "a characteristic marcescent stipule form, inflorescences that are capitate or branched to one or two orders with the flowers sessile to shortly pedicellate, lack of ethanol-soluble pigments in the testa, and aperturate pollen of a generalized form" (Taylor and Gereau 2013: 107). Based on these characters, they transferred numerous species to Carapichea, most of them from Psychotria subg. Heteropsychotria Steyerm. and the monospecific genus Stachyococcus Standl., and also described a new species. They recognized 23 species, all Neotropical, and divided the genus into seven informal groups: I. Ipecacuanha group; II. Carapichea group; III. Pacimonica group; IV. Altsonii group; V. Panurensis group; VI. Stachyococcus group; and VII. Bahia group. One new species, Carapichea cardenasiana, was later published [START_REF] Taylor | Rubiacearum Americanarum Magna Hama Pars XXXIX. New species of Carapichea and Rudgea (Palicoureeae) from Western Amazonia[END_REF], bringing the total to 24.

Since then, field and herbarium work by the authors -partly in the context of the Rubiaceae treatment for the Flora of the Guianas -allowed to critically compare our data with the literature. This has led us to reevaluate the infrageneric classification of [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF], the generic position of some species, and the delimitation of others, while significant range extensions have also been recorded. This paper presents new findings in the framework of a revision of Carapichea in the Guianas, providing two new combinations in Carapichea and two in Notopleura. A key to the Guianan species of Carapichea is presented, and synonymy, description, distribution, and specimens studied are provided for the species.

MATERIAL AND METHODS

This paper is based on a study of herbarium specimens from BBS, BM, BR, BRB, CAY, INPA, K, P, P-JJR, and U, as well as images of type specimens from B, F, G, K, MPU, NY, US, and VEN available at JSTOR Global Plants (https://plants.jstor.org/) and images from COL (http://www.biovirtual.unal.edu.co/en/collections/ search/plants/). We also consulted images available at speciesLink (https://specieslink.net/) and REFLORA Virtual Herbarium (http://reflora.jbrj.gov.br/reflora/ herbarioVirtual/). Carapichea adinantha, C. galbaoensis, C. guianensis, and C. ligularis were also studied in the field in French Guiana and Brazil. Descriptions are based on herbarium specimens, alcohol-preserved material, and field observations. All specimens cited have been examined, unless indicated by "n.v. ". For accurate citation of type specimens, the barcode number is cited between square brackets, or, when not available, the accession number is cited, preceded by "No. "

RESULTS AND DISCUSSION

The delimitation of Carapichea and transfer of three species to Notopleura

The revision of the genus Carapichea in the Guianas led us to reevaluate the delimitation of the genus, and in particular the position of the species included by [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] in their Altsonii group (Group IV). This group included four species from western Guyana and eastern Venezuela: C. altsonii (Sandwith) C.M.Taylor, C. nivea (Sandwith) C.M. Taylor, C. sandwithiana (Steyerm.) C.M.Taylor, and C. urniformis.

These four species have always been regarded as closely related. [START_REF] Steyermark | Rubiaceae. In: Maguire B & coll. (Eds) The Botany of the Guayana Highland -Part IX[END_REF] placed them in an infrageneric group of their own, Psychotria sect. Notopleura Benth. ser. Altsonii Steyerm., characterised by 1) terminal capitate inflorescences with conspicuous involucral bracts, 2) pedicellate flowers lacking bracts or bracteoles but subtended by glandular projections from the receptacle, and 3) stipules forming a truncate sheath (which is actually not the case in C. urniformis). [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] noted some differences between C. urniformis and the other three species, and the similarity of the latter with Notopleura, but opted to place all four species in Carapichea due to their terminal inflorescences, and "interpetiolar rather than sheathing stipules that lack glandular appendages" (Taylor and Gereau 2013: 107) as opposed to lateral inflorescences and stipules often sheathing, bearing glandular appendages, in Notopleura.

After a reexamination of herbarium material, we agree with [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] in positioning C. urniformis in Carapichea. This species has free stipules without appendages (actually not glandular), coriaceous leaves with conspicuous secondary and tertiary venation, and semi-circular pyrenes in cross-section; all these characters are shared with the type species C. guianensis. Therefore, C. urniformis appears to be correctly placed in this genus, in spite of its unique lyrate involucre.

The remaining three species of [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] Group IV, on the other hand, show several characters aberrant in Carapichea: 1) very fleshy stems and leaves, the latter with secondary venation not or hardly distinct (i.e. embedded within the lamina), 2) pyrenes dorso-ventrally flattened, and 3) stipules with basal interpetiolar tufts of appendages (early caducous, leaving distinct scars). These stipules are further unusual in being remarkably dimorphic: those of vegetative nodes consist of a truncate sheath with interpetiolar tufts of appendages, while those of flower-bearing nodes consist only of tufts of appendages, without a sheath. This very peculiar stipular dimorphism, which seems not to have been previously documented in Rubiaceae, also exists in Notopleura tapajozensis (Standl.) Bremek. (Fig. 1).

All above discussed characters of C. altsonii, C. nivea, and C. sandwithiana point to an affinity with Notopleura, a genus where interpetiolar stipular appendages are usually present, as well as fleshy leaves and dorsoventrally flattened pyrenes. The leaves of the Altsonii group closely resemble those of three Notopleura species, N. aneurophylla (Standl.) C.M.Taylor, N. aneurophylloides (Steyerm.) C.M.Taylor, and N. tapajozensis, in their obsolete or barely visible secondary venation (i.e. embedded within the lamina). Notopleura tapajozensis further resembles C. altsonii and similar species in its remarkable stipular dimorphism and in having terminal inflorescences and a bipartite disk (the latter was not checked in N. aneurophylla and N. aneurophylloides). These characters are unusual in Notopleura, which has the disk usually entire and lateral inflorescences, although some epiphytic species may have terminal inflorescences. The generic placement of N. tapajozensis has been confirmed by a phylogenetic study [START_REF] Bruniera | Sistemática e taxonomia de Rudgea Salisb[END_REF]: figures 1, 2), where it was found in a basal position in Notopleura (although the species sampling is very incomplete). The difference between the paniculate inflorescences with minute bracts of N. tapajozensis and the involucrate heads of C. altsonii, C. nivea, and C. sandwithiana is great, but in other characters their resemblance is so clear that a close affinity of these taxa leaves little doubt. Furthermore, N. aneurophylla and N. aneurophylloides show an intermediate condition between N. tapajozensis and the three species with involucrate solitary heads, in having a tricapitate inflorescence with large bracts.

We therefore transfer Carapichea altsonii, C. nivea, and C. sandwithiana to Notopleura. New combinations are necessary for the first two names, while N. sandwithiana (Steyerm.) C.M.Taylor is already available. These three species are placed in Notopleura subg. Notopleura, where the terrestrial species are grouped, while the epiphytic species are included in N. subg. Viscagoga (Baill.) C.M.Taylor [START_REF] Taylor | Overview of the Neotropical genus Notopleura (Rubiaceae: Psychotrieae), with the description of some new species[END_REF].

Revision of Carapichea in the Guianas

In the generic revision of [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF], six species of Carapichea were recorded from the Guianas: C. altsonii, C. guianensis, C. ligularis, C. nivea, C. tillettii, and C. urniformis. As a result of our work, and despite the exclusion of C. altsonii and C. nivea, the number of Carapichea species in the Guianas has increased to eight. A ninth species, C. araguariensis, though not recorded from the Guianas to date, is expected to occur there and is consequently included in this revision.

New combinations are made for two species, C. galbaoensis and C. squamelligera, based on Psychotria galbaoensis Steyerm. and P. squamelligera Steyerm., respectively. The former species was so far treated as a synonym of C. guianensis [START_REF] Delprete | Notes on some South American species of Psychotria subgenus Heteropsychotria (Rubiaceae), with observations on rubiaceous taxonomic characters[END_REF][START_REF] Delprete | Carapichea guianensis: the correct name for Psychotria carapichea (Rubiaceae), with a correction on the authority of P. carapichea, and a new combination in Carapichea[END_REF][START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF], which is here circumscribed in a narrower sense. The affinities of P. squamelligera were so far unclear and it is transferred to Carapichea due to its similarities with C. galbaoensis and C. guianensis. The first records of C. adinantha in French Guiana, which is a major range extension westwards, are documented. One species from Guyana is still imperfectly known and is treated here under the provisional name C. sp. A. Finally, the delimitation of C. ligularis is expanded to include C. pacimonica (Müll.Arg.) C.M.Taylor as a synonym.

The genus Carapichea now includes 22 species, plus an imperfectly known one. A closer study of Carapichea species in the Amazon basin, which is outside the scope of this paper, is likely to result in further taxonomic changes and novelties. In particular, it is likely that C. franquevilleana and C. klugii are synonymous; [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] already suggested this but provisionally kept them as distinct. TAXONOMIC TREATMENT Carapichea Aubl. (Aublet 1775: 167) [START_REF] Johansson | Pollen morphology in Psychotria (Rubiaceae, Rubioideae, Psychotrieae) and its taxonomic significance. A preliminary survey[END_REF]. The genus is rather variable in inflorescence and pyrene characters, as described by [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF]. Most species of Carapichea have a bipartite disk. This character was not emphasized by previous authors and is often not even mentioned in their descriptions, though it is well depicted on illustrations of C. tillettii (Steyermark 1972: 496, figure 70), C. urniformis (Steyermark 1972: 557, figure 74), and C. vasivensis and C. pacimonica (Steyermark 1974, figures 234, 240) [START_REF] Lachenaud | Six new and a little-known species of Rudgea (Rubiaceae -Palicoureeae) from the Guianas[END_REF].

Position within the tribe and delimitation of Carapichea.

Carapichea has been included in the Palicourea complex of the tribe Psychotrieae s.l. by [START_REF] Andersson | Re-establishment of Carapichea (Rubiaceae, Psychotrieae)[END_REF] and [START_REF] Lachenaud | Révision du genre Psychotria (Rubiaceae) en Afrique Occidentale et Centrale[END_REF]. This complex has been treated as the tribe Palicoureeae by [START_REF] Robbrecht | The major evolutionary lineages of the coffee family (Rubiaceae, angiosperms). Combined analysis (nDNA and cpDNA) to infer the position of Coptosapelta and Luculia, and supertree construction based on rbcL, rps16, trnL-trnF and atpB-rbcL data. A new classification in two subfamilies, Cinchonoideae and Rubioideae[END_REF], [START_REF] Razafimandimbison | Phylogeny and generic limits in the sister tribes Psychotrieae and Palicoureeae (Rubiaceae): evolution of schizocarps in Psychotria and origins of bacterial leaf nodules of the Malagasy species[END_REF], and [START_REF] Taylor | Rubiacearum Americanarum Magna Hama Pars XXXIX. New species of Carapichea and Rudgea (Palicoureeae) from Western Amazonia[END_REF] Description. Shrub or small tree up to 4 m tall, sparsely branched; terminal branchlets terete or slightly laterally compressed, 2.5-3.0 mm in diam., glabrous, soon covered with a greyish-beige thin bark. Stipules free, interpetiolar, 2-5 mm long, bifid at apex, each lobe deltoid to lanceolate, 1-2 mm wide at base, acute to acuminate, glabrous. Leaves with petioles 1.5-3.0 cm long, glabrous; blades elliptic, oblong-elliptic oblong-lanceolate, 19-34 × 5.5-15 cm, acute at base, acuminate at apex, chartaceous to papyraceous when dry, entirely glabrous, drying olivegreen; midrib and secondary veins slightly prominent on the upper side; secondary veins 9-12 on each side of the midrib, ascending and forming loops away from the margin; intersecondary veins 2-3 between each couple of secondary veins, terminating far from the margin; tertiary veins reticulate, sparse. Inflorescences terminal, erect, spiciform, 7-20 cm long, with (6-)7-9 multiflorous glomerules, rachis minutely puberulous; peduncle 1.0-8.7 cm long; bracts subtending each glomerule shallowly ovate to broadly deltoid, 1-2 mm long. Flowers 5-merous, sessile (heterostylous?). Hypanthium glabrous or sparsely puberulous. Disk 5-sulcate laterally. Calyx cupuliform, 1.0-2.2 mm long, minutely puberulous, margin truncate, undulate or shallowly denticulate, ciliate. Corolla hypocrateriform, white; tube cylindrical, 4.5-7.0 mm long, 1.5-1.7 mm wide, granular-puberulous outside, glabrous inside; lobes imbricate, 3.5-4.5 × 1.4-1.7 mm, reflexed at anthesis, internally appendiculate, granularpuberulous outside, glabrous inside; appendages linear, 1.5-2.0 mm long. Stamens inserted at 1.0-1.5 mm from base of corolla tube, included; filaments 1.1-1.3 mm long; anthers linear, 2.3-2.5 × 0.2-0.3 mm. Style as long as corolla tube (branch tips barely exserted), ca 4.5-7.0 mm long, style branches linear, 1.3-1.5 mm long. Fruits drupaceous, fleshy, red to pink when immature (one collection reported as "white"), turning dark blue to black at maturity, interior spongy-white, ellipsoid to subglobose, 13-14 × 9-10 mm when fresh, 7-16 × 5-12 mm when dry, glabrous, sessile. Pyrenes 2, thin-walled, plano-convex, elliptic to narrowly elliptic in outline, 9-10 × 7-8 mm, dorsal side smooth (i.e. without ridges), ventral side flat (without longitudinal furrow), with a pore with a raphal plug, sometimes with short basal marginal slits. Seeds with a deep T-shaped ventral groove in crosssection.

Distribution. This species occurs sporadically in Amazonian Peru, Colombia, Brazil (Acre, Amapá, Amazonas, Pará, Rondônia), and French Guiana. Ecology. In undercanopy of terra firme primary and secondary forest, often at river margins, on clay soil, at 100-200(-300) Notes. This species has recently been collected in French Guiana and the adjacent Brazilian state of Amapá, which represents a major eastern range extension. Although this species is known from numerous specimens, most of them are fruiting and we were able to dissect only a few flowers. In these flowers, the stamens are inserted at 1.0-1.5 mm from base of corolla tube and are included, while the style is as long as the corolla tube, with the branch tips barely exserted. Though we have not seen a short-styled form, we suspect the species may be heterostylous, as many of its congeners.

2. Carapichea araguariensis (Steyerm.) C.M.Taylor (Taylor and Gereau 2013: 120) Psychotria araguariensis Steyerm. (Steyermark 1972: 588) Type. BRAZIL -Amapá • Araguari River, Camp 12; 1°11'N, 52°08'W; 30 Sep. 1961;fl.;Murça Pires et al. 51371;holotype: NY [00132598]. Description. Shrub, 0.5-1.0 m tall, or treelet 2-5 m tall, glabrous; terminal branchlets terete, 2-4 mm in diam., soon covered with a greyish bark. Stipules shortly sheathing, truncate to very broadly ovate, 1-4 × 4.0-5.5 mm, entire and obtuse at apex, glabrous, soon corky and fragmenting. Leaves with petioles 0.7-2.5 cm long, glabrous; blades elliptic, to oblong-oblanceolate, 11-25 × 3.5-7.5 cm, acute-decurrent at base, acute and acuminate at apex, acumen narrowly triangular, 0.5-1.5 cm long, subcoriaceous to coriaceous when fresh, drying papyraceous to subcoriaceous, greygreen to olive brown, glabrous throughout; secondary veins 13-25 on each side of midrib, weakly ascending and hardly more prominent than the intersecondary veins, arching at 0.5-1.0 mm from the margin; intersecondary veins (1-)2-3 between each couple of secondary veins, terminating far from the margin; tertiary veins obsolete; domatia absent. Inflorescence compact-paniculate at early stage, expanding and becoming obviously paniculate at later stage, many-flowered, pedunculate; peduncles 4.5-9.5 cm long, glabrous or with distal portion puberulous, drying olive green to pale brown; secondary branches verticillate, 2-4 per node, 0.7-3.0 cm long, glabrous to puberulous, terminating into cymules; outer branches of cymules with bracts; bracts 2-5 in each cymule, subequal to unequal, narrowly elliptic to linear, longer ones 7-17 × 1-2 mm, shorter ones 4.0-4.5 × 0.7-1.2 mm, persistent or tardily caducous, drying olive green to brown, glabrous. Flowers 5-merous, (heterostylous?), sessile or with pedicel < 0.5 mm long during anthesis, elongating to 1-5 mm long at fruiting stage. Hypanthium narrowly obovoid, 0.7-1.0 mm long, glabrous. Disk bilobed to the base, 0.5 mm long, glabrous. Calyx cupular, 0.7-1.3 mm long, truncate or minutely denticulate, glabrous. Corolla infundibuliform, 14.5-16 mm long, glabrous, white ("orange" according to Pires et al. 51371), tube narrowly obconical, 11-12 mm long, 1.5 mm wide at base, 3.5-4.0 mm wide at mouth, glabrous outside and inside; lobes lanceolate, 3.0-3.5 × 1.2-1.3 mm, acute at apex, glabrous, bearing dorsal linear cornicula 0.3-0.7 mm long. Stamens inserted just below the corolla mouth, filaments 0.5 mm long, anthers subsessile, half-exserted, narrowly oblong, 2.3-3.0 × 0.3-0.6 mm. Style glabrous, barely exserted, 14.5-16.5 mm long. Fruits elliptic to ovoid, 6.5-10 × 5.5-8.5 mm, slightly costate when dry, green when young, orange or yellow at maturity. Pyrenes plano-convex, ellipsoid in outline, 6-9 × 4.5-5.5 mm, dorsal side with 3 prominent longitudinal ridges, ventral side with a shallow longitudinal groove. Seeds with a deep T-shaped ventral furrow. Distribution. Only known from northern Brazil (Amapá, Pará, and Amazonas states); to be expected in French Guiana. Ecology. In understory of moist, non-flooded forest, at 50-125 m elevation. Phenology. The flowering type specimen from Amapá state was collected in September; in Amazonas state, flowering specimens were collected in February and March, and fruiting specimens in February and August. Selected specimens examined. BRAZIL -Amazonas • Reserva Florestal Ducke, Estrada Manaus-Itacoatiara, km 26, floresta de campinarana; 2°53'S, 59°58'W; 13 Feb. 1996;fl., fr.;Campos et al. 481 Notes. No records of this species are known from the Guianas to date, but due to its occurrence in the Brazilian state of Amapá, it may well be found in adjacent French Guiana, and is therefore included in this treatment. Carapichea araguariensis resembles C. ligularis but has laxer inflorescences with the basal bracts inserted at the end of the secondary branches (vs at the end of the peduncle). The bracts are also usually smaller and more distinctly unequal than in C. ligularis, and the corolla tube is longer. This species also resembles C. necopinata from Brazil (Amazonas state), but the latter has an inflorescence with only three glomerules and outer bracts 20-25 mm long, while in C. araguariensis the inflorescence has at least five glomerules and outer bracts 7-17 mm long. See also the notes under C. sp. A below. Due to the rarity of flowering specimens, it is not known whether the flowers are heterostylous. In the few flowers that we analyzed, the stamens are included and the style is barely exserted, which is consistent with a long-styled form. Ecology. This species is locally frequent in the understory of terra firme rainforest, at 100-800 m elevation.

Carapichea galbaoensis

Phenology. This species flowers during the small dry season (March-early April) and produces fruits from mid-April to September. Specimens examined. FRENCH GUIANA 6). [START_REF] Delprete | Notes on some South American species of Psychotria subgenus Heteropsychotria (Rubiaceae), with observations on rubiaceous taxonomic characters[END_REF] showed that they represent separate species. [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF], in their revision of the genus, also recognized the two species as distinct, and even placed them in different groups, chiefly on account of their strikingly different leaf venation. On the other hand, C. guianensis closely resembles C. galbaoensis, which was until now considered a synonym [START_REF] Delprete | Notes on some South American species of Psychotria subgenus Heteropsychotria (Rubiaceae), with observations on rubiaceous taxonomic characters[END_REF][START_REF] Delprete | Carapichea guianensis: the correct name for Psychotria carapichea (Rubiaceae), with a correction on the authority of P. carapichea, and a new combination in Carapichea[END_REF][START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] and is here re-established as a distinct species, and C. squamelligera. The three species are almost identical in vegetative characters, and their differences are summarised in Description. Subshrub or shrub, 0.2-2(-4) m tall, singlestemmed or weakly branched (or rarely trees 7-10 m tall in the Brazilian and Venezuelan Amazon); terminal internodes terete, 2-3(-4) mm in diam., glabrous. Stipules shallowly sheathing, glabrous; sheath truncate to shallowly elliptic; lobes broadly triangular to broadly ovate, 2-5 mm long, persistent. Leaves with petioles 0.3-3.3 cm long, glabrous; blades elliptic, oblanceolate to narrowly oblongelliptic, (4-)8.5-21 × (1.5-)2-6.5 cm, acute-decurrent at base, acute and long-acuminate at apex, acumen narrowly triangular, 0.5-2.5 cm long, sometimes falcate, papyraceous to coriaceous, drying pale olive-green to dark grey-green, glabrous throughout (except sometimes the domatia); secondary veins 14-26 on each side of the midrib; intersecondary veins 2-3(-4) between each couple of secondary veins, terminating far from the margin; tertiary veins barely visible or obsolete; domatia absent, or present as a row of hairs on each side of the midrib below. Inflorescence capitate to subcapitate (shortly branched, ramifications ≤ 0.5 cm when present); peduncle 1.5-7.0 cm long, glabrous to shortly pubescent; bracts inserted at distal end of peduncle and sometimes also on inflorescence axes, shallowly connate or free at base, (2-)4-8, usually decussate, narrowly lanceolate to linear, (7-)12-30 × (1-)2-4 mm, pale green or pale orange-green, glabrous, persistent. Flowers 5-merous, heterostylous, sessile to subsessile, the pedicels elongating to 2-5 mm long at fruiting stage. Hypanthium obovoid, 0.6-1.0 mm long, glabrous. Disk bilobed to the base, 0.5 mm long, shorter than or as long as the calyx. Calyx cupular, glabrous, tube 0.5-1.0 mm long, truncate or with short triangular lobes < 0.5 mm long. Corolla hypocrateriform, orange to salmon colored, 7.5-13(-14) mm long, glabrous; tube subcylindrical, gradually wider towards the mouth, 6-10.4(-11) mm long, 0.8-1.0 mm wide at base, 1.5-1.6 mm wide at mouth, glabrous outside and inside; lobes oblong-ovate, 2-4 × 0.8-0.9 mm, acute at apex, not corniculate, glabrous. Short-styled flowers: Stamens inserted at distal 1/3 rd of the tube, anthers partially exserted beyond corolla mouth; filaments ca 0.3 mm long; anthers 2.3-2.5 × 0.5 mm; style included, 3.5-4.0 mm long; style branches narrowly oblong, 1.5 mm long. Longstyled flowers: stamens inserted at about the middle of the corolla tube, included; filaments 1.5 mm long, glabrous; anthers narrowly oblong, 2.5 × 0.3-0.4 mm, acute at both ends; style exserted just beyond the corolla mouth, 12.5 mm long (corolla tube 10.5 mm long), style branches obovate, ca 1 mm long. Fruits ellipsoid to oblong-ovoid, 6-12 × 4-9 mm, smooth (slightly costate when dry), orange to orange-red. Pyrenes plano-convex, elliptic in outline, 7-9 × 3-5 mm, dorsal side with 3 ridges, ventral side with a very shallow longitudinal groove. Seeds with a deep T-shaped ventral furrow. Distribution. This species occurs in southeastern Venezuela (state of Amazonas), French Guiana, and eastern Amazonian Brazil (states of Amapá, Pará, Roraima, and Amazonas). It is locally common, at least in French Guiana. It could be expected in Suriname and Guyana, but we have seen no collections of the species from these countries; the type specimen, reported to be from Guyana (Delprete 2001: 399;Taylor and Gereau 2013: 120) Notes. This species has often been confused with C. guianensis (see Notes under that name) but is very different in leaf venation and bract shape [START_REF] Delprete | Notes on some South American species of Psychotria subgenus Heteropsychotria (Rubiaceae), with observations on rubiaceous taxonomic characters[END_REF]Figs 5, 6). It is much more similar to C. araguariensis and C. sp. A, from which it differs by the characters mentioned in the key. Delprete (2001: 401) separated Carapichea ligularis from C. pacimonica by the lower habit, being a small, singlestemmed shrub 0.5-1.5 m tall (vs shrub up to 2 m tall or tree up to 10 m tall), the presence of corniculate appendages on the abaxial side of the corolla lobes (vs appendages absent), the subcapitate to sparsely cymose inflorescences (vs capitate), and the larger ovoid fruits (vs smaller, subspherical fruits). None of these characters proves to be reliable: the variation in habit and fruit seems to be continuous, the corolla appendages are commonly present in flower buds and usually fall off during or shortly after anthesis or during the preparation of herbarium specimens (Piero Delprete pers. obs.) and the inflorescences are usually capitate or nearly so at anthesis and often develop short ramifications in the fruiting stage. Taylor and Gereau (2013: 114) also kept the two species separate, based on other characters: "outermost inflorescence bracts at base straight-sided and not sheathing; plants often drying with a gray cast" in C. ligularis vs "outermost inflorescence bracts at base widened and shortly sheathing; plants often drying yellowish brown" in C. pacimonica. After a detailed study of numerous specimens throughout the geographic range, we observed that the inflorescence bracts can be shallowly sheathing or free at base in duplicate specimens of the same collection, and the colour of the dry plants may vary from greyish to olive green to pale brown to dark brown (possibly depending on the drying method). Taking into account all the above observations, we treat these two names as synonyms. Steyermark (1972: 589) regarded Psychotria necopinata Standl. as a synonym of P. pacimonica, but we follow [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] in recognizing the former as a distinct species, Carapichea necopinata, which is only known from the type and not recorded from the Guianas to date. The typification of C. ligularis has been a source of confusion. In his original description, Rudge (1806: 29, pl. 45) did not cite any gathering. Delprete (2001: 399) designated the lectotype as "Guyana, Rudge s.n. (P)", which was followed by Taylor and Gereau (2013: 120). However, this lectotypification is erroneous since no such specimen exists at P, and Rudge never travelled to South America; the material that he used to describe the taxa in his Plantarum Guianae Rariorum Icones et Descriptiones was in fact collected by Joseph Martin in French Guiana. The fate of Martin's specimens was described by [START_REF] Stearn | Martin's French Guiana plants and Rudge's "Plantarum Guianae Rariorum Icones[END_REF], and summarized by Stafleu and Cowan (1983: 971-972). Succinctly, in 1803, France and England were at war, and the specimens collected by Joseph Martin in French Guiana, originally intended for the Museum of Natural History of Paris, were confiscated by two British privateers and brought to London. About 400 of these specimens were bought by the British Museum (BM), and 772 were bought by Rudge and included in his own herbarium (Stafleu and Cowan 1983: 971-972). Also, Rudge gave a partial set of these specimens to Banks, whose herbarium became the founding collection of BM. After Rudge's death, his widow donated his herbarium to BM in 1847. Therefore, most of Martin's collections converged at BM, although some of his specimens are also reported to be at FI or FI-Webb (Stafleu and Cowan 1983: 971-972). These specimens have the penciled information "Guiana. Martin", which may give the false impression that they were collected in modern day Guyana, while they actually came from French Guiana. After an exhaustive search for original material of S. ligularis, we were unable to find any specimen at FI and FI-Webb. However, a specimen with the indication "Guiana, Martin" penciled on the upper corner of the sheet, exists at BM [BM001009103], and being the sole original material, should be regarded as the holotype. Müller (1881: 338), in the original description of Psychotria pacimonica, cited a single gathering, Spruce 3445, but he did not indicate the herbarium of deposit. Taylor and Gereau (2013: 120) assumed that the holotype is in M, but no specimen of Spruce 3445 was found there after an exhaustive search (Andreas Fleischmann pers. comm.). Specimens with this collection number are found in G, K, P, and W; the P sheet, which is the only one with open flowers, and has a label handwritten by Müller, is here designated as lectotype. The photograph of the fruits published by Campos and Brito (1999: 627) as Psychotria pacimonica appears to represent another species, probably not a Carapichea.

6. Carapichea sp. A.

Description. Shrub, up to 1 m tall, glabrous; terminal internodes terete, 3-4 mm in diam. Stipular sheath truncate to shallowly ovate, 2-4.5 mm long, glabrous, persistent. Leaves with petioles 1.6-2.7 cm long, glabrous; blades narrowly oblong-elliptic, oblong-lanceolate to oblongoblanceolate, 15-25 × 3.5-6.5 cm, acute-decurrent at base, acute and acuminate at apex, acumen narrowly triangular, 1.2-1.3 cm long, drying papyraceous and brown, glabrous throughout; secondary veins 17-22 on each side of midrib, with 2-3 intersecondary veins between each pair of secondary veins; tertiary venation reticulate; domatia absent. Inflorescence terminal, longpedunculate, distally short-trichotomous, peduncle 8.2-8.5 cm long, glabrous, drying brown; branches 4-6 mm long, thick-fleshy, puberulous, terminating into cymules; each cymule subtended by 1-2 bracts; bracts unequal within each cymule, narrowly lanceolate to narrowly oblonglanceolate, the longer ones 14 × 5.2 mm, the shorter ones 4.0 × 2.6 mm, persistent, drying brown. Hypanthium narrowly obovoid, 2.5 mm long. Calyx cupular, 0.7 mm long, undulate, glabrous. Corolla unknown (fallen off). Style 4 mm long, lobes narrowly elliptic (after corolla has fallen off). Fruits unknown. Distribution. Only known by two gatherings from the Kamoa Mountains, Guyana. Ecology. Understory of dense forest on brown sand with rocky outcrops, at 240-400 m elevation. Phenology. The two flowering collections were collected in November. Specimens examined. GUYANA • Upper Takutu-Upper Essequibo Region, Kamoa Mountains, 0-2 km N of camp on Kamoa River; 1°31'N, 58°49'W; 240 m; 11 Nov. 1996; fl.; Clarke 3081; MO, US • Upper Takutu-Upper Essequibo Region, Kamoa Mountains, 1.5 km S of Kamoa River; 1°31'N, 58°49'W; 400 m; 12 Nov. 1996; fl.; Clarke 3136; MO, US. Notes. This probably new taxon is only known from two gatherings, both of which have inflorescences but no corollas left (one specimen still has a fully elongated style) and no fruits. It closely resembles C. araguariensis and C. necopinata, both of which are so far only known from Brazil. The differences with the former species are indicated in the key. From the latter species, which is still only known from the type (Taylor and Gereau 2013: 120), it may be separated by its smaller bracts (the largest ones up to 14 mm long vs 20-25 mm long in C. necopinata), longer inflorescence peduncle (8.2-8.5 cm long vs 5.0-5.5 cm long), shorter calyx (0.7 mm long vs 1.5-2.0 mm long) and leaves with 17-22 (vs 22-24) secondary veins on each side of the midrib. With so little material, it is difficult to assess the significance of these characters, and better collections are required to establish the identity of this taxon. Description. Shrub up to 1.5 m tall, branched; terminal branchlets terete, 1.5-2.5 mm in diam., glabrous, soon covered with a greyish-brown corky bark. Stipules free, narrowly triangular to lanceolate, 6-8 × 1.5-4.0 mm, entire, glabrous outside, villose at the base inside, becoming corky and soon damaged. Leaves with petioles 0.5-1.4 cm, glabrous; lamina elliptic, 8-18.5 × 1.8-6.7 cm, attenuate and decurrent on the petiole at base, acuminate at apex, slightly coriaceous when dry, entirely glabrous, drying dark brown or dark olive-green; midrib and secondary veins prominent on the upper side; secondary veins 6-9 on each side of the midrib, strongly ascending, curving towards the margin and almost reaching it; 2-3 intersecondary veins between each pair of secondary veins; tertiary veins reticulate, dense and conspicuous in the dry state. Inflorescences terminal, apparently erect, in involucrate heads, few-flowered; peduncle 4-5 cm long, glabrous; involucre orange, consisting of two unequal, decussate pairs of almost free bracts, the outer pair broadly obovate, 12-17 × 9-12 mm, the inner pair narrowly elliptic, 11 × 2.5-4.0 mm, both pairs erect, obtuse at apex, entirely glabrous, persistent in the fruiting stage; interfloral bracts numerous and much narrower than the involucral ones, linear, acute, ca 9 × 0.5 mm, glabrous, persistent in the fruiting stage. Flowers unknown. Calyx (in fruit) shortly cupuliform, truncate, ca 1 mm long, glabrous, persistent on fruit. Disk bilobed to the base, ca 1 mm long. Fruits narrowly ovoid, 12 × 5 mm when dry, glabrous, sessile. Pyrenes plano-convex, narrowly elliptic to oblong in outline, 11-14 × 4-7 mm, acute at apex, dorsal side nearly smooth with 3 very vague indications of ridges, ventral side with a deep longitudinal narrow excavation, ± C-shaped in cross-section, opening by 3 dorso-basal slits running along the ridges. Seeds entire, C-shaped in cross-section. Distribution. Endemic to northeastern French Guiana, only known from the type specimen collected on Montagne Soufflet. Ecology. The type label only reports that the specimen was collected in forest, without indicating the elevation.

7.

Phenology. The only known specimen, with fruits, was collected in June.

Notes. This species is only known from the fruiting type specimen. Presumably due to the incompleteness of the material, [START_REF] Steyermark | New Rubiaceae from French Guiana[END_REF] noted that its immediate relationships were not evident. However, it closely resembles C. guianensis and C. galbaoensis in the marcescent stipules and in the shape, texture, and venation of the leaves -the three species being virtually indistinguishable in vegetative state -as well as in the shape and mode of opening of the pyrenes. The three species also share inflorescences with two pairs of involucral bracts -although [START_REF] Steyermark | New Rubiaceae from French Guiana[END_REF] described and illustrated only one pair of involucral bracts in C. squamelligera, two pairs are actually present -and the flowers are surrounded by numerous bracteoles. They can be separated by the characters summarized in Table 2. The above description is mostly based on the P isotype; the holotype in VEN was seen as a photograph only, and an isotype from CAY cited in the original description was sent on loan to VEN in 1981 but has not been returned. Description. Shrub or treelet 1-3 m tall; terminal branchlets terete to slightly quadrangular, 4.0-5.5 mm in diam., glabrous, soon covered with a buff corky bark.

Stipules shallowly sheathing at base, 3.5-8.0 × 5 mm, broadly ovate, glabrous, bilobed or irregularly fimbriate at apex, with lobes 1-5 mm long, soon corky and marcescent. Leaves opposite or ternate; petioles 1.5-3.0 cm long, glabrous; blades narrowly elliptic or oblong-elliptic, 14-25 × 5-10 cm, acute to decurrent at base, acute and longacuminate at apex, acumen narrowly triangular to linear, 1.0-2.5 cm long, coriaceous, drying olive green above and yellowish green below, glabrous throughout; primary and secondary veins prominent on both sides; secondary veins 8-12 on each side of midrib, curving towards the margin and almost reaching it; tertiary venation densely and prominently reticulate in the dry state; domatia absent. Inflorescence thyrsoid, rather narrowly pyramidal, longpedunculate (expanded at fruiting stage); peduncles 7.5-12.5 cm long, glabrous, drying brown; secondary branches whorled, 3-5 per node, spreading, 0.4-2.0 cm long (0.8-4.0 cm at fruiting stage), glabrous except fringes of hairs at nodes, terminating into cymules; cymules with 15-22 flowers; bracts 4-5 around each cymule, ovate to triangular, 1-4 × 0.7-2.0 mm, obtuse to rounded at apex, slightly concave, persistent, drying brown, glabrous except a row of hairs at the base inside. Flowers 5-merous, heterostylous, sessile. Hypanthium narrowly obovoid, 0.5-1.0 mm long, glabrous. Disk bilobed to the base, ca 1 mm long. Calyx cupular, 0.7-1.0 mm long, truncate or minutely denticulate, glabrous. Corolla hypocrateriform, 8-10 mm long, white, greenish-white or brownish-white; tube narrowly obconical to almost cylindrical, 6.0-6.5 mm long, 1.0-1.5 mm wide at base, 1.5-4.0 mm wide at mouth, glabrous outside, pubescent in distal portion inside; lobes lanceolate to triangular, 1.5-3.5 × 1.0- probably not distinct from each other) by its pyramidal inflorescence with ramifications shorter than rachis (vs not or hardly so), smaller and coriaceous bracts, corolla tube 6.0-6.5 mm long (vs 3 mm long), and sessile fruits (vs shortly pedicellate). As noted by [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF], the leaves of C. tillettii may be opposite or verticillate, sometimes with both conditions on the same branch, and the stipules can be bilobed or irregularly 3-5fid at apex. The locality, altitude, collection date, and field notes of the type collection have been wrongly cited both in the protologue (Steyermark 1972: 498) and in Taylor and Gereau (2013: 123). The type label actually reads "Tree to 3 m; flowers white; fruit dark red; occasional in wet forest along trail, ridge of Merume Mountain, elev. 1140 m. ". This species is not to be confused with Rudgea tillettii Steyerm. (Steyermark 1976: 416), which is a synonym of R. coussareoides (Standl.) C.M. [START_REF] Taylor | Taxonomic transfers in Neotropical Palicoureeae: new combinations in Rudgea and Palicourea[END_REF]. The latter somewhat resembles C. tillettii in general appearance, but its stipules are basally connate into a truncate sheath and bearing dorsal appendages (these soon caducous), its flowers are sparsely arranged (not densely crowded at the apex of inflorescence ramifications) and its disk is entire. 9. Carapichea urniformis (Steyerm.) C.M.Taylor (Taylor and Gereau 2013: 122) Psychotria urniformis Steyerm. (Steyermark 1972: 556, figure 74) Notes. This species is remarkable by its large, urn-shaped involucre, varying from bilobed to unequally 4-lobed, which is formed by the fusion of the external bracts. The involucre is reported to be red [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] although on the type label it is more precisely described as "green at base, shading through brown to pinkishred at tips". Carapichea urniformis was placed by [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] Notes. Sandwith (1939: 551-552), in the original description of Cephaelis nivea, cited the gathering Sandwith 1249 as the type, with no indication of the herbarium of deposit. Four sheets of this gathering have been found, three in K and one in NY, as well as one spirit collection (not seen by us) in K. According to the Code [START_REF] Turland | International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress[END_REF], all these samples represent original material. One of the K sheets, barcode K000174488, which has complete label data and bears the annotation "Typus" in Sandwith's handwriting, was cited as holotype by Taylor and Gereau (2013: 121-122). Being posterior to 2001, and not accompanied by the statement "here designated" or a similar expression, their citation cannot be treated as an inadvertent lectotypification (ICBN,Art. 7.11 & 9.10). Two sheets of Sandwith 1249 at K are labeled "Sheet I/II" (K000174488) and "Sheet II/II" (K000174489), and are physically kept together. Therefore, according Art. 8.3 of the Code [START_REF] Turland | International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress[END_REF], they should be treated as a single specimen with multiple preparations. This specimen, composed of two sheets, is here designated as the lectotype of Cephaelis nivea. Notopleura nivea differs from N. altsonii and N. sandwithiana by the characters summarized in Table 3, the most obvious of which -not mentioned by [START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] -is that the involucral bracts form a much longer basal cup than in the other two species. Taylor and Gereau (2013: 122) also reported differences in the colour of the bracts, which are apparently not constant, and in the colour of the corolla (supposedly purple in N. altsonii and yellow in N. nivea) but in both species the corolla is actually white outside and yellow inside. Notopleura sandwithiana (Steyerm.) C.M.Taylor (Taylor 2004: 663) Cephaelis sandwithiana Steyerm. (Steyermark 1967: 428, figure 42) Psychotria sandwithiana (Steyerm.) Steyerm. (Steyermark 1972: 555) Carapichea sandwithiana (Steyerm.) C.M.Taylor (Taylor and Gereau 2013: 122) Type. VENEZUELA -Bolívar • Along Río Framela, Cerro Pitón, Cordillera Epicara;400 m;3 Sep. 1962;[START_REF] Turland | International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress[END_REF]B. Maguire, J.A. Steyermark & C.K. Maguire 53562;holotype: VEN [No. 63029]. Description. Shrub 2.0-2.5 m tall, single-stemmed, glabrous; distal internodes terete, 10-12 mm in diam., somewhat succulent, glabrous. Stipules basally sheathing, truncate to shallowly elliptic, 2-3 × 10-12 mm, glabrous. Leaves with petioles 2.0-2.5 cm long, glabrous; blades elliptic to elliptic-obovate, 26-37 × 10-12 cm, acute at base, abruptly acuminate at apex, acumen narrowly triangular, 2.0-2.5 cm long, somewhat succulent to coriaceous when fresh, drying brownish-olive green above and pale olive green below ("silvery beneath" when fresh, fide Maguire et al. 53562), glabrous throughout; secondary veins 38-40 on each side on each side" without distinguishing secondary and intersecondary veins], barely visible in dry specimens; tertiary veins barely visible in dry specimens; domatia absent. Inflorescence terminal, long-pedunculate, capitate; peduncles 4.5 cm long, gradually wider towards the head, ca 3 mm wide just below the head, glabrous, drying dark brown; head multiflorous, subtended by cup-shaped structure made by 2 bracts connate at base, the cup-shaped base 3 mm long and 1.7 cm wide; the free portion of bracts subequal, broadly ovate, 1.8 × 2.6 cm, cream-white when fresh, pale brown when dry, persistent, drying pale brown, glabrous; internal bracts subtending fascicles of ca 3 flowers falcate-sublanceolate to oblong-lanceolate, 3.5-4.0 × 1.0-1.5 mm, entire of with a lateral tooth at base, glabrous; bracteoles absent [fide Steyermark]. Flowers 5-merous, pedicellate; pedicels 7-8 mm long, 1.5-2.0 mm thick, fleshy, angular. Hypanthium narrowly oblong-turbinate, size unknown, glabrous. Calyx cupular, 1.5-2.0 mm long, 5-toothed, teeth deltoid to triangular-subulate, 0.2-0.6 mm long, glabrous. Corolla hypocrateriform, 12-13 mm long, glabrous, white; tube narrowly cylindrical, 9-11 mm long, 1 mm wide, glabrous outside and inside; lobes ovate, 2 × 1.5 mm, subacute at apex, glabrous outside, papillose inside. Stamens inserted below the middle of the corolla base; anthers subsessile, included, narrowly oblong, 2 mm long. Disk and style unknown. Fruits ellipsoid, 4 x 2.5-3 mm. Pyrenes (seen through dried fruits) dorsally costate. Distribution. Only known from the holotype specimen, collected on Cordillera Epicara, state of Bolívar, southern Venezuela. Ecology. Evergreen forest, at ca 400 m elevation. Phenology. The only collection, with both flowers and fruits, was made in September.

Notes. The only collection of this species was seen on photograph, and as the high-resolution image available at JSTOR Global Plants (https://plants.jstor.org/). The above description is largely based on [START_REF] Steyermark | Rubiaceae. In: Maguire B & coll. (Eds) The Botany of the Guayana Highland -Part VII[END_REF][START_REF] Steyermark | Flora de Venezuela[END_REF]. Although Steyermark (1967: 428-430, figure 42) described and illustrated flowers, the holotype, which is evidently the specimen on which the drawing was based, has only fruits and no flower is left. Steyermark (1967: 430) separated Notopleura sandwithiana from N. altsonii by the "shorter peduncle, longer pedicels, absence of bracteoles, more conspicuously and unequally dentate calyx-lobes, and acute, ovate calyx-lobes". These two species are in fact quite similar. Notopleura sandwithiana apparently has only two involucral bracts (while there are four in N. altsonii) and also has a shorter peduncle and longer pedicels than those of N. altsoni. The taxonomic value of these characters is difficult to assess with so little material available. The length of the pedicels is of doubtful taxonomic value, since they most likely elongate during fruit ripening. [START_REF] Steyermark | Rubiaceae. In: Maguire B & coll. (Eds) The Botany of the Guayana Highland -Part VII[END_REF] mentioned, as additional differences, the shape of the calyx lobes, which is not a reliable character, and the absence of bracteoles subtending the individual flowers of N. sandwithiana, a feature that we could not check. For a morphological comparison of these two species see Table 3.

validating the documents necessary for the export permit. PD is also very grateful to the Environmental Protection Agency (E.P.A.), and especially Diana Fernandes for her assistance in the process of obtaining the collecting permit (No. 060716 BR004) and the export permit (No. 062716 SP007). OL wishes to thank Cony Decock and his parents Philippe and Isabelle Lachenaud for their assistance in the field. We are also grateful to Sophie Gonzalez (Herbier IRD de Guyane) and Sébastien Sant (Parc Amazonien de Guyane) for permission to reproduce their photographs.

Figure 1 .

 1 Figure 1. Stipular dimorphism in Notopleura tapajozensis. A. Vegetative node with a stipular sheath and a tuft of appendages on each side. B. Flower-bearing node with a tuft of appendages but no sheath. From Delprete & Benjamin 12849. Drawing by Olivier Lachenaud.

Figure 2 .

 2 Figure 2. Carapichea adinantha. A-B. Detail of inflorescence with buds and one open flower. C. Fruiting plant. D. Infructescence with mature fruits. E. Infructescence with mature fruits, some eaten by birds. A-B from Gonzalez et al. 3281; C-E from Gonzalez 1368. Photographs by Sophie Gonzalez.

Figure 3 .

 3 Figure 3. Carapichea galbaoensis. A. Stem with leaves, stipules, and inflorescences. B. Nodes with stipules. C. Flower bud and two floral bracts. D. Corolla at anthesis with partially exserted anthers, lateral view. E. Stamens, adaxial and lateral view. F. Infructescence surrounded by bracts, top view. G. Fruit (left) and seed (right). A-E from Mori & Pipoly 15511; F-G from Marshall & Rombold 102. Drawn by Bobbi Angell. Reproduced from Mori et al. (2002: figure 279) with permission.

Figure 4 .

 4 Figure 4. Carapichea galbaoensis. A. Inflorescence with open flowers, top view. B. Inflorescence with open flowers, side view. C. Flowering branch. D. Habit. A from plant not collected, Monts La Fumée, French Guiana; B-C from plant not collected, Mont Galbao, French Guiana; D from Lachenaud 1689. Photographs by Sébastien Sant (A-C) and Olivier Lachenaud (D).

Figure 5 .

 5 Figure 5. Carapichea guianensis. A. Habit. B. Detail of lower leaf surface. C. Inflorescence with long-styled flowers. D. Inflorescence with short-styled flowers. E. Infructescence. A from plant not collected, Mont Itoupé, French Guiana; B-C from Lachenaud 1725; D from plant not collected, Camopi, French Guiana; E from plant not collected, Bellevue de Papaïchton, French Guiana. Photographs by Sébastien Sant (A, D-E) and Olivier Lachenaud (B-C).

Figure 6 .

 6 Figure 6. Carapichea ligularis. A. Habit, with two inflorescences. B. Detail of lower leaf surface. C. Inflorescence with open flowers. D. Infructescence with immature fruits. A from plant not collected, Montagne des Gouffres, French Guiana; B from Lachenaud 1793; C-D from plants not collected, Saül region, French Guiana. Photographs by Sébastien Sant (A, C-D) and Olivier Lachenaud (B).

8.

  Carapichea tillettii (Steyerm.) C.M.Taylor(Taylor and Gereau 2013: 123) Psychotria tillettii Steyerm.(Steyermark 1972: 496, figure 70) Type. GUYANA • Upper Mazaruni River basin, Partang River, ridge of Merumé Mountains; 1140 m; 1 Jul. 1960; fr.; S.S. Tillett et al. 43946; holotype: NY [0013844], isotypes: COL [COL000004671], F [No. 1704833, 1704834], K [K000432817, K000432818], NY [0013845], P [P02428007], US [00131318, 00146632], VEN [No. 82283].

  [nom. rej. versus Cephaelis Sw. (Swartz 1788)]

	Chesnea Scop. (Scopoli 1777: 119), nom. superfl. illeg. -
	Type species: Carapichea guianensis Aubl.
	Ipecacuanha Arruda (in Koster 1816: 497), nom. nud. -
	Type species: Ipecacuanha officinalis Arruda, nom.
	illeg. [= Carapichea ipecacuanha (Brot.) L.Andersson]
	Nettlera Raf. (Rafinesque 1838: 147), nom. superfl. illeg.
	-Type species: Nettlera guianensis (Aubl.) Raf. [=
	Carapichea guianensis Aubl.]
	Ipecacuana Raf. (Rafinesque 1838: 147), orth. var. -
	Type species: Ipecacuanha fusca Raf. [= Carapichea
	ipecacuanha (Brot.) L.Andersson]
	Uragoga L. ex Baill. (Baillon 1879: 324), nom. superfl. illeg.
	Flowers bisexual, usually heterodistylous, protandrous,	Style included or partially
	(4)5-merous. Hypanthium ovoid to obovoid. Calyx tube	exserted, glabrous; branches 2, oblong or linear. Fruits
	extremely reduced or cup-shaped, truncate, undulate or	drupaceous, fleshy, variable in colour (yellow, orange-red,
	lobed, persistent, lobes (when present) small, broadly	red, purple, maroon, blue, white or cream-white, or black
	to narrowly triangular. Corolla tubular, funnelform to	at maturity) with 2 fibrous to woody pyrenes. Pyrenes
	hypocrateriform, actinomorphic, white, yellow, orange,	vertical, plano-convex, ovate to elliptic in outline, dorsal
	or salmon-pink, tube glabrous or pubescent inside;	side multi-costate, ventral side longitudinally sulcate or
	lobes valvate, oblong-ovate, margin entire, acute at apex,	rarely flat.
	thickened or sometimes with horn-like extensions (C.	Morphological characters. The main diagnostic
	araguariensis). Stamens included, partially exserted or	characters of Carapichea are the marcescent stipules
	well exserted beyond corolla mouth; filaments inserted	lacking dorsal appendages, lack of ethanol-soluble
		pigment in the seed testa, and aperturate pollen of a
		generalized form, corresponding to Types XIV and
		XVI of

-Lectotype species (designated by Petit 1964: 16): U. ipecacuanha (Brot.) Baill. [= Carapichea ipecacuanha (Brot.) L.Andersson] Stachyococcus Standl. (Standley 1936: 144) -Type species: Stachyococcus adinanthus (Standl.) Standl. [= Carapichea adinantha (Standl.) C.M.Taylor] Type species. Carapichea guianensis Aubl. Description of the genus. Subshrubs, shrubs, or small trees. Raphides present. Stipules interpetiolar, free or shortly connate at base, entire to bifid at apex, rarely multifid (C. ipecacuanha), lacking dorsal appendages, persistent on distal nodes, marcescent, and eventually falling off through fragmentation, leaving a welldeveloped scar or a persistent hardened basal portion. Leaves opposite, rarely ternate (C. tillettii), short-to long-petiolate; blades ovate, elliptic, obovate, oblong or narrowly elliptic, papyraceous to coriaceous; secondary veins strongly to weakly prominent; tertiary veins reticulate or subparallel; domatia absent, or present as a continuous line of pubescence along midrib. Inflorescence terminal, rarely pseudoaxillary, few-to many-flowered, variously capitate, subcapitate, spiciform, or thyrsoid and branched to 1 or 2 orders; bracts reduced to very large, free to variously connate, sometimes forming an involucre. at the basal (C. urniformis), median, or distal portion of the corolla tube, short, equal, glabrous; anthers narrowly elliptic or narrowly oblong or linear, round at base, round or apiculate at apex, dorsifixed near middle. Pollen aperturate. Ovary 2-locular, placenta reduced. Ovules 1 per locule, basally inserted, erect. Disk usually bilobed to the base, sometimes undivided and cylindrical or laterally 5-sulcate (C. adinantha).

Table 1 .

 1 Infrageneric groups in Carapichea. Species occurring in the Guianas are in bold.

  . Among species investigated for this character, the only exceptions are C. cardenasiana, C. ipecacuanha, and C. panurensis, which have an entire disk, and C. adinantha, which has an undivided, laterally 5-sulcate disk. We were not able to verify the shape of the disk in C. fimbriflora, C. lucida, C. maturacensis, and C. verrucosa. Among related genera, bipartite disks also occur in some species of Notopleura (see above) and in two recently described species of Rudgea Salisb.: R. glomerulata Zappi & O.Lachenaud and R. itoupensis O.Lachenaud

Table 1

 1 According to the present circumscription, Carapichea includes at least 22 species that are found mostly in the Amazon basin and the Guianas, with two species(C. affinis and C. ipecacuanha)

	extending to Central America, one (C. ipecacuanha)
	extending to the Cerrado Biome of Central Brazil, and a
	single species (C. lucida) endemic to the Atlantic Forest
	of Brazil. The species occur in lowland or lower montane
	forest habitats (up to 1570 m but mostly below 1000 m),
	either on drained or seasonally flooded soils. Eight species
	occur in the Guianas; a ninth one, C. araguariensis, is to
	be expected there and has been included in the treatment
	below.
	Infrageneric classification. No infrageneric taxa are
	formally recognised in Carapichea, but the species may be
	arranged in six groups (from
	Brazil, is very poorly known, but its leaf venation matches
	the Carapichea group, rather than the Ligularis group in
	which it was placed by Taylor and Gereau (2013).

, and includes Carapichea, Chassalia Comm. ex Poir., Eumachia DC. [= Chazaliella E.M.A.Petit & Verdc., Margaritopsis C.H.Wright], Geophila D.Don, Hymenocoleus Robbr., Notopleura, Palicourea Aubl. s.l., Puffia Razafim. & B.Bremer, and Rudgea. Phylogenetic studies strongly support the monophyly of Carapichea, albeit with limited sampling

[START_REF] Andersson | Re-establishment of Carapichea (Rubiaceae, Psychotrieae)[END_REF][START_REF] Razafimandimbison | Phylogeny and generic limits in the sister tribes Psychotrieae and Palicoureeae (Rubiaceae): evolution of schizocarps in Psychotria and origins of bacterial leaf nodules of the Malagasy species[END_REF][START_REF] Bruniera | Sistemática e taxonomia de Rudgea Salisb[END_REF]

. The genus appears either as sister to Eumachia

[START_REF] Andersson | Re-establishment of Carapichea (Rubiaceae, Psychotrieae)[END_REF][START_REF] Bruniera | Sistemática e taxonomia de Rudgea Salisb[END_REF] as Margaritopsis)

or as sister to a clade including Eumachia, Chassalia, Geophila, Puffia, and Hymenocoleus

[START_REF] Razafimandimbison | Phylogeny and generic limits in the sister tribes Psychotrieae and Palicoureeae (Rubiaceae): evolution of schizocarps in Psychotria and origins of bacterial leaf nodules of the Malagasy species[END_REF]

with low support in all cases. Distribution and ecology. ), separable by characters in the key below, and largely based on those proposed by

[START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF]

. We have usually kept the group names used by these authors, although their Pacimonica group is better called the Ligularis group now that C. pacimonica is a synonym of C. ligularis. Their Altsonii group is not maintained here, since, as discussed above, three of its species are transferred to Notopleura, while the last one, C. urniformis, is included in the Carapichea group of which it has all the key characters. Also transferred to the Carapichea group are C. tillettii (see Notes under this species) and C. fimbriflora. The latter species,

Key to infrageneric groups in Carapichea (the Roman numerals in parentheses refer to the numbering of the groups by Taylor and Gereau 2013)

  ....................................................................................................................................................................

	1.	Stipules multifid, with 3-8 linear lobes; rhizomatous undershrubs; pyrenes longitudinally twisted; inflorescence capitate and
		involucrate ..

Ipecacuanha group (I)

	-	Stipules entire or bilobed (rarely minutely fimbriate at apex in C. tillettii); erect shrubs; pyrenes straight; inflorescence variable .. 2
	2.	Leaves with secondary and tertiary veins ± parallel and hardly distinct from each other ....................................

Ligularis group (III) -

  Leaves with secondary veins much stronger than tertiaries, the latter either reticulate or invisible ...................................................... 3 .................................................................................................................................................................................

	3.	Secondary leaf veins forming conspicuous loops well before the margin; disk (when known) entire or crenate laterally, not bipartite;
		inflorescence glomerulate or spiciform (the ramifications < 5 mm long if present), with minute bracts ............................................. 4
	-	Secondary leaf veins curved towards the margin and almost reaching it; disk (when known) bipartite; inflorescence either broadly
		thyrsoid or capitate, the bracts usually well-developed .................................................................................................................................. 5
	4.	Stipules keeled dorsally; inflorescence glabrous; corolla glabrous outside; fruits pedicellate, crowned with persistent calyx tube;
		pyrenes strongly costate dorsally; leaves acute to obtuse at apex, secondary veins rather weak ....................... Panurensis group (V)
	-	Stipules not keeled; inflorescence puberulous; corolla (when known) puberulous outside; fruits sessile, not conspicuously crowned
		by the calyx; pyrenes smooth to verrucose dorsally, not ridged; leaves acuminate at apex, secondary veins very strong
		...

Stachyococcus group (VI) 5

  . Stipules bifid with linear lobes; inflorescence capitate, surrounded with two large green bracts ............................

Bahia group (VII) -

  Stipules entire or bilobed, the lobes if present rather broad; inflorescence thyrsoid to capitate ......................

Carapichea group (II) Key to species of Carapichea in the Guianas 1

  . Leaves with secondary and intersecondary veins ± parallel and hardly distinct from each other (Fig.6B); corolla lobes often corniculate dorsally .............................................................................................................................................................................................. 2 -Leaves with secondary veins thicker than tertiary veins; intersecondary veins absent; tertiary veins reticulate (Figs 3A, 5B); corolla lobes not corniculate (but sometimes with an internal appendage) ............................................................................................................ 4 2. Inflorescence with basal bracts inserted at the end of the peduncle; secondary branches absent or very short; corolla tube 6-9 mm long ....................................................................................................................................................................................................... -sulcate laterally; secondary veins arching far from the leaf margin ............ ............................................................................................................................................................................................................ ........................................................................................................................................................................................................ 6 6. Bracts fused for most of their length into an urn-shaped involucre ...................................................................................... C. urniformis -Bracts free or shortly connate at base ................................................................................................................................................................ ....................................................................................................................................................................... ........................................................................................................................................................................................................................

		C. ligularis
	-	Inflorescence basal bracts inserted at the end of the secondary branches, these always distinct; corolla tube 11-12 mm long
		(unknown in C. sp. A) ......................................................................................................................................................................................... 3
	3.	Inflorescence with 3 glomerules; secondary branches 0.4-0.6 cm long .......................................................................................... C. sp. A
	-	Inflorescence with 5-9(-13) glomerules (or shortly branched cymules); secondary branches 0.7-3.0 cm long ..... C. araguariensis
	4.	Inflorescence spiciform, with minute bracts; corolla puberulous outside, the lobes internally appendiculate (Fig. 2A); pyrenes
		ventrally flat and dorsally smooth; disk almost entire, 5C. adinantha
	-	Inflorescence capitate or thyrsoid; corolla (unknown in C. squamelligera) glabrous outside, the lobes not appendiculate; pyrenes
		ventrally sulcate, usually costate dorsally; disk bipartite; secondary veins arching near the leaf margin (almost reaching it) ......... 5
	5.	Inflorescence thyrsoid, with reduced bracts 1-4 mm long; corolla tube 6.0-6.5 mm long ...................................................... C. tillettii
	-	Inflorescence capitate, with bracts 12-90 mm long, forming an involucre; corolla tube 5-30 mm long (unknown in C.
		squamelligera) 7
	7.	Bracts orange, ± erect and not markedly unguiculate at base, the external pair 12-17 mm long; peduncle 4-6 cm long; pyrenes
		11-14 × 4-7 mm C. squamelligera
	-	Bracts mostly green, obviously unguiculate at base, the external pair 19-44 mm long; peduncle 1-3 cm long; pyrenes 8-9 × 3-4
		mm ..8
	8.	Corolla tube 5-8 mm long, conspicuously widening in upper part; involucral bracts in two very unequal pairs, the external one
		19-42 × 3-10 mm, the internal one 5-20 × 2-6 mm, both unguiculate at basal 4-6 mm; stipules 5-9 mm long ....... C. guianensis
	-	Corolla tube (15-)16-19 mm long, almost cylindrical; involucral bracts in two nearly equal pairs, the external one 27-44 × 9-25
		mm, the internal one 24-36 × 5-15 mm, both unguiculate at basal 9-10 mm; stipules 2-5 mm long ........................ C. galbaoensis
	1. Carapichea adinantha (Standl.) C.M.Taylor (Taylor
	and Gereau 2013: 123)
	Fig. 2
	Retiniphyllum adinanthum Standl. (Standley 1931: 355)
	Stachyococcus adinanthus (Standl.) Standl. (Standley
		1936: 144)
	Type. PERU -Loreto • near Iquitos, Mishayacu; 100 m;
	Feb.-Mar. 1930; fl.; Klug 988; holotype: F [No. 612612];
	isotypes: NY [00133128], US [0013820].

  m in elevation. Specimens examined. FRENCH GUIANA • Approuague River Basin, Rivière Kourouaï; 4°15'N, 52°01'W; 120 m; 9 Jul. 2008; fr.; Gonzalez 1368; BR, CAY, L, MO, NY, P, US • Monts Tumuc-Humac, Sommet en Cloche; 2°13'40"N, 54°28'10"W; 12 Mar. 2015; fl.; Gonzalez 3281; BR, CAY. BRAZIL -Acre • Mun. Cruzeiro do Sul, Rio Branco, ca 25 km from Rio Juruá, secondary forest with canopy 30-35 m tall; 7°45'S, 72°17'W; 240 m; 22 Nov. 2001; fl.; Delprete et al. 8045; NY, UFACPZ, additional duplicates at UFACPZ to be distributed • Mun. Cruzeiro do Sul, road Cruzeiro do Sul-Rio Branco, road to Canela Fina, ca 7 km from 3-way junction at 5 km N of Cruzeiro do Sul; 7°32'44"S, 72°44'31"W; 220 m; 23 Nov. 2001; fl.; Delprete et al. 8078; NY, UFACPZ, additional duplicates at UFACPZ to be distributed. -Amapá • Parque Nacional Montanhas do Tumucumaque, cabeceiras do Rio Amaparí, margem esquerdo do Rio Anacuí, trilha 1, floresta de terra firme; 6 Mar. 2006; fl.; Hamada et al. 139; INPA, MO n.v. -Amazonas • Distrito Agropecuário, Fazenda Porto Alegre, Reserva 3402 (Cabo Frio) of the WWF/INPA MCS project; 2°25'25"S, 59°54'38"W; 50-125 m; 1 Apr. COL, MO n.v. PERU -Loreto • Maynas Prov., Dtto. Amazonas, Explornapo Camp, cerca de Sucusari, a lo largo del Río Napo; 3°20'S, 72°55'W; 100-140 m; 15 Feb. 1991; fl.;

	Pipoly et al. 12979; MO • Chacra Canamá, Requena; 7 Dec.
	1962; fl.; Schunke Vigo 6243; K • Prov. Requena, Jenaro
	Herrera; 4°50'S, 73°45'W; 170 m; 5 Jul. 1981; fr.; Vásquez
	et al. 2192; MO • carretera Nauta-Iquitos km 5; 4°29'S,
	73°35'W; 28 Mar. 1987; fr.; Vásquez & Arevalo 8984; P •
	Iquitos, Asociación Agraria Paujil; 4°10'S, 73°20'W; 2
	Jul. 1988; fr.; Vásquez & Jaramillo 10837; P • Allpahuayo,
	Estación Experimental del Instituto de Investigaciones de
	la Amazonia Peruana; 2 Jun. 1990; fr.; Vásquez & Jaramillo
	14029; P.

1989; fr.; Aquino et al. 3; INPA • Basin of Rio Juruá, near mouth of Rio Embira (tributary of Rio Tarauacá); 7°30'S, 70°15'W; 6 Jun. 1933; fr.; Krukoff 4682; K, M, MO • Basin of Rio Juruá, near mouth of Rio Embira; 17 Jun. 1933; fr.; Krukoff 4900; M, MO, NY, US. -Pará • Igarapé das Pedras, Cataractam Furnas, Rio Tapajóz; 30 Dec. 1917; fl.; Ducke s.n. (MG 16859, RB 23125); B destroyed (photo at F), RB. -Rondônia • ca 4 km ENE along rd to São Sebastião off BR-364, terra firme forest; 24 May 1984; fr.; Frame 160; INPA, MO n.v., NY • Estrada Porto Velho-Cuiabá, BR-364, km 171, mata de terra firme; 6 Feb. 1983; fr.; Freitas et al. 13; INPA • Itapuã do Oeste, Floresta Nacional do Jamari, parcela convenio NYBG-RON; 9°15'6"S, 62°54'38"W; 23 Apr. 2015; fr.; Medeiros et al. 1692; MO n.v., NY, RON. COLOMBIA -Caquetá • Araracuara, trocha Yari; 0°25'S,

72°20'W; 200 m; 23 Jan. 1989; fl.; Gentry & Sánchez 65001;

  ; INPA, MO, NY • Mun. Manaus, Distrito Agropecuário da SUFRAMA, Rod. BR-174, km 72, depois 6 km W da BR, Fazenda Dimona, mata de terra firme sobre latosolo amarelo; 2°19'S, 60°5'W; 50-125 m; 25 Mar. 1992; fl.; Dick 62; INPA • estrada Manaus-Caracaraí, km 125, igarapé da Lage, terra

	• Reserva Florestal Ducke, Manaus-Itacoatiara km 26;
	2°53'S, 9°58'W; 16 Jun. 1994; fr.; Ribeiro & Assunção
	1325; K • Manaus, km 68 da Estrada Manaus-Itacoatiara,
	sub-bosque de terra firme; 19 Mar. 1963; fl.; Rodrigues
	4985; INPA • Reserva Florestal Ducke, Estrada Manaus-
	Itacoatiara, km 26, floresta de baixo, solo arenoso; 2°53'S,
	59°58'W; 3 Feb. 1995; fl.; Vicentini et al. 840; INPA, K,
	MG, MO, NY, U • ibid., 31 May 1995; fr.; Vicentini et al.
	984; K. -Pará • Mun. Anajas, opposite to town of Anajas,
	on Rio Anajas; 31 Oct. 1984; fl.; Sobel et al. 4943; MG,
	MO n.v.

firme;

13 Feb. 1974;[START_REF] Turland | International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress[END_REF] Loureiro et al. s.n. (INPA 47925)

;

INPA • ibid., 14 Feb. 1974; fl.; Loureiro et al. s.n

. (INPA 47947); INPA • Distrito Agropecuário, Reserve 1501 (km 41) of the Smithsonian/INPA Biological Dynamics of Forest Fragments Project; 2°24-2°25'S, 59°43'-59°45'W; 15 Jul. 1990; fr.; Mori & Costa Lima Assunção 21383; K

  Carapichea galbaoensis is closely related to C. guianensis and has been considered a synonym of the latter[START_REF] Delprete | Notes on some South American species of Psychotria subgenus Heteropsychotria (Rubiaceae), with observations on rubiaceous taxonomic characters[END_REF][START_REF] Delprete | Carapichea guianensis: the correct name for Psychotria carapichea (Rubiaceae), with a correction on the authority of P. carapichea, and a new combination in Carapichea[END_REF][START_REF] Boom | Guide to the Vascular Plants of Central French Guiana, Part 2, Dicotyledons[END_REF][START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] but is here reinstated as a distinct species. The two taxa are almost identical in vegetative characters (although C. galbaoensis has usually shorter stipules), fruits, pyrenes, and inflorescence structure, but the shape and dimensions of the involucral bracts differ, as well as the position of the stamens and style, and especially the length of the corolla tube, which is much longer in C. galbaoensis. Differences between them are summarised in Table2and illustrated in Figs 3, 5.

	Lachenaud 1689; BR, CAY • Montagne des Nouragues;	10-18 × 3.3-7.5 cm, attenuate and decurrent on the
	4°03'N, 52°42'W; Jun. 1989; fr.; Larpin 539; CAY [2	petiole at base, narrowly acuminate at apex, rather thick
	sheets] • Commune de Saül; 13 Jul. 1977; fr.; Moretti 752;	when fresh (becoming papyraceous when dry), entirely
	CAY [2 sheets] • Saül, Monts La Fumée; 3°37'N 53°12'W;	glabrous, drying dark grey-green to blackish; midrib and
	200-400 m; 30 Aug. 1982; fr.; Mori & Boom 14803; CAY,	secondary veins prominent on the upper side; secondary
	NY • Saül, La Fumée Oeste; 3°37'N, 53°12'W; 200-400 m;	veins 6-9 on each side of the midrib, strongly ascending,
	5 Apr. 1983; fl.; Mori & Pipoly 15511; CAY, NY • Saül, La	curving towards the margin and almost reaching it; with
	Fumée Mountain; 3°37'N, 53°12'W; 200-350 m; 22 May	2-3 intersecondary veins between each pair of secondary
	1986; fr.; Mori & Pennington 18097; CAY, P, U • Station	veins; tertiary veins reticulate, rather inconspicuous in
	des Nouragues; 4°03'N, 52°42'W; 20 Jul. 1989; fr.; Sabatier & Prévost 2547; CAY, U • Station des Nouragues, chemin ouest; 9 Mar. 1996; fl.; Solano K 282; CAY. BRAZIL -Amapá • Near Cachoeira Macacoara; 0°53'N, 53°21'W; 27 Aug. 1961; fr.; Egler & Irwin 46695 [Irwin's collection number]; U, US. Notes. superfl. Tapogomea carapichea Poir. (Poiret 1806: 587), nom. superfl. Carapichea aubletii DC. (Candolle 1830: 536), nom. superfl.	the fresh state, prominent when dry. Inflorescences in terminal involucrate heads, few-flowered; peduncle 1-3 cm long, erect to patent, glabrous; involucre green on both sides, sometimes with a reddish-brown basal part, consisting of two very unequal decussate pairs of bracts shallowly connate at base, the outer pair 19-42 × 3-10 mm, the inner pair 5-20 × 2-6 mm, both pairs with a basal stalk 4-6 mm long (that of the inner pair much narrower) and the medio-distal portion narrowly ovate to lanceolate, acute to round at apex, glabrous, persistent at fruiting stage; interfloral bracts much smaller than the involucral ones, elliptic, obovate to spathulate, 4-6 × 0.8-3.0 mm, obtuse to truncate at apex, glabrous, persistent in the fruiting stage. Flowers 5-merous, heterostylous, sessile. Hypanthium obovoid, 1.0-1.2 × 0.7 mm, glabrous. Disk bilobed to the base, 0.5-0.6 mm long. Calyx shortly cupuliform, truncate or undulate, 0.2 mm long, glabrous. Corolla hypocrateriform, white; tube infundibuliform, 5-8 mm long, 0.8-1.0 mm wide at base, 2.6-3.7 mm at mouth, glabrous outside, shortly pubescent inside around the insertion of the stamens; lobes ovate to deltoid, 2-3 × 1.5-2 mm, reflexed, glabrous throughout. Short-styled flowers: stamens inserted at distal 1/3 rd of the tube, exserted beyond corolla mouth; filaments ca 3 mm long; anthers 1.3-2.3 × 0.3-0.5 mm; style included, 5-6 mm long; style branches linear, 1.5-1.7 mm long. Long-styled flowers: stamens inserted at distal 1/3 rd of the tube, included; anthers subsessile, oblong-elliptic 1.5-1.8 × 0.3-0.4 mm; style exserted well beyond the mouth, 8.5-10
	Uragoga guianensis (Aubl.) Pulle (Pulle 1906: 446)	mm long, style branches obovate, 0.7-1.3 mm long. Fruits
	Cephaelis guianensis (Aubl.) Standl. (Standley 1929: 335) Psychotria ligularis var. carapichea Steyerm. (Steyermark 1972: 676) Psychotria carapichea (Steyerm.) Delprete (Delprete 2001: 401)	ellipsoid, 7-9 × 5-7 mm when dry (8-10 × 5-8 mm when fresh), vermillion, turning black at maturity, glabrous, sessile. Pyrenes plano-convex, narrowly elliptic in outline, 9 × 3.0-3.5 mm, dorsal side with 4 hardly distinct ridges, ventral side with a deep narrow excavation for the whole length, ± C-shaped in cross-section, opening by 4 dorso-
	Type. FRENCH GUIANA • "ad ripam amnis Galibiensis" [margin of the Galibi Creek]; s.d. [Apr.-May 1763]; Aublet s.n.; lectotype: P-JJR [8:266], designated by Lanjouw and	basal slits running along the ridges. Seeds entire, C-shaped in cross-section. Distribution. Endemic to the Guiana Shield, occurring
	Uittien (1940: 149); isolectotype: BM [BM001009105];	in French Guiana, Suriname, Guyana, and the contiguous
	see Delprete (2015).	Brazilian states of Amapá and Pará; locally common, but
	Description. Shrub 0.4-1.5 m tall, sparsely branched;	apparently absent from the range of C. galbaoensis (except
	terminal branchlets terete, 1-2 mm in diam., glabrous,	on the Montagne Bellevue de l'Inini).
	soon covered with a buff to pale grey corky bark. Stipules
	5-9 × 1.5-3.0 mm, free, triangular to lanceolate, entire or
	shortly bifid for up to 3 mm, glabrous outside, villose at the
	base inside, becoming corky and soon damaged. Leaves
	with petioles 0.8-3.0 cm long, glabrous; blade elliptic,

• Saül, layon blanc vers Limonade; 3°37'N, 53°12'E; 3 Aug. 1987; Allorge 311; P • Mont Bakra; 3°18'N, 52°57'W; 475 m; 16 Apr. 1993; fr.; Cremers 13141; CAY [2 sheets], U • Mont Chauve; 3°49'N, 52°44'W; 150 m; 16 Apr. 1997; Cremers & Crozier 15010; CAY • ibid.; 120 m; 21 Apr. 1997; fl.; Cremers & Crozier 15134; CAY • Saül, sur le layon vers La Fumée, 500 m avant le plateau La Douane; 3 Feb. 1981; imm. fr.; Fournet 62; CAY, P • Monts Galbao, à 10 km WNW de Saül; 10 May 1973; Granville 1584; CAY, COL, U • Saül, 1 km environ au S des Monts La Fumée; 6 Mar. 1975; fl.; Granville 2354; CAY • Saül, layon ORSTOM du plateau La Douane aux Monts La Fumée; 16 Jan. 1976; fl.; Granville 2664; CAY [2 sheets], P • Saül, antenne est La Fumée/Nouvelle France, pk 1.2; 5 Mar. 1977; fl.; Granville 2806; CAY • Sommet Tabulaire, versant sud; 500 m; 24 Aug. 1980; fr.; Granville 3596; CAY [2 sheets], P • Massif des Emerillons, zone sud; 250 m; 8 Sep. 1980; fr.; Granville 3773; CAY • tracé ORSTOM La Fumée, à 1,2 km environ au N du Plateau La Douane; 3 Apr. 1982; fl.; Granville 5050; CAY • env. 13 km SSW Saül, 2-3 km W of Crique Limonade, Montagne Périnette; 31 Mar. 1983; fl.; Granville 5435; CAY [2 sheets], P • Montagne Bellevue de l'Inini, zone centrale; 750-800 m; 23 Aug. 1985; fl., fr.; Granville 7761; CAY, INPA, MG, MO n.v., P, U • Station des Nouragues; 4°03'N, 52°42'W; 100 m; 6 Aug. 1989; fr.; Granville et al. 11071; CAY • Pic Coudreau, Monts Bakra; 3°18'N, 52°57'W; 710 m; 18 Apr. 1993; fl.; Granville & Cremers 11788; CAY • Monts Bakra, 1,5 km à l' ouest du Pic Coudreau; 3°18'N, 52°57'W; 520 m; 16 Jun. 2002; fr.; Granville et al. 14804; CAY • Saül, Monts La Fumée; 3°40'N, 53°10'W; 200-300 m; 26 Jul. 1987; fr.; Hahn 3634; CAY, U • Piste Sophie, Saül; 11 Sep. 1962; F. Hallé 818; P, U • Station des Nouragues; 4°03'N, 52°42'W; Jul. 1996; fr.; Hequet 222; CAY • Saül, Les Eaux Claires; 3°39'N, 53°12'W; 200-400 m; 5 Sep. 2000; fr.; Junikka & Nieminen 3031; CAY • Saül, Monts La Fumée; 17 Mar. 2014; 4. Carapichea guianensis Aubl. (Aublet 1775: 168) Fig. 5 Callicocca guianensis (Aubl.) J.F.Gmel. (Gmelin 1791: 372) Cephaelis involucrata Willd. (Willdenow 1798: 979), nom. Ecology. Commonly encountered in mature terra firme rainforest, or sometimes in seasonally flooded forest, from near sea level to 600 m elevation. Phenology. Flowering collections were made in January (once), March to early May (main flowering season), and

Table 2 .

 2 Diagnostic characters of Carapichea galbaoensis, C. guianensis, and C. squamelligera.

		C. galbaoensis	C. guianensis	C. squamelligera
	Stipules	2-5 mm long	5-9 mm long	6-8 mm long
	Peduncle	1-3 cm long, erect	1-3 cm long, erect to patent	4-5 cm long, apparently
				erect
	Involucre	Two pairs of subequal bracts	Two pairs of very unequal bracts Two pairs of unequal bracts
	Involucral bracts size	Outer bracts 27-44 × 9-25 mm,	Outer bracts 19-42 × 3-10 mm,	Outer bracts 12-14 × 9-11
		inner bracts 24-36 × 5-15 mm	inner bracts 5-20 × 2-6 mm	mm, inner bracts 11 ×
				2.5-4.0 mm
	Stalk of involucral bracts	Stalk 8-10 mm long	Stalk 4-6 long	Not markedly stalked
	Bracts colour	Green to white	Green (the base sometimes	Orange
			reddish-brown)
	Corolla tube length and	17-19 mm long, almost	5-8 mm long, conspicuously	Flowers unknown
	shape	cylindrical (slightly wider	wider towards the mouth
		towards the mouth)
	Anthers position in	Only tips exserted	Fully exserted	-
	short-styled flowers (style
	always included)
	Style position in long-	Just reaching corolla mouth	Well exserted	-
	styled flowers (anthers
	always included)
	Pyrene dimensions	8-9 × 3.5-4.0 mm	9 × 3.0-3.5 mm	11-14 × 4-7 mm

August (once). Fruiting specimens were collected from April to November. Local names. This species is called carapiche (hence the generic name) by the Karipuna ("Garipons"; Aublet 1775) and tapiiwapa' a by the Wayampi (Grenand 270). Specimens examined. GUYANA • Rupununi District, Kuyuwini landing, Kuyuwini River; 2°05'N, 59°15'W; 10 Oct. 1992; fr.; Jansen-Jacobs et al. 2844; CAY, U. SURINAME • Tumuc Humac Mountains, Talouakem, Litani River; 2°31'N, 54°45'W; 6 Aug. 1993; fr.; Acevedo-Rodriguez et al. 5924; CAY, U, US • Upper Litany River; 2°27'N, 54°48'W; 30 Jul. 1993; fr.; Granville et al. 11895; B n.v., BBS, CAY, MO n.v., P, U, US • Sipaliwini, vicinity of airstrip along Ulemari River, 71 km up Ulemari River from its confluence with Litani River; 3°6'17"N, 54°32'28"W; 150 m; 29 Apr. 1998; fr.; Hammel et al. 21736; MO n.v.of Tafelberg at the margin of Kappelsavanna; 10 Jun. 1963; Wessels Boer 1532; K, U. FRENCH GUIANA • Crique Grand Laussat; 5°25'N, 53°37'W; 11 Jul. 2004; fr.; Barrabé 77; CAY, NY, P • RN2, pk 50; 11 Jun. 1984; fr.; Feuillet 1405; B n.v., CAY, MO, P, U, US • Rivière Itany × Koulé-Koulé; 20 Jul. 1985; fr.; Feuillet 2480; CAY • ibid.; 21 Jul. 1985; fr.; Feuillet 2504; CAY, P • Montagne de l'Inini; 3°30'N 53°30'E; 9 Apr. 1986; fr.; Feuillet 3730; CAY, MO n.v., P • ibid.; 11 Apr. 1986; fl.; Feuillet 3843; CAY • Eastern Plateau of Montagne Tortue; 13 Jun. 1988; Feuillet 9974; K, U • Riviére Kourouaï, Bassin de l' Approuague; 4°15'N, 52°1'W; 120 m; 8 Jul. 2008; fr.; Gonzalez 1361; CAY • Mont Itupé-sommet tabulaire -15, versant W, layon D; 3°1'45"N, 53°5'28"W; 600 m; 15 Mar. 2010; fr.; Gonzalez 2158; CAY, MO, P, US • forêt à 1 km du saut "S" sur la rive droite du [fleuve] Grand Inini; 18 Aug. 1970; fr.; Granville C-1; CAY, COL, P, U • Monts Atachi Bacca; 5 Mar. 1971; fl.; Granville C-125; CAY, P • rive gauche du [Fleuve] Yaroupi, un peu audessus de Saut Ouaïmicouaré; 12 Apr. 1970; fl.; Granville 333; CAY [3 sheets], P, U • Crête des Monts Atachi Bacca; 5 Mar. 1971; fallen fl.; Granville 789; CAY, P • Route Apr. 1988; fl.; Granville 10279; CAY, P, U • Monts Atachi Bacca; 3°33'N, 53°55'W; 9 Jan. 1989; fl.; Granville et al. 10485; CAY, P, U • Camp Caïman, Montagne de Kaw; 4°32'N, 52°13'W; 11 Mar. 2004; fallen fl.; Granville & ; Cowan 38335; K, NY, U, US • ibid.; 17 Nov. 1954; fl.; Cowan 38384; U • ibid.; fl.; Cowan 38416; U • immediately east of Colonia Agricola do Oiapoque, about 4 km N of mouth of Cricu River; 3°43'N, 51°55'W; 14 Aug. 1960; fl., imm. fr.; Irwin et al. 47516; F, IAN, K, NY [2 sheets]. -Pará • Rio Trombetas, 4 km S of Cachoeira Porteira; 6 Jun. 1974; fr.; Campbell et al. P22528 [Prance's collection number]; INPA, NY, U, US.

Notes. The chaotic taxonomic history of this species has been summarized in

[START_REF] Delprete | Carapichea guianensis: the correct name for Psychotria carapichea (Rubiaceae), with a correction on the authority of P. carapichea, and a new combination in Carapichea[END_REF]

. It has often been confused with C. ligularis.

[START_REF] Steyermark | Rubiaceae. In: Maguire B & coll. (Eds) The Botany of the Guayana Highland -Part IX[END_REF]

, in particular, reduced it to a variety of the latter (as Psychotria ligularis var. carapichea). It is very strange that he arrived to this conclusion, since C. guianensis and C. ligularis are quite different in leaf venation (reticulate in the former and parallel in the latter) and bract shape (narrower and lacking an unguiculate basal part in C. ligularis)

(Figs 5,

Table 2

 2

	(see also Figs 3, 4).
	The dimensions of the involucral bracts are rather variable
	in C. guianensis. Two collections from French Guiana
	(Feuillet 3730 and 3843) have the bracts unusually broad,
	resembling C. galbaoensis, but their basal stalk is shorter
	than in the latter species, and the corolla is typical for C.
	guianensis. At the other extreme, Wessels Boer 719 from
	Suriname and Schnell 11468 from French Guiana have
	exceptionally narrow bracts, and also differ from the rest
	of the material in their shortly corniculate corolla lobes;
	they are provisionally referred to C. guianensis but might
	prove to be distinct.

  was actually collected in French Guiana (see Notes below). Ducke s.n.; INPA [No. 16533] • Nova Prainha, Rio Aripuanã, RADAM/BRASIL, SB-20-ZD, Ponto 10; 9 Jul. 1976; fr.; C.D.A. Mota s.n.; INPA [No. 60607] • Mun. Nova Olinda, Rio Paca, tributari of the Rio Mari Mari, terra firme forest, tree to 7 m tall; 2 Jul. 1983; fr.; Todzia et al. 2299; INPA, MO, NY, US • Mun. Axinim, basin of Rio Abacaxis, lower Rio Paca, ca 1 km from its confluence with Rio Mari Mari, forest on terra firme, shrub 2 m tall; 4°07'S, 58°58'W; 1 Jul 1983; fr.; Zarucchi et al. 2917; INPA, MO, NY. -Pará • Ilha de Marajó, Rio Jipurú, afluente do

		Pic Coudreau; 3°18'N, 52°57'W; 600 m; 18 Jun. 2002; fr.;
		Granville et al. 14864; B, CAY, MO, P • Monts Kotika,
		plateau latéritique sommital; 3°55'10"N, 54°11'10"W; 730
		m; 23 Feb. 2005; fr.; Granville et al. 16922; B, CAY, MO,
		NY, P, U • Route Cayenne-Régina, km 52; 4°34.638'N,
		52°23.873'W; 9 Jan. 2011; fr.; Lachenaud 1074; BR, CAY •
		route de Cacao; 4 May 2014; fr.; Lachenaud 1793; BR, CAY
		• Mont Itoupé, Sommet Tabulaire, Layon D, 2me sommet;
		3°02'35"N, 53°05'18"W; 760 m; 31 Mar. 2010; fr.; Molino
		& Sabatier 2843; BR, CAY, MO • Montagnes de la Trinité;
		4°36'30"N, 53°21'30"W; 12 July 2001; fl. buds, fr.; Poncy
		& Crozier 1458; CAY, NY, P, U, US • RN2 -Cayenne-
		Régina, km 85.5; 4°20'N, 52°18'W; 26 Jun. 2003; fl., fr.;
		Prévost & Sabatier 4742; CAY, MO • Rivière Itany, Crique
		Petit Marouini, près de son embouchure; 2 Sep. 1972; fr.;
		Sastre 1814; P.
	Ecology. In understory of non-flooded forest, usually on	BRAZIL -Amapá • Upper Jari River, foret primaire
	superficial soils (e.g. lateritic crusts, granitic boulders) at	humide; 2°28'N, 54°46'W; 420 m; 20 Aug. 1993; fl.;
	100-800 m elevation.	Granville et al. 12387; BBS, CAY [2 sheets], US. -
	Phenology. Flowering specimens were collected from	Amazonas • Rio Curicuriarí, afl. do Rio Negro; 21 Dec.
	August to January; and fruiting specimens throughout the year. Selected specimens examined. VENEZUELA -Amazonas • Cerro da Neblina, Río Yatua, along Upper Río Yatua between mouth of Río Yacibo and Piedra Arauicaua; 1 Feb. 1954; fl.; Maguire et al. 37411; F, US. FRENCH GUIANA • Route de l'Est, ca 25 km NW of Régina; 4°23'N, 52°19'W; 16 Mar. 1994; fr.; Andersson et al. 1989; BR, CAY • Cacao; 27 Oct. 1983; fl.; Billiet & Jadin 1877; BR • Route RN2 Cayenne-Régina, pk 67, Crique Tibourou; 4°29'N, 52°19'W; 10 Feb. 1993; fr.; Billiet & Jadin	1931; fl.; Rio Anajas; 20 Oct. 1987; fl.; A. Tavares & J. Cardoso
	5733; BR • Extension Nord-Ouest des Petites Montagnes	227; INPA, NY. -Roraima • Mun. Rorainópolis, Reserva
	Tortue; 4°20'29"N, 52°16'16"W; 30 m; 29 Sep. 2010; fl.,	Popular Xixuaú-Xiparina, ilha subindo o Rio Xipariná;
	fr.; Bordenave & Le Hir 9068; CAY, K, P, MO • Montagne	0°55'41"S, 61°50'26"W; 25 Aug. 2010; fl, young fr.; Zappi
	Tortue, Plateau Est; 4°17'N, 52°21'W; 460 m; 5 Oct. 2010;	et al. 2902; INPA, MIRR.
	fl.; Bordenave & Le Hir 9233; CAY • Upper Maroni River,
	layon de chasse au NE d' Antecume Pata, confluent de
	l'Itany et du Marouini; 19 Nov. 1977; fl.; Cremers 5077;
	CAY • Saül, piste du carbet; 31 Oct. 1984; fl.; Foresta 705;
	CAY • Monts Atachi Bacca, autour du Camp 3, entre les
	sommets 525 et 782 m; ca 400 m; 5 Mar. 1971; fr.; Granville
	C-128; CAY [2 sheets], P • Monts Atachi Bacca; 3 Mar.
	1971; fr.; Granville 752; P • Rivière Petite Ouaqui, rive
	droite, au niveau de l'ancien village Hubert; 20 Jul. 1973;
	fr.; Granville 1872; CAY, P • Sommet Tabulaire, zone nord;
	2 Sep. 1980; fr.; Granville 3710; P • Monts Bakra, 2,5 km
	à l' ouest du Pic Coudreau; 2 Oct. 1980; fr.; Granville 4065;
	CAY, P • Mont Bellevue de l'Inini, central zone, slope
	near the summit; 23 Aug. 1985; fr.; Granville 7762; CAY,
	F, INPA, MG, MO, NY, P, U • Région de l'Inini, Mount
	Atachi Bacca, NW summit, 6 km E of Gobaya Soula,
	near Camp 2; 10 Jan. 1989; fl.; Granville et al. 10541;
	CAY, MO n.v., P, U • Montagne de la Trinité, Bassin
	de la Mana, partie supérieure de la pente N du plateau
	tabulaire; 13 Mar. 1997; fr.; Granville 13318; CAY, K, MO,
	P, U, US • Montagne Lucifer, SW du plateau sommital;
	4°46'30", 53°56'30"W; 520 m; 18 Nov. 1999; fr.; Granville
	& Crozier 13869; CAY, MO • Monts Bakra, 1.5 km W du

  MO n.v., NY, US, Mount Ayanganna, east slope; 13 Mar. 2014;[START_REF] Turland | International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress[END_REF] Radosavljevic et al. 146;[START_REF] Bohn | [END_REF] Notes. This species was placed by[START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] in their Panurensis group, together with C. panurensis from Brazilian and Colombian Amazon, but the two differ in so many characters that a close relationship between them seems unlikely. In fact, C. panurensis has several aberrant characters within the genus: almost spiciform inflorescences (the lateral branches being extremely reduced), an entire disk, large mitriform stipules usually with a prominent midrib, and secondary leaf veins forming conspicuous loops far from the margin. On the other hand, C. tillettii has thyrsoid inflorescences, a bipartite disk, stipules smaller than those of C. panurensis and without prominent midrib, and secondary leaf veins looping near the margin. All these characters fit very well with[START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] Carapichea group, where C. tillettii most closely resembles C. franquevilleana and C. klugii. It differs from these species (which are

	Pakaraima Mountains, Mount Wokomung, summit
	plateau, from central plateau 1-2 km to escarpment;
	5°04'N, 59°52'W; 1500-1530 m; 19 Feb. 1993; fr.; Henkel
	& Chin 1483;
	1.3 mm, acute at apex, glabrous.
	Short-styled flowers: stamens exserted, filaments 2.5
	mm long, anthers narrowly oblong, 1.8 mm long; style
	included, 2.5 mm long, glabrous. Long-styled flowers:
	stamens included; style exserted, 8 mm long, shortly bifid,
	glabrous. Fruits ellipsoid to ovoid or subglobose, 7-10 ×
	5-8 mm, costate when dry, dark red or maroon (probably
	when immature) to purplish-black. Pyrenes plano-convex,
	elliptic to oblong in outline, 6-9 × 4.5-6.0 mm, dorsal
	side 3-4-costate, ventral side longitudinally sulcate. Seeds
	entire, C-shaped in cross-section.
	Distribution. Endemic to western Guyana, Potaro-
	Siparuni and Cuyuni-Mazaruni Regions, on Merume
	Mountains, Mount Ayanganna, and Mount Wokomung,
	which are the easternmost extensions of the Pakaraima
	Mountains.
	Ecology. Growing in dense scrub forest on tepui sandstone,
	at 1070-1570 m elevation.
	Phenology. Flowering specimens were collected in June
	and July, and fruiting specimens in February, March,
	June, and July.
	Selected specimens examined. GUYANA • Potaro-
	Siparuni Region, Ayanganna Slope; 2 Mar. 1960; fr.; R.
	Browne 118 (Forest Department of British Guiana No.
	7942); NY • Potaro-Siparuni Region, Mount Ayanganna,
	E face, camp above first of four escarpments; 5°20'19"N,
	59°56'46"W; 1070 m; 12 Jun. 2001; fl., fr.; Clarke et al.
	9062; MO n.v., US; ibid., Clarke et al. 9063; MO n.v., US
	• Potaro-Siparuni Region, Mount Ayanganna, E face,
	plateau above third of four escarpments; 5°23'12"N,
	59°58'36"W; 1570 m; 19 Jun. 2001; fr.; Clarke et al.
	9350; MO n.v., US • Potaro-Siparuni Region, Mount
	Ayanganna, E face; 5°23'05"N, 59°58'33"W; 1545 m; 26
	Jun. 2001; fl., fr.; Clarke et al. 9565; MO n.v., NY, US •
	Potaro-Siparuni Region, Mount Wokomung, easternmost
	pinnacle of massif; 5°05'34"N, 59°50'13"W; 1524 m; 30
	Jun. 2003; fl. buds; Clarke et al. 10341; MO n.v., NY, US
	• Potaro-Siparuni Region, Mount Wokomung, base of
	fourth escarpment; 5°05'39"N, 59°50'36"W; 1375 m; 4 Jul.
	2003; fr.; Clarke et al. 10508; MO n.v., NY, US • Potaro-
	Siparuni Region, Mount Wokomung, Little Ayanganna,
	summit of highest point of Mount Wokomung massif;
	5°04'53"N, 59°50'26"W; 1660 m; 6 Jul. 2003; fl.; Clarke
	et al. 10588; MO n.v., NY, US • Potaro-Siparuni Region,
	Mount Wokomung, summit; 5°04'3"N, 59°51'42"W; 1560
	m; 10 Jul. 2003; fl. buds; Clarke et al. 10713; MO n.v., US
	• Potaro-Siparuni Region, Mount Wokomung, summit;
	5°04'03"N, 59°51'42"W; 1560 m; 10 Jul. 2003; fl. buds;
	Clarke et al. 10757; MO n.v., US • Potaro-Siparuni Region,

  in their Altsonii group, together with C. altsonii, C. nivea, and C. sandwithiana. However, as discussed above, it is quite different from these three species (which are here transferred to the genus Notopleura) in characters of the stipules, leaf venation, and pyrenes, and is better placed in[START_REF] Taylor | The genus Carapichea (Rubiaceae, Psychotrieae)[END_REF] Carapichea group, alongside with C. guianensis and C. galbaoensis, which are quite similar in vegetative characters. Due to the small number of collections, it is not known whether the flowers of this species are heterostylous. In the few flowers that we analyzed, the stamens are included and the style is barely exserted, which is consistent with a long-styled form. Style glabrous, included, 8 mm long; branches oblong, ca 0.6 mm long. Fruits oblong-ovoid, 5-6 × 3-3.75 mm, smooth (slightly costate when dry), colour unknown (pale olive-green when dry). Pyrenes semi-ellipsoid, 5-6 × 2 mm, strongly compressed dorso-ventrally, dorsal side with 4 prominent ridges, ventral side with 2 longitudinal grooves separated by an irregular median crest prolonged at apex into a short spine. Seeds unknown. Distribution. Endemic to Guyana and reported from the Potaro River Basin (Kaieteur Savanna, Amatuk Portage, and Hoit Mountain), the Upper Mazaruni River Basin (Ayanganna Plateau), and the Cuyuni-Mazaruni Region (W branch of Eping River). Ecology. Growing in understory of primary and secondary forests, sometimes on sandstone or white sands, on lowland and mountain slopes, at 274-610 m elevation. Phenology. Flowering specimens were collected in February, April-May, and July-August; and specimens with fruits only once in August. Specimens examined. GUYANA • Kaieteur Savanna; 1936; fl.; Hollister s.n.; NY • Upper Mazaruni River Basin, Ayanganna Plateau, between Pong River (686 m) and G.S. Camp 13, in Dicymbe forest, on trail to Ayanganna; 25 Jul. 1960; fl.; Tillett et al. 44925; NY • Potaro River Gorge, Amatuk Portage; 27 Apr. 1944; fl.; Maguire & Fanshawe 23018; K, NY • Cuyuni-Mazaruni Region, W branch of Eping River; 7 Feb. 1991; fl.; McDowell & Stobey 3924; MO n.v., US • Potaro-Siparuni Region, black water creek near Camp NW along ravine, rocky ravine, mixed premontane wet forest; 5°12'N, 59°10'W; 274-610 m; 23 May 1991; fl.; McDowell et al. 4901; MO n.v., US • "Amutu" [Amatuk] Portage path; 20 Feb. 1879; fl.; im Thurn s.n.; K • Amatuk Falls, Potaro River; 26 Aug. 1933; fl.; Tutin 600; BM • Potaro district, Hoit Mountain; 2 Aug. 1959; fl.; Whitton 63; K.

	Psychotria altsonii (Sandwith) Steyerm. (Steyermark	Distribution. Only known from the foothills and higher
	1972: 555)	elevations of the Pakaraima Mountains, Guyana.
	Carapichea altsonii (Sandwith) C.M.Taylor (Taylor and Gereau 2013: 121)	Ecology. In understory of moist, non-flooded forest, commonly on boulders of sandstone or conglomerates; 85-1000 m elevation.
	Type. GUYANA -Cuyuni-Mazaruni Region • Macreba	Phenology. Specimens with flowers were collected from
	Falls, Kurupung River; Aug. 1925; fl. & fr. imm.; Altson	July to November, and with immature fruits in August.
	322; lectotype: K [K000174497], designated by Taylor and	Local names. The two following vernacular names are
	Gereau (2013: 121); isolectotypes: K [K000174398], NY	reported in Akawai (Guyana): duma-ek (Altson 522) and
	[00131011]; photo-K at NY.	waguk-wa-eba (Forest Department 2773).
	Species excluded from the genus Carapichea Palicourea hoffmannseggiana (Schult.) Borhidi (Borhidi 2011: 245) Carapichea patrisii DC. (Candolle 1830: 536) -Type: FRENCH GUIANA • s.loc.; s.d.; Patris s.n.; lectotype: G-DC [sheet N. 4], designated by Delprete and Kirkbride (2016: 422); possible isolectotype: G-DC [sheets N. 1, 2, and 3]. Cephaelis patrisii (DC.) D.Dietr. (Dietrich 1839: 773) -Type: same as for Carapichea patrisii. Uragoga carapichea Kuntze (Kuntze 1891: 955), non Uragoga patrisii (DC.) Kuntze (Kuntze 1891) -Type: same as for Carapichea patrisii. Type. BRAZIL -Pará • s.loc.; s.d.; Sieber s.n. [dedit J.C. Hoffmannsegg]; holotype: B [B-W4155]; photograph: NY. Specimens examined. GUYANA • near summit of Description. Shrub, 1.5-3 m tall, single-stemmed, glabrous; Kurupung Mts on trail from Macreba Falls to Kamarang; distal internodes terete, 4.5-8.0 mm in diam., somewhat 13 Sep. 1938; fl.; Forest Department 2773; K • Pakaraima secondary veins, terminating far from the margin; tertiary succulent, glabrous. Stipules consisting of a persistent, Mountains, Membaru-Kurupung trail, Dicymbe pole veins invisible, embedded within the lamina; domatia truncate sheath (absent in flower-bearing nodes), 2.5-3.5 forest; 1000 m; 29 Oct.-4 Nov. 1951; fl.; Maguire & absent. Inflorescence terminal, pedunculate, capitate; × 6.5-14 mm, glabrous, and a dense mass of basal linear Fanshawe 32353; NY • Upper Mazaruni River, Kurupung peduncles 2.5-5.3 cm long, glabrous, drying dark brown; appendages 1.0-2.5 mm long, very early caducous, leaving a River, Makreba Falls, on Kamarung trail; Sep. 1938-Feb. head multiflorous, subtended by a cupular structure scar. Leaves with petioles 1.0-5.5 cm long, glabrous; blades 1939; fl.; Pinkus 5; NY • Pakaraima Mountains, Kurupung 1.0-2.5 cm long, made of 4 bracts connate at base; free elliptic to oblanceolate, 14-35 × 4.5-10 cm, acute-decurrent River, Makreba Falls; 17 Jul. 1992; fl.; Hoffman et al. 2050; portion of bracts decussate, unequal to subequal, broadly at base, acute and long-acuminate at apex, acumen MO, US. ovate, 1.5-2.6 × 1.5-2.5 cm, acute to obtuse at apex, fleshy, narrowly triangular, 1-2 cm long, somewhat succulent Notes. This species closely resembles N. nivea and N. pure white, persistent, drying brown, with 3-5 ascending to fleshy when fresh, drying papyraceous, dark brown sandwithiana; differences between them are summarized veins; bracteoles triangular-ovate, 0.7-2.0 × 0.2-0.5 mm. above and pale brown to pinkish-brown below, glabrous in Table 3. Flowers 5-merous, pedicellate; pedicels 1.8-2.5 mm long, throughout; secondary veins 18-21 on each side of midrib; fleshy. Hypanthium truncate-obconical, 0.6-0.7 mm embedded in the lamina, barely visible above and obsolete Notopleura nivea (Sandwith) O.Lachenaud & Delprete, long, glabrous. Disk bilobed to the base, 0.5-1.0 mm long, below in dried specimens; intersecondary veins 1-3 comb. nov. bipartite. Calyx 0.7-1.3 mm long, shallowly denticulate, between each couple of secondary veins, terminating far urn:lsid:ipni.org:names:77302855-1 glabrous; teeth broadly to shallowly triangular, 0.4-0.5 from the margin; tertiary veins invisible, embedded within the lamina; domatia absent. Inflorescence terminal, long-pedunculate, capitate; peduncles 7.5-17 cm long, glabrous, drying brownish-black; head multiflorous, subtended by a cup-shaped structure 3-5 mm long made by 4 bracts connate at base; free portion of bracts in two decussate pairs, unequal to subequal, obovate to broadly ovate, 1.5-2.7 × 0.9-3.2 cm, pale greenish-white, white or pale violet, blades elliptic to oblanceolate, 18-29 × 5.7-11.6 cm, Stamens inserted 3-4 mm from corolla base, anthers with petioles 1.0-2.5 cm long, glabrous, drying black; 1-2 mm, acute at apex, glabrous outside, papillose inside. dense mass, very early caducous, leaving a scar. Leaves outside and inside; lobes triangular to ovate, 1.5-3.0 × basal linear appendages ca 2 mm long, aggregated into a subcylindrical, 9-12 mm long, 1-2 mm wide, glabrous nodes) 3-9 × 7-10 mm, glabrous, persistent, and numerous mm long, glabrous, white with lobes yellow inside; tube consisting of a truncate sheath (absent at flower-bearing 0.2 mm long, glabrous. Corolla hypocrateriform, 10.5-15 4-7 mm in diam., somewhat succulent, glabrous. Stipules mm long, truncate, slightly undulate or with minute teeth < (maybe more?), single-stemmed; distal internodes terete, base, 0.5-0.8 mm long, glabrous. Calyx cupular, 0.7-1.2 Description. Perennial herb or subshrub, 18-30 cm tall obovoid, 1.5-2.0 mm long, glabrous. Disk bilobed to the collection n.v., NY [00131025]. (to 3 mm long in fruiting stage). Hypanthium narrowly designated here; isolectotypes: K [K000662774], K spirit pedicellate; pedicels to 2 mm long at flowering stage 1249; lectotype: K [2 sheets, K000174488, K000174489], × 0.5-3.0 mm. Flowers 5-merous, subsessile to shortly River, Amatuk portage; 31 Aug. 1937; fl., fr.; Sandwith brown, glabrous; bracteoles subulate to lanceolate, 5-10 Type. GUYANA -Potaro-Siparuni Region • Potaro often tinged with pink or purple, persistent, drying dark mm long. Corolla hypocrateriform, 17.7-21.5 mm long, Cephaelis nivea Sandwith, Bulletin of Miscellaneous glabrous, white with lobes yellow inside; tube narrowly Information, Royal Gardens, Kew 1939: 551 (Sandwith cylindrical, 16-19 mm long, 1.3-1.5 mm wide, glabrous 1939) outside and inside; lobes ovate to triangular, 1-2.6 × 1-2.3 Psychotria nivea (Sandwith) Steyerm. (Steyermark 1972: mm, acute at apex, glabrous outside, papillose inside. 556) Stamens inserted at 9.5-12 mm from the base; filaments Carapichea nivea (Sandwith) C.M.Taylor (Taylor and extremely short; anthers included, narrowly oblong, 3 × Gereau 2013: 121) 0.3-0.4 mm.
	Species transferred to Notopleura subsessile, included, narrowly oblong, 2-3 × 0.4 mm. Style acute-decurrent at base, obtuse to acute and acuminate
	glabrous, 10 mm long, barely exserted; stigmas linear,	at apex, acumen deltoid to narrowly triangular, 0.5-1.0
	1 mm long. Fruits (slightly immature) ovoid to ellipsoid,	cm long, coriaceous to somewhat succulent when fresh,
	Notopleura altsonii (Sandwith) O.Lachenaud & 4-6 × 2.0-3.5 mm, slightly costate when dry, white to pale papyraceous when dry, drying greyish olive-green to
	Delprete, comb. nov. violet. Pyrenes plano-convex, elliptic in outline, 4-6 × brown above and pinkish to greyish pale brown or pale
	urn:lsid:ipni.org:names:77302854-1 2.0-3.5 mm, strongly compressed dorso-ventrally, dorsal olive green below, glabrous throughout; secondary veins
	side with 3 or 4 ridges, ventral side with 2 very shallow Cephaelis altsonii Sandwith, Bulletin of Miscellaneous 12-19 on each side of midrib, embedded within the depressions separated by a longitudinal median crest. lamina, barely visible above and below in dried specimens; Information, Royal Gardens, Kew 1931: 475 (Sandwith 1931) Seeds unknown. intersecondary veins (1-)2-3 between each couple of

Table 3 .

 3 Diagnostic characters of Notopleura altsonii,N. nivea, and N. sandwithiana.

		N. altsonii	N. nivea	N. sandwithiana
	Number of involucral bracts	Four	Four	Two
	Bracts	Connate at base for 0.3-0.5	Connate at base into an urn	Connate at base for 0.3 cm
		cm	1.0-1.8 cm deep
	Peduncle length	7.5-17 cm	2.5-5.3 cm	4.5 cm
	Pedicels	0-3 mm	1.8-2.5 mm	7-8 mm
	Corolla tube	9-11 mm long	16-19 mm long	9-11 mm long
	Distribution	Guyana	Guyana	Venezuela

ACKNOWLEDGEMENTS

We are grateful to the directors and curators of BR, F, L, NY, U, and US for large loans of specimens to CAY, which allowed considerable advancement of the Rubiaceae treatment for the Flora of the Guianas, and access to the material studied for this paper. The herbarium curators of BM, K, and P are thanked for their help while working in their institutes and/or for sending specimens on loan. We wish to thank Andreas Fleischmann, M herbarium curator, for confirming that there is no specimen of Spruce 3445 (type of Psychotria pacimonica) at M. Funds for field work by PD in the surroundings of the Chenapou Village, Upper Potaro River, Guyana, and along the Rio Negro Amazonas, Brazil, as well as for the study of herbarium specimens at the INPA herbarium (Manaus, Amazonas, Brazil) were provided by LABEX CEBA (Laboratory of Excellence -Center for the Study of Biodiversity in Amazonia; grant managed by the Agence Nationale de la Recherche, ANR-10-LABX-0025). PD would also like to thank Michael Hopkins, INPA herbarium curator, for assistance in the herbarium and for hosting him in his private residence during his stay in Manaus. During the expedition to the Chenapou Village, PD was accompanied by Paul Benjamin, member of the Chenapou community, who is acknowledged for his help during field work and processing of specimens. The staff of the Biodiversity Center of the University of Guyana, Georgetown, especially Kaslyn Holder-Collins and Elford Liverpool, is acknowledged for help with gathering field supplies, drying facilities, for their valuable collaboration, and for