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A taxonomic revision of the Rudgea hostmanniana complex in the Guiana shield region is presented. This group includes species with sheathing stipules bearing dorsal appendages, thick glabrous leaves lacking domatia, inflorescences thyrsoid to glomerulate but not regularly dichotomous, a calyx tube absent or very short, a corolla tube 2–7(–13) mm long, and fruits usually brownish when immature and turning red or orange at maturity. Eight species are recognized, among which Rudgea billietiae, endemic to Kaw Mountain in French Guiana, is newly described and illustrated; it is assessed as Endangered according to IUCN criteria. The other species are R. bolivarensis, R. cornigera, R. coussareoides, R. hostmanniana, R. maypurensis, R. pungens, and R. tanaosepala. A key to the species is presented, as well as a table summarising their diagnostic characters.

2. Rudgea bolivarensis Steyermark (1967: 415). Fig. 1A, 2A Type: — VENEZUELA. State of Bolivar, Gran Sabana, along wooded portion of quebrada tributary to Río Kukenan, south of Mount Roraima, 1005–1065 m, 2 October 1944 (fr.), J.A . Steyermark 59118 (holotype, VEN! [VEN15991]; isotype, NY! [NY00133207]). Much-branched shrub or tree 3–15 m tall; twigs 3–5 mm thick, glabrous, soon covered with a pale buffish-straw bark. Stipules 3.5–5 × 3–10 mm, glabrous, marcescent but soon damaged, consisting of a ± truncate sheath (usually split at flower-bearing nodes) bearing on each side of the node a dense mass of 10–25 thick aculeiform dorsal appendages 1–1.3 mm long, shorter than the sheath and inserted between 1/3 and 2/3 of its length from the base. Leaves opposite; petioles 0.3–2 cm long, glabrous; blades elliptic, 4–18.5 × 1.8–9 cm, acute to rounded at base, obtuse to hardly acuminate at apex, very coriaceous with thickened margin, entirely glabrous, drying olive brown to olive green; midrib slightly concave above; secondary veins 7–12 on each side of midrib, moderately ascending, forming irregular loops 1.5–8 mm from the margin; tertiary venation densely reticulate and prominent below (at least in the dry state); domatia absent. Inflorescences terminal, in rather lax panicles, 6.5–14 cm long, erect, minutely spreading-puberulous (the peduncle ± glabrous); peduncle terete, 4.5–8.5 cm long; branched portion 1.5–5.5 × 3.5–7.8 cm; secondary branches 3–4 per node, 0.4–2.8 cm long; bracts minute, ca. 0.7 x 0.5 mm long, triangular to linear, puberulous. Flowers sessile, 5(–6)- merous. Hypanthium cylindrical, 0.7 mm long, glabrous. Calyx tube extremely reduced, lobes triangular to narrowly elliptic, 0.5–1 × 0.3–0.5(–1) mm, obtuse to acute at apex, glabrous or minutely ciliate at apex. Corolla only known in young bud stage; tube glabrous outside, villose at distal portion inside; lobes not dorsally corniculate, glabrous outside. Stamens not seen. Disk shortly cylindrical, 0.3 mm long, glabrous. Style not seen. Fruits obovoid to ellipsoid, 5.5–9 × 4–6 mm when dry, green when immature, bright orange when mature, glabrous, sessile or with pedicel Pyrenes plano-convex, hemiobovoid, 6–8 × 4.5–5.5 mm, dorsal side faintly ridged with 3–4 dorsal and 2 lateral ridges, smooth between the ridges, ventral side smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —This species occurs in southeastern Venezuela (Mt. Roraima area) and adjacent Guyana and Brazil (Fig. 5), in forest/savanna edges, at 700–1100 m altitude. Phenology: —Specimens with flower buds were collected in December; and with fruits in June-July (immature), October and December (mature or nearly so). Notes: —The original description of this species was based on two specimens, one from Venezuela, and one from Brazil. Several additional specimens from Guyana have since been found, and represent a new record for the country. Strangely enough, two Guyanan specimens (Henkel et al. 5675, Henkel & Chin 5709) have coordinates falling in Brazil, which may be due to an error either of country or of coordinates reported on the label. Mature flowers are still unknown. Zappi & Steyermark (2004: 808) included Rudgea bolivarensis in synonymy under R. hostmanniana, which is similar in inflorescence and fruit characters. The stipules of R. bolivarensis, however, are very different and diagnostic, bearing only dorsal appendages that are shorter than the sheath (all other species of the complex have the stipular appendages exceeding the sheath, except R. coussareoides, which is easily recognized by its ternate leaves). Rudgea bolivarensis also differs from R. hostmanniana by having the corolla glabrous outside, leaves more coriaceous with a conspicuously thickened margin and an evident tertiary venation, and pyrenes with much less prominent dorsal ridges. The supposed differences in inflorescence structure, mentioned by Steyermark (1967), are actually not reliable; he described the pedicels of R. bolivarensis as 5–8 mm long, but the flowers are sessile and the fruits are sessile or subsessile (pedicel Additional Specimens Examined: — BRAZIL. Roraima [“ Rio Branco ”]: along igarapé 5 km SE of Serra Sabang, 720 m, 16–18 December 1954 (fl. buds & fr.), B . Maguire & C. K. Maguire 40276 (NY, VEN). GUYANA. Pakaraima Mountains, Upper Ireng R. watershed, Malakwalai-Tipu, 300 m downslope from NE escarpment, 4°48’N, 60°18’W, 1100 m, 17 July 1994 (imm. fr.), T. W . Henkel, M. Chin & L. Williams 5675 (CAY, U); Pakaraima Mountains, upper Ireng River watershed, E bank Kaalmang River at base of Achiknang, 4°59’N, 60°08’W, 700 m, 19 July 1994 (imm. fr.), T. W . Henkel & M. Chin 5709 (K); Pakaraima Mountains, upper Ireng River, hills 2–3 km E of Cipo settlement, 4°48’N, 60°2’W, 760 m, 12 October 1994 (fr.), P . Mutchnik, T. W. Henkel & L. Williams 1 (K); Northern Pakaraimas, Koa Valley, Annuyeng Creek from mouth to falls, 4°39’48”N, 59°48’33”W, 834 m, 10 June 1995 (imm. fr.), P . Mutchnick 1620 (K)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 226-227, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870, {"references":["Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B.,, Fidalgo, O. & Fidalgo, M. E. P. K. (eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436.","Zappi, D. & Steyermark, J. A. (2004) Rudgea. In: Steyermark, J. A., Berry, P. E., Yatskievytch, K. & Holst, B. K. (eds.) Flora of the Venezuelan Guayana, vol. 8, Poaceae - Rubiaceae. Missouri Botanical Garden Press, St Louis, pp. 805 - 816."]}

7. Rudgea pungens (Steyermark) C.M. Taylor, Bruniera & Zappi (2015: 45, p. 4). – Psychotria pungens Steyermark (1972: 677). (Fig. 7 D–F). Type: — FRENCH GUIANA. Without locality, s.d., F. M. R . Leprieur 118 (holotype, P! [P00837150]; probable isotype (unnumbered collection), G! [G00418599, G00418600]). Shrub 0.3–2 m tall, with nearly horizontal branches; twigs glabrous, 1–1.5 mm thick, soon covered with a pale strawcoloured bark. Stipules 8–16(–20) × 1–7 mm, glabrous, marcescent and soon corky, consisting of a narrow tubular sheath 5–9(–12) mm long (usually split at flower-bearing nodes) bearing 8–15 terminal linear appendages 3–8 mm long, and 5–7 dorsal linear appendages 0.5-3 mm long, forming a very short decurrent keel inserted 1–3 mm above the base of the stipule. Leaves opposite; petioles 0.2–0.6 cm long, glabrous; blades elliptic, 6.5–14.5(–16) × 1.8–4.5(–5.5) cm, acute or obtuse at base, gradually long-acuminate at apex, very thick, entirely glabrous, drying olive green to olive brown; midrib concave above; secondary veins 5–10 on each side of midrib, strongly ascending, forming an angle of 45–60° with the midrib; tertiary veins invisible in fresh leaves, sometimes slightly prominent when dry; domatia absent. Inflorescences terminal, subcapitate and involucrate, ca. 8-flowered, patent to sub-erect, glabrous, sessile or pedunculate; peduncle (when present) terete, to 1 cm long; flower-bearing portion 1.3–2.2 × 1.4–3.5 cm, secondary branches apparently absent or extremely reduced in flower, to 2 mm long in fruit; bracts pale green, numerous and imbricate in several rows, 10–20 × 2–7 mm, lanceolate, entire, acute at apex, glabrous or shortly ciliate, erect or patent, persistent in fruit. Flowers sessile, 5-merous, heterostylous. Hypanthium obovoid, 1.5 mm long, glabrous, Calyx tube 0.5–1 mm long, glabrous; lobes linear to narrowly triangular, 1.2–3.5 × 0.5–1 mm, glabrous or ciliate. Corolla white, tube narrow and almost cylindrical, 13 mm long, 1 mm wide at base, 1.5–2.5 mm wide at mouth, glabrous outside (inside not seen); lobes narrowly triangular, 2.5 × 1 mm, glabrous, with short, obtuse dorsal appendage 0.5 mm long. Stamens included in long-styled flowers, or exserted with filaments exceeding corolla throat by 1 mm in short-styled flowers; anthers 2.3 × 0.3 mm. Disk shortly cylindrical, 0.5 mm long, glabrous. Style exserted, exceeding corolla mouth by 1.5 mm in long-styled flowers, or included in short-styled flowers. Fruits ovoid with truncate apex, 11 × 7–8 mm when fresh, 7–10 × 5–7 mm when dry, dark red-brown and hard when immature, cherry red when mature, glabrous, sessile, crowned with persistent calyx 2.5–4 mm in diameter. Pyrenes plano-convex, hemi-obovoid to hemi-ellipsoid, 9 × 6.5 mm, dorsal side smooth, ventral side smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —This species is only known from French Guiana (Fig. 5), where it occurs on the main summits of the central Inini-Camopi chain (Mts Atachi Bacca to Mts Bakra) and disjunctly at low altitudes in the northeast; it should be expected in adjacent parts of Suriname and Brazil. It is apparently restricted to relatively low forests on rocky substrates, 5-800 m in elevation, and is locally abundant. Phenology: —Flowering collections were made in January, May and August; fruiting collections in January, March–April (full-sized, but still hard) and August (mature fruits). As in the similar Rudgea billietiae, the fruits probably take several months to mature. Notes: —This species, originally described in Psychotria (Steyermark 1972), was recently transferred to Rudgea (Taylor et al. 2015) but its affinities within the genus were still unclear. Taylor and Bruniera (2018) mentioned its similarity to the R. bracteata J.H. Kirkbride (1981: 97) group, especially because of the well-developed bracts, but members of that group have larger fruits, and stipules with a prominent dorsal keel and without marginal appendages. The involucrate inflorescences and narrowly tubular stipules of R. pungens are unusual characters in Rudgea, but the discovery of R. billietiae, with similar stipules and lax inflorescences, links it to R. cornigera, R. hostmanniana and R. tanaosepala, which have lax inflorescences and shorter stipular sheaths. The differences between these species are summarised in Table 1. The original description of the species was based on a single specimen without fruits or corollas, and is therefore incomplete on several points. The species is now known from additional complete material, which allows to present here a full description; only the interior of the corolla tube cannot be described, because the flowers are too few for a dissection to be made. The dimensions in brackets refer to a vegetative collection (Granville 4002) from Mount Bakra, French Guiana, which has slightly larger leaves and stipules; though it almost certainly belongs to this species, confirmation with reproductive material from the same area would be reassuring. Specimens Examined: — FRENCH GUIANA. Crique Kapiri – RN2, bassin de l’Approuague, 4°07’N, 52°05’W, 11 January 1991 (fallen fl.), G . Cremers 11474 (CAY); savane-roche [inselberg] Virginie, 4°11’N, 52°08’W, 18 February 2009 (fallen fl.), C . Delnatte, F. Billiet, J.- J. de Granville & B. Jadin 1682 (CAY); Fleuve Sinnamary, rive droite, layon ONF (n°7) direction Sud, à 7 km du fleuve, 1 May 1969 (fl.), J.- J . de Granville 128 (CAY); sommet des Monts Atachi Bacca, 4 March 1971 (fr.), J.- J . de Granville 765 (CAY, P); Monts Galbao, 10 km WSW Saül, 14 March 1973 (imm. fr.), J.- J . de Granville 1534 (CAY, P); Monts Bakra, versant Sud, 5 km WSW du pic Coudreau, 28 September 1980 (st.), J.- J . de Granville 4002 (CAY, P); Montagne Bellevue de l’Inini, zone centrale, 23 August 1985 (fl. & imm. fr.), J.- J . de Granville, L. Allorge, G. Cremers, A. R. A. Görts-van Rijn & J. F. Kodjoed 7770 (BR, CAY, P); Crique Gabaret, bassin de l’Oyapock, Saut Mérignan, 13 April 1988 (fallen fl.), J.- J . de Granville 10275 (CAY); Monts Atachi Bacca, versant Nord, 9 km au SE de Gobaya Soula, 3°33’N, 53°55’W, 12 January 1989 (fl.), J.- J . de Granville, G. Cremers, J. I. Hagemann, B. E. Leuenberger, R. W. Sanders & M. Sangrey 10616 (CAY, P); Monts Atachi Bacca, est du plateau sommital, 3°33’N, 53°55’W, 22 January 1989 (fr.), J.- J . de Granville, G. Cremers, J. I. Hagemann, B. E. Leuenberger, R. W. Sanders & M. Sangrey 10861 (CAY, P); layon Régina – Kaw, 4°21’W – 52°08’W, 7 August 1997 (fallen fl.), V . Hequet 688 (CAY); Route Régina – Saint Georges, piste de la savane-roche [inselberg] Virginie, 4°11’05”N, 52°08’13”W, 6 April 2014 (fr.), O . Lachenaud 1720 (BR, CAY, L, MO, P); without locality, s.d. (fl.), L.C.M. Richard s.n. (P [P04008549], mixed with R. billietiae); sommet nord du Mont Galbao, 29 January 1978 (fallen fl.), Tay 91 (MPU) .

4. Rudgea coussareoides (Standley) C.M. Taylor, Bruniera & Zappi (2015: 45, p. 4 [E-publication]). – Psychotria coussareoides Standley (1931: 441). – Palicourea coussareoides (Standley) Delprete & J.H. Kirkbride (2016: 417). Fig. 1B, 2B. Type: — VENEZUELA. Amazonas, slopes of Mount Duida, Agüita, 4000 ft [ca. 1220 m], 1928–29 (fl. buds), G. H. H . Tate 919 (holotype, NY! [NY00132650]). Rudgea tillettii Steyermark (1967: 416). Type: — GUYANA. Porkknocker Camp 2 on Partang River, about 19 miles from mouth, Merume Mountains, 625 m, 28 June 1960 (fl. buds), S. S . Tillett, C. L. Tillett & R. Boyan 43934 (holotype, NY! [2 sheets, NY00133238, NY00133239], isotypes K! [K000005080], VEN! [VEN 64649]). Psychotria coussareoides var. ciliata Steyermark (1972: 496). Type: — VENEZUELA. Bolivar: Cerro Venamo (parte SO), 950–1000 m, 10 January 1964 (fl. buds), J. A. Steyermark, G. C. K. Dunsterville & E. Dunsterville 92929 (holotype, VEN! [VEN 62981], isotype, NY! [NY00132651]). Shrub or tree 1.5–7 m tall; twigs glabrous, 1.5–3 mm thick, soon covered with a brown corky bark. Stipules 2.5–7 × 4.5–10 mm, glabrous, marcescent but soon damaged, consisting of a truncate sheath bearing 6 groups (2 between every pair of leaves) of ca. 10 thick aculeiform dorsal appendages 0.5–1 mm long, shorter than the sheath and inserted between 2/3 and 3/4 of its length from the base. Leaves ternate; petioles 0.8–2.5 cm long, glabrous; blades elliptic, 6.2–21 × 1.2–7 cm, decurrent on petiole at base, acuminate at apex, coriaceous with slightly recurved margin, entirely glabrous, drying olive brown to dark brown; midrib slightly prominent above; secondary veins 8–11 on each side of midrib, weakly to moderately ascending, forming irregular loops 0.5–2 mm from the margin; tertiary venation prominently reticulate (at least in the dry state) and concolorous below, the veins 1–2 mm apart; domatia absent. Inflorescences terminal, pale green to purplish (at least in the fruiting stage), in pyramidal panicles, 5.7–13.5 cm long, erect, minutely spreading-puberulous; peduncle terete, 2.2–4.2 cm long; branched portion 3.5–10.5 × 3–8 cm; secondary branches (2–)3–4 per node, 1.2–2 cm long; bracts of basal node resembling small leaves, 8–27 × 1–5.5 mm, narrowly elliptic, glabrous, the other bracts minutely triangular, 1–2.5 × 0.5–2 mm, entire or dentate, glabrous to puberulous. Flowers sessile, 5-merous, fragrant. Hypanthium cylindrical, 0.8–1 mm long, glabrous. Calyx shortly cupuliform, glabrous to puberulous; tube 0.3–0.7 mm long; lobes triangular, 0.2–0.3 mm long. Corolla white, hypocrateriform; tube funnelshaped, 2–2.5 × 1.2–1.8 mm, glabrous to papillose outside, densely villose inside except towards the base; lobes narrowly triangular, 2.5–3 × 1–1.5 mm, acute at apex, glabrous outside, papillose inside. Stamens mostly included with only the tips exserted, anthers 1 × 0.5 mm. Disk cylindrical, 0.3–0.7 mm long, glabrous. Style exserted, 4.5–5.5 mm long, bilobed, glabrous. Fruits ellipsoid to subglobose, 5.5–8 × 4–5.5 mm when dry, green or greyish-blue when immature, dark purple when mature, glabrous, sessile or with pedicel Pyrenes plano-convex, hemi-ellipsoid, 7–7.5 × 5–5.5 mm; dorsal side faintly and irregularly ridged, slightly granulose, ventral side smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —Occurring in western Guyana, eastern Venezuela, and northern Brazil (Amazonas) and apparently in disjunct populations in Amazonian Peru (Fig. 6); in tepui slope forests, sometimes on riverbanks, mostly 700–1500 m but as low as 300 m altitude in Peru. Phenology: —Flowers in October (buds), January-February, and June; fruits in January, April and October-November. Notes: —The systematic position of this species, which has only recently been transferred to Rudgea from Psychotria (Taylor et al. 2015) has long been unclear. It appears that it is part of the R. hostmanniana complex, and in particular closely resembles R. bolivarensis in stipule morphology. It is easily separated from the latter – and from all species of Rudgea in the Guiana Shield region – by its ternate, rather than opposite, leaves. Furthermore, its stipules bear two separate groups of appendages between every leaf pairs (as opposed to one in R. bolivarensis) and its inflorescences are also more slender than in R. bolivarensis. It is unclear whether the flowers of this species are heterostylous; only two specimens with open flowers have been seen, and both seem to pertain to the long-styled form. A specimen from Brazil, cited below, represents a new record for the country; though it was seen in photograph only, this identification is without any doubt. A previous record of this species from French Guiana (Funk et al. 2007, as Psychotria coussareoides) is presumably based on the specimen R.A.A. Oldeman & C. Sastre 88 (CAY), which is Coussarea sp. No authentic material of R. coussareoides from French Guiana has been seen by the authors. Additional Specimens Examined: — BRAZIL. Amazonas: Manacapuru, margen do lago grande, operação Radam, ponto 4, 9 October 1976 (fl. buds), T . R. Bahia 219 (F n.v., NY, photo K). GUYANA. Upper Mazaruni River region, Karowtipu Mountain, 5°45’N, 60°35’W, 1000 m, 25 April 1987 (fr.), B . M. Boom & D. Gopaul 7728 (MO); Pakaraima Mountains, Mount Wokomung, Wusupubaru Creek, 2 km from juncture with Suruwubaru Creek, 5°03’N, 59°53’W, 975–1125 m, 13 February 1993 (fl.), T . M. Henkel, M. Chin & W. Ryan 1326 (MO); Pakaraima Mountains, Mount Wokomung, headwaters of Wusupubaru Creek, 5°03’N, 59°53’W, 975–1125 m, 15 February 1993 (fl.), T . M. Henkel, M. Chin & W. Ryan 1385 (NY). PERU. Amazonas: Distrito Imaza, comunidad Aguaruna Putuim, Anexo de Yamayakat, 700–750 m, 21 January 1996 (fr.), C . Díaz S., H. Osores, H. Díaz & D. Díaz 7727 A (K). VENEZUELA. Amazonas: Distrito Atabapo, Cerro Marahuaca, 3°43’N, 65°30’W, 1200 m, 16 October 1988 (fr.), R . Liesner 24924 (K); Distrito Atabapo, Cerro Huachamacari, 3°49’N, 65°43’W, 800–1300 m, 5 November 1988 (fr.), R . Liesner 25881 (K)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 229-231, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870, {"references":["Taylor, C. M., Bruniera, C. P. & Zappi, D. C. (2015) Taxonomic transfers in Neotropical Palicoureeae: new combinations in Rudgea and Palicourea. Kew Bulletin 70: 45, 1 - 7 [E-publication]. https: // doi. org / 10.1007 / s 12225 - 015 - 9596 - 3","Standley, P. C. (1931) The Rubiaceae of Venezuela. Publications of the Field Museum of Natural History, Botanical Series 7 (4): 343 - 485. https: // doi. org / 10.5962 / bhl. title. 2347","Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B.,, Fidalgo, O. & Fidalgo, M. E. P. K. (eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436.","Steyermark, J. A. (1972) Rubiaceae. In: Maguire, B. & Steyermark, J. A. (eds.) The Botany of the Guayana Highland-Part IX. Memoirs of the New York Botanical Garden 23: 227 - 832.","Funk, V. A., Berry, P. E., Alexander, S., Hollowell, T. H. & Kelloff, C. L. (2007) Checklist of the Plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions of the United States National Herbarium 55: 1 - 584."]}

Key to the species 1. Leaves ternate; stipular appendages all dorsal and shorter than sheath (Fig. 1B) (Guyana, Venezuela, N. Brazil, Peru)..................................................................................................................................................................................................... R. coussareoides - Leaves opposite; stipular appendages exceeding sheath, except in R. bolivarensis...........................................................................2 2. Stipules narrowly tubular with sheath longer than broad (but often splitting very early at flower-bearing nodes); inflorescences glabrous, sometimes with large foliaceous bracts; corolla lobes corniculate, glabrous outside........................................................3 - Stipules with sheath broader than long; inflorescences puberulous (rarely glabrous in R. maypurensis), never with large foliaceous bracts; corolla lobes not corniculate, or, if corniculate, pubescent or puberulous outside at least near the apex..............................4 3. Stipules with dorsal appendages inserted near the base; inflorescence subcapitate with large foliaceous imbricate bracts 10–20 x 2–7 mm; corolla tube 13 mm long (French Guiana)......................................................................................................... R. pungens - Stipules with dorsal appendages inserted in upper half; inflorescence distinctly branched with small subulate bracts 3–5 mm long; corolla tube 6 mm long (French Guiana: Kaw Mountain)................................................................................................ R. billietiae 4. Stipules ± truncate with appendages shorter than sheath (Fig. 1A); leaves thickly coriaceous, with conspicuously reticulate tertiary venation; corolla lobes glabrous outside and not corniculate (Venezuela, Guyana, N Brazil)................................... R. bolivarensis - Stipules with appendages exceeding sheath; leaves with tertiary venation usually rather obscure; corolla lobes corniculate except in R. maypurensis...............................................................................................................................................................................5 5. Stipules with dorsal appendages recurved, free at base or shortly connate into a keel (Fig. 1E); leaves slightly cordate to rounded at base; corolla lobes not corniculate; pyrenes dorsally verrucose (Venezuela, N Brazil,?SE Colombia)................ R. maypurensis - Stipules with appendages all erect, the dorsal ones connate into a keel (Fig. 1C–D & F); leaves cuneate to obtuse at base, never cordate; corolla lobes corniculate; pyrenes not verrucose dorsally....................................................................................................6 6. Calyx lobes 3–5 mm long, linear (Fig. 2F); bracts 5–7 mm long, linear to narrowly lanceolate; corolla lobes usually with linear appendages (rarely these short and blunt); fruits ellipsoid to globose; leaf blade with secondary veins weakly ascending (Guyana)...................................................................................................................................................................... R. tanaosepala - Calyx lobes 0.7–2 mm long, semicircular to subulate (Fig. 2 C-D); bracts usually R. hostmanniana) then relatively broad; corolla lobes with short and blunt appendages; fruits obovoid, rarely subglobose; leaf blade with secondary veins usually strongly ascending...............................................................................................................................7 7. Stipules with central keel much exceeding the lateral appendages (Fig. 1D); leaves acute (rarely obtuse) at base, with petiole 0.5–2 cm long; inflorescence pyramidal or rarely hemispherical; calyx lobes triangular to semicircular (Fig. 2D); corolla tube 3–5 mm long (widespread).................................................................................................................................................... R. hostmanniana - Stipules with central keel roughly equalling the lateral appendages (Fig. 1C); leaves obtuse at base, with petiole 0.2–0.8 cm long; inflorescence hemispherical; calyx lobes subulate to narrowly triangular (Fig. 2C); corolla tube 5–7 mm long (Guyana, Suriname)........................................................................................................................................................................ R. cornigera, Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 222-223, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870

8. Rudgea tanaosepala Sandwith (1933: 334). Fig. 1F, 2F. Type: — GUYANA. Simuni Creek, Rupununi River, 25 August 1931 (fl.), T. A. W . Davis in Forest Department 2149 (holotype, K! [2 sheets, K000447201, K000447202]). Much branched shrub 2–7 m tall; twigs glabrous or minutely puberulous, 2–2.5 mm thick, soon covered with a pale straw-colored bark. Stipules 6.5–12 × 2–4.5 mm, glabrous or minutely puberulous, marcescent and soon corky, consisting of a short basal sheath 2–3.5 mm long (usually split at flower-bearing nodes) bearing on each side of the node one bifid lateral appendage 3–5.5 × 0.5 mm, and a central keel (4.5–)7–13 × 1–2 mm, the latter usually bifid for 2.5–4 mm long (rarely ± irregularly fimbriate) with each of the lobes divided in 3–5 linear appendages 1–2.5 mm long. Leaves opposite; petioles 0.1–0.7 cm long, glabrous or minutely puberulous; blades oblanceolate to narrowly elliptic, 7–20.7 × 1.8–7.2 cm, obtuse to rounded at base, gradually acuminate at apex, very thick, entirely glabrous, drying greyish-green; midrib concave or flat above; secondary veins 7–11 on each side of midrib, weakly ascending and forming loops 2–3 mm before margin; tertiary veins very lax and slightly prominent when dry (probably invisible when fresh); domatia absent. Inflorescences terminal, paniculate or rarely glomerulate, 0.8–4.4 cm long, erect, puberulous; peduncle terete, 0.1–2.4 cm long; flower-bearing portion 0.5–2.2 × 1.4–3.3 cm; secondary branches 3–4 per node, to 1.2 cm long, rarely absent; bracts 5–7 × 0.7–1.2 mm, linear to narrowly lanceolate, entire or with linear lateral teeth, sparsely and shortly puberulous outside. Flowers sessile, 5-merous, heterostylous, fragrant. Hypanthium obconical to almost cylindrical, 0.8 mm long, glabrous. Calyx tube extremely reduced, lobes linear, 3–5 × 0.7–1.2 mm, shortly and sparsely puberulous outside. Corolla white, hypocrateriform; tube narrowly infundibuliform, 6–7 mm long, 1–1.3 mm wide at base, 1.5–3 mm wide at mouth, glabrous outside, with a ring of dense short hairs at stamens insertion inside; lobes triangular to narrowly elliptic, 3.5–5 × 1–1.5 mm, puberulous outside at the apex, minutely papillose inside, with narrowly cylindrical (or rarely short and obtuse) dorsal cornicula (0.5–) 1–1.5 mm long. Stamens included in long-styled flowers, or exserted with filaments exceeding corolla mouth by 2–5 mm and anthers 2.2 × 0.4 mm in shortstyled flowers. Disk shortly cylindrical, 0.5 mm long, glabrous. Style exserted, 9–10 mm long in long-styled flowers, or included, 6 mm long, in short-styled flowers, lobes with papillose stigmatic surface. Fruits globose to ellipsoid, 7–11 × 7–11 mm when dry, dark brown (presumably immature) to red or orange (mature), glabrous, sessile, crowned with slightly accrescent persistent calyx 2.5 mm wide. Pyrenes plano-convex, hemi-ellipsoid or hemi-obovoid, 7–7.5 × 6.5 mm, dorsal side with 3 often very faint ridges mostly visible at the base, smooth between the ridges, ventral side smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —A species endemic to central Guyana (Fig. 5), locally common in the middle Essequibo River basin; it grows in lowland forest on white or brown sand, sometimes along streams, at 80–500 m elevation. Phenology: —Flowering specimens were collected in August and October–November; and fruiting specimens in February–April and September–October. Notes: —This species was for a long time only known from the type (cf. Steyermark 1967: 407). It is now fairly well represented in herbaria, where most of the specimens had so far been mistaken for the closely similar and much more widespread Rudgea hostmanniana. The species is distinctive, within the R. hostmanniana complex, by its long and very narrow calyx lobes (Fig. 2F; Table 1); the secondary leaf veins are also less ascending than in other species, though a few specimens of R. cornigera approach R. tanaosepala in this respect. The inflorescences in R. tanaosepala are quite variable; they are usually well-branched, but sometimes the ramifications are very short or absent (e.g. Hoffmann et al. 1488; Maas et al. 5885). Specimens from the south of the range, including the type, have usually glabrous vegetative parts (rarely the twigs are sparsely puberulous) and corolla lobes with narrowly cylindrical, 1–1.5 mm long dorsal cornicula. The more northern specimens (Ek et al. 602, 772; Maas et al. 5885) have minutely puberulous twigs, stipules and petioles, and corolla lobes with shorter and broader dorsal cornicula, ca. 0.5 mm long; it is possible that they represent a separate infraspecific taxon. Records of R. tanaosepala from French Guiana (Funk et al. 2007) are based on misidentified collections of the recently described R. glomerulata Zappi & O. Lachenaud (Lachenaud et al. 2022: 159). The latter differs from R. tanaosepala and other species of the R. hostmanniana complex by its stipules lacking dorsal appendages, large and deeply laciniate bracts 9–28 × 2.5–3.5 mm, longer corolla tube, 18 – 20 mm long that is entirely glabrous within, longer calyx tube 1.5 – 4 mm long, bipartite disk, yellow to yellow-orange fruits, and densely pubescent twigs, petioles and abaxial side of leaf veins. A collection from the Bakhuis Mountains, in Suriname, Bordenave, Doerga, van Troon & James 8619 (BBS n.v., CAY) resembles the variants of Rudgea tanaosepala with glomerulate inflorescences, but has smaller leaf blades, 5–7 × 0.8–1.7 cm, and stipules with four subequal dorsal appendages on each side and no dorsal keel. It presumably represents a new species, but the material is very poor, with only leaves, hypanthia and calyces. Specimens Examined: — GUYANA. Iwokrama Rainforest Reserve, Iwokrama Mts, 4°20’N, 58°50’W, 22 November 1995 (fl.), D. Clarke & B . Hoffmann 584 (CAY, U); Iwokrama Rainforest Reserve, Moco Moco II Creek, 20 mi. SW of Kurupukari on Kurupukari – Annai road, 4°25’N, 58°49’W, 24 March 1996 (fr.), D. Clarke 1431 (CAY, U); Iwokrama International Rainforest Reserve, Iwokrama Mts, 4°19’N 58°47’W, 22 September 1996 (fr.), D. Clarke 2492 (CAY, K, U); Iwokrama Rainforest Reserve, Iwokrama Mountains, 4°19’N, 58°47’W, 22 September 1996 (fr.), D. Clarke 2516 (CAY, U); Iwokrama Rainforest Reserve, N of Surama, 4°10’N, 59°03’W, 20 May 1995 (fr.), C . Ehringhaus 126 (CAY); Mabura region, West Pibiri compartment, 5°20’N, 58°30’W, 28 November 1992 (fl.), R . C. Ek 602 (U); Mabura region, Pibiri compartment, main road, 5°01.95’N, 58°37.73’W, 20 March 1993 (fr.), R . C. Ek, B. Gravendeel, B. Robers & M. Elsinga 772 (U); Region Potaro-Siparuni, Annai – Karupukari road, 18 km N of Surama village cut off, 0.5 km W of road, 4°14’N, 58°56’W, 29 April 1992 (imm. fr.), B . Hoffman, T. Pennington & C. Capellaro 1488 (CAY, U); Mabura Hill, 5°19’N 58°38’W, 27 November 1981 (fl.), P . J.M. Maas, A. Mennega & B. J. H. ter Welle 5885 (U); Iwokrama Reserve, Essequibo watershed, Georgetown–Lethem road, Mount Daniel transect, 4°28’26”N, 58°47’05”W, 27 February 1995 (fr.), P . Mutchnik 973 (K)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 237-238, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870, {"references":["Sandwith, N. Y. (1933) Contributions to the Flora of Tropical America: XVI. Bulletin of miscellaneous information, Kew 1933: 323 - 339. https: // doi. org / 10.2307 / 4113561","Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B.,, Fidalgo, O. & Fidalgo, M. E. P. K. (eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436.","Funk, V. A., Berry, P. E., Alexander, S., Hollowell, T. H. & Kelloff, C. L. (2007) Checklist of the Plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions of the United States National Herbarium 55: 1 - 584.","Lachenaud, O., Bruniera, C. P. & Zappi, D. C. (2022) Six new and a little-known species of Rudgea (Rubiaceae - Palicoureeae) from the Guianas. Phytotaxa 531 (3): 154 - 174. https: // doi. org / 10.11646 / phytotaxa. 531.3.1"]}

5. Rudgea hostmanniana Bentham (1850: 459) subsp. hostmanniana. Fig. 1D, 2D, 7A–C. Type: — SURINAME. Without locality, 1843 (fl.), F. W. R. Hostmann 548 (lectotype, first-step designated by Steyermark (1967: 413), second-step K! [K000579485], designated here; isolectotypes BM! [BM000832001], G! [G00436709, G00436710], GH! [GH00094434, 00094435], P! [P04008004]). Rudgea intercedens Müller Argoviensis (1881: 205), syn. nov. Type: — BRAZIL. [Pará]: Without locality, s.d. [1785] (fl.), L. C. M . Richard s.n. (lectotype, P! [P03985391], designated here; isolectotype, G! [G00436705]). Much-branched shrub 1–5(–8) m tall; twigs glabrous, 2–4 mm thick, soon covered with a pale buffish-straw bark. Stipules 7–13 × 3–8.5 mm, glabrous, marcescent and soon corky, consisting of a basal sheath 3–5 mm long (usually split at flower-bearing nodes) bearing on each side of the node 1–2 narrowly triangular lateral appendages 2–5 × 0.3–1 mm, and a central keel 5–10 × 1.2–2 mm, much exceeding the latter and divided into 2–8 terminal appendages 2–3 mm long, these often grouped in two phalanges. Leaves opposite; petioles 0.5–2 cm long, glabrous; blades elliptic to slightly oblanceolate, 8.5–20(–22) × 2–7.8(–12) cm, acute or rarely obtuse at base, obtuse to shortly acuminate at apex, very thick, entirely glabrous, drying yellowish-green to olive brown; midrib flat or concave above; secondary veins 8–12 on each side of midrib, rather strongly ascending, forming very inconspicuous loops 1.5–5 mm from the margin; tertiary venation invisible in the fresh state, sometimes prominent but very lax in the dry state; domatia absent. Inflorescences white, terminal, in lax to rather condensed, pyramidal or rarely hemispherical panicles, 5.2–18 cm long, erect, shortly spreading-puberulous, the peduncle often glabrescent; peduncle terete, 3–11 cm long; branched portion 1.7–11 × 2.4–8.5 cm; secondary branches (2–)3–4 per node, 0.5–3 cm long; bracts triangular to lanceolate, 2.5–10 × (0.7–) 1.5–3 mm, entire or the lower ones often dentate at base, acute at apex, glabrous to sparsely pubescent. Flowers sessile, 5-merous, heterostylous. Hypanthium obconical, 0.7 mm long, glabrous. Calyx tube extremely reduced, lobes triangular to semi-circular, 0.7–2 × 0.5–0.8 mm, acute to rounded at apex, glabrous to sparsely pubescent externally, ciliate on the margins. Corolla white, fragrant, hypocrateriform; tube narrowly funnel-shaped, 3–5 mm long, 1–1.2 mm at base, 2–3 mm wide at mouth, glabrous to densely villous outside, densely villous at upper half inside; lobes narrowly triangular, 3–4 × 1.2–1.5 mm, densely villous to sparsely pubescent outside at least near the apex, papillose inside, with hemispherical to conical dorsal cornicula ≤ 0.7 mm long. Stamens included and subsessile in long-styled flowers, or long exserted with filaments 3–4 mm long in short-styled flowers; anthers 1–1.2 x 0.3 mm. Disk shortly cylindrical, 0.6 mm long, glabrous. Style exserted, 5.5–6 mm long in long-styled flowers, or as long as corolla tube, 4.5–5 mm long in short-styled flowers; lobes 0.8–1.5 mm long, stigmatic surface papillose. Fruits obovoid or rarely subglobose, 5.5–10 × 5.5–7.5 mm when dry, green when immature, becoming orange-red later (said to turn black at full maturity), glabrous, sessile, crowned with persistent calyx 1–2 mm in diameter, i.e. not markedly accrescent. Pyrenes plano-convex, hemi-ellipsoid to hemi-obovoid, or rarely hemispherical, 5.5–9 × 4.5–8.5 mm, dorsal side with 3–4 prominent and 2 lateral ridges, smooth between the ridges, ventral side smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —Widespread and common in the three Guianas, occurring also locally in eastern Venezuela, and apparently disjunctly in the Brazilian state of Acre (Fig. 6); occurs mostly in riparian forest and in low coastal forest on white sands, occasionally also on granitic outcrops in the interior, from sea level to 850 m in elevation. Another subspecies, R. hostmanniana subsp. freemanii (Sprague & R.O.Williams ex R.O. Williams & Cheesman 1928: 39) Steyermark (1967: 414) is found in similar habitats in Trinidad, Tobago, Grenada, and the coastal cordillera of Venezuela. Phenology: —Flowering specimens were collected throughout the year; fruiting specimens throughout the year, except December–January, with a peak in August–September. Notes: —This species is here circumscribed in a narrower sense than previously (e.g., Zappi & Steyermark 2004; Bruniera 2015), and as thus includes only two subspecies, which distributions are discussed above. It is similar to Rudgea billietiae, R. bolivarensis, R. cornigera, R. maypurensis and R. tanaosepala, and has frequently been confused with all of them. Differences between these species are summarised in Table 1; especially diagnostic for R. hostmanniana is the shape of the stipules, with the dorsal keel long exceeding the lateral appendages (Fig. 1D). An illustration of R. hostmanniana (sensu stricto) may be found in the Flora de Venezuela (Steyermark 1974, fig. 166) and is quite accurate, although the tertiary leaf veins are usually less apparent than depicted on the figure. The above species description is based on subsp. hostmanniana. Subspecies freemannii is very similar to subsp. hostmanniana, differing only by the densely puberulous hypanthium and outer surface of the calyx, which is a rather slight but apparently constant character. The length of the peduncle, used as an additional diagnostic character by Steyermark (1967) actually shows much overlap between the two taxa. Not all authors have accepted the distinction of subsp. freemannii, for instance Acevedo-Rodriguez & Strong (2012: 842) considered it a synonym of R. hostmanniana. Zappi & Steyermark (2004) recognized three subspecies, although the third one, subsp. maypurensis (Standley) Zappi (in Zappi & Steyermark 2004: 808), is here reinstated in its original specific rank. The original description of Rudgea hostmanniana is based on several syntypes: Hostmann 548 from Surinam, Rob. Schomburgk 12 and Rich. Schomburgk 2 from Guyana (or more likely Roraima state, Brazil), and Lockhart s.n. from Trinidad, the latter belonging to subsp. freemannii. The second and third syntype actually represent the same collection (van Dam 2002: 100) since the Schomburgk brothers travelled together and often used separate numbers for the same gathering. Steyermark (1967: 413) cited Hostmann 548 as type, which is to be treated as a first-step lectotypification (Art. 9.10 of the Code) because he did not indicate the herbarium of deposit. The specimen of this collection in K, which is presumably the one studied by Bentham, is here selected as the second-step lectotype. Müller Argoviensis (1881: 205), in the original description of Rudgea intercedens, did not explicitly cite a type, and he only indicated “Habitat in Brasilia (ex hb. Juss., in hb. Rich., nunc in hb. Franq.)”. A specimen from the Richard herbarium in P, which is annotated R. intercedens probably in Müller’s hand, and bears the indication “Brasiliae – herb. Juss.”, is here selected as lectotype. This specimen matches R. hostmanniana in every detail; the synonymy of the two species, which Zappi (2003: 585) already noted to be strongly similar, was recognized by Bruniera (2015) in her thesis, and is here formally established. The fruits of Rudgea hostmanniana are usually red or orange; two specimen labels report that they turn black at maturity (Davidse 4110; Tostain et al. 1884) but this indication remains to be confirmed in the field.As far as is known, in all other species of the complex the mature fruits are red. A collection from Colombia, M.B. Monsalve 741 (GB), cited as Rudgea hostmanniana subsp. hostmanniana by Bruniera (2015: 137) probably represents a new species; its stipules lack the characteristic central keel of R. hostmanniana and its inflorescences have unusually large bracts. No authentic collection of R. hostmanniana from Colombia has been seen by the authors, although the species could be expected in the southeastern region of the country. Additional Specimens Examined: — BRAZIL. Amazonas: Km 1–5 road Boca do Acre – Rio Branco, 24 September 1966 (fl.), G . T. Prance, B. S. Pena, J. F. Ramos & E. R. Videcki Jr 2534 (U). FRENCH GUIANA. NE foot of Montagne des Pères, 5 km S of Kourou, 5°06’N, 52°36’W, 11 March 1994 (fl. & fr.), L. Andersson, C. Gustafsson, C. Persson & J. Rova 1954 (CAY, GB, K); Saut Dalles, Bassin du Sinnamary, 5°27’N, 53°01’W, 24 September 1992 (fl.), B. Bordenave 236 (CAY, P, U); Saut l’Autel, Bassin du Sinnamary, 4°42’N 52°58’W, 25 October 1992 (fl. buds), B. Bordenave 391 (CAY, P); ibid., 4°45’N, 53°06’W, 10 March 1994 (fl. & fr.), B. Bordenave 806 (CAY, P, U); Malmanoury, 5 February 1995 (st.), L. Cadamuro & F. Solacroup 242 (CAY); Fusées Sondes, 20 February 1995 (fr.), L. Cadamuro & F. Solacroup 292 (CAY); Diamant, 22 February 1995 (st.), L. Cadamuro & F. Solacroup 340 (CAY); Kikiwi, 2 March 1995 (st.), L. Cadamuro & F. Solacroup 349 (CAY) & 358 (CAY); Crique Canceler, 5°26’N, 53°02’W, 8 December 1996 (fl.), G. Cremers, F. Crozier & M. Hoff 14474 (P, U); Rivière de Kourou, March 1875 (fl. & fr.), J. Crevaux s.n. (P); village Boni de Assici, bassin du Maroni, 3°49’N, 54°12’W, 17 March 1989 (fl. & fr.), M. Fleury 773 (CAY); Piste de Saint-Elie, 20 February 1985 (fr.), P.-M. Forget 276 (CAY); Lieu-dit Maya, route de la Carapa, 4°56’59”N, 52°26’30”W, 1 March 2007 (fr.), S. Gonzalez 1095 (CAY); rive du Grand Inini vers Bicade, 21 August 1970 (fr.), J.-J. de Granville C-31 (CAY, P, U); îlets du Saut Emérillon sur le Grand Inini, 27 August 1970 (fr.), J.-J. de Granville C-107 (CAY, P); bordure du Grand Inini, à Dégrad Nicole, 4 September 1970 (fr.), J.-J. de Granville B-3657 (CAY, P); Rivière Petite Ouaqui, entre Saut Macaque et Saut Baille-Nom, 12 July 1973 (fl.), J.-J. de Granville B-4954 (CAY, P, U); Rivière Grande Ouaqui, à 8 km de son confluent avec la Petite Ouaqui, 13 July 1973 (st.), J.-J. de Granville 1814 (CAY, P); Rivière Petite Ouaqui, “Saut Baille Nom”, 14 July 1973 (st.), J.-J. de Granville 1829 (CAY, P, U); Fleuve Mana, Saut Fracas, 23 July 1981 (fl. buds), J.-J. de Granville 4640 (CAY, P); Rivière Mana, Gros Saut, 25 July 1981 (fr.), J.-J. de Granville 4654 (CAY, P, U); Rivière Grand Inini en aval et en amont de Dégrad Fourmi, 13 September 1985 (fr.), J.-J. de Granville 8180 (CAY, P, U); Site Ariane 4 - Crique Karouabo, C. S. G., 11 May 2007 (st.), J.-J. de Granville 17387 (CAY); Crique Arouany, 22 August 1962 (fl. & fr.), F. Hallé 632 (CAY, K, P); Saut Takari-Tanté, Bassin du Sinnamary, 4°37’N, 52°56’W, 16 November 1989 (fl.), M. Hoff 5864 (CAY, U); Saut Aïmara, Bassin du Sinnamary, 16 January 1992 (fl.), M. Hoff 7576 (CAY); Rivière Iracoubo, entre roche Hirondelle et Carbet Gendarmerie, 28 December 2010 (fl.), O. Lachenaud 981 (BR, CAY, MO); Kourou, Campus du CIRAD, 30 December 2010 (st.), O. Lachenaud 1011 (BR, CAY); “ Cayenne ”, no date (fl.), J.B. Leblond s.n. (G); without locality, 1859, F.M.R. Leprieur s.n. (G); Haute Mana, en amont de Gros Saut, 29 August 1981 (fr.), C. Moretti 1259 (CAY, P); entre Saut Vata et Saut Bérard, 22 September 1965 (fr.), R.A.A. Oldeman 1541 (CAY, P, U); Fleuve Sinnamary, ca. 4 km en amont sur la Crique Tigre, 4 August 1967 (fr.), R.A.A. Oldeman B-1153 (CAY, P); Fleuve Sinnamary, ca. 6 km en amont sur la Rivière Courcibo, 8 August 1967 (fr.), R.A.A. Oldeman B-1188 (CAY, P); Rivière Courcibo à ca. 1.5 km en amont su Saut Caouène, 10 August 1967 (fr.), R.A.A. Oldeman B-1202 (CAY, P, U); Fleuve Kourou, ca. 700 m en amont du Saut Léodate, 25 September 1967 (fr.), R.A.A. Oldeman B-1388 (CAY, P, U); Fleuve Sinnamary, Saut Bois Blanc, 22 April 1969 (fl.), R.A.A. Oldeman B-2305 (CAY, P, U); without locality, 1820 (fl.), G.S. Perrottet s.n. (G, P); without locality, July 1824 (fl.), P. A. Poiteau s.n. (K); Kourou, Campus Silvolab, 5°10’N, 52°39’W, 8 April 2001 (fl.), M.- F. Prévost & M. Fournier 4151 (BR, CAY, G, K, P, U); ibid., 16 April 2001 (fr.), M.-F. Prévost & D. Sabatier 4154 (CAY, G, K, MO, P); ad montem Macouriae, s.d. (fl.), L.C.M. Richard s.n. (P); près de Saut Macaque, rives du Grand Ouaqui, 12–14 September 1961 (fr.), R. Schnell 12153 (P); Crique Ouaqui, September 1961 (fr.), Service Forestier (BAFOG) 7769 (CAY, P, U), 7807 (CAY, P, U); Forêt de Maya, route de la Carapa, Macouria, 31 March 2008 (fr.), O. Tostain, T. Deville & V. Pelletier 1884 (CAY). GUYANA. Towakaima Falls, Barama River, 7°18’N 59°59’W, 3 October 1996 (fr.), T . R. van Andel, E. Samuels, N. George & M.A.J.P. Smeets 1692 (U); Mabaruma, Aruku River, 11 March 1945 (fl.), D. B . Fanshawe FD 5129 (K, U); Kaow Island, Essequibo River, 13 July 1943 (fr.), Forest Department 4132 (K); SE Kanuku Mountains, 3°03’N, 59°25’W, 25 June 1989 (fr.), L. J . Gillespie, D. Gopaul & Peterson 1824 (K, U); Surama Lake, 4 km NE of Surama Village, 3 May 1992 (fl.), B . Hoffman, D. Allicock & T. Allicock 1567 (CAY); Kanuku Mts, Moco Moco R., 3°18’N, 59°39’W, 19 July 1995 (fr.), M. J . Jansen-Jacobs, C. Simmons, A. Jacobs-Brouwer, V. James & R. Andrew 4574 (CAY, K, P, U); Upper Essequibo Region, Rewa River, Spider Mountains, 3°08’N, 58°32’W, 17 September 1999 (fl.), M. J . Jansen-Jacobs, B. J. H. ter Welle, P.P. Haripersaud, O. Muller & M. van der Zee 5955 (U); ibid., 20 September 1999 (fl.), M. J . Jansen-Jacobs, B. J. H. ter Welle, P.P. Haripersaud, O. Muller & M. van der Zee 6015 (CAY, K, P, U); Mazaruni River, September 1880 (fl.), G. S . Jenman 765 (K); Essequibo River, October 1881 (fl.), G. S . Jenman 1193 (K); Bartica, November 1888 (fl.), G. S . Jenman 4675 (K, U); Mazaruni River, August 1889 (fr.), G. S . Jenman 5440 (K); no locality, 1841 (fl.), Rob. Schomburgk 12 (BM, K, P); Isorova Hill, June 1912 (fr.), F. A . Stockdale 247 (K); Cuyuni River, Crab Fall, 30 April 1933 (fl.), T. G . Tutin 16 (BM, K). SURINAME. Without locality, 1841 (fl.), Berthoud-Coulon 197 (BM), 198 (BM); in districtu Surinamensi Para, February–April 1844 (fl.), A . Kappler 1455 (P); ibid., A . Kappler 1485 (G, P, U); Coppename River, near Raleigh Falls, 13 September 1933 (fr.), J . Lanjouw 814 (INPA, K, U); along Kort en Duur Creek, tributary of Perica River, 29 November 1953 (fl.), J . C. Lindeman 5117 (BR, U); Saramacca River Headwaters, Jacob Kondre, 16 June 1944 (fr.), B . Maguire 23833 (BR, K, P, U); Paramaribo, 23 May 1916 (fr.), J . S. Samuels 317 (K, L); in sylvis Paraensis prope Onoribo, 8 March 1838 (fl.), F . Splitgerber 665 (L); fluv. Corantijn, Kaboerie - Winana, 27 October 1916 (fr.), G . Stahel & J. W. Gonggrijp 2985 (U); Corantyne River, near Mac Claren, 23 January 1963 (fl.), J . G. Wessels Boer 555 (P, U); Paramaribo, 1851 (fr.), H . R. Wullschlaegel 242 (BR); “ Para ”, no date (fl.), H . R. Wullschlaegel 995 (BR). VENEZUELA. Miranda: 3 km SW of Araguita, along road between Caucagua and Altigracia de Orituco, 17 November 1973 (fr.), G . Davidse 4110 (L); Catalina, May 1896 (fl. buds), H . H. Rusby & R. W. Squires 209 (BM); ibid., May 1896 (fl.), H . H. Rusby & R. W. Squires 444 (K)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 231-234, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870, {"references":["Bentham, G. (1850) Plantae Regnellianae (Continuatio) Rubiaceae. Linnaea 23: 455 - 466.","Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B.,, Fidalgo, O. & Fidalgo, M. E. P. K. (eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436.","Muller Argoviensis, J. (1881) Flora Brasiliensis. Vol. VI Pars V. R. Fleischer, Leipzig [\" Lipsiae \"], 486 pp.","Williams, R. O. & Cheesman, E. E. (1928) Flora of Trinidad and Tobago. Vol. II. Part 1. Rubiales. Port of Spain, Government Printing Office, 48 pp.","Zappi, D. & Steyermark, J. A. (2004) Rudgea. In: Steyermark, J. A., Berry, P. E., Yatskievytch, K. & Holst, B. K. (eds.) Flora of the Venezuelan Guayana, vol. 8, Poaceae - Rubiaceae. Missouri Botanical Garden Press, St Louis, pp. 805 - 816.","Bruniera, C. P. (2015) Sistematica e taxonomia de Rudgea Salisb. (Palicoureeae, Rubiaceae) Unpublished Ph. D. Thesis, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, 283 pp. [http: // www. teses. usp. br / teses / disponiveis / 41 / 41132 / tde- 28072015 - 145432 / en. php]","Steyermark, J. A. (1974) Flora de Venezuela. Vol. IX, segunda parte. Rubiaceae. Instituto Botanico, Caracas, 498 pp.","Acevedo-Rodriguez, P. & Strong, M. T. (2012) Catalogue of Seed Plants of the West Indies. Smithsonian Contributions to Botany 98: 1 - 1192. https: // doi. org / 10.5479 / si. 0081024 X. 98.1","van Dam, J. A. C. (2002) The Guyanan Plant Collections of Robert and Richard Schomburgk. In: Flora of the Guianas, Supplementary Series 3. Royal Botanic Garden, Kew, 211 pp.","Zappi, D. (2003) Revision of Rudgea (Rubiaceae) in Southeastern and Southern Brazil. Kew Bulletin 58: 513 - 596. https: // doi. org / 10.2307 / 4111145"]}

6. Rudgea maypurensis Standley (1930b: 72). – Rudgea hostmanniana subsp. maypurensis (Standley) Zappi (in Zappi & Steyermark 2004: 808). (Figs. 1E, 2E) Type: — VENEZUELA. Amazonas: Maypures, June 1854 (fr.), R. Spruce 3615 (holotype, K! [K000447196]; isotype, P! [P04008962]; fragment, G! [G00436708]). R. corocoroensis Steyermark (1988: 349), syn. nov. Type: — VENEZUELA. Amazonas, Dpto. Atures, 5–8 km NW of settlement of Yutajé, 3 km W of Rio Coro-Coro, W of Serranía de Yutaje, 5°40’N, 66°09’W, 700–1000 m, 10 March 1987 (fl.), R. Liesner & B. Holst 21827 (holotype MO! [MO-2049858]; isotypes, F! [N °2030267], NY! [NY00133209], PORT! [PORT34149], U! [U0006286], US! [US 00153756]). Much-branched shrub 1–4 m tall; twigs densely patent-puberulous or more rarely glabrous, 1.5–3 mm thick, soon covered with a pale straw-coloured bark, becoming greyish on older stems. Stipules 3–10 x 3–5.5 mm, densely patentpuberulous to glabrous, marcescent and soon corky, consisting of a basal sheath 1–3 mm long (usually early split) bearing on each side of the node 4–6 erect linear terminal appendages 2–7 mm long, and 4–10 recurved dorsal appendages 1-3 mm long, these often connate at base into a short keel. Leaves opposite; petioles 0.1–0.7 cm long, patent-puberulous to glabrous; blades elliptic, 2.8–12 × 1–8.8 cm, slightly cordate to rounded at base, obtuse to hardly acuminate at apex, very thick, entirely glabrous, drying yellowish-green (or the young leaves blackish-green); midrib flat or concave above; secondary veins 6–11 on each side of midrib, rather strongly ascending, hardly prominent; tertiary venation not or hardly distinct; domatia absent. Inflorescences terminal, in rather condensed pyramidal panicles, 1.8–8.8 cm long, erect, shortly spreading-puberulous or more rarely glabrous; peduncle terete, 1–6.5 cm long; branched portion 0.8–3.5 × 1–3 cm; secondary branches (2–)3–4 per node, 0.4–2.7 cm long; bracts 1.5–5 × 0.7–2 mm, triangular to lanceolate, entire or often dentate at base, acute at apex, glabrous outside, pubescent inside. Flowers sessile, 5- merous, heterostylous. Hypanthium obconical, 0.7 mm long, glabrous. Calyx tube extremely reduced, lobes triangular, 0.5–1.8 × 0.5–0.7 mm, acute or obtuse at apex, densely ciliate. Corolla white (the lobes sometimes pale yellow), hypocrateriform; tube narrowly funnel-shaped, 3–4 mm long, 1–1.2 mm wide at base, 1.7–2 mm wide at mouth, glabrous outside, villose in the upper part inside; lobes narrowly triangular, 2.5 x 1.3 mm, glabrous to puberulous at apex outside, papillose inside, not corniculate dorsally. Stamens included, with subsessile anthers in long-styled flowers, or well-exserted, with filaments 3 mm long, in short-styled flowers; anthers 1.5 x 0.3 mm. Disk cylindrical to slightly conical, 0.5–0.8 mm long, glabrous. Style exserted, 6 mm long in long-styled flowers, or included, ca. 3.5 mm long in short-styled flowers, glabrous or densely pubescent in the distal half; lobes 0.5–1 mm long, stigmatic surface papillose. Fruits obovoid to subglobose, 4.5–6 × 4–5.5 mm when dry, green when immature, orange to red when mature, glabrous, sessile, crowned with persistent calyx 1–1.5 mm in diameter. Pyrenes plano-convex, hemi-obovoid, 5–5.5 × 4.2–5 mm, dorsal side with 2–4 prominent to very weak longitudinal ridges, slightly verrucose, ventral side ± smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —Restricted to southeastern Venezuela (Amazonas state) and adjacent northwestern Brazil (Amazonas state), and probably eastern Colombia (Fig. 6); occurs in dry forests bordering granitic rocks (“lajas”) where it is locally abundant, at 85–200 m in elevation. Phenology: —Specimens with flowers were collected in March–April, with immature fruits in April–May, and with mature fruits in June–August and once in November. Notes: —This taxon, treated as a subspecies of Rudgea hostmanniana by Zappi & Steyermark (2004), is distinct enough to retain its original species status. It differs from R. hostmanniana by its stipules (compare Fig. 1D & 1E), its corolla lobes that are not corniculate at apex, and its pyrenes that are dorsally verrucose (Table 1). The leaves are also more coriaceous and shiny than in R. hostmanniana, with an often slightly cordate base and a usually shorter petiole, and the fruits are generally smaller. An illustration of this taxon (as R. hostmanniana subsp. maypurensis) has been published by Zappi & Steyermark (2004: fig. 618). The original description of Rudgea maypurensis was published in Standley (1930b: 72), not in Standley (1931: 434) as incorrectly cited by Steyermark (1967: 411). The flowers are here described for the first time; they are mentioned neither in the protologue, nor in any of the subsequent descriptions (Standley 1930b: 72; Steyermark 1967: 411, 1974: 1070-1071; Zappi & Steyermark 2004: 808–809). The type specimen of Rudgea corocoroensis was only seen in photograph, which is sufficient to establish that it agrees with R. maypurensis in all essential characters, particularly the diagnostic stipules, although the petioles are longer than usual for the species; the two names are therefore synonymized here. Steyermark (1988: 350) described the stipules of R. corocoroensis as having “5-7 rigid aculeae arising at or just below the sheath summit”, apparently omitting the dorsal appendages that are clearly present, and the calyx tube as 2 mm long, which is instead much smaller. He considered R. corororoensis as related to Rudgea morichensis Steyermark (1967: 424) but the latter is quite different, e.g. in its deeply cupular calyx and prominent tertiary leaf veins, and does not seem to belong to the R. hostmanniana complex. The types of both Rudgea maypurensis and R. corocoroensis have glabrous twigs, petioles and inflorescences; in all other collections seen these parts are shortly patent-puberulous. A collection from Brazil, cited below, is a new record for the country; although it is sterile and was only seen on photograph, its identification is without any doubt. A specimen from Colombia, Cuatrecasas 4052, referred to this species with some doubt by Steyermark (1967: 411), has not been seen for this revision. Additional Specimens Examined: — BRAZIL. Amazonas: vicinity of Pico Rondon, Perimetral Norte Highway km 211, 1°32’N 62°48’W, 2 February 1984 (st.), G . T. Prance, I. L. do Amaral, J. J. Pipoly, A. S. Tavares, C.D.A. da Mora & A. Cress 28731 (NY). VENEZUELA. Amazonas: 8 km S of Puerto Ayacucho, estación de piscicultura, 5°36’N, 67°37’W, 13–15 April 1978 (fl.), G . Davidse & O. Huber 14888 (K); Raudal d’Atures, 1 August 1887 (fr.), Gaillard 36 (P); Estación de Piscicultura de Puerto Ayacucho, 5°37’N, 67°37’W, 15 April 1977 (st.), O . Huber 617 (K); ibid., 15 April 1977 (fl.), O . Huber 617a (K); ibid., 15 April 1977 (imm. fr.), O . Huber 617b (K); 1–2 km E of Hotel Amazonas, Puerto Ayacucho, 8 November 1953 (fr.), B . Maguire, J. J. Wurdack & G. S. Bunting 36034 (K); 6 km N of Puerto Ayacucho on road to El Burro, 26 April 1984 (fl.), T . Plowman 13733 (K); ibid., T . Plowman 13742 (K); Cerro Piapoco, cerca del km 12–13 de la carretera Puerto Ayacucho – Sanariapo, 31 July 1967 (fr.), L . Ruiz-Terán 4444 (BR, K); Puerto Ayacucho, 18 May 1940 (fr.), L . Williams 12972 (K)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 235-236, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870, {"references":["Standley, P. C. (1930 b) Studies of American Plants - III. Publications of the Field Museum of Natural History, Botanical Series 8 (1): 1 - 73. https: // doi. org / 10.5962 / bhl. title. 2342","Zappi, D. & Steyermark, J. A. (2004) Rudgea. In: Steyermark, J. A., Berry, P. E., Yatskievytch, K. & Holst, B. K. (eds.) Flora of the Venezuelan Guayana, vol. 8, Poaceae - Rubiaceae. Missouri Botanical Garden Press, St Louis, pp. 805 - 816.","Steyermark, J. A. (1988) Flora of the Venezuelan Guayana - IV. Annals of the Missouri Botanical Garden 75: 311 - 351. https: // doi. org / 10.2307 / 2399477","Standley, P. C. (1931) The Rubiaceae of Venezuela. Publications of the Field Museum of Natural History, Botanical Series 7 (4): 343 - 485. https: // doi. org / 10.5962 / bhl. title. 2347","Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B.,, Fidalgo, O. & Fidalgo, M. E. P. K. (eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436.","Steyermark, J. A. (1974) Flora de Venezuela. Vol. IX, segunda parte. Rubiaceae. Instituto Botanico, Caracas, 498 pp."]}

1. Rudgea billietiae O. Lachenaud, sp. nov. Fig. 3 & 4 Foliis glabris crassis venulis inconspicuis, stipulis basi connatis et dorso appendiculatis, fructibusque statu immaturo brunneis deinde rubris Rudgeae pungenti, R. hostmannianae, R. cornigerae et R. tanaosepalae affinis. A Rudgeae pungenti differt bracteis multo minoribus, 3–5 mm longis, inflorescentiis manifeste ramosis et corollae tubo multo breviore, 6 mm longo (vs. 13 mm longo); ab alteris speciebus stipulis longiore tubulosis in vaginam 7.5–12 mm longis (vs. 2–5 mm longis) connatis, praeterea inflorescentiis glabris (vs. puberulis) distinguitur. Type: — FRENCH GUIANA. Route de Kaw, pk 33, sentier vers les grottes, 4 December 2000 (fl.), F. Billiet & B. Jadin 7456 (holotype, BR! [BR000000907382]; isotypes, CAY! [CAY014905, CAY014906], MO n.v., NY n.v.). Much-branched shrub 1.75–3 m tall; trunk pale grey; branches ascending; twigs glabrous, 1–2 mm thick, soon covered with a pale straw-coloured bark. Stipules 9–15 × 2.5–4.5 mm, glabrous, marcescent and soon corky, consisting of a narrow tubular sheath 7.5–12 mm long (usually split at flower-bearing nodes) bearing ca. 12 terminal linear appendages 1.5–4 mm long, and 6–8 early caducous dorsal appendages 1 mm long, forming a very short decurrent keel inserted in the upper half of the stipule. Leaves opposite; petioles 0.2–1 cm long, glabrous; blades elliptic, 4.7–13.5 × 1.3–5.5 cm, decurrent on petiole at base, gradually acuminate at apex, very thick and easily cracking in the fresh state, entirely glabrous, drying greyish-green to yellowish; midrib concave above; secondary veins 6-11 on each side of midrib, strongly ascending, forming an angle of ca. 45° with the midrib; tertiary veins invisible in fresh material, sometimes faintly prominent in dry material; domatia absent. Inflorescences terminal in rather contracted panicles, 1–4 cm long, erect, white in flower and green in fruit, glabrous; peduncle terete, 0.5–1.6 cm long; branched part 0.5–2.4 × 1.3–3.5 cm; secondary branches 3–4 per node, 0.2–1 cm long; bracts linear or narrowly lanceolate, 3–5 × 0.3–1.5 mm, entire or dentate in lower half, shortly ciliate in upper part. Flowers sessile, 5-merous. Hypanthium obconical, 0.5–0.8 mm long, glabrous. Calyx tube extremely reduced, lobes linear to narrowly triangular, 1.5–2 × 0.3–0.6 mm, very acute at apex, glabrous or ciliate. Corolla white, hypocrateriform; tube funnel-shaped, 6 mm long, 0.5–1 mm wide at base, 2–3 mm wide at mouth, glabrous outside, pubescent inside at the distal portion below the mouth; lobes triangular, 4–5 × 2 mm, glabrous on both sides, with broad, rounded dorsal cornicula 0.7–1 mm long. Stamens white, exserted; filaments exserted beyond the corolla mouth by 3 mm; anthers 1.2 × 0.3 mm. Disk shortly cylindrical, 0.7 mm long, glabrous. Style included, 5 mm long, glabrous; lobes 0.8 mm long, stigmatic surface papillose. Fruits ellipsoid, 6–7 × 5–6 mm when dry, dark red-brown and hard when immature, bright cherry red and soft when mature, glabrous, sessile or with very short pedicel ≤ 0.5 mm long, crowned with persistent calyx 1.5 mm in diameter. Pyrenes plano-convex, hemiellipsoid, 6.5 × 5 mm, dorsal side with 3 hardly distinct longitudinal ridges, smooth between the ridges, ventral side smooth. Seeds with a deep T-shaped ventral furrow. Distribution and ecology: —This species is endemic to the Kaw mountain in northeastern French Guiana (Fig. 5) where it occurs exclusively in low, open canopy forest on lateritic crust, 150 – 300 m in elevation. It is locally abundant in this habitat, which only covers small areas on the ridge of the mountain. Phenology: —Flowers have been collected in December (beginning of first rainy season) which is probably the main flowering period; a single inflorescence was also seen in June, among a mostly fruiting population (Lachenaud 1878), which is probably a case of flowering asynchrony. Fruits have been collected in November (once, immature) and were seen in abundance from February to late June at least (OL pers. obs.). They apparently take about six months to mature. They are already full-sized in February, but still hard and brown, and remain so for a long time; they start to turn soft and red in late June (OL pers. obs.). Eponymy: —This species is named after the Belgian botanist Frieda Billiet, collector of the type. Together with her husband Bernard Jadin (1948–2012), she made important plant collections in French Guiana between 1981 and 2009, several of which have been described as new species, e.g., Philodendron billietiae Croat (1995: 24), Oryctanthus guianensis Kuijt (2011: 465) and Rudgea jadinii O. Lachenaud (Lachenaud et al. 2022: 168). Conservation status assessment: —Endangered [EN B1 ab(iii) & 2ab(iii)]. Rudgea billietiae has a very restricted range, being endemic to the ridge of the Kaw Mountain in French Guiana, where it grows in low stunted rainforest on lateritic crust. Its extent of occurrence (EOO) is calculated to be 68 km ², which falls within the limit for Critically Endangered under criterion B1, while its area of occupancy (AOO) is estimated to be 24 km ², within the limit for Endangered status under criterion B2 (the actual AOO is however certainly Notes: —Specimens of Rudgea billietiae have for a long time been confused in herbaria with either R. hostmanniana or R. pungens. In the vegetative state R. billietiae is very similar to the latter, which also has stipules connate into a long narrow tube (a character not always easy to see on herbarium specimens, because the stipules of the flower-bearing nodes tend to split early) and can only be separated by the position of the dorsal appendages of the stipules, which are inserted in their upper half in R. billietiae and near their base in R. pungens. The two species differ markedly in their inflorescences, especially in the development of their bracts (much larger in R. pungens) and in the size of the corolla tube, which is about twice longer in R. pungens. They also seem to have different ranges, R. pungens being apparently absent from the Kaw Mountain. In inflorescence structure and corolla length R. billietiae resembles R. hostmanniana, R. cornigera and R. tanaosepala, but these have much shorter stipular sheaths and puberulous inflorescence axes (glabrous in R. billietiae). Differences between all these species are summarised in Table 1. It is not known whether the flowers of R. billietiae are heterostylous: only two collections with open flowers have been seen (the type and Martin s.n.) and both are of the brevistylous form. Three specimens collected in the late XVIIIth or early XIXth centuries, Leblond s.n., Martin s.n. and Richard s.n., have no precise locality; the two former are labelled “Cayenne”, but this indication was often used for the whole of French Guiana at that time. They may well have come from the Kaw Mountain, which is only about 50 km from Cayenne and was already accessible at that time. The Richard collection is a mixture, including two branches of R. pungens (one flowering, one fruiting) and one of R. billietiae (fruiting), collected at different times of the year and probably in different places. Additional Specimens Examined (paratypes): — FRENCH GUIANA. Montagne de Kaw, E end ca. 10 km from end of road, 4°32’N 52°07’W, 10 March 1994 (fr.), L . Andersson, C. Gustafsson, C. Persson & J. Rova 1949 (CAY); Montagne de Kaw, 13 May 1985 (fr.), C . Feuillet 2244 (CAY, P); Montagne de Kaw, forêt dense sur versant nord, à proximité de la route, 5 km E de Camp Caïman, 14 June 1979 (st.), J.- J . de Granville 2978 (CAY, P); Montagne de Kaw, extrémité est, versant sud, 3 November 1985 (imm. fr.), J.- J . de Granville 8244 (CAY, P, U); Montagne de Kaw, sentier des grottes, 23 February 2014 (imm. fr.), O . Lachenaud 1643 (BR, CAY, MO, P); same locality, 4°33’15”N 52°10’18”W, 28 June 2014 (fl. buds & fr.), O . Lachenaud 1878 (BR, CAY, MO, P); Montagne de Kaw, Amazon Lodge, 4°33’35.1”N 52°12’20.7”W, 23 August 2017 (st.), O . Lachenaud 2639 (BR, CAY, MO); Montagne de Kaw, au pk 30.5, à gauche de la route, 3 January 2019 (st.), O . Lachenaud 2689 (BR, CAY); Route de Fourgassié, peu après le croisement de la route de Kaw, 3 January 2019 (fallen fl.), O . Lachenaud 2695 (BR, CAY); “Cayenne”, no date [ca. 1800], J.B. Leblond s.n. (G); “Cayenne”, no date [ca. 1800] (fl.), J . Martin s.n. (BM [3 sheets], K [2 sheets]); no locality or date (fr.), L. C. M . Richard s.n. (P [P04008549], mixed with R. pungens; see notes above)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, The Rudgea hostmanniana complex (Rubiaceae) in the Guiana Shield region, pp. 219-242 in Phytotaxa 561 (3) on pages 223-226, DOI: 10.11646/phytotaxa.561.3.1, http://zenodo.org/record/7069870, {"references":["Croat, T. B. (1995) Two new species of Araceae for the Guianas. Novon 5: 24 - 29. https: // doi. org / 10.2307 / 3391825","Kuijt, J. (2011) Two new species of Oryctanthus (Loranthaceae) from Colombia and French Guiana. Novon 21 (4): 463 - 467. https: // doi. org / 10.3417 / 2010106","Lachenaud, O., Bruniera, C. P. & Zappi, D. C. (2022) Six new and a little-known species of Rudgea (Rubiaceae - Palicoureeae) from the Guianas. Phytotaxa 531 (3): 154 - 174. https: // doi. org / 10.11646 / phytotaxa. 531.3.1"]}

Rudgea graniticola O. Lachenaud, sp. nov. (Fig. 5) Ramis minute puberulis, foliis coriaceis, fructibus aurantiacis ellipsoideis, pyrenis dorso costatis, corolla tubo breve (4.5-5 mm) et intus villoso, stipulisque parvis (3-4.5 mm) dorso carina appendiculata munitis Rudgeae crassilobae (Bentham 1841: 233) B.L. Robinson (1910: 408) affinis, sed differt foliis glaberrimis (vs. in axillis nervorum pubescentibus) costa mediana supra prominente (vs. impressa) venulis laxiore reticulatis, corollaque lobis intus glabris (vs. villosis). Type: ��� FRENCH GUIANA. Montagne des Nouragues, 4��03���N, 53��42���W, October 1989 (fl.), D.Larpin 722 (holotype,CAY [CAY078936]; isotype, P [P04008509]). Shrub, much-branched, or small tree, to 10 m tall, to 15 cm at dbh; main trunk often leaning or branched from base, the branches erect; bark grey, rugose; terminal branchlets 1 ��� 1.5 mm thick, minutely puberulous (appearing glabrous to the naked eye), soon covered with a grey bark. Stipules minutely puberulous, 3 ��� 4.5 �� 0.8 ��� 3 mm, consisting of a persistent truncate to round sheath 1 ��� 2 mm long, bearing on each side a dorsal keel with 3 ��� 5 terminal, corky and soon caducous appendages 1 ��� 2 mm long. Leaves opposite, petiolate; petioles 0.3 ��� 0.8 cm long, minutely puberulous; blades elliptic or slightly obovate, 2.7 ��� 10 �� 1 ��� 5 cm, decurrent on petiole at base, shortly acuminate at apex, coriaceous with recurved margin, entirely glabrous, dark green above and paler green below in life, variously drying green, yellowish or brown; midrib convex on both sides; secondary veins 5 ��� 7 on each side of midrib, moderately ascending, forming conspicuous loops 0.5 ��� 3 mm from the margin; tertiary veins conspicuously and rather laxly reticulate; domatia absent. Inflorescences terminal, white, trichotomously paniculate or rarely reduced to a glomerule, 0.8 ��� 6.8 cm long, minutely puberulous; peduncle terete, 0.5 ��� 3 cm long; flowering portion 0.3 ��� 3.8 �� 0.5 ��� 3.5 cm; main ramifications, if present, 2 per node, 0.1 ��� 1.2 cm long; bracts shortly triangular, up to 1.5 mm long, acute, corky, whitish. Flowers sessile, 4-merous, heterostylous. Hypanthium tronco-conical, 0.5 mm long, puberulous. Calyx cupuliform, 0.5 ��� 1 mm long, minutely puberulous, cleft at least to the middle (often nearly to the base) into elliptic to triangular lobes, these acute or obtuse at apex. Corolla white; tube narrowly cylindrical, 4.5 ��� 5 �� 1 ��� 1.5 mm, glabrous outside, pubescent at the distal portion inside; lobes narrowly triangular, 2.5 ��� 3.5 �� 1 ��� 1.3 mm, apex with dorsal cornicula Stamens half-exserted in long-styled flowers, or fully exserted, with filaments exserted 2 mm beyond corolla mouth in short-styled flowers; anthers elliptic, 1 ��� 1.3 �� 0.3 mm, dorsifixed. Disk hemispherical, entire, 0.5 mm, glabrous. Style exserted, 7 mm long in long-styled flowers, or included, 4 mm long in short-styled flowers, papillose towards apex; branches c. 0.7 mm long. Fruits ellipsoid, 7 ��� 9 �� 5 ��� 6 mm when dry, green when immature, orange when mature, glabrous, sessile, the calyx not enlarging. Pyrenes ellipsoid, 6.5 ��� 7 �� 4.5 mm, with 3 ��� 4 strong dorsal ridges; seeds with a deep T-shaped ventral furrow. Distribution and ecology: ��� This species has a scattered distribution in northern and central French Guiana (Chutes Voltaire, Nouragues, la Trinit��, Massif des Emerillons) and northern Suriname (Bakhuis Mts) (Fig. 2.) It grows exclusively in low forest on granitic inselbergs, at 150 ��� 410 m elevation, and is locally very common in this habitat. Phenology: ��� Flowering specimens were collected in March and October; specimens with immature fruits were collected in January, and with mature ones from February to April. Conservation status assessment: ��� Endangered [B1ab(iii)+2ab(iii)]. Rudgea graniticola occurs on inselbergs in French Guiana and Suriname, and is known from 17 collections representing five occurrences. Its extent of occurrence (EOO) is calculated to be 44,550 km ��, exceeding the limit for Vulnerable status under subcriterion B1, and its area of occupancy is estimated to be 24 km ��, within the limit for Endangered status under subcriterion B2. Of the four occurrences in French Guiana, three occur in protected areas (Parc Amazonien de Guyane, R��serve Naturelle des Nouragues, and R��serve Naturelle de la Trinit��); the fourth one (near Chutes Voltaire) may be impacted by touristic activities but the current level of threat is rather low. The only occurrence in Suriname occurs in a mining concession, which leads us to expect a decline in habitat extent and quality. The five occurrences represent five locations in the sense of IUCN, and the species qualifies for Endangered status according to the conditions B1ab(iii)+B2ab(iii). Considering however that its range and habitat are not well prospected, the species may be more widespread than records suggest, in which case this assessment will have to be revised. Notes: ��� This species has previously been confused with the more widespread and rather similar Rudgea crassiloba. Both have relatively small and coriaceous leaves, minutely puberulous twigs, very short stipules with a dorsal keel bearing terminal appendages, usually paniculate inflorescences (sometimes glomerulate in R. graniticola), a short corolla tube, orange to red ellipsoid fruits, and dorsally costate pyrenes. Both are also commonly associated with granitic rocks (hence the specific name of R. graniticola) although R. crassiloba may also be found on other substrates. They differ by the characters given in the diagnosis. Rudgea graniticola also resembles R. ayangannensis (see above) in vegetative and inflorescence characters. Differences between the three species are summarised in Table 3. The distribution of R. graniticola is separate from both R. ayangannensis, which is endemic to Guyana, and R. crassiloba, which is mostly found in the Amazon/Orinoco basin and only reaches the southern part of Suriname and French Guiana. According to Bruniera (2015), Rudgea crassiloba belongs to the informal ��� lanceifolia clade���, being closely related to R. reticulata Bentham (1850: 458). In view of its morphology, R. graniticola probably also belongs to this clade. The inflorescences are very variable in this species. Collections from the Nouragues inselberg (including the type specimen) have lax and many-flowered panicles, while those from Chutes Voltaire and Massif des Emerillons have few-flowered glomerules. Specimens from Bakhuis Mountains and La Trinit�� are intermediate, and since no other differences have been found, we recognize only one variable taxon. Characters in bold are diagnostic for the species The oldest collection of this species (Collector unknown in A.P. de Candolle & C.L. L���H��ritier s.n.) is labelled ���Cayenne���, which at that time often referred to the whole of French Guiana. In view of its habitat, the species is unlikely to occur in the close vicinity of Cayenne. Neither Candolle nor L���H��ritier ever visited French Guiana, so they must have obtained this material from another, unknown collector. Additional specimens examined (paratypes): ��� FRENCH GUIANA. ���Cayenne��� (see notes), no date [c.1800] (fr.), Collector unknown in A.-P. de Candolle & C.L. L���H��ritier s.n. (G); Massif des Emerillons, zone nord, savane roche au sommet d���une colline sur la gauche de la Haute Approuague, 29 September 1980 (fallen fl.), J.J. de Granville 3949 (CAY, P); Montagnes de la Trinit��, sommet nord, 12 January 1984 (imm. fr.), J.J. de Granville, C.C. Berg, M.J. Jansen-Jacobs & J. van Setten 5909 (BR, CAY, K, P, U) & 5938 (BR, CAY, K, P, U); Montagnes de la Trinit��, inselberg Nord-Ouest, 16 January 1984 (fallen fl.), J.J. de Granville, C.C. Berg, M.J. Jansen-Jacobs & J. van Setten 6016 (CAY, U); Station des Nouragues, 4��03���N, 52��42���W, 22 February 1991 (fr.), J.J. de Granville 11164 (CAY, P, U); Montagne des Nouragues, 4��03���N, 52��42���W, April 1989 (fr.), D. Larpin 496 (CAY, P); Inselberg des Chutes Voltaire, 20 December 2015 (fallen fl.), O. Lachenaud 2166 (BR, CAY); without locality, 1842 (fr.), E. M��linon s.n. (P); Nouragues Field Station, 4��05.289���N, 52��40.774���W, 16 February 2002 (fr.), S.A. Mori, F. Blanchard & T.A. Lobova 25381 (CAY); R��serve des Nouragues, Station de recherches, 4��03���N, 52��42���W, 14 March 2004 (fl. & fr.), O. Poncy, L. Barrab��, P. Petronelli, J.Y. Serein & J.G. Jourget 1878 (CAY, P); Montagne des Nouragues, 4��03���N, 52��42���W, 17 April 1988 (fr.), C. Sarthou 275 (CAY); Station des Nouragues, 13 March 1996 (fr.), P. Solano K331 (CAY); same locality, 14 March 1996 (fr.), P. Solano K394 (CAY); inselberg Trinit��, 4��35���N, 53��21���W, 9 May 1998 (st.), J.F. Villiers & C. Sarthou 6293 (CAY). SURINAME. Bakhuis Mountains, concession BMS, zone 22, centre, 4��29���45���N, 57��00���59���W, 5 April 2006 (fr.), B. Bordenave, S. Dourga, F. van Troon, I. van Troon & J. James 8365 (CAY)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas, pp. 154-174 in Phytotaxa 531 (3) on pages 163-166, DOI: 10.11646/phytotaxa.531.3.1, http://zenodo.org/record/5886210, {"references":["Bentham, G. (1841) Schomburgk's Guiana Plants In: Hooker, W. J. (Ed.) Journal of Botany 3: 226 - 233.","Robinson, B. L. (1910) XVII. II Spermatophytes, new or reclassified, chiefly Rubiaceae and Gentianaceae. Proceedings of the American Academy of Arts and Sciences 45: 394 - 412. https: // doi. org / 10.2307 / 20022574","Bruniera, C. P. (2015) Sistematica e taxonomia de Rudgea Salisb. (Palicoureeae, Rubiaceae) Unpublished Doctoral Thesis, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, 283 pp. [http: // www. teses. usp. br / teses / disponiveis / 41 / 41132 / tde- 28072015 - 145432 / en. php]","Bentham, G. (1850) Plantae Regnellianae (Continuatio) Rubiaceae. Linnaea 23: 455 - 466.","Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B. et al. (Eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436."]}

Rudgea leucocarpa O. Lachenaud, sp. nov. (Fig. 10) Fructibus albis vel luteis apice truncato calyce remnante in diametro valde accrescente, pyrenis dorso leviter verrucosis, stipulis brevibus (2-4 mm longis) margine fimbriatis dorso exappendiculatis, foliis glaberrimis costa mediana supra prominente, inflorescentiis glomerulatis vel brevissime ramosis, inter congeneribus satis recedit. Type: ��� FRENCH GUIANA: Sommet Tabulaire [Mount Itoup��], versant sud,> 50 km SE de Sa��l, 24 August 1980 (fr.), J.J. de Granville 3587 (holotype, P [P06800583]; isotypes, CAY [CAY079068, CAY079069], NY n.v.). Rudgea sp. B auct. (Boom & Delprete 2002: 646). Shrub 0.2 ��� 2 m tall, much branched; terminal branchlets 1.5 ��� 2 mm thick, glabrous, long remaining green. Stipules consisting of two flat interpetiolar portions, ovate to triangular, 2 ��� 4 �� 2 ��� 4 mm, irregularly laciniate with 7 ��� 10 marginal appendages 0.6 ��� 1.5 mm long and lacking dorsal appendages, glabrous, marcescent and soon corky, surrounding a very fugacious internal sheath 1 ��� 2(��� 3) mm long. Leaves opposite, petiolate; petiole 0.6-1.5 cm long, glabrous; blades elliptic, 8 ��� 18.5 �� 3 ��� 7.3 cm, acute at base, narrowly and usually long acuminate at apex, coriaceous or papyraceous when dry, entirely glabrous, drying olive green with shiny underside, midrib convex on both sides, secondary veins 7 ��� 10 on each side of midrib, weakly to moderately ascending, forming very conspicuous loops 2 ��� 7 mm from the margin; tertiary veins prominent at least when dry, laxly reticulate, forming areolae ca. 3 mm in diameter; domatia absent. Inflorescences terminal, glomerulate or sometimes very shortly branched, few-flowered, erect, glabrous; peduncle terete, 3 ��� 9 mm long; main ramifications (when present) 2 per node, 0 ��� 2 mm long; bracts shortly triangular, 1.5 mm long, glabrous. Flowers sessile, 5-merous, whether distylous unknown. Hypanthium tronco-conical, 1.5 mm long, glabrous. Calyx tube 0.7 mm long, glabrous; lobes triangular, 1 ��� 1.5 mm long, acute at apex, patent. Corolla unknown. Fruits subglobose to broadly ovoid, 13 ��� 20 mm in diam. when fresh, 11���16 �� 11���15 mm when dry, green when immature, white to yellow when mature, glabrous, sessile; calyx scar much enlarging on fruit, 5���9 mm wide. Pyrenes ellipsoid to ovoid, 10.5 ��� 12 �� 8 ��� 9 mm, slightly verrucose, with one weak dorsal ridge; seeds with a deep, �� T���shaped ventral groove. Distribution and ecology: ���This species occurs in northern Brazil (Reserva Biologica Uatum�� in Amazonas state), extreme southeastern Suriname (Tumuc Humac Mts) and French Guiana, where it is found mostly in the central and southern hill ranges (Tumuc Humac Mountains, Montagne Bellevue de l���Inini, Mount Galbao, Mount Itoup��) with one record in the Approuague River basin in the northeast (Fig. 9). It occurs in lowland forests, and mostly in submontane forests on ridges where locally not uncommon, at 350���730 m in elevation. Phenology: ���Immature fruits collected in January and March; mature fruits in August and early September. Conservation status assessment:���Least Concern (LC). Rudgea leucocarpa occurs in French Guiana, Suriname and northern Brazil (state of Amazonas), and is known from 13 collections representing eight occurrences. Its extent of occurrence (EOO) is calculated to be 39,915 km ��, exceeding the limit for Vulnerable status under subcriterion B1, and its area of occupancy is estimated to be 32 km ��, within the limit for Endangered status under subcriterion B2. Four of its occurrences are protected in the Parc Amazonien de Guyane, French Guiana, and in the Reserva Biologica Uatum��, Amazonas state, Brazil. The other occurrences are in areas sparsely populated and difficult to access, where there is no evidence of a particular threat. Consequently, the species is here assessed as LC. Notes: ���This species is quite distinct from all other so far recorded in the Guianas, though in the absence of flowers it is difficult to suggest any affiliation with other species of the genus. The large white to yellow fruits with a conspicuously enlarged calyx scar are especially diagnostic (species of Rudgea occurring in the Guianas have mostly red or orange fruits, or if white, much smaller). In this character, and also in the leaves, pyrenes, seeds and inflorescences, Rudgea leucocarpa resembles Carapichea verrucosa C.M. Taylor (in Taylor & Gereau 2013: 124) from western Amazonia (Peru, Colombia, Brazil), although the latter has even larger fruits (2.5 ��� 3 �� 2 ��� 2.5 cm) with a distinctly warty surface, and its stipules are distinctly longer than broad and apparently not fimbriate. Both species are known from fruiting material only, and have never been studied phylogenetically, so it is unclear whether their resemblance is superficial or indicative of a true relationship. Carapichea verrucosa is unusual in its genus and its position remains uncertain, although the stipules seem to exclude it from Rudgea. In vegetative characters R. leucocarpa also shows a superficial resemblance to R. graciliflora, but the latter has smaller fruits with the calyx not enlarged and a purplishbrown (when immature) or orange-red (when mature) colour, the leaf midrib concave above, and the stipules with a dorsal keel bearing apical appendages, while those of R. leucocarpa are dorsally flat and marginally fimbriate. This species was already recognized as probably new by Boom & Delprete (2002: 646). The inflorescences, described by these authors as paniculate, are in fact more commonly glomerulate (the ramifications, if present, being very short). The internal sheath of the stipules apparently tends to disintegrate early in their development, and is often not visible on herbarium material. According to the label of Bordenave 1248, the texture of the fruits is reminiscent of marshmallow paste (���p��te de guimauve���). Additional specimens examined (paratypes): ��� FRENCH GUIANA. Saut Tortue, 4��11���N, 52��24���W, 17 November 1994 (fr.), B. Bordenave 1248 (P); Tumuc Humac, for��t dense sur la cr��te du Mitaraka sud, �� 1,5 km environ �� l���ouest du sommet, 17 August 1972 (fr.), J.J. de Granville 1280 (CAY); Monts Galbao, �� 10 km WSW de Sa��l, 14 March 1973 (imm. fr.), J.J. de Granville 1536 (CAY, P); Pente NE des Monts Galbao, 11 March 1975 (imm. fr.), J.J. de Granville 2386 (CAY, P). Montagne Bellevue de l���Inini, extr��mit�� SW, versant NW, 15 August 1985 (fr.), J.J. de Granville, L . Allorge, G. Cremers, A.R.A. G��rts-van Rijn & J.F. Kodjoed 7525 (CAY); Montagne Bellevue de l���Inini, zone orientale, versant sous le vent, 28 August 1985 (fr.), J.J. de Granville, L . Allorge, G. Cremers, A.R.A. G��rts-van Rijn & J.F. Kodjoed 7858 (CAY); Montagne Bellevue de l���Inini, zone centrale, 8 September 1985 (fr.), J.J. de Granville, L . Allorge, G. Cremers, A.R.A. G��rts-van Rijn & J.F. Kodjoed 8109 (CAY); Mont Galbao, secteur Sud, 3��35���N, 53��20���W, 22 January 1986 (imm. fr.), J.J. de Granville, C. Feuillet, L . Hollenberg, O. Poncy & H. Sangray 8912 (CAY); same locality, 27 January 1986 (imm. fr.), J.J. de Granville, C. Feuillet, L . Hollenberg, O. Poncy & H. Sangray 9002 (CAY). SURINAME. Tumuc Humac Mts, Talouakem, 2��31���N, 54��45���W, 9 August 1993 (fr.), P. Acevedo-Rodriguez, J.J. de Granville, L . Hollenberg, A. Joly & C. Avril 5969 (CAY); Tumuc Humac, fronti��re Br��sil ��� Surinam, brousse et for��t basse sur le sommet du Paloulouim��enpeu, 2 August 1972 (fr.), J.J. de Granville 1087 (CAY). BRAZIL. Amazonas. Mun. Presidente Figueiredo, Rebio [Reserva Biologica] Uatum��, grade do PPBio, Baixio, 10 July 2008 (fr.), J.F. Stancik, S. Sakagawa, M.S. Tavares, R. L. da Silva & F.C. Costa 449 (K)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas, pp. 154-174 in Phytotaxa 531 (3) on pages 171-173, DOI: 10.11646/phytotaxa.531.3.1, http://zenodo.org/record/5886210, {"references":["Boom, B. M. & Delprete, P. G. (2002) Rubiaceae. In: Mori, S. A., Cremers, G., Gracie, C. A., Granville, J. - J. de, Heald, S. V., Hoff, M. & Mitchell, J. D. (Eds.) Guide to the Vascular Plants of Central French Guiana. Part 2. Dicotyledons. Memoirs of the New York Botanical Garden 76 (2): 606 - 647.","Taylor, C. M. & Gereau, R. E. (2013) The genus Carapichea (Rubiaceae, Psychotrieae). Annals of the Missouri Botanical Garden 99 (1): 100 - 127. https: // doi. org / 10.3417 / 2011064"]}

Rudgea glomerulata Zappi & O. Lachenaud, sp. nov. (Fig. 3) Stipulis et bracteis valde laciniatis, calyce tubo conspicuo 1.5 ��� 4 mm longo lobisque triangularibus, inflorescentiis pendulis et disco bipartito Rudgeae itoupensi valde similis, sed differt bracteis externis maioribus et angustioribus, 9 ��� 28 �� 2.5 ��� 3.5 mm (vs. ca. 7 �� 7 mm) apice longe acuminatis et calycibus valde excedentibus, corollaque tubo 18 ��� 20 mm longo (vs. 11.5 mm) Type: ��� FRENCH GUIANA. Montagne de Kaw, route de Kaw, p.k. 46, 4��33���N, 52��09���W, 14 December 1986 (fl.), J.J. de Granville 9075 (holotype, MO [MO-2365274]; isotypes, CAY [CAY079168], P [P03985332], U [U 0130743]). Shrub, 1 ��� 3 m tall, much-branched; terminal branchlets 2���3 mm thick, densely pubescent with appressed hairs, soon covered with a pale brownish bark. Stipules consisting of two flat interpetiolar portions, narrowly ovate, 8 ��� 13 �� 2 ��� 5 mm, deeply laciniate with 15 ��� 20 marginal appendages 1.5���5 mm long and lacking dorsal appendages, surrounding an internal sheath 2 ��� 3.5 mm long bearing ca. 10 lateral appendages 1.5 ��� 3 mm long in the axils of the petioles, pubescent, soon corky and eventually caducous. Leaves opposite, petiolate; petioles 0.5 ��� 2 cm long, pubescent like the stems; blades elliptic to narrowly obovate, 10.5 ��� 24 �� 3 ��� 8 cm, acute to obtuse at base, acuminate at apex, the margin slightly revolute, papyraceous to slightly coriaceous when dry, glabrous above, pubescent below with sparse appressed hairs 0.2 mm long, drying olive green to olive brown; midrib convex on both sides, secondary veins 7 ��� 14 on each side of midrib, weakly ascending, forming loops at 1���5 mm from the margin; tertiary veins concolorous, prominently and rather laxly reticulate in the dry state; domatia absent. Inflorescences terminal, capitate to shortly branched, 4.5 ��� 15 cm long, many-flowered, �� pendulous, densely appressed hairy; peduncle terete, 2.5 ��� 13.5 cm long; flowering portion 1 ��� 3 �� 2 ��� 5.5 cm; ramifications absent or very short (��� 28 �� 2.5 ��� 3.5 mm, deeply and irregularly laciniate towards their base, the upper ones similar but narrower, 9 ��� 11 �� 1.5 ��� 2 mm, all densely appressed hairy outside and glabrous inside. Flowers (4)5-merous, whether distylous unknown, sessile. Hypanthium tronco-conical to subcylindrical, 1 ��� 1.5 mm long, densely villose. Calyx deeply cupuliform, pubescent outside; tube 1.5 ��� 4 mm long; lobes triangular, 1 ��� 2 mm long, the apex acute but often damaged. Corolla white; tube narrow and almost cylindrical, 18 ��� 20 �� 1.5 mm, glabrous throughout; lobes triangular, 4.5 ��� 6 mm long, broadly corniculate at apex, glabrous on both sides except the cornicula minutely puberulous. Stamens fully exserted; filaments exserted 3 mm beyond corolla mouth; anthers 2.5 �� 0.4 mm, dorsifixed. Disk bipartite, cylindrical, 1 mm long, glabrous. Style included, 5 mm long, branches 1 mm long. Fruits ellipsoid to ovoid or rarely subglobose, 15 ��� 18 �� 10 ��� 15 mm when fresh, 7.5 ��� 17 �� 7.5 ��� 13 mm when dry, dark green when immature, pale yellow to yellow-orange when mature, glabrous, on short accrescent pedicel 2 ��� 5 mm long, crowned with persistent calyx tube; mesocarp yellowish-white. Pyrenes ellipsoid, 10.5 ��� 13.5 �� 8 ��� 9 mm, the dorsal surface minutely verruculose, smooth; seeds with a deep T-shaped ventral furrow. Distribution and ecology: ���Only known from northern French Guiana, where it occurs mostly in the northeastern hill ranges (Montagne des Trois Pitons, Nouragues, and eastern side of the Kaw mountain) with an isolated record further west in the Sinnamary basin (Fig. 4). The species occurs in the undergrowth of mature forest, at 90���400 m elevation. Phenology: ���The species was collected with flowers in November and December, corresponding to the beginning of the first rainy season. Immature fruits have been found in February (end of first rainy season) and mature ones from April to July (second rainy season). Conservation status assessment:���Vulnerable (VU) (B1ab(iii)+2ab(iii)). Rudgea glomerulata is endemic to French Guiana, and is known from 16 collections, representing eight occurrences. Its extent of occurrence (EOO) is calculated to be 5,026 km ��, within the limit for Vulnerable status under subcriterion B1, and its area of occupancy is estimated to be 40 km ��, within the limit for Endangered status under subcriterion B2. Two of its occurrences are protected in the R��serve Naturelle des Nouragues and R��serve Naturelle des Marais de Kaw-Roura. The other occurrences are unprotected but most of them are found in remote areas with currently low level of threat. However, one on the Kaw Mountain is in an area experiencing some level of timber exploitation, and also harboring important bauxite and gold deposits; mining projects there were abandoned in 2008, but may resurface in the future. Another potential threat to this occurrence comes from touristic development and related infrastructure. We may therefore project a decline in habitat extent and quality. The eight occurrences represent eight locations in the sense of IUCN, and the species qualifies for Vulnerable status under the conditions B1ab(iii)+2ab(iii). Notes: ���This species is similar to R. itoupensis, described below; the two are remarkable in the genus by their deeply laciniate bracts and stipules (shallowly laciniate in most other species), �� pendulous inflorescences, welldeveloped calyx tube, and bipartite disk (the disk in Rudgea species is usually entire). It differs from R. itoupensis by its larger and narrower bracts that long exceed the calyces, and its longer corolla tube (Table 2). The two species also have different geographical and altitudinal ranges. In view of their stipule morphology and fruit colour, these species probably belong to the informal ��� lanceifolia clade��� defined by Bruniera (2015). Another species resembling R. glomerulata is Palicourea yneziae C.M.Taylor (Taylor 2015: 81) [Syn. Rudgea mexiae Standley (1936: 165)] from Peru and southern Colombia. This species was recently transferred from Rudgea to Palicourea, although it is unusual in the latter genus ��� especially due to its fimbriate stipules ��� and its placement there is provisional (Taylor 2015: 59���60). Its stipules, bracts and calyces are quite similar to those of R. glomerulata, although the leaf venation, disk, and fruit colour are different in both taxa (Table 2). In the absence of phylogenetical analysis including these species, it is unclear whether the resemblance between them is due to convergence or indicative of an affinity. There is also some resemblance between R. glomerulata and R. lanceifolia, especially in the shape of the stipules, but the differences are numerous: R. lanceifolia is a taller plant 3-10 m high, with more strongly ascending secondary leaf veins, an erect and usually branched inflorescence, entire or shortly dentate bracts, a calyx divided almost to the base, a usually longer (17-80 mm) corolla tube that is pubescent on both sides, and fruits dark red when immature and black when mature, crowned with an entire markedly accrescent disk 2-3 mm long and 5-10 mm in diameter. The inflorescences of Rudgea glomerulata are often capitate, but may have short ramifications, especially at the fruiting stage, as the rachis might expand after anthesis. It is not known whether the flowers are heterostylous; the only open flowers seen (on the type) are short-styled. Two collections, one from the Pakaraima Mountains in Guyana, K.M. Redden, R. Williams, C. Perry, C. Paul & M. Lyle 1927 (P [P01019851]) and the another from Araracuara in Colombia, H. Vester & A. Matapi 754 (L.4195512]), very closely resemble R. glomerulata in characters of the bracts, inflorescences and stipules, as well as in leaf shape and venation, but have a hirsute (rather than appressed) indumentum on the petioles and lower surface of leaf veins. The former specimen also has a much shorter peduncle, while the latter appears to have an erect inflorescence. More material is required to decide whether these collections are conspecific or not with R. glomerulata. Additional specimens examined (paratypes): ��� FRENCH GUIANA. Nouragues, 13 May 1985 (fr.), A. Cockle 101 (CAY); Eastern Plateau of Montagne Tortue, 11 km WNW of Approuague River, 4��18���N, 52��22���W, 12 June 1988 (fr.), C. Feuillet 10045 (K); Station des Nouragues, 4��03���N, 52��42���W, 22 November 1989, (fl. buds), G. Cremers 10928 (CAY); for��t sur la rive gauche de l���Arataye, �� environ 2 km du Saut Parar��, 14 February 1969 (imm. fr.), J.J. de Granville 82 (CAY, P); estuaire de l���Oyapock, entre le village de Petit Toucouchi et la montagne des Trois Pitons, 20 January 1981 (fallen fl.), J.J. de Granville 4279 (CAY, P, U); Kaw: Montagne Favard, 20 April 1984 (fr.), J.J. de Granville 6880 (CAY, MO); Montagne de Kaw, extr��mit�� est, versant sud, 3 November 1985 (fallen fl.), J.J. de Granville 8245 (CAY, MO); Station des Nouragues, 4��03���N, 52��42���W, 23 February 1991 (imm. fr.), J.J. de Granville 11173 (CAY, MO); D.Z. de Crique Jupiter, bassin du Sinnamary, 24 April 1991 (imm. fr.), J.J. de Granville, C. Roesel & L. Brothers 11497 (CAY); Station des Nouragues, 4��03���N, 52��42���W, June 1989 (fr.), D. Larpin 643 (CAY); mont��e du Pic des Trois Pitons, 9 June 1980 (fr.), C. Moretti 1166 (CAY, P); Nouragues Field Station, 4��05���N, 52��40���W, 27 February 2002 (imm. fr.), S.A. Mori, F. Blanchard & T.A. Lobova 25483 (CAY, NY, P); Arataye (affluent de l���Approuague) au saut n��1, rive droite, 10 February 1969 (imm. fr.), R.A.A. Oldeman B-2127 (CAY); Fleuve Arataye, Saut Parar��, 27 July 1984 (fr.), B. Ri��ra 659 (CAY); station des Nouragues, bassin de l���Arataye, 4��03���N, 52��42���W, 11 July 1989 (fr.), D. Sabatier & M.-F. Pr��vost 2533 (CAY, P, U)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas, pp. 154-174 in Phytotaxa 531 (3) on pages 159-162, DOI: 10.11646/phytotaxa.531.3.1, http://zenodo.org/record/5886210, {"references":["Bruniera, C. P. (2015) Sistematica e taxonomia de Rudgea Salisb. (Palicoureeae, Rubiaceae) Unpublished Doctoral Thesis, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, 283 pp. [http: // www. teses. usp. br / teses / disponiveis / 41 / 41132 / tde- 28072015 - 145432 / en. php]","Standley, P. C. (1936) Studies of American Plants VI. Publications of the Field Museum of Natural History, Botanical Series 11 (5): 145 - 276. https: // doi. org / 10.5962 / bhl. title. 2345"]}

Rudgea jadinii O. Lachenaud, sp. nov. (Fig. 7-8) Stipulis brevis apice fimbriatis dorso exappendiculatis, foliis glaberrimis venulis inconspicuis et corollae tubo longe et anguste cylindrico Rudgea graciliflora Standley (1936: 262) similis, sed differt inflorescentiis glomerulatis (vs. breviter ramosis), staminibus et stylo in corollae tubo inclusis, et calyci lobis majoribus 1.5-2.5 mm longis (vs. 0-1 mm longis). Type: ��� FRENCH GUIANA: Route de Kaw, pk 33, sentier vers les grottes, 4��33���N, 52��13���W, 4 December 2000 (fl.), F. Billiet & B. Jadin 7459 (holotype, CAY [CAY014902]; isotype, BR [BR000000907386]). Shrub, much branched, or small tree, to 6 m tall; terminal branchlets 1 ��� 1.5 mm thick, glabrous. Stipules 2 ��� 3 �� 2 ��� 3 mm, glabrous, marcescent and soon corky, consisting of a truncate to round basal sheath 0.5 ��� 2 mm long (usually broken at flower-bearing nodes), bearing on each side a dorsal keel with 4 early caducous terminal appendages Leaves opposite, petiolate; petioles 0.3 ��� 1 cm long, glabrous; blades elliptic, 6.5-12.2 �� 3-6.4 cm, obtuse to acute at base, abruptly acuminate for 0.5 ��� 1.5 cm at apex, coriaceous, entirely glabrous, drying olive green to blackish; midrib concave or flat above, secondary veins 7 ��� 10 on each side of midrib, flat above, weakly ascending, forming conspicuous loops 1.5 ��� 4 mm from the margin, tertiary veins inconspicuous or very lax; domatia absent. Inflorescences terminal, glomerulate, 5 ��� 15-flowered, sessile or with peduncle ��� 3.5 �� 0.7 ��� 1.5 mm, entire or minutely toothed, glabrous at base and shortly pubescent at apex. Flowers sessile, 5-merous, whether distylous unknown. Hypanthium tronco-conical, 0.5 mm long, glabrous. Calyx lobed nearly to the base; lobes narrowly ovate, 2 ��� 2.3 �� 0.5 ��� 0.8 mm, glabrous at base, minutely and sparsely pubescent at apex, alternating with minute colleters in their sinuses. Corolla white; tube very narrowly cylindrical, 6.5 ��� 10 �� 0.1 ��� 0.15 cm, glabrous outside, pubescent at distal 1.5 cm inside; lobes narrowly elliptic, 8 ��� 12 �� 1.5 mm, with short rounded dorsal appendages near apex, glabrous outside except the minutely puberulous appendages, papillose inside. Stamens included, inserted 4 mm below corolla mouth, subsessile; anthers narrowly elliptic, 3 �� 0.4 mm. Disk cylindrical, 0.5 mm long, glabrous. Style included, glabrous, 6.5 cm long, branches 2.5 mm. Fruits unknown. Distribution and ecology: ���Endemic to northeastern French Guiana, where known from the Kaw Mountain Range and the Nouragues inselberg (Fig. 9); in the former locality it occurs in low forest on lateritic crust, in the latter presumably in a similar habitat but on granite. The species appears to be very rare; the first author was unable to find it in the type locality, despite visiting the site on several occasions at various times of the year. Phenology: ���Flowering specimens were collected twice in December, at the beginning of the first rainy season, which represents a flowering peak for most shrubby Rubiaceae in French Guiana. Conservation status assessment: ���Endangered [B2ab(iii)]. Rudgea jadinii is endemic to French Guiana and is known from two collections representing two occurrences. Its extent of occurrence (EOO) is therefore not calculable, and its area of occupancy is estimated to be 8 km ��, within the limit for Critically Endangered under subcriterion B2. One of its occurrences is protected in the R��serve Naturelle des Nouragues. The other, on the Kaw Mountain, has no official protection status, and is potentially threatened by touristic development, forest exploitation and/or mining activities (mining projects in the area were abandoned in 2008 but may resurface in the future), which leads us to anticipate a decline in habitat extent and quality. The two occurrences represent two locations in the sense of IUCN, and the species qualifies for Endangered status under the conditions B2ab(iii) Notes: ��� Rudgea jadinii resembles R. graciliflora, from which it differs by the glomerulate inflorescence, flowers with anthers and style both included, and calyx lobes 1.5 ��� 2.5 mm long, while R. graciliflora has shortly branched inflorescences, distylous flowers with either style or anthers exserted, and the calyx truncate or with lobes R. jadinii are 6.5 ��� 10 cm long, which are much longer than in specimens of R. graciliflora from the Guianas, which are 2 ��� 5.2 cm long and represent the typical form of the species. However, some collections of R. graciliflora from western Amazonia have the corolla tube length comparable to that of R. jadinii. These have been originally described as R. klugii Standley (1936: 164); their status may have to be re-assessed, but this is outside the scope of this paper. Rudgea graciliflora belongs to the informal ��� lanceifolia clade��� (Bruniera 2015) which includes species from the Guianas and the in Amazon basin, and R. jadinii probably belongs there as well. It is not known whether R. jadinii is heterostylous; the only flowers seen have the style and anthers both included, the latter well above the former. Eponymy: ���This species is named in memory of the Belgian botanist Bernard Jadin (1948���2012), collector of the type and of many other French Guianan plants, together with his wife Frieda Billiet - who is remembered in Philodendron billietiae Croat (1995: 24). Additional specimen examined (paratype): ��� FRENCH GUIANA. Station des Nouragues, entre les croisements H XXII et I XXII, 27 December 1988 (fl.), D. Loubry 100 (CAY)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas, pp. 154-174 in Phytotaxa 531 (3) on page 168, DOI: 10.11646/phytotaxa.531.3.1, http://zenodo.org/record/5886210, {"references":["Standley, P. C. (1936) Studies of American Plants VI. Publications of the Field Museum of Natural History, Botanical Series 11 (5): 145 - 276. https: // doi. org / 10.5962 / bhl. title. 2345","Bruniera, C. P. (2015) Sistematica e taxonomia de Rudgea Salisb. (Palicoureeae, Rubiaceae) Unpublished Doctoral Thesis, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, 283 pp. [http: // www. teses. usp. br / teses / disponiveis / 41 / 41132 / tde- 28072015 - 145432 / en. php]","Croat, T. B. (1995) Two new species of Araceae for the Guianas. Novon 5: 24 - 29. https: // doi. org / 10.2307 / 3391825"]}

Lachenaud, Olivier, Bruniera, Carla P., Zappi, Daniela C. (2022): Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas. Phytotaxa 531 (3): 154-174, DOI: 10.11646/phytotaxa.531.3.1

Rudgea approuaguensis O. Lachenaud, sp. nov. (Fig. 1) Inflorescentiis glomerulatis et breviter pedunculatis, fructibus rubris et ellipsoideis, foliis glabris crassis venulis laxissimis, stipulisque margine fimbriatis dorso exappendiculatis et applanatis Rudgeae bremekampianae Steyermark (1967: 407) valde affinis, sed habitu humiliore (35 ��� 40 cm nec 1 ��� 4 m alto), caule breviter pubescente (nec glabro), foliis minoribus nervis lateralibus paucioribus, inflorescentiis ebracteatis (nec bracteis foliaceis conspicuis munitis), calyce breviore lobato et corolla breviore (tubo 6 mm longo nec 10 ��� 16 mm) conspicue differt. Type: ��� FRENCH GUIANA. Haute Approuague, sur la Crique Calebasse au village Germain, 25 August 1968 (fr.), R.A.A. Oldeman B-1828 (holotype, P [P06839007]; isotypes, CAY [CAY021301], K [K001301358], L n.v., NY [NY03120147]). Subshrub, 35 ��� 40 cm tall, sparsely branched; terminal branchlets 2.5 ��� 3 mm thick, densely and shortly pubescent with patent hairs 0.3 mm long, soon covered with a corky buff-grey bark. Stipules consisting of two flat interpetiolar portions, broadly elliptic, 7 ��� 12 �� 4 ��� 6 mm, with numerous marginal appendages 1.5 mm long and lacking dorsal appendages, externally puberulous, soon corky and easily damaged, hiding an internal sheath 1-3 mm long. Leaves opposite, petiolate; petioles 0.2 ��� 0.7 cm long, densely puberulous; blades elliptic or obovate, 7 ��� 13 �� 2.6 ��� 4.6 cm, acute at base, shortly acuminate at apex, coriaceous, glabrous on both sides, drying olive green to greyish; midrib flat or concave above; secondary veins 9 ��� 12 on each side of midrib, rather ascending, not very prominent, forming inconspicuous loops 1.5 ��� 3 mm from the margin; tertiary veins laxly reticulate, slightly prominent when dry (possibly invisible in the fresh state); domatia absent. Inflorescences terminal, glomerulate, ca. 20-flowered, erect; peduncle terete, puberulous, 0.5 cm in flower, 1.5 cm in fruit; flowering part 0.8 cm in diameter in flower; no distinct ramifications or bracts. Flowers 5-merous, whether distylous unknown, sessile. Hypanthium tronco-conical, 0.3 mm, glabrous. Calyx cupuliform, 1 mm long, divided in short, round lobes for 1/3 or 1/2 of its length, densely ciliate. Corolla white; tube nearly cylindrical, 6 �� 3 mm, glabrous outside; lobes triangular, 3.5 mm, not or hardly corniculate, minutely puberulous outside at the apex, otherwise glabrous. Stamens exserted just beyond the corolla mouth; filaments barely exceeding corolla mouth; anthers elliptic, 2 �� 0.25 mm. Disk hemispherical, 0.5 mm long. Style included, 4.5 mm long, glabrous; branches 1.2 mm. Fruits ellipsoid, 12���13 �� 8���9 mm when dry, round at apex, red, glabrous, sessile; calyx scar not enlarging. Pyrenes ellipsoid, 11.5 �� 6.8 mm, dorsally smooth except for a slight median ridge in the lower half; seeds with a deep T-shaped ventral furrow. Distribution and ecology: ���Endemic to French Guiana; apparently restricted to the upper Approuague River (Fig. 2). Occurs presumably in lowland forest, ca. 100 m in elevation, although the habitat is not precisely recorded. Phenology: ���Flowers were collected only once in July, at the end of the second rainy season, and fruits only once in August. Conservation status assessment: ���Endangered [EN B2ab(iii)]. Rudgea approuaguensis is endemic to French Guiana and is known from two specimens, representing two occurrences. Its extent of occurrence (EOO) is therefore not calculable, and its area of occupancy is estimated to be 8 km ��, within the limit for Critically Endangered under subcriterion B2. The area where it occurs is not protected, and illegal gold mining occurring along the Approuague River is a potential threat, which leads us to predict a decline in the extent and quality of habitat. The two occurrences represent two locations in the sense of IUCN, and the species qualifies for Endangered status according to the conditions B2ab(iii). Notes: ���This species closely resembles Rudgea bremekampiana Steyermark (1967: 407), from French Guiana and northern Brazil (state of Amap��), and has been previously confused with it in herbaria. Both species have glomerulate inflorescences with a short peduncle, red, ellipsoid fruits, thick, glabrous leaves, and flat stipules (i.e., lacking a dorsal keel) with marginal appendages. Rudgea bremekampiana differs from R. approuaguensis by the glabrous stems, conspicuous bracts, and greater dimensions of all its organs (differences are summarised in Table 1). It is not known whether the species is heterostylous, since only one collection bears flowers; these are too few to dissect, and the interior of the corolla tube is therefore not described. Additional specimen examined (paratype): ��� FRENCH GUIANA. Fleuve Approuague, au Saut Grand Canori, 12 July 1968 (fl.), R.A.A. Oldeman 2765 (CAY, P)., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas, pp. 154-174 in Phytotaxa 531 (3) on pages 155-157, DOI: 10.11646/phytotaxa.531.3.1, http://zenodo.org/record/5886210, {"references":["Steyermark, J. A. (1967) Rubiaceae. In: Maguire, B. et al. (Eds.) The Botany of the Guayana Highland-Part VII. Memoirs of the New York Botanical Garden 17: 230 - 436."]}

Rudgea itoupensis O. Lachenaud, sp. nov. (Fig. 6) Inflorescentiis pendulis dense globosis bracteis valde laciniatis munitis discoque bipartito ab omnibus congeneribus differt Rudgea glomerulatae excepta, a quae distinguitur bracteis externis brevioribus et latioribus 7 �� 7 mm (vs. 9 ��� 28 �� 2.5 ��� 3.5 mm) calycibus non excedentibus et corollae tubo minore 11.5 mm longo (vs. 18 ��� 20 mm). Type: ��� FRENCH GUIANA: Sommet Tabulaire [Mount Itoup��], zone sud, ca. 50 km SE de Sa��l, 22 August 1980 (fl.), J.J. de Granville 3550 (holotype, CAY [CAY077712]; isotype, CAY [CAY077713]). Shrub 3 m tall, much-branched; terminal branchlets 1.5 ��� 2 mm thick, densely appressed-pubescent. Stipules consisting of two flat interpetiolar portions, narrowly ovate, 9 ��� 16 �� 3 ��� 4 mm, deeply laciniate with ca. 13 marginal appendages 4 ��� 9 mm long and lacking dorsal appendages, pubescent, soon corky and eventually caducous, surrounding an internal sheath 2 mm long bearing ca. 4 lateral appendages 2 mm long in the axils of the petioles. Leaves opposite, petiolate; petioles 0.5 ��� 1 cm long, appressed pubescent; blades narrowly elliptic, 9.5 ��� 15.5 �� 1.9 ��� 3.7 cm, acute at base, gradually long-acuminate at apex, papyraceous when dry, glabrous above and shortly appressed hairy beneath with hairs denser on the veins, drying dark grey-brown, midrib convex on both sides, secondary veins 9 ��� 12 on each side of midrib, weakly ascending, forming rather conspicuous loops 1 ��� 2 mm from the margin, tertiary veins prominent and forming a moderately lax reticulum (ca. 2 mm) in the dry state; domatia absent. Inflorescences terminal, nodding, capitate and hemispherical, many-flowered, densely appressed-pubescent; peduncle terete, 1.3 ��� 4.5 cm long; flowering portion 1.1 ��� 1.5 cm in diam., no distinct ramifications; bracts green, not exceeding the calyces, the basal ones 7 �� 7 mm, deeply and irregularly laciniate for about 2/3 of their length, the upper ones almost linear and pinnatifid, 3.5 �� 0.4 mm, all densely appressed hairy outside and glabrous inside. Flowers sessile, (4 ���)5-merous, apparently distylous (see Notes). Hypanthium tronco-conical, 1 mm, pubescent. Calyx deeply cupuliform, pubescent outside, tube 2.5 ��� 4 mm long, lobes initially triangular and acute at apex, 1 mm long, but very soon damaged, becoming �� round and scariose at margin. Corolla white; tube narrowly cylindrical, 11.5 �� 1 mm, glabrous outside and inside; lobes triangular, 3 mm, broadly corniculate at apex, glabrous on both sides except the cornicula minutely puberulous. Stamens included with their apex just reaching corolla mouth in long-styled flowers; anthers linear, 3.5 �� 0.5 mm, dorsifixed. Disk bipartite, cylindrical to slightly conical, 1 ��� 1.3 mm long, glabrous. Style included, glabrous, 10 mm long and almost reaching corolla mouth in long-styled flowers, or 3.8 mm long in short-styled flowers, branches 0.8 ��� 1 mm long. Fruits unknown. Distribution and ecology: ���Endemic to French Guiana (Fig. 4); apparently restricted to Mount Itoup�� (also known as Sommet Tabulaire) in the central Inini-Camopi chain. Collected only once in submontane forest, at 750 m elevation. Phenology: ���Flowers collected once in August. Conservation status assessment:���Vulnerable (VU) (D2). Rudgea itoupensis is known from a single collection near the summit of Mount Itoup�� in south-central French Guiana, where it occurs in submontane forest. Its extent of occurrence (EOO) is therefore not calculable, and its area of occupancy (AOO) is estimated to be 4 km ��, within the limit for Critically Endangered under subcriterion B2. Its only known location, which lies in the Parc Amazonien de Guyane, is very remote and difficult to access, and unlikely to be directly impacted by human activities. However, considering its very restricted range and particular habitat, the species may be at risk from any stochastic events (and possibly from future climatic changes). Being known from a single location, it qualifies for Vulnerable status under criterion D2. Notes: ���This species is very close to Rudgea glomerulata, described above, from which it differs by the shorter and relatively much broader bracts, not exceeding the calyces, and a shorter corolla tube; both species are apparently endemic to French Guiana, but have different ranges and habitats (Table 2). They share dense and more or less pendulous inflorescences with deeply laciniate bracts, narrowly elliptic leaves with a sparsely hairy lower surface, a well-developed calyx tube, and a bipartite disk, while all other Rudgea species in the Guianas (and, as far as we know, elsewhere) have an entire disk. The only collection seen includes both long- and short-styled flowers (the latter with the corollas fallen off and only the style remaining) that are on separate branches, which were presumably collected from different individuals., Published as part of Lachenaud, Olivier, Bruniera, Carla P. & Zappi, Daniela C., 2022, Six new and a little-known species of Rudgea (Rubiaceae-Palicoureeae) from the Guianas, pp. 154-174 in Phytotaxa 531 (3) on pages 166-167, DOI: 10.11646/phytotaxa.531.3.1, http://zenodo.org/record/5886210

The Global Strategy for Plant Conservation (GSPC) was established by the Conference of Parties in 2002 to decrease the loss of plant diversity, reduce poverty and contribute to sustainable development. To achieve this overarching goal, the GSPC has established a series of targets, one of which is to ensure that plant diversity is well understood, so that it can be effectively conserved and used in a sustainable manner. Brazil hosts more than 46,000 species of plants, algae and fungi, representing one of the most biodiverse countries on Earth, and playing a key role in the GSPC. To meet the GSPC goals of Target 1 and facilitate access to plant diversity, Brazil committed to preparing the List of Species of the Brazilian Flora (2008-2015) and the Brazilian Flora 2020 (2016-present). Managing all the information associated with such great biodiversity has proven to be an extremely challenging task. Here, we synthesize the history of these projects, focusing on the multidisciplinary and collaborative approach adopted to develop and manage the inclusion of all the knowledge generated though digital information systems. We further describe the methods used, challenges faced, and strategies adopted, as well as summarize advances to date and prospects for completing the Brazilian flora in 2020.

Examination of type specimens at the P herbarium showed that Almeidea longifolia A. St.-Hil. (Rutaceae) is an illegitimate substitute name for A. affinis A. St.-Hil. The latter name is proposed here as a heterotypic synonym of A. rubra A. St.-Hil. O estudo dos tipos nomenclatórios no herbário P mostrou que Almeidea longifolia A. St.-Hil. (Rutaceae) é um nome substituto ilegítimo de Almeidea affinis A. St.-Hil., e este último nome é aqui considerado sinônimo heterotípico de A. rubra A. St.-Hil.