Carapichea ligularis (Rudge) Delprete (Delprete 2003: 89)

Lachenaud, Olivier & Delprete, Piero, 2022, Revision of Carapichea (Rubiaceae - Psychotrieae) in the Guianas, with two new combinations and transfer of three species to Notopleura, Plant Ecology and Evolution 155 (2), pp. 275-300 : 275

5. Carapichea ligularis (Rudge) Delprete (Delprete 2003: 89)

Fig. 6

Schradera ligularis Rudge ( Rudge 1806: 29, plate 45) - Type: same as for Carapichea ligularis .

Cephaelis ligularis (Rudge) A.Rich. ex DC. ( Candolle 1830: 533) - Type: same as for Carapichea ligularis .

Psychotria ligularis (Rudge) Steyerm. ( Steyermark 1972: 675) - Type: same as for Carapichea ligularis .

Psychotria pacimonica Müll.Arg., syn. nov. ( Müller 1881: 337) - Type: VENEZUELA - Amazonas • "in regione superiore Rio Negro ad flumen Pacimoni"; Feb. 1854; fl.; Spruce 3445; lectotype (designated here): P [P00837131]; isolectotypes: G [G00300288], K [K000432842], W [1889-0014290].

Uragoga pacimonica ( Müll.Arg.) Kuntze ( Kuntze 1891: 961) - Type: same as for Psychotria pacimonica .

Carapichea pacimonica ( Müll.Arg.) C.M.Taylor ( Taylor and Gereau 2013: 120) - Type: same as for Psychotria pacimonica .

Type.

FRENCH GUIANA [as “Guiana”] • s.loc.; s.d.; fl.; Martin s.n.; holotype: BM [BM001009103].

Description. Subshrub or shrub, 0.2-2(-4) m tall, single-stemmed or weakly branched (or rarely trees 7-10 m tall in the Brazilian and Venezuelan Amazon); terminal internodes terete, 2-3(-4) mm in diam., glabrous. Stipules shallowly sheathing, glabrous; sheath truncate to shallowly elliptic; lobes broadly triangular to broadly ovate, 2-5 mm long, persistent. Leaves with petioles 0.3-3.3 cm long, glabrous; blades elliptic, oblanceolate to narrowly oblong-elliptic, (4-)8.5-21 × (1.5-) 2-6.5 cm, acute-decurrent at base, acute and long-acuminate at apex, acumen narrowly triangular, 0.5-2.5 cm long, sometimes falcate, papyraceous to coriaceous, drying pale olive-green to dark grey-green, glabrous throughout (except sometimes the domatia); secondary veins 14-26 on each side of the midrib; intersecondary veins 2-3(-4) between each couple of secondary veins, terminating far from the margin; tertiary veins barely visible or obsolete; domatia absent, or present as a row of hairs on each side of the midrib below. Inflorescence capitate to subcapitate (shortly branched, ramifications ≤ 0.5 cm when present); peduncle 1.5-7.0 cm long, glabrous to shortly pubescent; bracts inserted at distal end of peduncle and sometimes also on inflorescence axes, shallowly connate or free at base, (2-)4-8, usually decussate, narrowly lanceolate to linear, (7-)12-30 × (1-) 2-4 mm, pale green or pale orange-green, glabrous, persistent. Flowers 5-merous, heterostylous, sessile to subsessile, the pedicels elongating to 2-5 mm long at fruiting stage. Hypanthium obovoid, 0.6-1.0 mm long, glabrous. Disk bilobed to the base, 0.5 mm long, shorter than or as long as the calyx. Calyx cupular, glabrous, tube 0.5-1.0 mm long, truncate or with short triangular lobes <0.5 mm long. Corolla hypocrateriform, orange to salmon colored, 7.5-13(-14) mm long, glabrous; tube subcylindrical, gradually wider towards the mouth, 6-10.4(-11) mm long, 0.8-1.0 mm wide at base, 1.5-1.6 mm wide at mouth, glabrous outside and inside; lobes oblong-ovate, 2-4 × 0.8-0.9 mm, acute at apex, not corniculate, glabrous. Short-styled flowers: Stamens inserted at distal 1/3rd of the tube, anthers partially exserted beyond corolla mouth; filaments ca 0.3 mm long; anthers 2.3-2.5 × 0.5 mm; style included, 3.5-4.0 mm long; style branches narrowly oblong, 1.5 mm long. Long-styled flowers: stamens inserted at about the middle of the corolla tube, included; filaments 1.5 mm long, glabrous; anthers narrowly oblong, 2.5 × 0.3-0.4 mm, acute at both ends; style exserted just beyond the corolla mouth, 12.5 mm long (corolla tube 10.5 mm long), style branches obovate, ca 1 mm long. Fruits ellipsoid to oblong-ovoid, 6-12 × 4-9 mm, smooth (slightly costate when dry), orange to orange-red. Pyrenes plano-convex, elliptic in outline, 7-9 × 3-5 mm, dorsal side with 3 ridges, ventral side with a very shallow longitudinal groove. Seeds with a deep T-shaped ventral furrow.

Distribution.

This species occurs in southeastern Venezuela (state of Amazonas), French Guiana, and eastern Amazonian Brazil (states of Amapá, Pará, Roraima, and Amazonas). It is locally common, at least in French Guiana. It could be expected in Suriname and Guyana, but we have seen no collections of the species from these countries; the type specimen, reported to be from Guyana ( Delprete 2001: 399; Taylor and Gereau 2013: 120) was actually collected in French Guiana (see Notes below).

Ecology.

In understory of non-flooded forest, usually on superficial soils (e.g. lateritic crusts, granitic boulders) at 100-800 m elevation.

Phenology.

Flowering specimens were collected from August to January; and fruiting specimens throughout the year.

Selected specimens examined.

VENEZUELA - Amazonas • Cerro da Neblina, Río Yatua, along Upper Rio Yatua between mouth of Rio Yacibo and Piedra Arauicaua ; 1 Feb. 1954; fl.; Maguire et al. 37411; F , US.

FRENCH GUIANA • Route de l'Est , ca 25 km NW of Régina; 4°23'N, 52°19'W; 16 Mar. 1994; fr.; Andersson et al. 1989; BR, CAY • Cacao; 27 Oct. 1983; fl.; Billiet & Jadin 1877; BR • Route RN 2 Cayenne-Régina, pk 67, Crique Tibourou; 4°29'N, 52°19'W; 10 Feb. 1993; fr.; Billiet & Jadin 5733; BR • Extension Nord-Ouest des Petites Montagnes Tortue ; 4°20 ’29” N, 52°16 ’16” W; 30 m; 29 Sep. 2010; fl., fr.; Bordenave & Le Hir 9068; CAY, K, P, MO • Montagne Tortue, Plateau Est; 4°17'N, 52°21'W; 460 m; 5 Oct. 2010; fl.; Bordenave & Le Hir 9233; CAY • Upper Maroni River , layon de chasse au NE d’Antecume Pata, confluent de l’Itany et du Marouini; 19 Nov. 1977; fl.; Cremers 5077; CAY • Saül, piste du carbet; 31 Oct. 1984; fl.; Foresta 705; CAY • Monts Atachi Bacca, autour du Camp 3, entre les sommets 525 et 782 m; ca 400 m; 5 Mar. 1971; fr.; Granville C-128; CAY [2 sheets], P • Monts Atachi Bacca; 3 Mar. 1971; fr.; Granville 752; P • Rivière Petite Ouaqui, rive droite, au niveau de l’ancien village Hubert; 20 Jul. 1973; fr.; Granville 1872; CAY, P • Sommet Tabulaire, zone nord; 2 Sep. 1980; fr.; Granville 3710; P • Monts Bakra, 2,5 km à l’ouest du Pic Coudreau ; 2 Oct. 1980; fr.; Granville 4065; CAY, P • Mont Bellevue de l'Inini , central zone, slope near the summit; 23 Aug. 1985; fr.; Granville 7762; CAY, F, INPA, MG, MO, NY, P, U • Région de l’Inini, Mount Atachi Bacca, NW summit, 6 km E of Gobaya Soula, near Camp 2; 10 Jan. 1989; fl.; Granville et al. 10541; CAY, MO n.v., P, U • Montagne de la Trinité, Bassin de la Mana, partie supérieure de la pente N du plateau tabulaire; 13 Mar. 1997; fr.; Granville 13318; CAY, K, MO, P, U , US • Montagne Lucifer, SW du plateau sommital; 4°46 ’30”, 53°56 ’30” W; 520 m; 18 Nov. 1999; fr.; Granville & Crozier 13869; CAY, MO • Monts Bakra, 1.5 km W du Pic Coudreau ; 3°18'N, 52°57'W; 600 m; 18 Jun. 2002; fr.; Granville et al. 14864; B, CAY, MO, P • Monts Kotika, plateau latéritique sommital; 3°55 ’10” N, 54°11 ’10” W; 730 m; 23 Feb. 2005; fr.; Granville et al. 16922; B, CAY, MO, NY, P, U • Route Cayenne-Régina, km 52; 4°34.638'N, 52°23.873'W; 9 Jan. 2011; fr.; Lachenaud 1074; BR, CAY • route de Cacao; 4 May 2014; fr.; Lachenaud 1793; BR, CAY • Mont Itoupé, Sommet Tabulaire, Layon D, 2me sommet; 3°02 ’35” N, 53°05 ’18” W; 760 m; 31 Mar. 2010; fr.; Molino & Sabatier 2843; BR, CAY, MO • Montagnes de la Trinité; 4°36 ’30” N, 53°21 ’30” W; 12 July 2001; fl. buds, fr.; Poncy & Crozier 1458; CAY, NY, P, U , US • RN2 - Cayenne-Régina, km 85.5; 4°20'N, 52°18'W; 26 Jun. 2003; fl., fr.; Prévost & Sabatier 4742; CAY, MO • Rivière Itany, Crique Petit Marouini, près de son embouchure; 2 Sep. 1972; fr.; Sastre 1814; P .

BRAZIL - Amapá • Upper Jari River , foret primaire humide; 2°28'N, 54°46'W; 420 m; 20 Aug. 1993; fl.; Granville et al. 12387; BBS, CAY [2 sheets] , US. - Amazonas • Rio Curicuriarí, afl. do Rio Negro; 21 Dec. 1931; fl.; Ducke s.n.; INPA [No. 16533] • Nova Prainha , Rio Aripuana , RADAM / BRASIL, SB-20-ZD, Ponto 10; 9 Jul. 1976; fr.; C.D.A. Mota s.n.; INPA [No. 60607] • Mun. Nova Olinda, Rio Paca , tributari of the Rio Mari Mari , terra firme forest, tree to 7 m tall; 2 Jul. 1983; fr.; Todzia et al. 2299; INPA, MO, NY , US • Mun. Axinim, basin of Rio Abacaxis, lower Rio Paca , ca 1 km from its confluence with Rio Mari Mari , forest on terra firme, shrub 2 m tall; 4°07'S, 58°58'W; 1 Jul 1983; fr.; Zarucchi et al. 2917; INPA, MO, NY. - Pará • Ilha de Marajo , Rio Jipuru , afluente do Rio Anajas ; 20 Oct. 1987; fl.; A. Tavares & J. Cardoso 227; INPA, NY. - Roraima • Mun. Rorainópolis, Reserva Popular Xixuaú-Xiparina, ilha subindo o Rio Xiparina ; 0°55 ’41” S, 61°50 ’26” W; 25 Aug. 2010; fl, young fr.; Zappi et al. 2902; INPA, MIRR .

Notes.

This species has often been confused with C. guianensis (see Notes under that name) but is very different in leaf venation and bract shape ( Delprete 2001; Figs 5 , 6 ). It is much more similar to C. araguariensis and C. sp. A, from which it differs by the characters mentioned in the key.

Delprete (2001: 401) separated Carapichea ligularis from C. pacimonica by the lower habit, being a small, single-stemmed shrub 0.5-1.5 m tall (vs shrub up to 2 m tall or tree up to 10 m tall), the presence of corniculate appendages on the abaxial side of the corolla lobes (vs appendages absent), the subcapitate to sparsely cymose inflorescences (vs capitate), and the larger ovoid fruits (vs smaller, subspherical fruits). None of these characters proves to be reliable: the variation in habit and fruit seems to be continuous, the corolla appendages are commonly present in flower buds and usually fall off during or shortly after anthesis or during the preparation of herbarium specimens (Piero Delprete pers. obs.) and the inflorescences are usually capitate or nearly so at anthesis and often develop short ramifications in the fruiting stage. Taylor and Gereau (2013: 114) also kept the two species separate, based on other characters: "outermost inflorescence bracts at base straight-sided and not sheathing; plants often drying with a gray cast" in C. ligularis vs "outermost inflorescence bracts at base widened and shortly sheathing; plants often drying yellowish brown" in C. pacimonica . After a detailed study of numerous specimens throughout the geographic range, we observed that the inflorescence bracts can be shallowly sheathing or free at base in duplicate specimens of the same collection, and the colour of the dry plants may vary from greyish to olive green to pale brown to dark brown (possibly depending on the drying method). Taking into account all the above observations, we treat these two names as synonyms.

Steyermark (1972: 589) regarded Psychotria necopinata Standl. as a synonym of P. pacimonica , but we follow Taylor and Gereau (2013) in recognizing the former as a distinct species, Carapichea necopinata , which is only known from the type and not recorded from the Guianas to date.

The typification of C. ligularis has been a source of confusion. In his original description, Rudge (1806: 29, pl. 45) did not cite any gathering. Delprete (2001: 399) designated the lectotype as "Guyana, Rudge s.n. (P)", which was followed by Taylor and Gereau (2013: 120). However, this lectotypification is erroneous since no such specimen exists at P, and Rudge never travelled to South America; the material that he used to describe the taxa in his Plantarum Guianae Rariorum Icones et Descriptiones was in fact collected by Joseph Martin in French Guiana. The fate of Martin’s specimens was described by Stearn and Williams (1957), and summarized by Stafleu and Cowan (1983: 971-972). Succinctly, in 1803, France and England were at war, and the specimens collected by Joseph Martin in French Guiana, originally intended for the Museum of Natural History of Paris, were confiscated by two British privateers and brought to London. About 400 of these specimens were bought by the British Museum (BM), and 772 were bought by Rudge and included in his own herbarium ( Stafleu and Cowan 1983: 971-972). Also, Rudge gave a partial set of these specimens to Banks, whose herbarium became the founding collection of BM. After Rudge’s death, his widow donated his herbarium to BM in 1847. Therefore, most of Martin’s collections converged at BM, although some of his specimens are also reported to be at FI or FI-Webb ( Stafleu and Cowan 1983: 971-972). These specimens have the penciled information "Guiana. Martin", which may give the false impression that they were collected in modern day Guyana, while they actually came from French Guiana. After an exhaustive search for original material of S. ligularis , we were unable to find any specimen at FI and FI-Webb. However, a specimen with the indication "Guiana, Martin" penciled on the upper corner of the sheet, exists at BM [BM001009103], and being the sole original material, should be regarded as the holotype.

Müller (1881: 338), in the original description of Psychotria pacimonica , cited a single gathering, Spruce 3445, but he did not indicate the herbarium of deposit. Taylor and Gereau (2013: 120) assumed that the holotype is in M, but no specimen of Spruce 3445 was found there after an exhaustive search (Andreas Fleischmann pers. comm.). Specimens with this collection number are found in G, K, P, and W; the P sheet, which is the only one with open flowers, and has a label handwritten by Müller, is here designated as lectotype.

The photograph of the fruits published by Campos and Brito (1999: 627) as Psychotria pacimonica appears to represent another species, probably not a Carapichea .

Figures

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

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