Passiflora riparia was incompletely described by Masters, who cited specimens of Martius and Spruce. While Spruce 2191, the unique syntype with an observable corona, exhibits a reduced outermost series of filaments, the accompanying iconography represents two equal outer series. Later descriptions have neither added significant information nor corrected the inconsistency in the corona description, so that four closely related species have been distinguished on the basis of traits not properly documented for P. riparia: P. emiliae (unequal outer series of filaments), P. crenata (bract color), P. pergrandis (flower size and sepal awn length), and P. fernandezii (hypanthium pubescence and shape). The present study compares (i) the descriptions of the above-mentioned taxa and (ii) 43 associated vouchers, as well as live specimens from two associated P. crenata populations. These and other specimens were georeferenced for a comparison of their distribution and habitat. Of the five P. riparia descriptions found in floras, only that of the Flora of Ecuador appears clearly divergent, corresponding in fact to P. tolimana. Those of the four other taxa only differ by unequal corona filaments (except for P. crenata) and the pubescence of floral parts. However, 22 vouchers associated with all these descriptions (including 16 for P. riparia), as well as the live specimens, share both these traits; the other 21 vouchers were uninformative and/or could not be assigned to any of the five species. The wider sample of 62 specimens indicates no significant differences in either geographic or in climatic distribution (lowlands of the Amazon basin), and a marked preference for riparian habitats. Thus, their very close morphology and ecology justify the placement of P. emiliae, P. crenata, P. pergrandis and P. fernandezii as synonyms of P. riparia, designating Spruce 2191 as epitype. The most similar species, P. ambigua (20 specimens mapped), differs in corolla and bract color, as well as a distribution centered along the tropical Andes of South America and in Central America, in more diverse habitats.

Passifloraceae , subgenus Passiflora, series Laurifoliae

Five subgenera are currently recognized in the genus Passiflora L.: Passiflora, Astrophea (DC.) Mast., Decaloba (DC.) Rchb., Deidamioides (Harms) Killip and Tetrapathea (DC.) P. S. Green. Subgenus Passiflora, rich with more than 250 species, is characterized by large flowers with a corona made of several rows of filaments. Its supersection Laurifoliae (Cervi) Feuillet & J. M. MacDougal includes several series, organized around a clear morphological pattern (Rome and Coppens d’Eeckenbrugge 2017). In particular, series Laurifoliae Killip (1938) ex Cervi (1997) forms a very homogenous morphological group, with a very difficult taxonomy (Killip 1938, Holm-Nielsen et al. 1988). With the recently described P. gustaviana Ocampo & Molinari (Ocampo and Molinari 2017), it is composed of 25 species including glabrous to pubescent plants, whose stems are terete to angular and sometimes corky on old parts; leaves are unlobed, oblong to oblong-lanceolate, not peltate, with entire to glandular-serrulate margins, biglandular petioles; and stipules are setaceous or linear, and early deciduous. Their three bracts, free at base, with entire or serrulate-glandular margins, are more than 1 cm long. Their flowers are pendent, usually large and showy, often fragrant, with a short hypanthium and two campanulate series of long external filaments, and a variable number of series of reduced internal filaments (Rome and Coppens d’Eeckenbrugge 2017).

According to Killip (1938), Passiflora riparia Martius ex Masters appears to be the commonest representative of the series Laurifoliae in the upper and middle Amazon basin. Its original description by Masters (1872), in Flora Brasiliensis, refers to two syntypes: Martius 3228, from the mouth of the Madeira River in the Upper Amazonas, and Spruce 2191 (the latter cited by Masters with an exclamation mark), collected from São Gabriel da Cachoeira, near the Rio Negro. The number of series of external corona filaments is not mentioned in Masters’ description. In the accompanying botanical iconography, flowers appear to have one series of long filaments, whereas a schematic longitudinal section shows two equal series of external filaments. This detail has become problematic since the relative length of the two external filament series became an important diagnostic trait when Killip (1938) created the series.

Martius’ specimen consists of two samples with much deteriorated flowers, which do not allow assessing the length of filament series. However, one of them holds a brief note in Latin handwritten by Martius, with incomplete information about the filaments: “Corona exterior serie simplici constat filorum alborum quae vittis quadratis violaceis picta sunt atque compressa; interior similis crassior”. This indicates the existence of two outer series, the inner one with stronger filaments, but gives no direct information on their relative length. The flower of Spruce 2191, the specimen examined by Masters, has two very unequal series of external filaments, the outermost being reduced, with few short filaments. Thus, the illustration accompanying Masters’ description of Passiflora riparia reflects misinterpretation of the syntypes concerning an important diagnostic trait in the series. However, this anomaly was not corrected in the successive re-descriptions of the species (Killip 1938, Tillett 2003, Cervi 1997). The resulting confusion is responsible for the creation of several species that are closely similar or identical to the syntypes of Passiflora riparia.

Thus, in 1966, Sacco described Passiflora emiliae Sacco in the Boletim do Nacional Museum (Rio de Janeiro), from two Kuhlmann specimens that had been first determined as P. riparia. Indeed, as in Spruce 2191, the Kuhlmann specimens correspond to Masters’ drawing of P. riparia, with the exception of their two unequal outer series of filaments. This is why Sacco (1966) provided a key where P. riparia is differentiated from P. emiliae by two subequal outer series of filaments.

Passiflora crenata Feuillet and Cremers (1984) was described as a French Guiana endemic, very similar to P. riparia. It was differentiated by the white color of its bracts (a trait not precisely described for P. riparia) and by its solitary flowers, in contrast to those of P. riparia, which are clustered on short lateral stems. The original description of P. crenata mentions two outer series of the same length as the sepals, however both the holotype and the illustration provided by Feuillet and Cremers (1984) show two very unequal outer series of filaments, the outermost appearing much shorter and thinner.

In the Flora of Ecuador, Holm-Nielsen et al. (1988: 116) designated Martius 3228 as the lectotype of P. riparia and referred the other syntype, Spruce 2191, to P. ambigua Hemsl. With two glands placed above the middle of petiole (at the apex in the reference vouchers), their description of P. riparia is clearly different from the syntypes and the original description. Moreover, their description indicates a confusion between the innermost series of filaments and the operculum. Indeed, the authors describe “a nectar ring 0.5 cm long, recurved margin minutely sinuate to lobulate” and an “operculum 1 cm, erect, entire, borne just below corona”. In fact, the “nectar ring” corresponds to the operculum as observed in the series Laurifoliae; and the part named “operculum” by Holm-Nielsen et al. is the same innermost series of filaments, oriented towards the androgynophore and covering the entrance of the hypanthium, that exists in P. tolimana Harms, P. kapiriensis Rome and Coppens, and other species of the series (Rome and Coppens d’Eeckenbrugge 2017). In the herbarium specimen Knapp 6242 cited by Holm-Nielsen et al. (1988), these inner filaments appear more or less fused together, forming an erect tubular structure. On the next page, Holm-Nielsen et al. described P. ambigua, including P. emiliae as a synonym of this species. Furthermore, in their determination key, they distinguished P. pergrandis Holm-Nielsen and Lawesson (1987), another species of series Laurifoliae, from P. riparia by the awn of the sepal not exceeding its apex (vs. an awn exceeding the sepal apex). However, this trait is found in Spruce’s syntype of P. riparia (which they assigned to P. ambigua instead of P. riparia).

In 1989, Escobar described Passiflora fernandezii Escobar as a close Bolivian relative of P. riparia: “P. fernandezii most closely ressembles P. riparia […], but differs from it by the pubescence and shape of the hypanthium. Both of these characters are variable in collections ascribed to that species […] so that further study of the Passiflora riparia complex is needed.”

Here, we revise the different treatments of P. riparia and compare them with the descriptions and types of P. emiliae, P. crenata, P. pergrandis, and P. fernandezii, and re-examine the other vouchers associated to them by the different authors of these species. Thus, we can demonstrate that, based on the textual descriptions of P. riparia by Martius and Masters and the observation of both syntypes, Martius 3228 and Spruce 2191, as well as the polymorphism observed on the reference vouchers mentioned in the different treatments, there is no ground for differentiating these five species. We also verify that none of these five taxa can be differentiated by its adaptation, using label information and a multivariate analysis of climates in their respective ranges, while observing that P. ambigua is marginally sympatric with them, less strongly related to riparian habitats, and adapted to a wider range of climates, particularly in tropical highlands. Consequently, we place P. emiliae, P. crenata, P. pergrandis and P. fernandezii in synonymy of P. riparia, and provide a more complete and precise description that takes into account the pan-Amazonian geographical distribution of this species and its attendant polymorphism.

A first comparison confronts the descriptions of P. riparia by Masters (1872), Killip (1938), Cervi (1997), Tillett (2003), Holm-Nielsen et al. (1988), the original description of P. emiliae by Sacco (1966), that of P. crenata by Feuillet and Cremers (1984), that of P. pergrandis by Holm-Nielsen and Lawesson (1987), that of P. fernandezii by Escobar (1989). This comparison is focused on traits relative to stems, stipules, petiole, leaves, inflorescences, bracts, hypanthium, sepals, petals, corona (outer and inner series of filaments), operculum, androgynophore, ovary, and fruit. It has been extended to the description of the very similar P. ambigua by Hemsley (1902).

A second comparison bears on all available vouchers (43 specimens; see Table 2) associated with the descriptions of any of the first five species by these same authors, focusing on the following traits: petiolar gland position, presence of pseudoracemes, plant pubescence, bract size, color of bracts and perianth, when discernible or mentioned on the voucher label. We also include our own field observations of P. crenata in French Guiana, on living material collected from localities indicated in the original description.

Other 94 herbarium specimens have been observed and determined, for a comparative study of the habitat of the same taxa. Some sheets were studied from scans provided by the host herbaria. The complete list, comprising 137 materials initially classified under P. riparia or one of its four presumed synonyms or under P. ambigua, is listed in the appendix. For the distribution study, only fertile samples allowing unambiguous determination were retained, as well as other good quality reports (including photographs); the P. crenata sample from French Guiana was reduced to 11 specimens, to avoid an excessive weight for this taxon. When label information allowed, geographic coordinates were assigned to the voucher using Google Earth and gazetteers, the collecting sites were mapped and a distribution model was developed, using the MAXENT software and 19 bioclimatic variable layers from the Worldclim database at a 2'30" grid resolution (corresponding roughly to 4.4 × 4.6 km at Equator; http://www.worldclim.org/current) (Hijmans et al. 2005). MAXENT identifies potential distribution areas based on their similarity in climatic conditions compared to those at the sites where the species has already been observed, hence modeling where conditions are suitable for their development. It infers the probability distribution of maximum entropy (i.e., closest to uniform) subject to the constraint that the expected value of each environmental variable (or its transform and/or interactions) under this estimated distribution matches its empirical average (Phillips et al. 2006). A logistic threshold value equivalent to the 10^th percentile training presence was retained to separate climatically favorable areas from marginally fit areas. Thresholds of 33 and 67% training presence were used to discriminate “very good” and “excellent” climates.

Furthermore, 19 bioclimatic variables corresponding to the collection sites were extracted from the Worldclim database (Hijmans et al. 2005), and those variables that most contributed to the model were submitted to a Principal Component Analysis (PCA) for a comparative analysis of climatic adaptation.

Most of the confusion in the definition of P. riparia has arisen from the incomplete description by Masters (1872), in the very difficult context of the highly uniform series Laurifoliae, and from the mention of two equal series of outer corona filaments by Killip (1938), which was reproduced in the following descriptions of Cervi (1997) and Tillett (2003) (although the latter mentioned the occasional loss of the outermost series). This ‘two equal outer series’ problem originated from the ambiguous iconography accompanying the description of Masters (1872), which is inconsistent with the syntype Spruce 2191, and all other interpretable specimens mentioned in the literature. Indeed, our study of the specimens examined by Killip (1938) and Cervi (1997) shows that, whenever the relative length of outer corona filaments is observable, i.e. in 12 of them, their outer series is reduced, or even absent in nine of them. This clearly indicates that Killip (1938) and Cervi (1997) had not observed themselves the combination of two equal series of outer filaments with multiple series of very short inner filaments and other typical traits as white to red bracts and white corollas.

In 1966, Sacco realized the inconsistency between Killip’s description of P. riparia and two Kuhlmann specimens of this species where he could observe the reduction of the outermost series of filaments. But he used them as types for the description of a new species, P. emiliae, logically differentiated from P. riparia by the outer corona structure. It seems that this new species description remained confidential and only attracted the attention of Cervi, who reclassified a few other exemplars from P. riparia to P. emiliae, such as Ducke 21311, Kuhlmann sn (R136313). Later, in 1997, he changed his mind and followed Holm-Nielsen et al. (1988) in considering it a synonym of P. ambigua. Table 1 shows that the description of P. emiliae only differs from those of P. riparia by the pubescence of the flower; however, this trait is also found in many examined specimens of P. riparia, including the syntypes (Table 2), as well as in the other presumed synonyms, P. crenata, P. fernandezii and P. pergrandis.

Originally, P. crenata was described as an endemic of French Guiana, distinguished from P. riparia by its white bracts and the type of inflorescence. However, bract color at anthesis varies between white and deep red in populations from French Guiana and from Brazil (Figure 1, Table 2) and the type specimens of P. emiliae also present white bracts. In the examined specimens from Ecuador and Peru, bract color varied from red to purple and violet. The presence of single axillary flowers or of pseudoracemes is also a variable trait, even at the intra-individual level, as noted in the descriptions of P. riparia by Tillett (2003), P. emiliae by Sacco (1966), and observed by ourselves in P. crenata. Furthermore, other species of series Laurifoliae present both solitary flowers and pseudoracemes, as P. ambigua, P. popenovii Killip, P. gabrielliana Vanderpl., P. capparidifolia Killip, P. killipiana Cuatrec., P. phellos Feuillet, P. laurifolia. Thus, no traits allow distinguishing P. crenata from P. riparia and P. emiliae.

The two more recent descriptions of P. pergrandis and P. fernandezii did not present new variations in any of the characters discussed above. Finally, we can conclude that the differences in the descriptions of P. riparia, P. emiliae, P. crenata, P. pergrandis and P. fernandezii are either related to the confusion introduced by the iconography associated with the original description or with the imprecisions related to the initial observation of two dry specimens. More details and variation have logically been documented after the examination of specimens from a wider geographic range, as well as the direct observations on living materials. The variations observed among specimens of P. riparia encompass the variations between this taxon on one hand and P. crenata, P. emiliae, P. fernandezii or P. pergrandis on the other hand. Furthermore, most specimens from the latter taxa have been collected under very similar habitats, dominated by lowland tropical climates in riparian habitats (Table 3, Figure 3), further justifying their placement as synonyms of P. riparia. Only the description of P. riparia in the Flora of Ecuador is clearly different, as it includes the mention of a lilac corolla, two extremely long series of outer filaments and the presence of an inner series of 1 cm erect filaments, a trait combination indicating that Holm-Nielsen et al. (1988) described P. tolimana instead of P. riparia.

Passiflora ambigua appears very similar to P. riparia, mostly differing in the color of bracts (green) and the corolla (lilac), but its distribution and ecology are different, with a much lesser frequency of riparian habitats and a capacity to thrive at much higher elevations, up to hillsides in the Andes of tropical South America and Central America.

Gathering P. riparia, P. crenata, P. emiliae, P. fernandezii and P. pergrandis into a single species imposes a new description, taking into account the reduction of the outermost series of corona filaments and variation observed in other traits, on our whole sample. In the following treatment, the lectotype and isotype chosen by Lawesson are logically retained, however, as its damaged flowers do not allow the observation of the corona structure, we choose as epitype the syntype Spruce 2191, i.e., the much better-preserved specimen that was observed by Masters for the original description.

Following our morphological and ecological analyses, P. emiliae, P. pergrandis, P. fernandezii and P. crenata are placed as synonyms of P. riparia, which reduces the current number of species belonging to series Laurifoliae to 21. Like Passiflora nitida, P. laurifolia or P. ambigua, P. riparia is a new example of a very widely distributed species in this series. In fact, its variability appears relatively limited in the context of its wide distribution.

The description of P. riparia in the Flora of Ecuador corresponds to P. tolimana; Holm-Nielsen 1040 and Knapp 6242 are the only known specimens from Ecuador for this species, hitherto considered endemic to Colombia. The determination of Spruce 2191 under P. ambigua by Lawesson, endorsed by Holm-Nielsen et al. (1988), cannot be accepted, given that this specimen has white petals and sepals (vs. red to purple in P. ambigua).

This research has been carried out with support from the Herbarium of Pontificia Universidad Católica del Ecuador and the Ecuadorian Ministry of the Environment. We thank especially Daniela Cevallos and Katya Romoleroux for their help during our expedition in Ecuador. The first author also thanks the Missouri Botanical Garden, Jim Solomon, and John MacDougal for their help during the week he spent in the herbarium at St. Louis, Mathew Rees (Kew Garden) for his help during Panama and Ecuador trips, Jean-Jacques de Granville for the exceptional welcome in French Guiana, Rich Hoyer (Birdernaturalist) and Luiz Otavio Adão Teixeira for their stunning pictures of P. riparia, and the different herbaria with which we have collaborated for this work: Field Museum herbarium, Paris herbarium, Lyon botanical garden (Frédéric Danet), Cayenne herbarium, New York Botanical Garden, Kew Garden, Munich herbarium (Hajo Esser). Finally, we thank Ana Carolina Mezzonato-Pires and John MacDougal for their comments and corrections to improve the manuscript.

Examined specimens

P. ambigua

BRAZIL. Amazonas: Esperança, mouth of Javari River, non-flood forest, 4-2-1942, Ducke 878 (MO). COLOMBIA. Antioquia: San Luis, 26 Jul. 1981, Hoyos 132 (JAUM). Municipality of Frontino, Las Orquídeas National Park, 31 Jan. 1995, Pipoly 18177 (JAUM). Municipality of Turbo, road of Tapón del Darién, 28 Feb. 1984, Brand 947 (JAUM). Chocó: area of Baudó, 4 Feb. 1967, Fuchs 21744 (COL). Huila: Municipality of Gigante, farm of Adonai Moreno, 20 Dec. 1976, Escobar 435 (MO). Meta: Río Bravo village, La Cristalina stream, El Pital, 4 Mar. 1986, Devia 1121 (MO). COSTA RICA. Puntarenas: Osa Península, 6 Jan. 1994, Aguilar 2974 (CR). Cartago: Orosi, Navarro del Muñeco, 12 Apr. 1998, Blanco 805 (USJ). San José: Vicinity of El General, Jan. 1939, Skutch 3037 (US). ECUADOR. Esmeraldas: El Timbre, near Esmeraldas, 6 Aug. 1962, Jativa 439 (US). Napo: Road Hollín-Loreto-Coca, Chaluayacu Community, 23 Dec. 1988, Cerón 5772 (MO). Pichincha: Alluriquin, 19 Oct. 1921, Werling 458 (QCA). Zamora-Chinchipe: Municipality of Zamora, Road Zamora-Loja, 29 Oct. 1991, Palacios 8811 (MO). Palanda, Río Vergel Valley, 14 Nov. 2006, Werff 22075 (MO). GUATEMALA. Secanquim, 14 May 1914, Cook 79 (US). HONDURAS. Machaca, 8 Feb. 1934, Schipp 1302 (F). MEXICO. Veracruz: Biological station of Los Tuxtlas, 7 Apr. 1971, Calzada 230 (MEXU). Municipality of San Andrés Tuxtla, south of Ebitrolotu, 30 APr. 1973, Villegas Herrera 112 (MEXU). PANAMA. Colón: Santa Rita Ridge, east of transisthmian highway, 16 Dec. 1972, Gentry 6561 (MO). Panamá: just before la Eneida along new trail which begins exactly beside López House, 8 March 1968, Correa 825 (PMA). El Llano-Carti Road, 18 Apr. 1981, Sytsma 4027 (MO). PERU. Loreto: Maynas, near Iquitos, in 1996, Grandez 7837 (MO).

P. laurifolia

BRAZIL. Bahia: Barra, Spruce 1394 (P). Pará: Santarem, 19 Jan. 1934, Swallen 3309 (US). COLOMBIA. Vaupés: Vicinity of Mitú, Rio Vaupés, 22 Feb. 1945, Allen 3340 (US). GUYANA. Western extremity of Kanuku Mountains, in drainage of Takutu River, 4 March 1938, Smith 3157 (U).

P. phellos

BRAZIL. Amazonas: Uanadona, near mouth of Rio Dimiti, 10 May 1948, Schultes 9900 (K). VENEZUELA. Río Pacimini, Feb. 1854, Spruce 3390 (K).

P. riparia

BOLIVIA. La Paz: Prov. Iturralde, Siete Cielos, Rio Manupare, 4 June 1987, Solomon 16923 (MO). BRAZIL. Acre: Municipality of Plácido de Castro, Rio Abunã, 10 Jan. 1995, Figueiredo 506 (NY). Amapá: National park of Tumucumaque, Rio Jari, 21 Feb. 2013, Hopkins 2284 (INPA). Municipality of Monção, basin of the Rio Turiaçu, Ka’apor Indian Reserve, 12 Feb. 1985, Balée 794 (MO), Balée 810 (MO). Amazonas: Presidente Figueredo, Nazaré locality, Rio Uatumã between Rio Pitinga and Rio Uatumã, 18 March 1986, Ferreira 6808 (NY). Amazonas: Mouth of Rio Madeira, Martius 3228 (M). São Gabriel da Cachoeira near the Rio Negro, Apr. 1852, Spruce 2191 (P). Pará: Ruropolis Presidente Medici, 9 Feb. 1976, Bamps 5341 (K). Municipality of Almeirim, Mte. Dourado, road Mte Dourado-Munguba, 10 Feb. 1986, Pires 784 (INPA), Mt. Dourado, 20 June 1988, Pires 2227 (NY). Banks of Rio São Manoel, 14 Jan. 1952, Pires 3931. Belem, 29 Dec. 1922, Ducke 17338 (US). Rio Xingu, Island of Piracui, 22 Oct. 1986, Souza 449. Goias: ca. 2 km S. of Guará, 19 March 1968, Irwin 21451 (NY). Mato Grosso: Rio Arinos, Dec. 1914, Kuhlmann 1066 (R), Kuhlmann 1064 (R). Novo Mundo, Forest Reserve of Rio Cristalino, 11 Feb. 2008, Zappi 1194, margin of Rio Cristalino, 29 Nov. 1996, Dubs 2335 (K). Alta floresta, private property of environmental preservation, 10 Nov. 2006, Sasaki 1087 (K), 13 Dec. 2006, Sasaki 1199 (INPA), 23 Jan. 2007, Sasaki 1390 (INPA). COLOMBIA. Guainía: confluence of Ríos Guaviare, Atabapo and Inirida, 23 Aug. 1975, García-Barriga 20929 (COL). ECUADOR. Sucumbios: Cuyabena faunistic reserve, Rio Cuyabeno from outlet of Laguna Grande and 5 km upstream, 2 Apr. 1989, Balslev 84715 (QCA). Orellana: National park Yasuní, south of Río Napo, 31 Jan. 1998, Burnham 1628 (QCA), Río Tivacuno, 0.5 km upstream of confluence with Río Tiputini, 24 March 1998, Burnham 1690 (QCNE). National Park Yasuní, 16 Feb. 2002, Pérez 392 (QCA). Zamora-Chinchipe: Road Zamora-Gualaquiza, ca 5 km north of Cumbaraza, 20 Apr. 1974, Harling 13771 (GB). Nangaritza, Río Tzenganga, 6 June 2005, Quizhpe 1201 (MO). Hill about 2 km downstream from Campamento Shaime along Río Nangaritza, 15 Feb. 1994, Werff 13084 (QCNE). FRENCH GUIANA. Road to Montsinery, 7 March 2017, Rome 567 (P). Road between Cayenne and Cacao, 25 Feb. 2017, Rome 559 (P). Road to Régina, 17 May 2008, Rome 118 (LYJB), Rome 119 (LYJB), Rome 120 (LYJB), Rome 121 (LYJB), Rome 122 (LYJB). Road between Régina and St Georges, 10 Apr. 2008, Rome 45 (LYJB), 19 May 2008, Rome 129 (LYJB), Rome 136 (LYJB), Rome 138 (LYJB), Rome 142 (LYJB). Road of Kaw Mountain, 24 Nov. 2009, Rome 201 (LYJB), Rome 202 (LYJB), 29 Jan. 2013, Rome 400 (LYJB). Road to Tonnegrande, 30 Jan. 2013, Rome 405 (LYJB), Rome 406 (LYJB), Rome 407 (LYJB), Rome 408 (LYJB), Rome 409 (LYJB). Kotika mount, 21 Feb. 2005, Granville (de) 16896 (CAY). Municipality of Saint-Georges, Oyapock basin, 15 Aug. 1997, Berton 250 (CAY). Kamuyene Kamuyene, 12 Apr. 2005, Bourdy 3128 (CAY). Road of Kaw mountain, 24 Jan. 1983, Feuillet 573 (CAY), 1 Jan. 1986, Feuillet 2975 (CAY), 25 Aug. 2009, Feuillet 17078 (CAY). Path between Roura and the Gabrielle Creek, 21 Apr. 1979, Prevost 563 (CAY). Road to Régina, near Camp Hervo, 14 March 2009, Vanderplanck 1605 (CAY). PERU. Huánuco: Prov. Pachitea, region of Pucallpa, western part of the “Sira montains” and adjacent lowland, 31 July 1988, Wallnöfer 12-31788 (MO). Along Río Pachitea, 12 July 1967, Schunke 2112 (US). Junín: Chanchamayo prov., El Bocaz, 20 Nov. 1982, Vargas 64 (USM). Loreto: in lower forest at Neshuya, 2 Oct. 1965, Schunke 907 (US). Madre de Dios: National Park Manu, Río Manu, 1 Oct. 1986, Foster 11539 (F), Foster 11558 (F). Tambopata prov., Las Piedras, Cusco Amazónico, 7 Oct. 1991, Timaná 2457 (MO). San Martín: Mariscal Cáceres, mouth of Río Tocache, 6 Nov. 1969, Schunke 3579 (US). San Martín, Boca Toma del Shicayo, N of Tarapoto along Río Shilcayo, 27 May 1986, Alcorn 10 (MO). Juan Jui, Alto Río Huallaga, Oct. 1934, Klug 3897 (MO). Chazuta, Río Huallaga, March 1935, Klug 4037 (K). Mariscal Cáceres, Canyon of Huaquisha, right margin of Río Huallaga, 30 June 1974, Schunke 7083 (MO). Amazonas: between Uchpayaco and Rimachi, banks of Río Pastaza, 30 July 1979, Díaz 1283 (MO). Pasco: Oxapampa, Puerto Bermúdez, 14 July 1929, Killip 26683 (US). Oxapampa, ca. 1 km from division of Villa Rica-Pto. Bermúdez road and Villa Rica-Palcazu road, on Palcazu branch, 15 Aug. 1984, Knapp 6633 (MO). Loreto: Maynas, Iquitos, July 1929, Williams 1440 (US), March 1930, Williams 7996 (US). Yurimaguas, lower Río Huallaga, March 1930, Williams 7876 (US). Maynas, district of Iquitos, Caserío, 25 Feb. 1976, Revilla 241 (MO), Río Nanay near Astoria, 15 March 1973, Rimachi 126 (USM). Tarapoto, Nov. 1902, Ule 6547 (K). Ucayali: Pucapanga, margin of Río Ucayali, 20 Aug. 1965, Sagástegui 5739 (HUT). SURINAME. Sipaliwini, vicinity of airstrip along Ulemari river, 1 May 1998, Evans 2971 (MO).

P. sp. (impossible or doubtful identification)

BRAZIL. Amazonas: Lake of Aleixo, in riparian forest, 3 Apr. 1932, Ducke 24044 (US). Bahia, Barra, March 1851, Spruce 1172 (P). ECUADOR. Morona Santiago: Shuar Pampants center, 10 Sept. 1986, Warush RBAE86 (QCNE). PERU. Amazonas: Río Cenepa, 2 June 1973, Ancuash 506 (US). Junín: Junín, Puerto Yesup, 10 July 1929, Killip 26307 (US). Loreto: Alto Amazonas, Yurimaguas, lower Río Huallaga, 22 Aug. 1929, Killip 28214 (US). Near Iquitos, July 1929, Williams 1392 (US). La Victoria on the Amazon River, Aug. 1929, Williams 3126 (US). Santa Rosa, Lower Río Huallaga below Yurimaguas, 1 sept. 1929, Killip 28940 (US). Wooded banks of lower Río Huallaga, 5 Sept. 1929, Killip 29012 (US). San Martín: Alto Río Huallaga, Dec. 1929, Williams 6300 (US). Tarapoto, Alto Río Huallaga, Dec. 1929, Williams 5848 (F).

P. tolimana

ECUADOR. Napo: 2 km W of Archidonia, 27 June 1968, Holm-Nielsen 1040 (AAU). Morona-Santiago: ca. 32.5 km S. of Gualaquiza on road to Zamora, 4 Feb. 1984, Knapp 6242 (US).

P. venusta

PERU. San Martín: Lamas, Dec. 1929, Williams 6378 (US). Tarapoto, Alto Río Huallaga, Dec. 1929, Williams 5637 (F).

Index to Numbered Collections

Aguilar, R. 2974 (ambigua)

Alcorn, P. 10 (riparia)

Allen, P. H. 3340 (laurifolia)

Ancuash, E. 506 (sp.)

Balée, W. L. 794, 810 (riparia)

Balslev, H. 84715 (riparia)

Bamps, P. 5341 (riparia)

Berton, M. E. 250 (riparia)

Blanco, M. 805 (ambigua)

Bourdy, G. 3128 (riparia)

Brand, J. 947 (ambigua)

Burnham, R. J. 1628, 1690 (riparia)

Calzada, J. I. 230 (ambigua)

Cerón, C. E. 5772 (ambigua)

Cook, O. F. 79 (ambigua)

Correa, M. D. 825 (ambigua)

Devia, W. 1121 (ambigua)

Díaz, C. 1283 (riparia)

Ducke, A. 17338 (P. riparia), 24044 ( sp.), 878 (ambigua)

Dubs, B. 2335 (riparia)

Escobar, L. 435 (ambigua)

Evans, R. 2971 (riparia)

Feuillet, C. 573, 2975, 17078 (riparia)

Ferreira, C. A. C. 6808 (riparia)

Figueiredo, C. 506 (riparia)

Foster, R. B. 11539 (riparia)

Fuchs, H. P. 21744 (ambigua)

García-Barriga, H. 20929 (riparia)

Gentry, A. 6561 (ambigua)

Grandez, C. 7837 (ambigua)

Granville (de), J. J. 16896 (riparia)

Harling, G. 13771 (riparia)

Holm-Nielsen, L. 1040 (tolimana)

Hopkins, M. J. G. 2284 (riparia)

Hoyos, S 132 (ambigua)

Irwin, H. S. 21451 (riparia)

Jativa, C. D. 439 (ambigua)

Killip, E. P. 24044, 26307, 28214, 28940, 29012 ( sp.), 26683 (riparia)

Klug, G. 3897, 4037 (riparia)

Knapp, S. 6633 (P. riparia), 6242 (tolimana)

Kuhlmann, J. G. 1066, 1064 (riparia)

Martius, C. F. P. 3228 (riparia)

Palacios, W. 8811 (ambigua)

Pérez, A. J. 392 (riparia)

Pipoly, J. 18177 (ambigua)

Pires, M. J. P. 784, 2227, 3931 (riparia)

Prevost, M. F. 563 (riparia)

Quizhpe, W. 1201 (riparia)

Revilla, J. 241 (riparia)

Rimachi, M. 126 (riparia)

Rome, M. 567, 559, 118, 119, 120, 121, 122, 45, 129, 136, 138, 142, 405, 406, 407, 408, 409 (riparia)

Sagástegui, A. 5739 (riparia)

Schunke, J. V. 907, 2112, 3555, 3579, 7083 (riparia)

Sasaki, D. 1087, 1199, 1390 (riparia)

Schipp, W. A. 1302 (ambigua)

Schultes, R. E. 9900 (phellos)

Skutch, A. F. 4037 (ambigua)

Smith, A. C. 3157 (laurifolia)

Solomon, J. C. 16923 (riparia)

Souza (de), S. A. M. 449 (riparia)

Spruce, R. 1172 (sp.), 1394 (laurifolia), 3390 (phellos), 2191 (riparia)

Swallen, J. R. 3309 (laurifolia)

Sytsma, K. J. 4027 (ambigua)

Timaná, M. 2457 (riparia)

Ule, E. 6547 (riparia)

Vanderplanck, R. J. R. 1605 (riparia)

Vargas, M. A. 64 (riparia)

Villegas Herrera, A. 112 (ambigua)

Wallnöfer, B. 12-31788 (riparia)

Warush, A. 86 (sp.)

Werff (van der), H. 13084 (riparia), 22075 (ambigua)

Werling, L. 458 (ambigua)

Williams, L. 1392, 3126, 5848, 6300 (sp.), 5637, 6378 (venusta), 1440, 7876, 7996 (riparia)

Zappi, D. 1194 (riparia)