Towards a monophyletic Licania: a new generic classification of the polyphyletic Neotropical genus Licania (Chrysobalanaceae)

Summary

Licania, currently the most species-rich genus of Chrysobalanaceae, is confined to the Neotropics, and is highly polyphyletic in molecular phylogenetic analyses. These studies show that the three subgenera of Licania, as well as sections within them, are not supported as monophyletic. Here we re-delimit the genus using a molecular phylogenetic analysis as a framework. We recognise the following eight genera: core Licania, Moquilea, Leptobalanus, Hymenopus, Microdesmia, Parinariopsis, Geobalanus; and Cordillera gen. nov. is described to accommodate Licania platycalyx. New combinations, as well as a description of each genus and a key to the species are provided, in addition to a key for all the Neotropical genera of the family, including those presented here. Chrysobalanaceae now comprise sixteen Neotropical genera and the redefined Licania, with c. 100 species, is then the second largest genus in the family after Hirtella (107 species).

Introduction

Licania Aubl. (Aublet 1775), as currently delimited, with over 214 species endemic to the Neotropics (Prance & Sothers 2003a; Sothers et al. 2014), is the largest genus of Chrysobalanaceae. Its distribution spans the southern USA, Mexico, Central America, the Caribbean and tropical South America. Although at the alpha-taxonomic level the genus, and family as a whole is considered well known, recent molecular studies have highlighted the polyphyly of Licania (Bardon et al. 2016; Sothers 2010; Sothers et al. 2014). Prance & White (1988) divided Licania into five subgenera, Afrolicania, Angelesia, Licania, Moquilea and Parinariopsis. Recently the African and Malesian subgenera, Afrolicania and Angelesia, respectively, have been re-instated at generic rank (Prance & Sothers 2003a; Sothers & Prance 2014, respectively), rendering Licania entirely Neotropical. This paper addresses the polyphyly of the remaining three subgenera.

Licania and its subgenera have a complex taxonomic history. Ever since Aublet (1775) described Licania and Moquilea, among other Neotropical genera of Chrysobalanaceae such as Acioa and Couepia, in his Histoire des Plantes de la Guyane, taxonomists have disagreed over their circumscription. Aublet (1775) distinguished Licania from Moquilea by the length of the stamens in relation to the corolla: exserted in Moquilea vs included in Licania. He placed Couepia near Moquilea, presumably based on the exserted stamens and their number, despite differences in the position of the ovary on the receptacle (basal in Moquilea and Licania vs at the mouth in Couepia and Acioa). After his death in 1778, Aublet’s herbarium was dispersed and as a result, during the 19th century, confusion prevailed over the delimitation of Licania and Moquilea. Subsequent taxonomists either accepted Licania and Moquilea as separate genera (Martius & Zuccarini 1827; Zuccarini 1832; Meisner 1837a & b; Bentham 1840; Endlicher 1840, 1842; Hooker 1865, 1867; Fritsch 1888), or considered Moquilea as a synonym of Licania (Grisebach 1857; Baillon 1869; Eichler 1878; Fritsch 1889); or treated Couepia as a synonym of Moquilea (Martius & Zuccarini 1827; Walpers 1843; Mueller 1857). Others accepted all four genera cited above (Jussieu 1789; Brown 1818; Candolle 1825; Lindley 1836; Fritsch 1888). Grisebach (1857) was the first to merge Moquilea into Licania.

Bentham (1840) accepted Moquilea, Licania and Couepia as distinct genera (he did not see Acioa and therefore did not consider it). He delimited Licania as displaying the following characters: 4 – 5-dentate or shortly pentafid calyx, apetalous or with 4 – 5 petals, fewer than 15 fertile stamens, and ovary at the base of the receptacle; he divided the genus into five sections: Batheogyne, Leptobalanus, Microdesmia, Eulicania, and Hymenopus. Bentham defined Moquilea as having a pentafid calyx, with petals present, more than 30 fertile stamens, and ovary at the base of the receptacle, characters which overlapped with those of Licania. Hooker (1867) accepted both Licania and Moquilea as genera but redefined Bentham’s sections of Licania: Licania with two sections (Eulicania and Hymenopus) and Moquilea with three (Eumoquilea, Leptobalanus and Microdesmia). Thus, both Licania and Moquilea included apetalous and petalous taxa. Fritsch (1888) recognised the four genera of Aublet (Acioa, Couepia, Licania and Moquilea) but later merged Moquilea with Licania (Fritsch 1889), following Grisebach (1857). This concept of Moquilea as a synonym of Licania was followed in most subsequent classifications (Hallier 1903; Prance 1963, 1972a, 1989; Prance & White 1988; Prance & Sothers 2003a) and in the 20th century only Hutchinson (1964) treated Licania and Moquilea as separate genera.

When Prance & White (1988) divided Licania into five subgenera, Licania, Moquilea, Parinariopsis, Afrolicania and Angelesia, this was based largely on characters of the ovary (unilocular and at or near the base of the receptacle) and other characters such as stamen exsertion; subgenus Licania was subdivided into five sections (sects. Hirsuta, Hymenopus, Cymosa, Pulverulenta, Licania) and subgenus Moquilea into three (sects. Moquilea, Leptobalanus, Microdesmia). Molecular phylogenetic analyses retrieved species in Licania subgen. Moquilea as sister to core Couepia (Bardon et al. 2013; Sothers 2010; Sothers et al. 2014) and further indicated that the sections of subgen. Licania and subgen. Moquilea were not supported as monophyletic (Sothers unpublished data; Bardon et al. 2013). These results paved the way for a reassessment of the monophyly of Licania.

Here we propose a new generic classification for Licania s.l. based on molecular phylogenetic evidence supported by morphological characters. We divide the genus into eight genera, five of the eight sections in subgen. Licania and subgen. Moquilea are recognised as genera: Moquilea, Leptobalanus, Microdesmia, Hymenopus and Licania; the taxa in subgen. Licania sect. Hirsuta are incorporated into genus Hymenopus, and subgen. Licania sects. Cymosa and Pulverulenta remain in core Licania and are not recognised as sections; subgen. Parinariopsis is upgraded to generic rank, Geobalanus is re-instated at generic rank from the synonymy of subgen. Moquilea to accommodate three species previously in sect. Moquilea (L. michauxii, L. riverae and L. retifolia). A new genus, Cordillera, is described to accommodate Licania platycalyx, which was previously transferred to Licania from Couepia (Sothers et al. 2014: 193).

Materials and Methods

Methodology for DNA extraction, sequencing and matrix alignment, and phylogenetic analyses follows that of Sothers et al. (2014). We include the taxa and markers of Sothers (2010) (85 taxa, 4524 bp, two plastid and two nuclear markers) with additional taxa and one additional marker added since that study (184 taxa, 5348 bp, 3 plastid and 2 nuclear). Results for the phylogenetic analyses are presented in Fig. 1 (combined strict consensus tree with bootstrap support); Appendix 1 includes the taxa used for the molecular phylogenetic analyses presented here, and the markers used; the ITS and Xdh datasets are missing several taxa.

Fig. 1

Strict consensus tree of the combined analysis (plastid, ITS and Xdh), highlighting clades with taxa of Licania s.l. and the new generic names; bootstrap support is shown above branches. Clades of all other Neotropical genera and the outgroup are also cited.

Results

Fig. 1 is a strict consensus tree of the combined dataset of the five markers, three plastid (matK, ndhF and rbcL) and two nuclear (ITS rDNA and Xdh). The type species for seven of the eight sections of Licania subgen. Licania and subgen. Moquilea have been included in the analysis presented here (Table 1). Further comments on the recognition and relationships among the genera are included under each genus in the section on taxonomy.

Table 1. Taxonomic representation of the progression of changes from the previous classification of Licania s.l., sensu Prance & White (1988) and Prance & Sothers (2003a), and the current classification presented here. Species in bold have been sequenced and are included in Fig. 1.

Species of Licania s.l. were retrieved in four major clades: 1) a large clade composed of species in subgen. Licania sect. Licania, sect. Cymosa and sect. Pulverulenta (= genus Licania s.s.), with moderate support and sister to Hirtella; 2) a large clade, which includes Gaulettia, sister to L. arborea (= genus Microdesmia) and L. platycalyx (= genus Cordillera) and with species from subgen. Licania sect. Hymenopus retrieved in two sub-clades (= genus Hymenopus): one also including L. latifolia (previously in subgen. Licania sect. Hirsuta) and L. amapaensis (previously subgen. Moquilea sect. Microdesmia), sister to Afrolicania, and the other sub-clade including the type of sect. Hymenopus (L. divaricata) sister to L. licaniiflora (= genus Parinariopsis), with moderate support; 3) a highly supported clade of species from subgen. Moquilea sect. Leptobalanus (= genus Leptobalanus) sister to Gaulettia; and 4) species in subgen. Moquilea sect. Moquilea retrieved in two lineages sister to Couepia (= genus Moquilea).

Licania michauxii and L. riverae (= genus Geobalanus), previously of subgen. Moquilea sect. Moquilea, are retrieved in a fifth clade together with the Paleotropical genera Grangeria and Parastemon, distantly related to Licania s.l. but without support.

Analyses for the ITS, Xdh and plastid datasets displayed similar results to the combined analysis presented here. Our results are also broadly supported by other phylogenetic analyses (Bardon et al. 2016; Sothers et al. 2014; Sothers 2010).

Discussion

This study presents a new classification for Licania s.l. given its rampant polyphyly as evidenced in phylogenetic studies involving the Chrysobalanaceae (Bardon et al. 2013; Sothers 2010; Sothers et al. 2014; Yakandawala et al. 2010). Moreover, morphological characters that were previously held as central to the taxonomy of subgenera and sections, do not support the previous classification. Table 1 outlines the taxonomic changes made here as compared with the previously accepted delimitation of Licania s.l. and Table 2 summarises the key morphological characters for the genera treated here.

Table 2. Selected key morphological characters of genera previously in Licania s.l. treated here.

The unexpected revelation of a close relationship between Couepia and Licania subgen. Moquilea uncovered by molecular phylogenetic studies (Sothers 2010; Sothers et al. 2014), highlighted the need to re-evaluate the monophyly of Licania s.l. Within Chrysobalanaceae, genera have often been defined by suites of overlapping characters. Among these characters, however, the position of the ovary on the receptacle has been emphasised as pre-eminent and is generally constant within genera. Couepia and Licania were separated mainly because of the differing position of the ovary, despite sharing other characters. In Licania s.l. the ovary was typically at, or near, the base of the receptacle, except in Licania subgen. Parinariopsis (= genus Parinariopsis) and Licania platycalyx (= genus Cordillera), which were unique within Licania s.l. because of the ovary positioned laterally rather than at the base of the receptacle. Outside Licania s.l., only in Parinari and Hirtella does the ovary occur in two different positions within the same genus, on the wall or at the mouth of the receptacle. Despite this, the pantropical genus Parinari is highly supported in phylogenetic analyses, as are the Neotropical Hirtella.

The redelimitation of the former subgen. Licania and subgen. Moquilea into six segregate genera follows a major split among taxa with petals (genera Moquilea, Hymenopus, Microdesmia, Geobalanus) and apetalous taxa (genera Leptobalanus and Licania). In previous classifications (Prance & White 1988; Prance & Sothers 2003a), both subgen. Moquilea and subgen. Licania included sections with and without petals. Thus, subgen. Moquilea comprised sects Moquilea, Microdesmia (with petals) and Leptobalanus (apetalous); subgen. Licania consisted of apetalous sects. Licania, Cymosa and Pulverulenta, sect. Hirsuta with petals, and sect. Hymenopus with both apetalous and petalous taxa (Table 1). Apetaly is present in only one other genus, the monospecific Afrolicania, endemic to and widespread in West and western Central Africa; its presence in the large clade of Neotropical taxa of Chrysobalanaceae (c. 400 spp.) means that this clade is not exclusively Neotropical as stated by Sothers et al. (2014). Our new classification indicates that apetaly has arisen more than once in Chrysobalanaceae.

Other characters that had been used to separate species into subgenera and sections of Licania s.l. include number of stamens, their length relative to the calyx (far-exserted vs equal or included), presence of stomatal cavities on the lower surface of the leaf and leaf pilosity. Generally, the delimitation of sections as defined by Prance & White (1988) and Prance & Sothers (2003a) are retrieved as well-supported clades in the phylogeny presented here, but not as sections of Licania s.l. Thus, the generic status conferred on Moquilea and Licania by Aublet (1775), as well as Bentham’s (1840) suggestion that his sections of Licania (e.g. Leptobalanus, Microdesmia, Hymenopus) were worthy of generic status, hold true.

We took a conservative approach with respect to accepting genera, particularly where the placement of species is not well-supported by either morphology or phylogenetic results. This is the case for Hymenopus as defined here, for which two separate (but closely placed) clades are retrieved, but no morphological characters have been found to circumscribe them as separate genera. Our data do not refute or support species of Hymenopus as a single genus. Similarly in Moquilea, two species (M. egleri and M. minutiflora) are retrieved in a separate clade, but we keep them within the concept of Moquilea for now. Further changes may be needed if these now separate clades do not form a single clade with more morphological and molecular studies, but at present giving these distinct generic recognition seems unwarranted. Other genera that are here weakly supported in the phylogenetic analyses are Parinariopsis, Cordillera, Microdesmia and Geobalanus, but delimitation of these genera is well-supported by morphological and geographic evidence.

In the molecular phylogeny presented here, the segregate genera of Licania s.l. are variously allied to Couepia, Gaulettia and Hirtella. Chrysobalanus is distantly related to these genera even though there have been suggestions of merging it with Licania (Gómez de la Maza 1887; Morales 1887; Chappill 1992). Relationships among Geobalanus, Microdesmia and Moquilea remain unclear but they appear to be phylogenetically distinct. Prance (pers. obs.) regarded sect. Microdesmia (of subgen. Moquilea) as artificial; and a much reduced Microdesmia is maintained as a genus here, albeit further sampling is needed. The two species display a unique geographic disjunction and suggest ecological niche adaptation. Geobalanus displays few morphological characters to distinguish it from Moquilea, Microdesmia and Chrysobalanus, although Microdesmia has deeply reticulate leaves often with stomatal cavities, characters that are lacking in Geobalanus, Moquilea and Chrysobalanus.

Despite the generally well-supported molecular results for Licania s.l. and its segregates, the morphological characters that distinguish the eight genera discussed here are comparatively few and not well studied. Characters that merit further investigation include leaf pilosity, presence of stomatal cavities, inflorescence type, sessile vs pedicellate flowers, receptacle morphology, stamen disposition, degree of fusion of the stamens, and fruit characters. Future work should focus on a greater understanding of the morphological differences between the apetalous genera Leptobalanus and Licania, and more taxon sampling within Moquilea, including taxa formerly in sect. Microdesmia. Relationships among the two clades of Hymenopus should also be resolved. Additional sampling may support further changes, but there is strong support for the phylogenetic framework presented here: Moquilea is sister to Couepia, core Licania appears in a separate lineage from Leptobalanus, Microdesmia and Hymenopus, as well as from the monospecific Cordillera and Parinariopsis, and Geobalanus.

Taxonomy

Below we present our concept of the genera derived from Licania s.l., based on molecular phylogenetic analyses supported by morphological evidence. All species and names previously in Licania s.l. cited in Prance & Sothers (2003a) are included here, in addition to five others described since then: Licania condoriensis Prance (2013: 72), L. palcazuensis Prance (2014: 2), L. apiknae Prance (2014: 4), L. monteagudensis Prance (2014: 6), and L. arachicarpa N. Zamora (2013: 1). A key to the Neotropical genera of Chrysobalanaceae, together with brief generic descriptions, keys to species, new combinations, synonymy, geographic distribution, and habitat where known are included. Genera are presented in order of increasing number of species, beginning with the monospecific genera Cordillera, gen. nov., and Parinariopsis, followed by Microdesmia (2 spp.), Geobalanus (3 spp.), Hymenopus (28 spp.), Leptobalanus (31 spp.), Moquilea (54 spp.) and Licania s.s. (c. 100 spp.). For species descriptions of all except five recently described species, see Prance & Sothers (2003a, b). All specimens cited have been seen by either the first and/or and second authors, apart from specimens of L. arachicarpa.

Key to Neotropical genera

Roman numerals refer to genera treated here.

• 1. Ovary bilocular

• 2. Stamens exserted, 20 – 60, in a complete circle; ovary at mouth of receptacle …………...………...Maranthes

• 2. Stamens 6 – 9, unilateral; ovary at side of receptacle or at mouth

• 3. Leaves with reticulate venation or with stomatal cavities, lanate (never glabrous); epicarp verrucose…………...………...Parinari

• 3. Leaves not reticulate, venation indistinct, without stomatal cavities, leaf undersurface glabrous or with whitish pubescence; epicarp smooth………………………………………………….Exellodendron

• 1. Ovary unilocular

• 4. Ovary at base of receptacle

• 5. Petals absent, stamens 2 – 14 (– 22)

• 6. Stamens exserted, always in a complete circle, (7 –) 8 – 10 (14 – 22)…………...………...VI. Leptobalanus

• 6. Stamens included, unilateral or less frequently in a complete circle, (2 –) 3 – 7 (11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . VIII. Licania

• 5. Petals present, stamens 3 – 90

• 7. Stamens included, 3 – 10, leaf undersurface smooth, glabrous or hirsute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .V. Hymenopus

• 7. Stamens exserted (or less often equalling), 11 – 90 (– 300), if only 8 – 14 then leaves deeply reticulate or with stomatal cavities

• 8. Stamens equalling calyx in length, 8 – 14, leaves deeply reticulate or with stomatal cavities ………………………………...III. Microdesmia

• 8. Stamens far exserted, 11 – 90, leaves not deeply reticulate, without stomatal cavities

• 9. Filaments partially fused in groups, pubescent; fruit longitudinally ridged……………….………………………………………………………………………...Chrysobalanus

• 9. Filaments free almost to base, glabrous; fruit not longitudinally ridged

• 10. Inflorescence of cymose or compact panicles 1 – 5 cm long; stamens 14 – 17……...…………………………………………………………………………IV. Geobalanus

• 10. Inflorescence racemose, panicles or racemose panicles, usually exceeding 5 cm in length; stamens 11 – 90……...…………………………………………………………………………VII. Moquilea

• 4. Ovary at mouth of receptacle or laterally on wall of receptacle

• 11. Receptacle flattened, almost solid; Andean, montane……...…………………………………………………………………………I. Cordillera

• 11. Receptacle short or cylindrical, hollow with a distinct cavity; lowland or rarely montane

• 12. Stamens united into a ligule for at least ½ their length, or inflorescence flagelliflorous………………………………………………………………..Acioa

• 12. Stamens free almost to base, inflorescence not flagelliflorous

• 13. Stamens 3 – 9, unilateral; leaves glabrous or hispid, without stomatal cavities; bracteoles often glandular …………………………………Hirtella

• 13. Stamens 12 – 300, unilateral or in a complete circle; leaves glabrous or pubescent, but not hispid, or with stomatal cavities; bracteoles not glandular

• 14. Stamens exserted barely beyond calyx; ovary inserted laterally in mid receptacle; bracteoles enclosing flower buds………………..…………………………………………………...II. Parinariopsis

• 14. Stamens far exserted; ovary inserted at mouth of receptacle; bracteoles usually not enclosing flower bud except in some Couepia

• 15. Leaves lanate pubescent, or hairs only between reticulations or within stomatal cavities; deeply reticulate and usually with stomatal cavities………………..…………………………………………………...Gaulettia

• 15. Leaves lanate, tomentose, hirsute or glabrous, without stomatal cavities

• 16. Calyx lobes with 2 glands on exterior………………..…………………………………………………...Acioa edulis

• 16. Calyx lobes without glands on exterior

• 17. Stamens c. 17; fruit small with bony endocarp…….……………………………………. Hirtella recurva

• 17. Stamens 20 – 300; fruit with hard woody endocarp……………………………………………Couepia

I. Cordillera Sothers & Prance gen. nov. Type: Couepia platycalyx Cuatrec.

http://www.ipni.org/urn:lsid:ipni.org:names:77156115-1

Trees. Leaves with caducous stipules; petioles 7.8 mm long, glabrous, terete; lamina obovate to obovate-orbicular, 8.5 – 12 × 7.9 cm, rounded at base and apex, glabrous above, with very sparse short appressed pubescence beneath; primary veins 10 – 12 pairs, prominent beneath. Inflorescences terminal little-branched panicles; rachis and branches with a few minute appressed hairs. Bracts and bracteoles to 1.5 mm long, caducous. Receptacle broadly turbinate, flattened, oblique, almost solid, 3 mm long, appressed pubescent on exterior, with dense deflexed hairs at throat within, filling the very small cavity; pedicels to 1.5 mm long. Calyx lobes rounded. Petals 5, unequal in length, glabrous, white. Stamens c. 54, inserted in a complete circle; filaments glabrous, exserted, connate at base. Ovary villous, on wall of receptacle. Style pubescent at base. Fruit ovoid, 8 × 5.5 cm; epicarp glabrous, smooth or with a few large scars or sparse lenticels; mesocarp thin, fleshy; endocarp thin, bony, glabrous within. Fig. 2.

Fig. 2

Cordillera platycalyx. A habit; B fruit; C style and ovary; D half flower; E flower; F petal. A, C – F drawn from G. Herrera 5375; B drawn from O. Vargas 175. drawn by j. m. fothergill.

recognition. Cordillera shares affinities with both Couepia and Licania; it has a greater number of stamens (c. 54) as in Couepia (10 – over 300), but the ovary is positioned at the side of the receptacle, differing from both Licania s.s. (at the base) and Couepia (at the mouth). It also differs from Couepia by the turbinate and almost solid receptacle (vs hollow in Couepia). It has petals, unlike Licania s.s.

distribution. A monospecific genus restricted to the Neotropics, from Costa Rica and Panama to northern South America, in Colombia, Ecuador and Venezuela. Occurs in higher altitude Andean and Central American forests, from 1000 – 2700 m, and has been recorded as low as 250 m.

etymology. The epithet Cordillera alludes to the locality where José Cuatrecasas collected the type specimen, in the Cordillera Oriental, as cited on the label. This name is appropriate since it also highlights the distribution and habitat of the genus: high altitudes along the Andes mountain range.

notes. The generic position of Cordillera platycalyx has always been problematic. Cuatrecasas (1950) stated that it was ‘an extraordinary member of Couepia and of the Chrysobalanoideae’, and Prance (1972a) was uncertain of its affinities to Couepia because of the ovary and fruit; he saw only the type and one other specimen so maintained the species in Couepia. Sothers et al. (2014: 193) transferred Couepia platycalyx to Licania but stated that its inclusion in Licania was tentative, primarily to effect the monophyly of Couepia. Here we provide further evidence from molecular phylogenetic analyses and incorporating more sampling of Licania that do not place Cordillera platycalyx in a clade with Licania s.s. Cordillera platycalyx shares a similar receptacle morphology to Licania but differs in the position of the ovary, which is on the wall of the receptacle rather than at its base. Parinariopsis licaniiflora also has a laterally inserted ovary, but not on the wall of the receptacle, and differs from Cordillera platycalyx in other characters, such as the number of stamens (54 in Cordillera; 18 – 25 in Parinariopsis), the receptacle which is almost solid in Cordillera (a character found elsewhere only in Maranthes), and in fruit characters. In addition, the two genera are separated geographically and altitudinally; Cordillera platycalyx is restricted to the highlands of western Amazonia, the Andes and southern Central America, whereas P. licaniiflora has an eastern Amazonian and Guianan distribution, occurring in lowland areas. Cordillera also differs from Couepia and Licania s.l. by the leafless flowering branches, an unusual character in Chrysobalanaceae.

Cordillera platycalyx (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156116-1

Couepia platycalyx Cuatrec., Fieldiana, Bot. 27: 66 (Cuatrecasas 1950); Licania platycalyx (Cuatrec.) Sothers & Prance (Sothers et al. 2014: 193). Type: Colombia, Cundinamarca, Cordillera Oriental, Dintel, entre Facatativa y La Vega, Cuatrecasas & Pérez-Arbeláez 15336 (holotype F; isotypes COL, US).

distribution and habitat. Costa Rica and Panama to northern South America, in Colombia, Ecuador and Venezuela. Occurs in higher altitude Andean and Central American forests, from 1000 – 2700 m, and has been recorded as low as 250 m.

II. Parinariopsis (Huber) Sothers & Prance stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156120-1

Licania subg. Parinariopsis Huber, Bol. Mus. Goeldi Hist. Nat. Ethnogr. 5: 368 (1909). Type: Licania parinarioides Huber.

Moquilea subg. Microbalanus Blume (1856 [1852]: 92). Type: Moquilea parviflora Blume (=Licania obtusifolia Fritsch).

Trees. Leaf lower surface deeply reticulate and lanate. Petiole usually with two large glands. Inflorescence panicles. Bracts and bracteoles large (3 – 6 mm) and enclosing groups of flowers. Petals 5. Stamens 16 (18) – 25, inserted in a complete circle; filaments almost free, slightly exceeding calyx lobes, hirsutulous. Ovary villous, inserted at side of receptacle. Fruit oblong, 6 – 8 × 3 – 4.5 cm, stipitate when young, becoming sessile by expansion of the stipe; epicarp conspicuously lenticellate; endocarp glabrous within. Fig. 3F.

Fig. 3

Transverse section of selected flowers of the segregates of Licania sensu lato. A Licania kunthiana; B Leptobalanus sclerophyllus; C Moquilea longicuspidata; D Hymenopus macrophyllus; E Geobalanus oblongifolius; F Parinariopsis licaniiflora. adapted from the artwork of j. dyer (a , b , d , e), j. m. fothergill (c) and r. wise (f).

distribution. Guianas, Venezuela, Colombia, Peru and Amazonian Brazil (Acre, Amazonas, Pará).

notes. Parinariopsis, a monospecific genus, was previously included as a subgenus of Licania. Huber (1909) described it under Licania but noted it as distinct from other Licania. The position of Parinariopsis lacks support but it is morphologically distinct from all other genera included in that clade, composed of genera with morphologically diverse features. It differs from core Licania, Moquilea, Leptobalanus, Geobalanus, Microdesmia and Hymenopus by the ovary inserted at the side of the receptacle (vs at the base) and from most species of Licania s.l. by the pair of bracts enclosing the flower buds, a feature common in Parinari and a few species of Couepia; it differs from Cordillera by the hollow receptacle (vs almost solid in Cordillera). In addition, Parinariopsis and Cordillera are allopatric; Parinariopsis is a species from the eastern Amazon region and the Guianas in lowland rainforest, whereas Cordillera is found in northwestern Andean South America, extending into Central America, in higher elevation forests.

Parinariopsis licaniiflora (Sagot) Sothers & Prance stat. & comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156118-1

Moquilea licaniiflora Sagot, Ann. Sci. Nat., Bot., Ser. VI, 15: 308 (1883), as licaniaeflora; Licania bracteosa Fritsch (1889: 54), nom. superfl. illegit; Licania licaniiflora (Sagot) S. F. Blake (1917: 66). Type: French Guiana, Rio Oiapoque, J. Martin s.n. (lectotype P, fide Prance 1972a: 91; isolectotype K).

Moquilea parviflora Blume (1856 [1852]: 92); Licania obtusifolia Fritsch (1889: 53); Licania parviflora (Blume) Lemée (1952: 23), nom. illegit., non L. parviflora Benth. (Bentham 1840), synon. nov. Type: French Guiana, Hort. Bot. Paris s.n. (not traced; PCU missing, n.v.).

Licania parinarioides Huber (1909: 368). Type: Brazil, Pará, rio Mapuera, A. Ducke s.n. (holotype MG 8961; isotypes F, RB 15180).

Licania parinarioides var. latifolia Maguire (Cowan 1952: 397). Type: Brazil, Pará, Cametá, A. Ducke s.n. (holotype US; isotypes BM, MG 16299, RB).

Licania capinensis Huber (1910: 71). Type: Brazil, Pará, Capim, J. E. Huber s.n. (holotype MG 946; isotype BM).

Licania huberiana Maguire (Cowan 1952: 398). Type: Brazil, Pará, Belém, rio Guamá, A. Ducke s.n. (holotype US; isotypes P, RB 18809, S).

Licania obovatifolia Maguire (Cowan 1952: 398). Type: Brazil, Amazonas, rio Negro, Padauari, R. L. Fróes 22566 (holotype NY, isotypes COL, IAN, SP).

Licania wilson-brownei Maguire (Cowan 1952: 399). Type: Guyana, Iramaipong, Kanuku Mts, G.Wilson-Browne 482 (F. D. 5888) (holotype NY; isotypes K, US).

distribution and habitat. Widespread in the Guianas, Venezuela, Colombia, Peru and Amazonian Brazil (Acre, Amazonas, Pará). Open forest at river margins, savanna margins and in secondary forest.

notes. We have been unable to locate a type specimen for Moquilea parviflora, renamed as Licania obtusifolia by Fritsch (1889). It was described from very poor material and Prance (1972a: 194) placed it among imperfectly known species. We have placed it in synonymy because the brief description fits that of Parinariopsis licaniiflora well, but a further search for the type would be worthwhile.

III. Microdesmia (Benth.) Sothers & Prance stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156124-1

Licania sect. Microdesmia Benth., J. Bot. (Hooker) 2: 212 (Bentham 1840). Type: Licania rigida Benth.

Trees. Leaf lower surface lanate and deeply reticulate; stomatal cavities present. Petiole with two glands. Bracteoles small, persistent. Inflorescence racemose panicles. Bracts and bracteoles persistent, 1.5 – 2.5 mm long. Petals 5. Stamens 8 – 14, inserted in a complete circle, equalling calyx lobes; filaments connate for half their length, densely pubescent. Ovary villous to densely villous, at base of receptacle. Fruit ovoid, 4 – 5.5 cm long; epicarp smooth, glabrous; endocarp puberulous or glabrous within.

distribution. Two species, Microdesmia arborea widely distributed from Mexico and Central America to northwestern South America (Venezuela, Colombia, Bolivia, Peru, Ecuador) and in Brazil in the state of Acre, and M. rigida restricted to northeastern Brazil.

notes. Licania sect. Microdesmia was treated as artificial by Prance (pers. obs.) and of the 11 species listed in Prance & Sothers (2003a), one is transferred here to Hymenopus, seven to Moquilea and only two species remain in Microdesmia, M. rigida and M. arborea. These two species have deeply reticulate lower leaf surfaces with stomatal cavities and are allopatric. Both species produce a seed oil extracted for use in paints, soaps, candles and as grease; their wood is durable and used for building and construction purposes. As a result they have been cultivated and moved around by indigenous and local people for their useful products, especially the oil. Both species occur in dry habitats (caatinga, gallery, scrub, dry plains). The combination of deeply reticulate leaf venation and stomatal cavities in species occurring in drier habitats as seen in M. arborea and M. rigida suggest that this genus has a similar adaptation to Gaulettia (Sothers et al. 2014), a genus mostly restricted to white sand habitats.

Key to species of Microdesmia

• 1. Stamens 8 – 12; pubescence of inflorescence light grey; Mexico and Central America to western and central South America…………………………………………..1. M. arborea

• 1. Stamens c. 14; pubescence of inflorescence ferrugineous; restricted to northeastern Brazil…………………………………………..2. M. rigida

1. Microdesmia arborea (Seem.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156125-1

Licania arborea Seem., Bot. Voy. Herald 3: 118, t. 25 (Seemann 1853). Type: Panama, H. Cuming 1109 (holotype BM; isotypes E, K, MO; phototype US).

Licania seleriana Loes. (Loesener 1911: 55). Type: Mexico, Oaxaca, S. Bartolo Yauhtepec, G. E. Seler 1660 (lectotype GH, fide Prance & Sothers 2003a: 91; isolectotype US).

Licania retusa Pilg. (Pilger 1914: 137). Type: Brazil, Acre, E. Ule 9568 (holotype B, destroyed; lectotype designated here, K; isotypes L, RB, U).

Licania bullatifolia Cuatrec. (Cuatrecasas 1950: 63). Type: Colombia, Putumayo, J. Cuatrecasas 10865 (holotype F; isotypes COL, NY, P, US).

distribution and habitat. From Mexico through Central America to Bolivia, Colombia, Peru, Ecuador, Venezuela and northern Brazil. Certainly native in Mexico and Central America and N South America, possibly introduced further south into Ecuador, Peru and Brazil for the oil bearing seeds. Dry plains, slopes and scrub forest.

2. Microdesmia rigida (Benth.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156126-1

Licania rigida Benth., J. Bot. (Hooker) 2: 220 (Bentham 1840). Type: Brazil, Ceará, G. Gardner 1592 (holotype K; isotypes BM, F, G, GH, NY, OXF, P, US, W).

Pleragina umbrosissima Arruda ex H. Kost. (Koster 1816: 500), nom. inval., fide G. T. Prance 1966: 134.

distribution and habitat. Native to northeastern Brazil, cultivated in Puerto Rico, Trinidad and Florida, and perhaps naturalised elsewhere in the Neotropics. Dry forest and gallery forest.

IV. Geobalanus Small (1913: 80). Type: Geobalanus oblongifolius (Michx.) Small.

Suffrutex with underground stems or small trees. Leaves glabrous or lanate. Petioles glandular or eglandular. Inflorescences panicles or cymose panicles. Bracts and bracteoles minute, persistent or caducous. Petals 5, not clawed (vs clawed in Chrysobalanus). Stamens 14 – 17, filaments exceeding to far-exceeding calyx lobes, in a complete circle, connate at base or to $\raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right.$, glabrous. Ovary glabrous or sparsely villous, at base of receptacle. Fruit ovoid, to 3 cm long; epicarp smooth, glabrous; endocarp hard, fibrous, pubescent within. Fig. 3E.

distribution. Three species distributed from southeastern United States, to Mexico, El Salvador and Costa Rica.

notes. Geobalanus differs from Licania and Leptobalanus by the presence of petals (vs apetalous in the other two genera). It is distinguished from Hymenopus by the more numerous stamens and the filaments exceeding the calyx lobes (vs 3 – 10 stamens and included in Hymenopus). It differs from Chrysobalanus by the stamens (slightly united in Chrysobalanus vs markedly united at the base in Geobalanus), the inflorescence and the unridged fruit in Geobalanus (vs ridged in Chrysobalanus). The three species of Geobalanus occupy distinct habitats: G. oblongifolius is found in pine barrens, sand dunes and oak scrubland; G. retifolius in gallery and non-flooded forests and G. riverae is found in transition forests, between humid and dry forests, from 600 – 900 m. The position of Geobalanus in the phylogenetic tree presented here lacks support, appearing as an early diverging genus sister to Parastemon and Grangeria, both paleotropic genera. However, other phylogenetic analyses place G. oblongifolius (as L. michauxii) sister to Licania s.l., Couepia, Hirtella, and Gaulettia (Bardon et al. 2016), genera to which it holds greater affinities.

Key to species of Geobalanus

• 1. Leaves oblong-elliptic; filaments connate for $\raisebox{1ex}{$1$}\!\left/ \!\raisebox{-1ex}{$3$}\right.$ of length……………………3. G. riverae

• 1. Leaves lanceolate to oblong-lanceolate; filaments connate only at base…………………….2

• 2. Tree; inflorescence short and dense with rachis glabrous; petioles 4 – 6 mm long; Mexico and El Salvador……………………2. G. retifolius

• 2. Suffrutex; inflorescence lax with rachis puberulous; petioles 1 – 3 mm long; SE USA……………………1. G. oblongifolius

1. Geobalanus oblongifolius (Michx.) Small (1913: 81, 200).

Chrysobalanus oblongifolius Michx. (Michaux 1803: 283); Chrysobalanus incanus Raf. (Rafinesque 1836: 26); Licania michauxii Prance (1970: 526). Type: U.S.A., Georgia, A. Michaux s.n. (holotype P).

Chrysobalanus retusus Raf. (Rafinesque 1836: 26). Type: U.S.A., L. W. Nuttall (in Herb. Collins) s.n. (holotype BM).

Geobalanus pallidus Small (1913: 81, 200); Chrysobalanus pallidus (Small) L. B. Sm. (Smith 1946: 136). Type: Florida: J. K. Small & J. J. Carter 711 (lectotype NY, fide Prance 1972a: 42).

distribution and habitat. SE United States. Pine barrens, sand dunes and oak scrubland.

2. Geobalanus retifolius (S. F. Blake) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156130-1

Licania retifolia S. F. Blake, Contr. Gray Herb. 52: 66 (1917). Type: Mexico, Guerrero, Cerro de los Cayones, E. Langlassé 992 (holotype GH; isotypes K, P, US).

distribution and habitat. Southwestern Mexico and El Salvador. Gallery and non-flooded forests; rare.

3. Geobalanus riverae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156131-1

Licania riverae Prance, Kew Bull. 50: 711 (1995). Type: Costa Rica, Guanacaste, P. N. Rincón de La Vieja, San Jorge, G. Rivera E. 1296 (holotype K; isotypes INB, MO).

distribution and habitat. Costa Rica. Transition between humid and dry forest, 600 – 900 m.

V. Hymenopus (Benth.) Sothers & Prance stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156133-1

Licania sect. Hymenopus Benth., J. Bot. (Hooker) 2: 212 (Bentham 1840). Type: Licania divaricata Benth.

Licania sect. Hirsuta Prance (1967: 226), synon. nov. Type: Licania latifolia Benth.

Trees. Leaf lower surface glabrous or hirsute, never tomentose, pulverulent, farinaceous nor with stomatal cavities. Petiole with pair of glands present or not. Inflorescences terminal and subterminal panicles or racemose panicles. Bracts and bracteoles small, persistent to subpersistent. Petals 4 – 5. Stamens 3 – 10, unilateral, in ¾ of a circle or in a complete circle, equal or shorter than calyx; filaments glabrous, free, connate either at base only, to ½ length or fully connate (only in H. conferruminatus). Ovary villous, tomentose, pilose or velutinous; at base of receptacle. Fruit globose, ovoid, ellipsoid, pyriform, to 9 × 8 cm, epicarp costate, smooth, glabrous, lenticellate, or pilose; endocarp glabrous or sparsely puberulous within. Fig. 3D.

distribution. Twenty-eight species from Central America (Costa Rica and Panama), Trinidad and Tobago, to northern South America (Colombia, Peru, Ecuador, Bolivia, Venezuela, the Guianas, and in Brazil mainly in the Amazonian region. Absent from Mexico.

notes. The genus Hymenopus includes species formerly in Licania sect. Hymenopus and sect. Hirsuta of subgen. Licania, and one species formerly in subgen. Moquilea sect. Microdesmia; the apetalous species previously in sect. Hymenopus are transferred to Licania s.s.; these taxa also have unilateral stamens, a predominant character in Licania s.s. Hymenopus differs from Licania by the presence of petals and by the mostly glabrous or hirsute leaves. They share the ovary at the base of the receptacle and reduced number of stamens. Taxa of Hymenopus sequenced here are retrieved in two subclades that do not appear to have any clear morphological synapomorphies to distinguish them. Our analyses do not refute or support the two clades of Hymenopus as distinct from each other. We have chosen to maintain all the taxa as a single genus. Further work may prove that there is more than one distinct genus within our concept of Hymenopus.

Key to species of Hymenopus

• 1. Leaf lower surface sparsely hirsute along venation or hispid at least on midrib

• 2. Inflorescence and flowers sparsely hirsutulous; leaf lamina appearing deeply rugose above; primary veins and venation deeply impressed on upper surface

• 3. Primary veins 13 – 17; petioles 10 – 16 mm; stamens 10………………………….2. H. amapaensis

• 3. Primary veins 7 – 10; petioles 6 – 10 mm; stamens 3 or 7 – 9

• 4. Leaf oblong, apex acuminate; Brazil (Amazonia)………………………………11. H. hirsutus

• 4. Leaf ovate to ovate-elliptic, apex rounded to obtuse; Central America……..7. H. costaricensis

• 2. Inflorescence and flowers densely pubescent; leaf lamina not rugose, primary veins plane or impressed and venation more or less plane on upper surface

• 5. Young stems, leaf midrib, and lower inflorescence branches hispid; Venezuela (Amazonas)…………...........12. H. hispidus

• 5. Young stems puberulous; leaf midrib hirsute; inflorescence and exterior of flowers with short grey pubescence or densely ferrugineous-tomentose

• 6. Inflorescence and exterior of flowers with a short grey pubescence; Trinidad and Amazonian Venezuela, Guianas, Brazil………….10. H. heteromorphus

• 6. Inflorescence and exterior of flowers densely ferrugineous-tomentose

• 7. Leaf chartaceous, apex acuminate; Bolivia and Amazonia (Brazil, Ecuador, Peru)…………. . . . 14. H. krukovii

• 7. Leaf thick-coriaceous, apex rounded to apiculate

• 8. Primary leaf veins 6 – 9, prominulous above; petioles 3 – 4 mm long; Guyana, Venezuela………………..…………………………………………………...16. H. lasseri

• 8. Primary leaf veins 12 – 20, slightly impressed above; petioles 7 – 15 mm long; Guyana and Amazonia………………..…………………………………………………...17. H. latifolius

• 1. Leaf lower surface glabrous

• 9. Flowers in small cymules on short distinct secondary branches of inflorescence at least 1 mm long

• 10. Leaves with a finely pointed acumen 15 – 30 mm long; flowers c. 1 mm long; ovary and style pubescent; Colombia (Valle)………………………………………21. H. minutiflorus

• 10. Leaves rounded or acute to broadly acuminate, the acumen 2 – 10 mm long; flowers 2 – 5 mm long; ovary and style glabrous or pubescent

• 11. Inflorescence compact and triangular, to 8 cm long; rachis and branches glabrous, the rachis not lenticellate; peduncles 2 – 4 mm long; exterior of flowers glabrous; stamens 5; Costa Rica………………………………………………………………....…………………...24. H. operculipetalus

• 11. Inflorescence lax and spreading, over 8 cm long; rachis and branches sparsely puberulous to tomentellous; peduncles sessile to c. 1 mm long; exterior of flowers puberulous, tomentellous or tomentose; stamens 5 – 7

• 12. Flowers 4 – 5 mm long, inflorescence rachis 2 – 3 mm thick; leaves thickly coriaceous; Venezuela………………..…………………………………………………...24. H. operculipetalus

• 11. Inflorescence lax and spreading, over 8 cm long; rachis and branches sparsely puberulous to tomentellous; peduncles sessile to c. 1 mm long; exterior of flowers puberulous, tomentellous or tomentose; stamens 5 – 7

• 12. Flowers 4 – 5 mm long, inflorescence rachis 2–3 mm thick; leaves thickly coriaceous; Venezuela………………………………………………………………....…………………...25. H. pakaraimensis

• 12. Flowers 1.5 – 3 mm long; inflorescence rachis 1 mm thick; leaves chartaceous or thinly coriaceous

• 13. Leaf apex rounded; reticulation intricate; Trinidad and Amazonian Venezuela, Guianas, Brazil………………..…………………………………………………...10. H. heteromorphus

• 13. Leaf apex acuminate; reticulation lax; Amazonia

• 14. Petioles 7 – 12 mm long, with 2 glands near junction of lamina; inflorescence rachis lenticellate; stamens 6 – 7; endocarp longitudinally striate; Amazonia……………….27. H. reticulatus

• 14. Petioles 4 – 6 mm long, eglandular; inflorescence rachis not lenticellate; stamens 5; endocarp not striate; Brazil (C Amazonia)..…………..……1. H. adolphoduckei

• 9. Flowers borne mainly on primary branches of inflorescence or on secondary branches, but not in pedunculate cymules

• 15. Leaves narrowly oblong with nearly parallel sides; stamens usually slightly exceeding calyx lobes; mouth of receptacle filled by a dense lanate mass; petals slightly unguiculate

• 16. Receptacle narrowly urceolate, densely arachnoid-pubescent; flowers in dense glomerules on primary branches; Guyana and western Amazonia..4. H. arachnoideus

• 16. Receptacle campanulate, puberulous; flowers not densely glomerulate

• 17. Flowers 1.5 – 2 mm long; leaves to 17 cm long, usually smaller; stipules to 5 mm, caducous; Brazil (Amazonia)…………………………22. H. oblongifolius

• 17. Flowers 2.5 – 3 mm long; leaves usually exceeding 16 cm in length (to 40 cm); stipules to 15 mm, subpersistent; Guianas, Venezuela, eastern brazilian Amazonia………………..…………………………………………………...19. H. macrophyllus

• 15. Leaves ovate to oblong-lanceolate, but sides converging; stamens shorter than calyx lobes; mouth of receptacle with short deflexed hairs only; petals with broad simple bases

• 18. Exterior of flowers and rachis and branches of inflorescence glabrous or sparsely hirsutulous; filaments connate for at least ½ length

• 19. Leaf apex caudate to cuspidate

• 20. Leaves elliptic, coriaceous, the apex caudate; petioles 5 – 7 mm long; French Guiana, Brazil (Amazonia), Colombia, Peru……………………5. H. caudatus

• 20. Leaves narrowly oblong, chartaceous, the apex cuspidate; petioles 1 – 3 mm long; Brazil (Mato Grosso)………………..…………………………………………………...20. H. miltonii

• 19. Leaf apex acute to acuminate

• 21. Stipules large and foliaceous, caducous; exterior of flowers glabrous; French Guiana, Venezuela………………..…………………………………………………...18. H. latistipulus

• 21. Stipules small, lanceolate, persistent; exterior of flowers hirsutulous

• 22. Leaves 20 – 31 × 6.5 – 11.5 cm, primary veins and midrib slightly impressed above; petioles 10 – 13 mm; Brazil (Rondônia)………………..…………………………………………………...6. H. conferruminatus

• 22. Leaves up to 27 × 10 cm usually much smaller, primary veins and midrib plane or impressed above; petioles 2 – 9 mm

• 23. Leaves 9 – 27 cm long, thick-coriaceous, the apex abruptly short-acuminate; primary veins plane above; Guianas, Brazil (Pará)………………..…………………………………………………...8. H. divaricatus

• 23. Leaves 7 – 11 cm long, membranous, the apex with a well-developed acumen 5 – 9 mm long; primary veins slightly impressed above; Costa Rica, Venezuela, Brazil (Pará)………………..…………………………………………………...9. H. glabriflorus

• 18. Exterior of flowers and usually rachis and branches of inflorescence densely puberulous to tomentose; pubescence completely covering exterior of calyx; filaments free to base

• 24. Midrib broad towards base, 2 – 3.5 mm thick; leaves very thick-coriaceous; stipules to 15 mm long, subpersistent; anthers deltoid or nearly so; Amazonia………………..…………………………………………………...13. H. intrapetiolaris

• 24. Midrib narrower towards base, 1 – 2 mm thick; leaves membranous to coriaceous; stipules to 8 mm long, persistent to caducous; anthers reniform

• 25. Calyx lobes and petals 4; stamens 3…………………………...3. H. arachicarpus

• 25. Calyx lobes and petals 5; stamens 5 – 7

• 26. Leaf apex long acuminate, base cuneate, petioles eglandular, leaf base confluent into petiole

• 27. Leaves 9 – 18 × 4.2 – 7 cm; flowers brown pubescent; Guianas, Brazil (Amazonia)………………..…………………………………………………...15. H. laevigatus

• 27. Leaves 4 – 5 × 2.5 – 3.8 cm; flowers grey pubescent; Brazil (Amazonia) . . . 23. H. occultans

• 26. Leaf apex retuse to bluntly acuminate, base subcordate, rounded, subcuneate; petioles usually glandular, leaf base not confluent; Trinidad and Amazonia

• 28. Stipules 5 – 6 mm long, caducous; primary veins prominulous above; petioles 8 – 13 mm, with 2 prominent raised glands; Brazil (C Amazonia)………………..…………………………………………………...28. H. sothersiae

• 28. Stipules 1 – 3.5 mm long, persistent or caducous; primary veins plane or slightly impressed above; petioles 3 – 12 mm, eglandular or with 2 ± sessile glands

• 29. Primary veins plane above, secondary venation plane or only slightly raised; petioles usually glandular, endocarp usually smooth, not angular (ridged in var. glabrus); Amazonia………………..…………………………………………………...10. H. heteromorphus

• 29. Primary veins slightly impressed above, secondary venation prominently reticulate; petioles eglandular; endocarp angular but not ridged; Venezuela, Brazil (C Amazonia)………………..…………………………………………………...26. H. prismatocarpus

1. Hymenopus adolphoduckei (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156951-1

Licania adolphoduckei Prance, Novon 11: 325 (2001). Type: Brazil, Amazonas, Reserva Florestal Ducke, Manaus – Itacoatiara road, km 26, P. A. C. L. Assunção 502 (holotype INPA; isotype K).

distribution and habitat. Brazil, central Amazonia. Beside rivers and streams in forest, on sandy soil.

2. Hymenopus amapaensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156952-1

Licania amapaensis Prance, Fl. Neotrop. Monogr. 9: 174 (1972a). Type: Brazil, Amapá, rio Araguari, Serra do Navio, J. M. Pires, W. A. Rodrigues & G. C. Irvine 51198 (holotype NY; isotypes IAN, INPA, K, MG).

distribution and habitat. Northern Brazil and French Guiana. Non -flooded forest.

3. Hymenopus arachicarpus (N. Zamora) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156953-1

Licania arachicarpa N. Zamora, Phytoneuron 26: 1 (2013). Type: Costa Rica, Heredia, Sarapiquí, Finca La Selva, the OTS Field Station on the Rio Viejo just E of its junction with the Rio Sarapiquí, about 100 m, B. Hammel 11149 (holotype CR; isotypes CAS, LSCR, MO, NY).

distribution and habitat. Costa Rica (Osa Peninsula). Tropical wet forest formations.

4. Hymenopus arachnoideus (Fanshawe & Maguire) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156954-1

Licania arachnoidea Fanshawe & Maguire, Bull. Torrey Bot. Club 75: 318 (Maguire 1948a). Type: Guyana, Bartica-Potaro road, D. B. Fanshawe 1495 (F.D.4231) (holotype NY; isotypes IAN, K).

distribution and habitat. Guyana, Amazonian Brazil and western Amazonia in Colombia, Ecuador and Peru. Non-flooded forest.

5. Hymenopus caudatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156955-1

Licania caudata Prance, Fl. Neotrop. Monogr. 9: 100 (1972a). Type: Brazil, Amazonas, Manaus, D. Coelho INPA 3946 (holotype NY; isotypes IAN, INPA, MG, RB).

distribution and habitat. Widespread in Amazonia (Guianas, Colombia, Ecuador, Peru, Brazil). Flooded as well as non-flooded forest.

6. Hymenopus conferruminatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156956-1

Licania conferruminata Prance, Kew Bull. 49: 362 (1994). Type: Brazil, Rondônia, Ji-Paraná, 45 km from town on line 56, C. A. Cid Ferreira 9032 (holotype INPA; isotypes CAS, K, MO).

distribution and habitat. Brazil; known only from the type collection. Forest on terra firme.

7. Hymenopus costaricensis (Standl. & Steyerm.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156957-1

Licania costaricensis Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 335 (Standley & Steyermark 1940). Type: Costa Rica, Alajuela, San Carlos, Vila Guesada, Au. Smith 1779 (holotype F; isotypes MO, NY, US; phototype K).

distribution and habitat. Costa Rica and Panama. Collected in hill pastures in the Alajuela province of Costa Rica and in lowland forest in Panama.

8. Hymenopus divaricatus (Benth.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156958-1

Licania divaricata Benth., J. Bot. (Hooker) 2: 221 (Bentham 1840); Licania heteromorpha Benth. var. divaricata (Benth.) Fritsch (1889: 45). Type: Guyana, R. & R. Schomburgk 463 (holotype K; isotypes BM, CGE, G, L, NY, OXF, P, W).

distribution and habitat. Guianas and northern Brazil. Forest on terra firme and savanna margins.

9. Hymenopus glabriflorus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156959-1

Licania glabriflora Prance, Fl. Neotrop. Monogr. 9: 104 (1972a). Type: Venezuela, Bolívar, Sierra Imataca, Río Reforma, J. A. Steyermark 87907 (holotype NY).

distribution and habitat. Costa Rica, Colombia, Venezuela, Guianas and Brazil (Amapá, Pará). Terra firme forest.

10. Hymenopus heteromorphus (Benth.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156960-1

Licania heteromorpha Benth., J. Bot. (Hooker) 2: 221 (Bentham 1840). Type: Guyana, R. & R. Schomburgk 873 (holotype K; isotypes BM, F, GH, L, NY, OXF, P, US, W).

Licania guianensis Klotzsch in M. R. Schomb. (Schomburgk 1848: 1199), nom. nud.

distribution and habitat. Central and South America. A variable and widespread species which we divide into five varieties. For distribution see under each variety.

10a. Hymenopus heteromorphus var. heteromorphus

Licania benthamii Hook. f. (Hooker 1867: 12). Type: Venezuela, Amazonas, Vasiva and Pacimoni, R. Spruce 3278 (lectotype K, fide Prance 1972a: 106; isolectotypes BM, BR, C, CGE, GH, GOET, LD, LE, NY, OXF, P, RB, TCD).

Licania biglandulosa Griseb. ex Urb. (Urban 1908: 354). Type: Trinidad, H. Crueger 105 (lectotype GOET, fide Prance & Sothers 2003a: 114; isolectotype K).

distribution and habitat. Trinidad to Colombia and Peru, the Guianas, Venezuela, Brazil and Bolivia. Commonest in periodically flooded forest, but occurring elsewhere.

10b. Hymenopus heteromorphus var. glabrus (Mart. ex Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156962-1

Licania glabra Mart. ex Hook. f., Fl. Bras. 14 (2): 10 (Hooker 1867); Licania heteromorpha var. glabra (Mart. ex Hook. f.) Prance (1972a: 108). Type: Venezuela, Amazonas, Río Guainia, mouth of Río Casiquiare, R. Spruce 3503 (lectotype K, fide Prance 1972a: 108; isolectotypes NY, TCD, P).

Licania costata Spruce ex Hook. f. (Hooker 1867: 10), nom. nud. in synon.

distribution and habitat. Colombia, Venezuela, Guyana, French Guiana, Brazil, Bolivia and Peru. Riverine forest mainly in W Amazonia, but scattered throughout the basin.

10c. Hymenopus heteromorphus var. subcordatus (Fritsch) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156963-1

Licania heteromorpha var. subcordata Fritsch, Ann. K. K. Naturhist. Hofmus. 4: 45 (1889); Licania subcordata Fritsch (1894: 18). Type: Brazil, Amazonas, J. E. Pohl 4402 (lectotype W, fide Prance 1972a: 108).

distribution and habitat. Amazonian Venezuela and Brazil. Riverine forest.

10d. Hymenopus heteromorphus var. perplexans (Sandwith) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156964-1

Licania heteromorpha var. perplexans Sandwith, Bull. Misc. Inform., Kew 1931: 371 (1931). Type: Guyana, Roraimas, R. & R. Schomburgk 824 (holotype K; isotypes NY, W).

distribution and habitat. Guyana. Forest.

10e. Hymenopus heteromorphus var. revolutus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156968-1

Licania heteromorpha var. revoluta Prance, Acta Amazon. 13 (1): 24 (1983). Type: Brazil, Amazonas, 20 km NW of Manaus, Tarumã, B. W. & S. P. Nelson 1058 (holotype INPA; isotypes K, NY, RB).

distribution and habitat. Brazil; known only from the type collection. Forest on terra firme, clay soil.

11. Hymenopus hirsutus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156974-1

Licania hirsuta Prance, Fl. Neotrop. Monogr. 9: 92 (1972a). Type: Brazil, Amazonas, rio Tefé, R. L. Fróes 26137 (holotype NY; isotypes IAN, NY).

distribution and habitat. French Guiana, Peru and Brazilian Amazonia. Flooded or non-flooded forest.

12. Hymenopus hispidus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156975-1

Licania hispida Prance, Fl. Neotrop. Monogr. 9S: 37 (1989). Type: Venezuela, Amazonas, Dept. Río Negro, Cerro Aratitiyope, J. A. Steyermark, P. E. Berry & F. Delascio 130185 (holotype NY; isotype MO).

distribution and habitat. Venezuela; known only from the type. Forest at base of high bluffs of an igneous rock mountain.

13. Hymenopus intrapetiolaris (Spruce ex Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156976-1

Licania intrapetiolaris Spruce ex Hook. f., Fl. Bras. 14 (2): 11 (Hooker 1867). Type: Venezuela, Amazonas, Río Guainia, R. Spruce 3539 (lectotype K, fide Prance 1972a: 105; isolectotypes BM, BR CGE, E, LE, NY, OXF, P, TCD, W ).

Licania heteromorpha var. grandifolia Benoist (1919: 513). Type: French Guiana: Cayenne, J. Martin s.n. (syntype K).

Licania intrapetiolaris var. brevis J. F. Macbr. (Macbride 1934: 369). Type: Peru: Mishuyacu, near Iquitos, G. Klug 612 (holotype F).

distribution and habitat. Guianas, N Brazil and Amazonian Colombia, Peru and Venezuela. Lowland forest in open places.

14. Hymenopus krukovii (Standl.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156977-1

Licania krukovii Standl., Field Mus. Nat. Hist., Bot. Ser. 17 (3): 256 (Standley 1937). Type: Brazil, Amazonas, near mouth of rio Embira, B. A. Krukoff 5070 (holotype F; isotypes A, BM, K, LE, M, MICH, MO, NY, S, US).

distribution and habitat. Western Amazonia in Bolivia, Ecuador, Peru, Brazil. Forest on high ground, and upland in Bolivia.

15. Hymenopus laevigatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156981-1

Licania laevigata Prance, Fl. Neotrop. Monogr. 9S: 41 (1989). Type: Brazil, Amazonas, Manaus-Caracaraí Rd., km 45, M. F. Silva & L. Coelho 80 (holotype INPA35442; isotype NY).

distribution and habitat. Venezuela, Guianas, Brazil and Peru. Terra firme forest.

16. Hymenopus lasseri (Maguire) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156982-1

Licania lasseri Maguire, Fieldiana, Bot. 28: 253 (1952). Type: Venezuela, Bolívar, Kavanayen, T. Lasser 1730 (holotype NY; isotype F).

distribution and habitat. Colombia, Guyana and adjacent Venezuela. Open forest on slopes, and savanna and river margins.

17. Hymenopus latifolius (Benth. ex Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156992-1

Licania latifolia Benth. ex Hook. f., Fl. Bras. 14 (2): 11 (Hooker 1867). Type: Brazil, Pará, Óbidos, R. Spruce 457 (lectotype K, fide Prance 1972a: 96; isolectotypes CGE, NY, OXF, P).

Licania obovata Benth. ex Hook. f. (Hooker 1867: 11). Type: Brazil, Amazonas, Manaus, R. Spruce 1569 (lectotype K, fide Prance 1972a: 95; isolectotypes BM, C, CGE, E, GH, GOET, K, LD, LE, M, NY, OXF, P, TCD).

distribution and habitat. Guianas, Venezuela and Amazonian Ecuador, Peru and Brazil. Non-flooded forest.

18. Hymenopus latistipulus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77156996-1

Licania latistipula Prance, Fl. Neotrop. Monogr. 9: 103 (1972a). Type: Venezuela, Delta Amacuro, east of Río Grande, ENE of El Palmar, near boundary of Bolívar State, L. Marcano-Berti 346 (holotype VEN; isotype VEN).

distribution and habitat. French Guiana and Venezuela. Forest on high ground.

19. Hymenopus macrophyllus (Benth.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77157006-1

Licania macrophylla Benth., Hooker’s J. Bot. Kew Gard. Misc. 2: 240 (Bentham 1850). Type: Brazil, Pará, near Belém, R. Spruce 139 (lectotype K, fide Prance 1972a: 100; isolectotypes CGE, NY, OXF, P, W s.n., probable isotype).

distribution and habitat. Guianas, Venezuela, Peru and Amazonian Brazil. Periodically flooded forest.

20. Hymenopus miltonii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472813-2

Licania miltonii Prance, Acta Amazon. 13 (1): 24 (1983). Type: Brazil, Mato Grosso, Aripuanã, BR-164, Núcleo Juína, M. G. Silva & A. Pinheiro 4296 (holotype MG; isotype NY).

distribution and habitat. C & W Brazil (Mato Grosso, Acre). Forest on terra firme, clay soil.

21. Hymenopus minutiflorus (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472814-2

Licania minutiflora Cuatrec., Fieldiana, Bot. 27: 64 (Cuatrecasas 1950), non L. minutiflora (Sagot) Fritsch (1889); Licania minuscula Cuatrec. (Cuatrecasas 1951: 113). Type: Colombia, Chocó, J. Cuatrecasas 21546 (holotype F; isotypes COL, P).

distribution and habitat. Colombia; known only from the original collections. Gallery forest.

22. Hymenopus oblongifolius (Standl.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472815-2

Licania oblongifolia Standl., Field Mus. Nat. Hist., Bot. Ser. 17 (3): 257 (Standley 1937). Type: Brazil, Amazonas, Humaitá, B. A. Krukoff 6812 (holotype F; isotypes A, B, BM, BR, K, LE, MICH, MO, NY, RB, S, US).

distribution and habitat. Amazonia: Venezuela, Peru and Brazil (Amazonas, Pará). Non-flooded forest.

23. Hymenopus occultans (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472816-2

Licania occultans Prance, Fl. Neotrop. Monogr. 9S: 42 (1989). Type: Brazil, Amazonas, Manaus-Itacoatiara Rd., km 31, W. C. Steward & J. F. Ramos P17669 (holotype INPA; isotypes MO, NY).

distribution and habitat. Central Amazonia and French Guiana. Forest on terra firme.

24. Hymenopus operculipetalus (Standl. & L. O. Williams) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472817-2

Licania operculipetala Standl. & L. O. Williams, Ceiba 3: 46 (Standley & Williams 1952). Type: Costa Rica, Puntarenas, Palmar, S of Osa, P. H. Allen 6030 (holotype EAP, n.v.; isotypes GH, US; phototype K).

distribution and habitat. Known only from Costa Rica. Forested hills.

25. Hymenopus pakaraimensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472818-2

Licania pakaraimensis Prance, Brittonia 28: 218 (1976). Type: Venezuela, Bolívar, Sierra Pakaraima, headwaters of Río Paragua, J. A. Steyermark 107357 (holotype NY; isotypes MO, VEN).

distribution and habitat. Venezuela; known only from the type collection. Submontane forest at 1400 m.

26. Hymenopus prismatocarpus (Spruce ex Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472819-2

Licania prismatocarpa Spruce ex Hook. f., Fl. Bras. 14 (2): 19 (Hooker 1867). Type: Venezuela, Amazonas, San Carlos, Río Negro, R. Spruce 3490 (lectotype K, fide Prance 1972a: 106; isolectotypes BM, BR, CGE, GH, LD, LE, NY, OXF, P, TCD, W).

distribution and habitat. Central Amazonian Brazil and Venezuela. Forest on terra firme.

27. Hymenopus reticulatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472820-2

Licania reticulata Prance, Fl. Neotrop. Monogr. 9: 97 (1972a). Type: Brazil, Amazonas, Manaus, Parque 10, F. Mello & L. Coelho s.n. (holotype NY; isotypes IAN, INPA 4141, MG).

distribution and habitat. French Guiana, western (Colombia, Ecuador, Peru) and Brazilian Amazonia. Non-flooded and periodically flooded forest.

28. Hymenopus sothersiae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472821-2

Licania sothersiae Prance, Novon 11: 328 (2001). Type: Brazil, Amazonas, Manaus, Reserva Florestal Ducke, J. E. L. S. Ribeiro 1142 (holotype INPA; isotype K).

distribution and habitat. Brazil, central Amazonia. Forest on terra firme.

VI. Leptobalanus (Benth.) Sothers & Prance stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472823-2

Licania sect. Leptobalanus Benth., J. Bot. (Hooker) 2: 212 (Bentham 1840). Type: Licania apetala (E. Mey.) Fritsch.

Licania sect. Batheogyne Benth. (Bentham 1840: 212). Type: L. turbinata Benth.

Trees. Leaf lower surface glabrous, tomentose or lanate, stomatal cavities present or absent. Petiole with or without glands. Inflorescence panicle, racemose panicle or panicle of cymes. Bracteoles small. Petals absent. Stamens (7 –) 8 – 10 (12 – 14) (19 in L. joseramosii; 22 in L. calvescens), filaments usually far exceeding calyx lobes (equalling or shorter in L. emarginatus), in a complete circle; filaments free or less often slightly connate at base, glabrous. Ovary villous or bearing some form of pilosity, or in some species glabrous above and densely villous or tomentose at base only, rarely entirely glabrous (L. undulatus); inserted at base of receptacle. Fruit globose to ellipsoid, 5.5 × 2 – 2.5 cm; epicarp smooth, lenticellate or pubescent; endocarp pubescent or rarely glabrous (L. granvillei and L. sparsipilis). Fig. 3B.

distribution. Thirty-one species, from Mexico, Central America and the Caribbean through to northern South America, the Guianas, Peru, Ecuador, Bolivia and SE Brazil.

notes. Leptobalanus includes species previously in Licania sect. Leptobalanus. The genus is apetalous as in core Licania, but differs by the mostly far-exserted to exserted stamens (vs included in Licania) and the slightly more numerous stamens (vs 3 – 8 in Licania), although there is an overlap with some species of Licania displaying 8 – 11 stamens. Leptobalanus shares a similar number of stamens with Hymenopus but differs by the absence of petals (vs presence of petals in Hymenopus) and the exserted stamens (vs included in Hymenopus). In Leptobalanus the stamens are in a complete circle (vs mostly unilateral in Licania). In molecular phylogenetic analyses Leptobalanus is retrieved in a clade with Microdesmia, Afrolicania, Parinariopsis, Cordillera, Gaulettia and Hymenopus.

Key to species of Leptobalanus (key does not include L. mexicanus as it is imperfectly known)

• 1. Leaf lower surface glabrous or lanate, the venation not prominent, stomatal cavities absent; stamens 8 – 14 (–22) (19 in L. joseramosii; 22 in L. calvescens)

• 2. Stamens exserted well beyond calyx lobes

• 3. Most flowers borne in small groups or cymules on distinct secondary branches (peduncles) of inflorescence; peduncles over 2 mm long

• 4. Rachis of inflorescence densely yellow-villous-tomentose; Amazonia (Brazil, Colombia, Venezuela)………………..…………………………………………………...31. L. wurdackii

• 4. Rachis of inflorescence grey-puberulous

• 5. Inflorescence short and compact, 3 – 5 cm long; flowers 4 – 5 mm long; leaves ovate; NE Brazil………………..…………………………………………………...29. L. turbinatus

• 5. Inflorescence longer and spreading, 6 – 20 cm long; flowers 2 – 3.5 mm long; leaves oblong to elliptic

• 6. Flowers 3.5 – 5 mm long; petioles deeply canaliculate; with 2 prominent glands at lamina base; rachis arachnoid-pubescent when young; Amazonia………………………..17. L. latus

• 6. Flowers 2 – 3 mm long; petioles terete to shallowly canaliculate; lamina base eglandular; rachis puberulous; Guianas, Brazil to S Amazonia…………………………..2. L. apetalus

• 3. Flowers borne on primary branches of inflorescence; peduncles usually sessile

• 7. Leaf apex predominantly obtuse to rounded

• 8. Rachis of inflorescence densely villous-tomentose; flowers c. 4 mm long; NW Amazonia (Colombia, Brazil, Venezuela)………………………………………………31. L. wurdackii

• 8. Rachis of inflorescence grey-puberulous; flowers 3 mm long; Guianas, Brazil (Amazonia)………………..…………………………………………………...23. L. parvifolius

• 7. Leaf apex distinctly acuminate (rarely only bluntly acuminate in L. jefensis)

• 9. Flowers 4 – 5 mm long; receptacle cupuliform (except in L. diegogomezii)

• 10. Leaves with hispid pubescence on venation otherwise glabrous; receptacle campanulate; leaf base subcordate; Costa Rica………………..…………………………………………………...8. L. diegogomezii

• 10. Leaves glabrous or with short waxy lanate-pulverulent pubescence beneath; receptacle urceolate; leaf base usually rounded to subcuneate

• 11. Leaves glabrous beneath, inflorescence grey-villous-tomentose; Brazil (Mato Grosso)………………..…………………………………………………...19. L. maguirei

• 11. Leaves with short persistent waxy lanate-pulverulent pubescence beneath; inflorescence brown-puberulous to tomentellous when mature; Amazonian Colombia, Peru, Brazil………………..…………………………………………………...17. L. latus

• 9. Flowers 2 – 3.5 mm long; receptacle urceolate or campanulate

• 12. Stipules persistent on young branches; petioles densely tomentose when young, 2.5 – 3.5 mm thick; upper portion of ovary glabrous; central Brazil………………..…………………………………………………...12. L. gardneri

• 12. Stipules usually caducous; petioles sparsely lanate-pubescent, puberulous, or glabrous, to 2 mm thick; upper portion of ovary pubescent

• 13. Inflorescence sparsely hirsutulous or yellow-arachnoid-tomentose; flower exterior yellow-brown-tomentose; petioles 2 – 3 mm long, canaliculate

• 14. Inflorescence sparsely hirsutulous; leaf apex finely acuminate, the acumen 5 – 10 mm long; northern Colombia……………………………7. L. cuspidatus

• 14. Inflorescence arachnoid-tomentose; leaf apex acute or bluntly acuminate, the acumen 3 – 5 mm long; Panama………………………………..15. L. jefensis

• 13. Inflorescence puberulous to tomentose; flower exterior grey-tomentose; petioles 3 – 11 mm long, terete or shallowly canaliculate towards base

• 15. Leaf bases subcordate; Panama…………………………………21. L. morii

• 15. Leaf bases cuneate or subcuneate to rounded

• 16. Leaf lower surface rufous-brown-pubescent, with deeply reticulate venation; petioles 8 – 11 mm long; leaf apex acuminate or cuspidate; inflorescence brown-tomentellous or tomentellous

• 17. Leaf apex acuminate, the acumen 5 – 10 mm; stamens 7 – 8; Costa Rica………………..…………………………………………………...28. L. stevensii

• 17. Leaf apex cuspidate, the acumen 10 – 15 mm long; stamens 10 – 12; Colombia………………..…………………………………………………...6. L. cuatrecasasii

• 16. Leaf lower surface glabrous or rarely white-arachnoid-pubescent, venation smooth, not deeply reticulate; petioles 3 – 8 mm long (9 – 11 in L. undulatus); leaf apex acuminate; inflorescence brown or grey- puberulous to tomentose

• 18. Rachis and branches of inflorescence tomentose; petioles usually bearing 2 distinct glands; Central America………………..…………………………………………………...26. L. sparsipilis

• 18. Rachis and branches of inflorescence puberulous; petioles eglandular

• 19. Leaves oblong lanceolate, margins undulate; Colombia (Caquetá)………………..…………………………………………………...30. L. undulatus

• 19. Leaves elliptic to oblong, margins entire

• 20. Leaves bullate; stamens 8; Peru……………3. L. bullatus

• 20. Leaves not bullate; stamens 14

• 21. Inflorescence and flower exterior sparsely puberulous; leaves thickly coriaceous, the apex finely attenuate with acumen 10 – 25 mm long; Guianas, N Amazonian Brazil………………..…………………………………………………...13. L. granvillei

• 21. Inflorescence usually densely grey-puberulous; leaves chartaceous, the apex acuminate with acumen 3 – 15 mm long; Colombia, Peru, Venezuela to southern Brazil………………..…………………………………………………...2. L. apetalus

• 2. Stamens not or scarcely exceeding calyx lobes

• 22. Leaf lower surface lanate; flowers sessile on primary branches of inflorescence; stamens c. 22; Colombia (Valle)…………………………………………………………………………4. L. calvescens

• 22. Leaf lower surface glabrous; flowers in groups on distinct secondary branches of inflorescence; stamens 10 – 19; Amazonia

• 23. Stamens c. 19; leaves 15 – 20 cm long, with long acumen 11 – 19 mm long; flowers c. 5 mm long; Brazil (N Amazonia)……………………………………………………..16. L. joseramosii

• 23. Stamens 10 – 11; leaves 3 – 8.5 cm long, with acumen 0 – 6 mm long; flowers 2 – 3 mm long; Peru, Venezuela, Amazonian Brazil……………………………………………9. L. emarginatus

• 1. Leaf lower surface with stomatal cavities or very deeply cut reticulation resembling stomatal cavities, lanate-pubescent in mouth of cavities; stamens 8 – 13

• 24. Inflorescence with secondary branches; flowers on distinct secondary branches of inflorescence that are at least 2 mm long

• 25. Flowers in small distinct groups or in cymules on short secondary branches (peduncles); leaf upper surface smooth, the base rounded to cuneate

• 26. Inflorescence 15 – 22 cm long, ferrugineous-arachnoid; leaf lower surface brown when dry; Guyana………………..…………………………………………………...24. L. persaudii

• 26. Inflorescence 6 – 8 cm long, grey-arachnoid or puberulous; leaf lower surface grey when dry; Guianas, Brazil (northern Amazonia)………………………………………….27. L. sprucei

• 25. Flowers more or less clustered in large groups on primary and secondary branches of inflorescence, but without distinct peduncles; leaf upper surface usually papillose, the base often cordate; Brazil (Amazonia and south)……………………………………………………….25. L. sclerophyllus

• 24. Inflorescence with primary branches only; flowers on primary branches of inflorescence, sessile or rarely with short peduncles not exceeding 1 mm in length

• 27. Pedicels 2 mm long (in bud), to 4 mm in flower; Guianas……………………..1. L. albiflorus

• 27. Pedicels not exceeding 2 mm, often shorter

• 28. Leaf reticulation extremely prominent on upper surface; fruit exterior appressed-sordid- yellow-velutinous

• 29. Leaves deeply cordate; Colombia, Venezuela…………………..5. L. cardiophyllus

• 29. Leaves rounded to subcuneate at base

• 30. Petioles 5 – 13 mm long; flowers c. 3 mm long, borne on slender branchlets 0.5 – 1 mm thick; receptacle campanulate; flowers and inflorescence grey-tomentose; Panama, Venezuela, Guianas, western and central Amazonia (Brazil, Ecuador, Peru)……………………………………………….….18. L. longistylus

• 30. Petioles 15 – 20 mm long; flowers c. 4 mm long, borne on branchlets c. 2 mm thick; receptacle cupuliform; flowers and inflorescence brown-tomentose; Colombia (Pacific)………………..…………………………………………………...11. L. fuchsii

• 28. Leaf reticulation prominulous on upper surface; fruit exterior glabrous or brown-tomentose

• 31. Young branches densely tomentellous, with a thick corky bark; central Brazil………………..…………………………………………………...14. L. humilis

• 31. Young branches glabrous or nearly so, the bark thin

• 32. Flowers 3.5 – 5 mm long; leaf apex mucronate; Guyana……...10. L. foveolatus

• 32. Flowers 2 – 3 mm long; leaf apex rounded to acuminate

• 33. Leaf apex finely acuminate, leaves narrowly oblong; Central America………………..…………………………………………………...15. L. jefensis

• 33. Leaf apex acute to bluntly acuminate, leaves ovate to oblong; Brazil (Amazonia), Venezuela, Guianas………………..…………………………………………………...22. L. octandrus

1. Leptobalanus albiflorus (Fanshawe & Maguire) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472825-2

Licania albiflora Fanshawe & Maguire, Bull. Torrey Bot. Club 75: 318 (Maguire 1948a). Type: Guyana, Madray-Bubu trail, Essequibo-Demerara Region, D. B. Fanshawe 1678 (F.D. 4414) (holotype NY; isotype K).

distribution and habitat. Guianas, rare but widespread. Primary forest.

2. Leptobalanus apetalus (E. Mey.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472826-2

Hirtella apetala E. Mey., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 803 (Meyer 1825); Licania apetala (E. Mey.) Fritsch (1889: 54). Type: Suriname, Meyer s.n. (holotype GOET; isotype M).

2a. Leptobalanus apetalus (E. Mey.) Sothers & Prance var. apetalus. Type: Suriname, Meyer s.n. (holotype GOET; isotype M).

Licania pendula Benth. (Bentham 1840: 218); Moquilea pendula (Benth.) Hook. f. (Hooker 1867: 22); Licania apetala var. pendula (Benth.) Fritsch (1889: 55). Type: Brazil, lagoons of rio Negro, R. & R. Schomburgk 906 (holotype K; isotypes BM, CGE, E, F, GH, LE, NY, OXF, P, TCD, US, W).

Licania floribunda Benth. (Bentham 1840: 219); Moquilea floribunda (Benth.) Hook. f. (Hooker 1867: 21). Type: Guyana, without locality, R. & R. Schomburgk 897 (holotype K; isotypes BM, CGE, E, F, GH, L, OXF, P, US, W).

Moquilea orinocensis Rusby (1920: 27). Type: Venezuela: Punta Piedra Is., H. H. Rusby & R. W. Squires 426 (holotype NY; isotypes F, K, M, MO, Z).

Licania dahlgrenii Standl. (Standley 1937: 255). Type: Brazil, Piauí, Pedro Segundo, B. E. Dahlgren 876 (holotype F; isotypes P, S, US).

Licania hylaea Cuatrec. (Cuatrecasas 1956: 198). Type: Colombia, Amazonas, Río Apaporis between Río Kananari and Pacoa, H. García-Barriga 14100 (holotype US; isotypes COL, NY).

distribution and habitat. Throughout the Guianas, Venezuela, Bolivia, Ecuador, Peru, Colombia, and in Amazonian, northeastern, central and southeastern Brazil. Gallery forest, periodically flooded forest, river margins, savanna margins, and open river beaches on sandy soil.

2b. Leptobalanus apetalus var. apertus (Benth.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472827-2

Licania aperta Benth., J. Bot. (Hooker) 2: 218 (Bentham 1840); Licania apetala var. aperta (Benth.) Prance (1972a: 68). Type: Guyana, R. & R. Schomburgk 593 (holotype K; isotypes BM, BR, CGE, E, GH, L, NY, OXF, P).

Licania pubiflora Benth. (Bentham 1840: 219). Type: Guyana, R. & R. Schomburgk 136 (holotype K; isotypes BM, CGE, NY, OXF, P, TCD).

Licania caracasana Klotzsch ex Hook. f. (Hooker 1867: 25), nom. nud. in synon.

Licania affinis Kuntze (1891: 217), nom. illegit., non Fritsch (1889).

Licania kuntzeana Urb. (Urban 1908: 353); Moquilea kuntzeana (Urb.) R. O. Williams (1932: 314). Type: Trinidad-Tobago, Trinidad, C. E. O. Kuntze 1014 (lectotype K, fide Prance & Sothers 2003a: 70; isolectotype US).

distribution and habitat. Trinidad, Venezuela, Guianas and in Amazonian Bolivia, Colombia, Ecuador, Peru, and Brazil. Riverine forest, riverbanks and fresh water beaches, and savanna margins.

3. Leptobalanus bullatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472828-2

Licania bullata Prance, Kew Bull. 50: 143 (1995). Type: Peru, Huánuco, Prov. Pachitea, region of Pucallpa, Sira Mts, B. Wallnöfer 13-41088 (holotype K; isotype W).

distribution and habitat. Peru. Lower montane forest around 800 m.

4. Leptobalanus calvescens (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472829-2

Licania calvescens Cuatrec., Fieldiana, Bot. 27: 64 (Cuatrecasas 1950). Type: Colombia, Valle, Río Yurumangui, Veneral, J. Cuatrecasas 15835 (holotype F; isotype COL).

distribution and habitat. Colombia (Chocó and Valle). Pacific coastal forests.

5. Leptobalanus cardiophyllus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472830-2

Licania cardiophylla Prance, Kew Bull. 47: 637 (1992). Type: Venezuela, Amazonas, Dept. Atabapo, Caño Yagua, Cucurital de Yagua, G. Davidse, O. Huber & S. S. Tillett 17437 (holotype MYF; isotypes K, MO, NY).

distribution and habitat. Venezuela and Colombia. Gallery forest.

6. Leptobalanus cuatrecasasii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472831-2

Licania cuatrecasasii Prance, Acta Amazon. 8 (4): 577 (1978). Type: Colombia, Valle, Alto Yunda, Río Anchicayá, S. Hilty 0-1 (holotype US; isotype NY).

distribution and habitat. Known only from Colombia (Valle) and Ecuador. Upland and lowland forest.

7. Leptobalanus cuspidatus (Rusby) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472832-2

Moquilea cuspidata Rusby, Descr. S. Amer. Pl. 27 (1920); Licania cuspidata (Rusby) Prance (1972a: 72). Type: Colombia, Magdalena, Santa Marta, near Las Partidas, H. H. Smith 1773 (holotype NY; isotypes A, BM, BR, F, GH, K, MICH, P, US).

distribution and habitat. Colombia; known only from the type collection. Montane forest.

8. Leptobalanus diegogomezii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472833-2

Licania diegogomezii Prance, Kew Bull. 50: 717 (1995). Type: Costa Rica, Puntarenas, Res. Forestal Golfo Dulce, 10 km from Chacarita, R. Aguilar 898 (holotype MO; phototype K).

distribution and habitat. Costa Rica; known only from two collections. Lowland forest, 100 m.

9. Leptobalanus emarginatus (Spruce ex Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472834-2

Licania emarginata Spruce ex Hook. f., Fl. Bras. 14 (2): 15 (Hooker 1867). Type: Brazil, Amazonas, rio Uaupés, near Panuré, R. Spruce 2699 (lectotype K, fide Prance 1972a: 74; isolectotypes BM, BR, CGE, F, GH, GOET, LD, LE, NY, OXF, P, RB, W).

Licania petrensis Prance (1995: 713). Type: Venezuela, Amazonas, Dept. Atures, Puerto Ayacucho-El Burro, Km 48, A. Gröger 690 (holotype VEN; isotype K).

distribution and habitat. Venezuelan Guayana and Amazonian Brazil and Peru. Non-flooded forest, especially on granitic outcrops.

10. Leptobalanus foveolatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472835-2

Licania foveolata Prance, Fl. Neotrop. Monogr. 9: 80 (1972a). Type: Guyana, Upper Mazaruni R., Aga Creek, R. Boyan 28 (F.D. 7852) (holotype K; isotypes NY, U).

distribution and habitat. Collected only in Guyana. Riverine forest.

11. Leptobalanus fuchsii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472836-2

Licania fuchsii Prance, Fl. Neotrop. Monogr. 9: 79 (1972a). Type: Colombia, Chocó, Rio Baudó, 13.5 km above mouth, H. P. Fuchs 22036 (holotype NY; isotypes K, MO, US).

distribution and habitat. Colombia. Pacific coastal forest.

12. Leptobalanus gardneri (Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472848-2

Moquilea gardneri Hook. f., Fl. Bras. 14 (2): 21 (Hooker 1867); Licania gardneri (Hook. f.) Fritsch (1889: 56). Type: Brazil, Minas Gerais, near São Romão, G. Gardner 4539 (holotype K; isotypes BM, GH).

Licania mattogrossensis Pilg. (Pilger 1923: 540); Moquilea mattogrossensis (Pilg.) Malme (1930: 11). Type: Brazil, Mato Grosso, Coxim, F. C. Hoehne 4262 (holotype B, destroyed; lectotype R, fide Prance & Sothers 2003a: 74; phototype NY).

distribution and habitat. Bolivia, and in C, NE, SE and N Brazil. Riverine forest bordering cerrado, and in cerrado itself.

13. Leptobalanus granvillei (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472849-2

Licania granvillei Prance, Proc. Kon. Ned. Akad. Wetensch. C. 89: 114 (1986). Type: French Guiana, Saül, Monts La Fumée, S. A. Mori & B. M. Boom 14764 (holotype NY; isotypes CAY, P).

distribution and habitat. From Panama, Ecuador and Colombia to French Guiana, Guyana, Venezuela and Brazil (Amazonas). Upland forest on terra firme and along riverbanks.

14. Leptobalanus humilis (Cham. & Schltdl.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472850-2

Licania humilis Cham. & Schltdl., Linnaea 2: 549 (Chamisso & Schlechtendal 1827); Moquilea humilis (Cham. & Schltdl.) Hook. f. (Hooker 1867: 26); Chrysobalanus humilis (Cham. & Schltdl.) Kuntze (1893: 76). Type: Brazil, Minas Gerais, F. Sellow s.n. (holotype LE; isotype BR; isotype fragments F, M, P).

Chrysobalanus sublanatus Kuntze (1893: 76). Type: Brazil, Mato Grosso, C. E. O. Kuntze s.n. (holotype NY).

Licania ulei Taub. (Taubert 1896: 428). Type: Brazil, Minas Gerais, rio Paranaíba, E. Ule 3187 (holotype B, destroyed; lectotype designated here, HBG; isotype R).

distribution and habitat. Bolivia and Brazil. In cerrados.

15. Leptobalanus jefensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472851-2

Licania jefensis Prance, Brittonia 28: 215 (1976). Type: Panama, Summit of Cerro Jefe, J. D. Dwyer et al. 5047 (holotype NY; isotype MO).

distribution and habitat. Known only from Costa Rica and Panama. Cloud forests.

16. Leptobalanus joseramosii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472852-2

Licania joseramosii Prance, Acta Amazon. 8 (4): 581 (1978). Type: Brazil, Amazonas, Manaus-Caracaraí Rd., Km 130, O. P. Monteiro & J. F. Ramos 29 (holotype INPA 54340; isotype NY).

distribution and habitat. Brazil; known only from the type collection. White sand forest or campina.

17. Leptobalanus latus (J. F. Macbr.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472853-2

Licania lata J. F. Macbr., Candollea 5: 369 (Macbride 1934). Type: Peru, Loreto, Mishuyacu, near Iquitos, G. Klug 560 (holotype F; isotype NY).

distribution and habitat. Guianas, Venezuela, C & W Amazonia (Colombia, Ecuador, Peru and Brazil). Non-flooded forest and secondary scrub on sandy soil.

18. Leptobalanus longistylus (Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472856-2

Moquilea longistyla Hook. f., Fl. Bras. 14 (2): 24 (Hooker 1867); Licania longistyla (Hook. f.) Fritsch (1889: 56). Type: Venezuela, Amazonas, Río Orinoco below Esmeralda, R. Spruce 3232 (lectotype K, fide Prance 1972a: 78; isolectotypes BM, BR, E, GH, LE, NY, OXF, P, TCD, W).

distribution and habitat. Panama, Venezuela, Guianas, and W & C Amazonia in Colombia, Ecuador, Peru, and in C and N Brazil. Periodically flooded forest.

19. Leptobalanus maguirei (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472857-2

Licania maguirei Prance, Fl. Neotrop. Monogr. 9: 70 (1972a). Type: Brazil, Mato Grosso, rio Juruena, Brasília to Acre Highway, B. Maguire, J. M. Pires, C. Maguire & N. T. da Silva 56470 (holotype NY).

distribution and habitat. Brazil (known only from Mato Grosso and Mato Grosso do Sul). Riverine forest.

20. Leptobalanus mexicanus (Lundell) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472858-2

Licania mexicana Lundell, Wrightia 5: 40 (1974). Type: Mexico, Sinaloa, between Rancho Del Piño and Chele, C. L. Lundell 13023 (holotype LL; isotypes CAS, MICH).

distribution and habitat. Known only from the type. Habitat unknown.

21. Leptobalanus morii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472869-2

Licania morii Prance, Brittonia 28: 215 (1976). Type: Panama, El Llano-Cartí Rd, 12 km from Inter American Hwy, S. A. Mori & J. A. Kallunki 4665 (holotype NY; isotype MO).

distribution and habitat. Panama; known only from the type collection. Moist forest at 350 m.

22. Leptobalanus octandrus (Hoffmanns. ex Roem. & Schult.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:604728702

Hirtella octandra Hoffmanns. ex Roem. & Schult., Syst. Veg. ed. 9, 5: 274 (Roemer & Schultes 1819); Licania turiuva Cham. & Schldtl. (Chamisso & Schlechtendal 1827: 550), nom. superfl. illegit.; Moquilea turiuva (Cham. & Schldtl.) Hook. f. (Hooker 1867: 25), nom. superfl. illegit.; Licania octandra (Hoffmanns. ex Roem. & Schult.) Kuntze (1891: 217). Type: Brazil, Pará, F. W. Sieber s.n. (holotype B, Herb. Willdenow 4851; phototype NY).

22a. Leptobalanus octandrus subsp. octandrus

Licania bothynophylla Mart. (Martius 1841: 15); Moquilea bothynophylla (Mart.) Hook. f., (Hooker 1867: 26). Type: Brazil, Rio de Janeiro, Campos Bravos, C. F. P. Martius 449 (holotype M; isotypes BR, F, GH, K, LE, M, NY, P, W).

Moquilea utilis Hook. f. (Hooker 1867: 24); Licania utilis (Hook. f.) Fritsch (1889: 56). Type: Brazil, Pará, Santarém, R. Spruce 877 (lectotype K, fide Prance 1972a: 83; isolectotypes LD, NY, P).

Licania sellowiana Klotzsch ex Hook. f. (Hooker 1867: 26), nom. nud. in synon.

Licania hookeri var. obtusa Huber (1909: 368). Type: Brazil, Pará, Faro, A. Ducke MG 8371 (holotype MG; isotypes BM, G).

Licania takutuensis Standl. (Smith 1939: 182). Type: Guyana, W extremity of Kanuku Mts, A. C. Smith 3302 (holotype F; isotypes A, IAN, K, LE, NY, P, S, US).

distribution and habitat. Widely distributed from N Venezuela through the Guianas, Peru, Colombia, Bolivia and E & C Amazonia to Northeast, East-central & Southeast Brazil. Forest, both on flooded and non-flooded ground but especially in open riverine habitats and at savanna margins, and in the gallery forest of the Planalto of central Brazil.

22b. Leptobalanus octandrus subsp. pallidus (Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472871-2

Moquilea pallida Hook. f., Fl. Bras. 14 (2): 25 (Hooker 1867); Licania hookeri Fritsch (1889: 56); Licania pallida (Hook. f.) Kuntze (1891: 217) nom. illegit., non L. pallida Spruce ex Sagot (1883: 306); Licania octandra subsp. pallida (Hook. f.) Prance (1972a: 84). Type: Venezuela, Amazonas, Río Casiquiari, Vasiva or Pacimoni, R. Spruce 3302 (lectotype K, fide Prance 1972a: 84; isolectotypes BM, BR, CGE, E, GH, GOET, LD, LE, P, OXF, RB, TCD).

Licania egensis Fritsch (1889: 58). Type: Brazil, Amazonas, Tefé, E. Poeppig 2531 (lectotype W, fide Prance 1972a: 84; isolectotypes G, LE).

Licania stenocarpa Standl. (Standley 1937: 258). Type: Brazil, Amazonas, São Paulo de Olivença, B. A. Krukoff 8553 (holotype NY; isotypes A, BM, BR, F, K, MO, P, S, US).

distribution and habitat. Amazonia: Guyana, Venezuela, Bolivia, Colombia, Ecuador, Peru and Brazil. Non-flooded forest.

22c. Leptobalanus octandrus subsp. grandifolius (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472872-2

Licania octandra subsp. grandifolia Prance, Acta Amazon. 4 (1): 18 (1974). Type: Brazil, Amazonas, Rio Javari, behind Estirão do Equador, E. Lleras, W. C. Steward, J. C. Ongley et al. P17270 (holotype INPA; isotypes K, MO, NY, P).

distribution and habitat. Colombia, Peru and Brazil. Forest on terra firme, especially on sandy soils.

23. Leptobalanus parvifolius (Huber) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472873-2

Licania parvifolia Huber, Bol. Mus. Paraense Hist. Nat. 5: 367 (1909). Type: Brazil, Pará, rio Mapuera [infra Taboleiro Grande], A. Ducke s.n. (holotype MG 8979; isotypes F, G, RB).

Licania arachnites Standl. (Smith 1939: 181). Type: Guyana, Essequibo R., near Onoro Creek, A. C. Smith 2696 (holotype F; isotypes A, IAN, K, MAD, MO, NY, P, U, US).

distribution and habitat. Guyana, Venezuela, Bolivia and the southern part of Amazonian Brazil. Open riverine localities, especially river banks and islands.

24. Leptobalanus persaudii (Fanshawe & Maguire) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472874-2

Licania persaudii Fanshawe & Maguire, Bull. Torrey Bot. Club 75: 375 (Maguire 1948b). Type: Guyana, Hope, A. C. Persaud 95 (lectotype NY, fide Prance 1972a: 75; isolectotypes F, K, US).

distribution and habitat. Confined to the Guianas. Primary forests.

25. Leptobalanus sclerophyllus (Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472875-2

Moquilea sclerophylla Hook. f., Fl. Bras. 14 (2): 23, t. 7 (Hooker 1867); Moquilea sclerophylla var. pohliana Hook. f. (Hooker 1867: 24), nom. inval.; Licania sclerophylla (Hook. f.) Fritsch (1889: 56). Type: Brazil, Minas Gerais, C. F. P. Martius 1467 (lectotype M, fide Prance 1972a: 77; isolectotype BR).

Moquilea sclerophylla var. myristicoides Hook. f. (Hooker 1867: 24); Licania myristicoides Benth. (Hooker 1867: 24), nom. inval. in synon.; Licania sclerophylla var. myristicoides (Hook. f.) Fritsch (1889: 56). Type: Brazil, Pará, Santarém, R. Spruce 987 (lectotype K, fide Prance & Sothers 2003a: 84; isolectotypes NY, P).

Moquilea sclerophylla var. scabra Hook. f. (Hooker 1867: 24); Licania scabra Benth. ex Hook. f. (Hooker 1867: 24), nom. inval. in synon.; Licania sclerophylla var. scabra (Hook. f.) Fritsch (1889: 56). Type: Brazil, Pará, Santarém, R. Spruce 840 (lectotype K, fide Prance & Sothers 2003a: 84; isolectotypes NY, P).

Licania aspera Standl. (Standley 1937: 254). Type: Brazil, Pará, R. C. Monteiro da Costa 279 (holotype F; isotypes IAN, US).

distribution and habitat. Bolivia, Colombia, Peru, eastern and southern Amazonia and the planalto of central Brazil. Non-flooded forest, riverine forest or cerrado.

note. The collection Martius 1467b is the type of Swartzia lomatopus Mart. and has been erroneously cited as the type of Leptobalanus sclerophyllus.

26. Leptobalanus sparsipilis (S. F. Blake) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472876-2

Licania sparsipilis S. F. Blake, Contr. Gray Herb. 52: 67 (1917). Type: Belize, Sittee R., M. E. Peck 858 (holotype GH; isotypes K, NY).

distribution and habitat. From southern Mexico to Colombia. Forest on low wet ground.

27. Leptobalanus sprucei (Hook. f.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472877-2

Moquilea sprucei Hook. f., Fl. Bras. 14 (2): 22 (Hooker 1867); Licania sprucei (Hook. f.) Fritsch (1889: 55). Type: Brazil, Amazonas, Manaus, R. Spruce 1801 (lectotype K, fide Prance 1972a: 76; isolectotypes BM, CGE, E, GH, GOET, LD, LE, M, NY, P, RB, TCD, W).

distribution and habitat. Guianas, Venezuela and in the Amazon region of Brazil. Primary forest on non-flooded ground and secondary forest.

28. Leptobalanus stevensii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472878-2

Licania stevensii Prance, Kew Bull. 47: 637 (1992). Type: Costa Rica, Limón, 2 km S of Río Colorado, W. D. Stevens, G. Herrera & O. M. Montiel 25100 (holotype K; isotype MO).

distribution and habitat. Costa Rica. Lowland evergreen forest, often on slopes.

29. Leptobalanus turbinatus (Benth.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472885-2

Licania turbinata Benth., J. Bot. (Hooker) 2: 218 (Bentham 1840). Moquilea turbinata (Benth.) Hook. f. (Hooker 1867: 23). Type: Brazil, Pernambuco, G. Gardner 1149 (holotype K; isotypes BM, CGE, E, GH, NY, OXF, P, S, SP, US, W).

distribution and habitat. NE Brazil (Bahia and Pernambuco). Coastal forest and restinga forest.

30. Leptobalanus undulatus (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472886-2

Licania undulata Prance, Kew Bull. 50: 715 (1995). Type: Colombia, Caquetá, Araracuara region, Quebrada El Engaño, D. Restrepo & A. Matapí 542 (holotype MO; phototype K).

distribution and habitat. Colombia; known only from the type collection. Habitat unknown.

31. Leptobalanus wurdackii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472887-2

Licania wurdackii Prance, Fl. Neotrop. Monogr. 9: 63 (1972a). Type: Venezuela, Amazonas, Río Atabapo, Caño Guasariapuna, 1 km above San Fernando, B. Maguire, J. J. Wurdack & G. S. Bunting 36238 (holotype NY; isotypes K, P).

distribution and habitat. Amazonian regions of Venezuela, Colombia and Brazil (Amazonas). Riverine forest and river banks.

VII. Moquilea Aubl., Hist. Pl. Guiane 1: 521, t. 208 (Aublet 1775).

Dahuronia Scop. (Scopoli 1777: 219), nom. superfl. illegit; Licania subg. Moquilea (Aubl.) Prance (1967: 224); Licania sect. Moquilea (Aubl.) Prance (1970: 527). Type: Moquilea guianensis Aubl.

Trees. Leaf lower surface glabrous or lanate, never with stomatal cavities. Petiole with or without glands. Inflorescence panicles, racemes or racemose panicles. Bracteoles small. Petals 4 – 5. Stamens 11 – 60 (– 90), in 1 – several rows (as in Couepia), inserted in a complete circle; filaments glabrous, usually far exceeding calyx lobes, less frequently only slightly exceeding or rarely equalling the calyx lobes, free or slightly united at base. Ovary lanate, tomentose or pilose, inserted at base of receptacle. Fruit globose to ellipsoid, to 11 – 15 × 10 cm, epicarp glabrous, smooth, lenticellate, crustaceous or pubescent; endocarp glabrous or sparsely pubescent within. Fig. 3C.

distribution. A genus with 54 species distributed from Mexico, Central America (Panama, Nicaragua, Costa Rica), the Leeward and Windward Islands, and throughout South America (Guianas, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil).

notes. Bentham (1840) maintained Moquilea as a distinct genus and stated that it had similar flowers to Chrysobalanus (all fertile stamens and in a complete circle), differing mostly in the inflorescence type. Moquilea shares several morphological characters with Couepia, such as number of stamens, exserted stamens, and leaf pilosity, but differs in the placement of the ovary. Moquilea is retrieved as sister to Couepia in all our analyses (combined, plastid, ITS and Xdh) and in a separate lineage from Licania s.s., Leptobalanus, Hymenopus and Chrysobalanus. It appears in two lineages (M. egleri and M. minutiflora distinct from the rest of Moquilea) but morphologically they are not distinguishable and so we have maintained them as Moquilea pending further analyses.

Key to species of Moquilea

• 1. Stamens exserted beyond calyx lobes, 12 – 90

• 2. Leaf lower surface with an appressed lanate-arachnoid pubescence

• 3. Inflorescence of fasciculate, short, dense-flowered racemes; primary veins impressed above; Panama……………..18. M. fasciculata

• 3. Inflorescence of lax, branched, racemose panicles or cymose panicles, or a raceme with long pedicels; primary veins usually prominulous or plane above

• 4. Inflorescence a few flowered (3 – 4) raceme; pedicels 5 – 7 mm long, thin; leaves long-cuspidate with thin acumen 15 – 25 mm long; Ecuador……………..32. M. longicuspidata

• 4. Inflorescence of lax branched racemose panicles or cymose panicles; pedicels sessile or rarely to 5 mm long, thick; leaves acuminate

• 5. Primary leaf veins 8 – 15 pairs, the lamina usually oblong-lanceolate

• 6. Leaves 4 – 11 × 1.1 – 4 cm, oblong-lanceolate

• 7. Inflorescence of once-branched racemose panicles; stamens c. 25; receptacle sessile……………..5. M. boliviensis

• 7. Inflorescence of much-branched panicles with many short branches bearing 1 – 3 flowers; stamens 12 – 14; receptacle shortly pedicellate……………..41. M. palcazuensis

• 6. Leaves 15 – 20 × 5.5 – 8.3 cm, oblong

• 8. Petioles 13 – 15 mm long, slightly winged; lamina coriaceous; Venezuela . . . 8. M. cariae

• 8. Petioles 5 – 8 mm long, not winged; lamina chartaceous, leaf apex cuspidate, acumen 5 – 10 mm; Ecuador……………..10. M. celiae

• 5. Primary leaf veins 16 – 29 pairs, the lamina oblong to elliptic

• 9. Inflorescence ramiflorous, borne on young woody twigs well below shoot apex

• 10. Flowers (calyx and receptacle) 6 – 7 mm long, young leaf lower surface with shaggy dense ferrugineous pubescence; leaves 27 – 47 cm long; Colombia, Ecuador

• 11. Leaves elliptic, 22 – 26 cm broad; petioles 20 – 25 mm long; stipules 10 – 12 mm long; Colombia (Valle)……………..21. M. gentryi

• 11. Leaves oblong, 9 – 10.5 cm broad; petioles 14 – 15 mm long; stipules 5 – 7 mm long; Ecuador (Napo)……………..26. M. hedbergii

• 10. Flowers (calyx and receptacle) 2 – 5.5 mm long, leaf lower surface with compact appressed pubescence or with caducous pubescence, becoming glabrous when mature; leaves 16 – 33 cm long

• 12. Leaf lower surface glabrous when mature, with a lanate caducous pubescence on youngest leaves; flowers c. 2 mm long; Colombia (Valle, 0 – 50 m)……………..53. M. velata

• 12. Leaf lower surface with a compact persistent grey-lanate or brown pubescence; flowers 3 – 5.5 mm long

• 13. Leaf lower surface ferrugineous to brown-tomentose, secondary venation not conspicuously reticulate

• 14. Stamens 13 – 15; stipules linear, 4 – 7.5 cm long; petioles canaliculate; Costa Rica……………..4. M. belloi

• 14. Stamens 25 – 30; stipules triangular-lanceolate, 3 – 7 mm long; petioles terete; Colombia, Ecuador, Peru……………..35. M. macrocarpa

• 13. Leaf lower surface grey-tomentellous, conspicuously reticulate with parallel secondary venation; leaves thickly coriaceous

• 15. Inflorescence 15 – 30 cm long, much-branched, flowers densely packed, bracteoles caducous; Colombia, Ecuador……………..15. M. durifolia

• 15. Inflorescence 8 – 15 cm long, unbranched or with a few branches; bracteoles persistent, membranous; Ecuador ……………..23. M. grandibracteata

• 9. Inflorescence terminal or axillary, but on leafy shoot apex

• 16. Flowers 1.5 – 2 mm long; leaves 12 – 19 × 3 – 6 cm, brown-pubescent beneath, without conspicuously reticulate venation; Colombia……………..54. M. veneralensis

• 16. Flowers 3 – 5 mm long; leaves 15 – 35 × 6 – 13 cm, grey-pubescent beneath, often with conspicuously reticulate parallel tertiary venation

• 17. Inflorescence and flowers with light-brown-tomentellous pubescence, leaves gradually tapering from mid-point to acuminate apex; centre primary veins far apart (18 – 22 mm), the secondary venation obscure; Colombia (Chocó)……………...37. M. maritima

• 17. Inflorescence and flowers with ferrugineous pubescence; leaves tapering from well above midpoint, centre primary veins 5 – 15 mm apart, with conspicuous parallel secondary venation

• 18. Leaves 6 – 9.5 × 2 – 3 cm; petioles 4 – 5 mm long; Venezuela (Lara)……………..40. M. montana

• 18. Leaves 12 – 46 × 3 – 26 cm; petioles 6 – 15 mm long

• 19. Leaf bases subcordate, with 2 stalked glands near junction of petiole; Costa Rica……………..13. M. corniculata

• 19. Leaf bases rounded to subcuneate, without stalked glands

• 20. Leaves oblong-lanceolate 20 – 37 cm long, primary veins impressed above

• 21. Primary leaf veins 21 – 24 pairs; petioles 6 – 8 mm long, terete; secondary venation conspicuously reticulate with parallel secondary veins; bracts 25 – 35 mm, persistent; Peru (San Martín, Pasco)……………..19. M. filomenoi

• 21. Primary leaf veins 34 – 40 pairs; petioles 11 – 15 mm long, canaliculate; secondary venation inconspicuous and covered by lanate pubescence, bracts caducous; Peru……………..52. M. vasquezii

• 20. Leaves oblong, 12 – 18 cm long; primary veins plane to prominulous above

• 22. Leaf apex cuspidate; petioles 6 – 8 mm long; flowers sessile; Ecuador……………..10. M. celiae

• 22. Leaf apex acuminate; petioles 10 – 12 mm long; flowers on pedicels 0.5 mm long; Colombia (Antióquia)……………..7. M. cabrerae

• 2. Leaf lower surface glabrous when mature (sometimes lanate when young)

• 23. Exterior of flowers and inflorescence glabrous or sparsely puberulous

• 24. Leaf broadly acuminate, the acumen 4 – 8 mm long; flowers 2.5 – 3 mm long; pedicels 2 – 5 mm long; Brazil (Bahia)……………..45. M. salzmannii

• 24. Leaf with well-developed finely pointed acumen 8 – 12 mm long; flowers 4 – 4.5 mm long; pedicels 1 – 3 mm long

• 25. Inflorescence and exterior of flowers puberulous; petioles 10 – 12 mm long; leaves elliptic, 5.5 – 9 cm broad; Peru (Loreto)…………………………..........……………….......…….30. M. klugii

• 25. Inflorescence and exterior of flowers glabrescent; petioles 5 – 7 (– 9) mm long; leaves oblong to oblong-elliptic, 2 – 5.5 cm broad; Guianas, Peruvian Amazonia …………….……25. M. guianensis

• 23. Exterior of flowers and usually inflorescence densely tomentose

• 26. Petals unguiculate

• 27. Flowers 5 – 6 mm long; pedicels 2 – 2.5 mm long; ovary pilose; Brazil (Amazonia)……………..51. M. unguiculata

• 27. Flowers 3 mm long; pedicels 0 – 0.25 mm long; ovary glabrescent; Amazonia (Brazil, Bolivia, Peru)……………..34. M. longipetala

• 26. Petals not clawed

• 28. Leaves 30 – 40 × 20 – 25 cm……………..38. M. megalophylla

• 28. Leaves smaller, 2 – 15 (30) × 1.5 – 10 (14)

• 29. Leaf veins and midrib impressed on upper surface

• 30. Petioles 10 – 13 mm long; leaves 10 – 14 × 2.5 – 5.2 cm; primary veins 11 – 17 pairs; stamens 17; western Amazonia……………..1. M. angustata

• 30. Petioles 5 – 7 mm long; leaves 14 – 25 × 4.5 – 8 cm; primary veins 18 – 22 pairs; stamens 12 – 14; Brazil (Pará)……………..2. M. anneae

• 29. Leaf veins plane or prominulous on upper surface

• 31. Leaves with a farinaceous pubescence when young

• 32. Inflorescence racemose or little-branched; petioles 4 – 6 mm long……………..50. M. tomentosa

• 32. Inflorescence much-branched; petioles 8 – 16 mm long……………..43. M. pyrifolia

• 31. Leaves glabrous when young

• 33. Inflorescence with flowers borne in cymules on short tertiary branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48. M. tachirensis

• 33. Inflorescence not in cymules

• 34. Pedicels 3 – 7 mm long; leaves with finely pointed acumen 1.5 – 2.5 mm long……………..33. M. longipedicellata

• 34. Pedicels 0.5 – 2.5 mm long; leaf acumen rather blunt or cuspidate, less than 10 mm long

• 35. Petioles 16 – 22 mm long

• 36. Petioles terete, thick, leaf apex rounded; stamens c. 22……………..28. M. jaramilloi

• 36. Petioles canaliculated, thin; leaf apex acuminate; stamens c. 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. M. cecidiophora

• 35. Petioles 2 – 15 mm

• 37. Stamens 25 – 30

• 38. Inflorescence a much-branched panicle……………..14. M. dodsonii

• 38. Inflorescence a racemose panicle

• 39. Leaves 21 – 26 cm long; primary veins 15 –19 pairs; petioles c. 10 mm long……………..17. M. espinae

• 39. Leaves 6 – 14 cm long; primary veins 8 – 11 pairs; petioles 4 – 6 mm long……………..31. M. leucosepala

• 37. Stamens 11 – 23

• 40. Leaves oblong-lanceolate, 13 – 22 cm long……………..42. M platypus

• 40. Leaves elliptic to oblong, usually smaller (15 – 18.5 in M. maranhensis; 10 – 21 in M. guatemalensis)

• 41. Leaf base subcordate; stamens 16 – 33

• 42. Leaves 15 – 18.5 cm long……………..36. M. maranhensis

• 42. Leaves 8 – 15.5 cm long……………..6. M. brittoniana

• 41. Leaf base rounded to cuneate; stamens 11 – 21

• 43. Leaves 2 – 4 × 1.1 – 2.7 cm……………..27. M. imbaimadaiensis

• 43. Leaves 4.5 – 16 × over 2 cm broad Brazil, Colombia, Peru

• 44. Flowers 4 – 5 mm long; stipules linear 5 – 7 mm……………..22. M. gonzalezii

• 44. Flowers 2 – 3.5 mm long; stipules to 2 mm long

• 45. Petioles 2 – 5 mm long; leaves obovate to elliptic

• 46. Inflorescence once-branched; leaves obovate to elliptic, 4.5 – 9 cm long…………….. . . . . . . . . . . . . . . . . . . . . . . . 16. M. egleri

• 46. Inflorescence twice-branched; leaves elliptic, 10 – 21 cm long……………..24. M. guatemalensis

• 45. Petioles 6 – 14 mm long; leaves oblong to oblong-elliptic

• 47. Leaf apex cuspidate; stamens 11 – 12……………..29. M. kallunkiae

• 47. Leaf apex acuminate; stamens 16 – 21

• 48. Inflorescence rachis and branches sparsely grey-pubescent……………..20. M. fritschii

• 48. Inflorescence rachis and branches brown or yellow-brown tomentose

• 49. Inflorescence little-branched; flowers 3 – 4 mm; ovary and petioles glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11. M. chiriquiensis

• 49. Inflorescence much-branched; flowers 2 – 3 mm; ovary pubescent; petioles tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39. M. minutiflora

• 1. Stamens equalling or shorter than calyx lobes, 10 – 17 (– 25 in L. salicifolia and L. silvatica)

• 50. Leaves 15 – 36 cm long; stipules 12 – 22 mm long; inflorescence borne on woody defoliated stalks ……………..53. M. velata

• 50. Leaves less than 14 cm long; stipules 2 – 10 mm long; flowers in terminal or axillary inflorescences on young branchlets

• 51. Inflorescence of unbranched subspikes or racemes

• 52. Flowers c. 1.5 mm long, borne in clusters along rachis, the rachis 4 – 15 mm long; Colombia (Boyacá)…………………………………………………......…………………47. M. subarachnophylla

• 52. Flowers c. 2.5 mm long, not clustered, the rachis 2 – 4 mm long; Peru (Madre de Dios) …………………………………………………..…………………………………49. M. tambopatensis

• 51. Inflorescence of panicles, or a panicle of cymules

• 53. Leaf lower surface densely lanate-pubescent

• 54. Leaf apex with a well-developed acumen 3 – 5 mm long; stamens c. 25; exterior of receptacle ferrugineous; Colombia (Antióquia)……………….………….………………...44. M. salicifolia

• 54. Leaf apex obtuse to broadly acuminate, the acumen not exceeding 1 mm; stamens c. 15; exterior of receptacle brown; Brazil (Goiás)………..……………………….........3. M. araneosa

• 53. Leaf lower surface glabrous

• 55. Leaf acumen 3 – 7 mm long; leaves 5 – 10 cm long; stamens c. 25; flowers sessile on primary branches of inflorescence, densely brown-tomentose; eastern Brazil……………..46. M. silvatica

• 55. Leaf acumen 10 – 15 mm long; leaves 7.5 – 14 cm long; stamens 12 – 14; flowers predominantly on short secondary branches of inflorescence, short grey-tomentose; Colombia (Valle)……………..12. M. chocoensis

1. Moquilea angustata (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472888-2

Licania angustata Prance, Fl. Neotrop. Monogr. 9: 53 (1972a). Type: Brazil, Amazonas, Mun. de São Paulo de Olivença, near Palmares, B. A . Krukoff8445 (holotype NY; isotypes A, BM, BR, F, K, LA, LE, MO, P, S, US).

distribution and habitat. Western Amazonia: Ecuador, Peru and Brazil. Lowland non-flooded forests.

2. Moquilea anneae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472889-2

Licania anneae Prance, Brittonia 31: 250 (1979c). Type: Brazil, Pará, Cuiabá-Santarém Hwy., km 1305, vicinity of igarapé José Preto, G. T. Prance, A. S. Silva, M. J. Balick et al. 25652 (holotype MG; isotypes AAU, FHO, K, MO, NY, P, RB, U, US).

distribution and habitat. Brazil; known only from the type collection. Disturbed forest beside road on marginally flooded and non-flooded areas.

3. Moquilea araneosa (Taub.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472890-2

Licania araneosa Taub., Bot. Jahrb. Syst. 21: 428 (Taubert 1896). Type: Brazil, Goiás, Serra Dourada, E. Ule 2885 (holotype B, destroyed; lectotype HBG, fide Prance 1972a; isolectotypes P, R, fragment F).

distribution and habitat. Brazil (planalto central). Cerrado and gallery forest.

4. Moquilea belloi (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472891-2

Licania belloi Prance, Kew Bull. 50: 709 (1995). Type: Costa Rica, Alajuela, Reserva Biológica Monteverde, Río Peñas Blancas, E. Bello 1321 (holotype K; isotypes CR, MO).

distribution and habitat. Costa Rica and Nicaragua. Lowland and premontane moist tropical forest, 0 – 800 m.

5. Moquilea boliviensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472892-2

Licania boliviensis Prance, Fl. Neotrop. Monogr. 9: 43 (1972a). Type: Bolivia, Dept. La Paz, Larecaja Province, 10 km S of Mapiri, B. A. Krukoff 111233 (holotype NY; isotypes A, F, K, LA, MICH, MO, S, US).

distribution and habitat. Bolivia. Terra firme forest.

6. Moquilea brittoniana (Fritsch) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60473137-2

Licania brittoniana Fritsch, Oesterr. Bot. Z. 42: 6 (1892), as britteniana. Type: Bolivia, Río Beni, near junction with Madre de Dios, H. H. Rusby 2442 (holotype NY; isotypes BM, E, F, GH, K, MICH, P, US).

Licania pallida Britton, Bull. Torrey Bot. Club 17: 9 (1890), non L. pallida Spruce ex Sagot (1883: 306).

Moquilea elata Pilg. (Pilger 1914: 136); Licania elata (Pilg.) Pilg. ex L. O. Williams (1936: 174). Type: Brazil, Acre, Seringal São Francisco, E. Ule 9446 (holotype B, lost; lectotype MG 14292, fide Prance & Sothers 2003a: 63; isolectotype K).

distribution and habitat. W Amazonia in Bolivia, Colombia, Ecuador, Peru and Brazil. Non-flooded and periodically flooded forest.

7. Moquilea cabrerae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472893-2

Licania cabrerae Prance, Brittonia 28: 210 – 212 (1976). Type: Colombia, Antióquia, Estación Experimental Florestal, Piedras Blancas, I. Cabrera R. 94 (holotype COL).

distribution and habitat. Colombia. Montane and cloud forests, at 2200 – 2550 m.

8. Moquilea cariae (Cardozo) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472898-2

Licania cariae Cardozo, Ernstia n.s. 1: 143 (1992), as carii. Type: Venezuela, Aragua, Parque Nacional Henri Pittier, A. Cardozo & D. Hidalgo 1666 (holotype MY).

distribution and habitat. Venezuela; known only from the type location. Cloud forest.

9. Moquilea cecidiophora (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472899-2

Licania cecidiophora Prance, Biotropica 10: 85 (1978). Type: Peru, Amazonas, E of Chávez Valdivia Military Post, E. Ancuash 752 (holotype NY; isotypes F, MO).

distribution and habitat. Peru. Rainforest on terra firme.

10. Moquilea celiae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472900-2

Licania celiae Prance, Mem. New York Bot. Gard. 64: 198 (1990). Type: Ecuador, Napo, Parque Nacional Yasuní, C. E. Cerón & F. Hurtado 3843 (holotype K; isotypes AAU, F, GB, MO, NY).

distribution and habitat. Ecuador and Peru. Lowland rainforest and premontane wet forest to 800 m.

11. Moquilea chiriquiensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472902-2

Licania chiriquiensis Prance, Brittonia 29: 154 (1977). Type: Panama: Chiriquí, Cerro Colorado, S. A. Mori & R. L. Dressler 7778 (holotype NY; isotype MO).

distribution and habitat. Panama. Cloud forest over 1200 m.

12. Moquilea chocoensis (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472903-2

Licania chocoensis Cuatrec., Fieldiana, Bot. 27: 65 (Cuatrecasas 1950). Type: Colombia, Chocó, Río San Juan, Quebrada del Taparal, J. Cuatrecasas 21455 (holotype F; isotype P).

distribution and habitat. Confined to Colombia. Pacific coastal forests.

13. Moquilea corniculata (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472904-2

Licania corniculata Prance, Kew Bull. 50: 141 (1995). Type: Costa Rica, Puntarenas, Cantón de Osa, Rancho Quemado, J. Marín 458 (holotype K; isotypes CR, MO).

distribution and habitat. Costa Rica (Puntarenas and Osa Peninsula). Lowland rainforest.

14. Moquilea dodsonii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472905-2

Licania dodsonii Prance, Fl. Neotrop. Monogr. 9S: 27 (1989). Type: Ecuador, Esmeraldas, Mun. de Lita, Lita-San Lorenzo km 19, P. Acevedo R. & D. C. Daly 1657 (holotype NY; isotype K).

distribution and habitat. Panama and Ecuador. Tropical rainforest.

15. Moquilea durifolia (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472906-2

Licania durifolia Cuatrec., Fieldiana, Bot. 27: 61 (Cuatrecasas 1950). Type: Colombia, Valle, Cordillera Occidental, Monte de La Guarida, J. Cuatrecasas 22251 (holotype F; isotypes COL, MAD, P).

distribution and habitat. Colombia, Ecuador and Peru. Montane and cloud forests, 500 – 2000 m.

16. Moquilea egleri (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472909-2

Licania egleri Prance, Fl. Neotrop. Monogr. 9: 56 (1972a). Type: Brazil, Pará, Tapajós, rio Cururu, W. A. Egler 887 (holotype NY; isotypes IAN, MG 23789).

distribution and habitat. Amazonian Brazil and northern limits of the planalto of central Brazil and in Bolivia, Colombia, Peru and Venezuela. Open non-flooded forest, beside rivers and in tall cerrado (cerradão).

17. Moquilea espinae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472908-2

Licania espinae Prance, Kew Bull. 47: 633 (1992). Type: Colombia, Chocó, Bahia Solano Mun., Corregimiento El Valle between El Valle and Cabañas Punta R oca, J. Espina, F. Garcia & S. Pino 2879 (holotype CHOCO; isotype MO).

distribution and habitat. Colombia; known only from the type collection. Rainforest.

18. Moquilea fasciculata (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472910-2

Licania fasciculata Prance, Acta Amazon. 8 (4): 579 (1978). Type: Panama, Colón, Zona de Santa Rita, M. D. Correa A. & R. L. Dressler 1815 (holotype MO; isotype NY).

distribution and habitat. Panama. Forests.

19. Moquilea filomenoi (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472912-2

Licania filomenoi Prance, Fl. Neotrop. Monogr. 9S: 20 (1989). Type: Peru, Prov. Mariscal Cáceres, San Martín, 12 km W of Tocache Nuevo, Río Tocache, 500 m, A. H. Gentry et al. 25665 (holotype NY; isotypes F, MO).

distribution and habitat. Peru (San Martín and Pasco). Mature flatland forest on lateritic soil.

20. Moquilea fritschii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472913-2

Licania fritschii Prance, Fl. Neotrop. Monogr. 9: 59 (1972a). Type: Brazil, Pará, Belém, Reserva Mocambo, J. M. Pires & N. T. da Silva 6731 (holotype NY; isotypes IAN, UB).

distribution and habitat. Amazonian Brazil. Periodically flooded forest.

21. Moquilea gentryi (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472914-2

Licania gentryi Prance, Fl. Neotrop. Monogr. 9S: 24 (1989). Type: Colombia, Valle, Bajo Calima, 15 km N of Buenaventura, A. H. Gentry et al. 40355 (holotype MO; isotype NY).

distribution and habitat. Colombia. Lowland rainforest.

22. Moquilea gonzalezii (Miranda) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472915-2

Licania gonzalezii Miranda, Bol. Soc. Bot. México 29: 36 (1965). Type: Mexico, Nayarit, J. G. Ortega 41 (holotype MEXU; isotype US).

distribution and habitat. Southwestern Mexico. Gallery forest, 780 – 800 m alt.

23. Moquilea grandibracteata (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472916-2

Licania grandibracteata Prance, Fl. Neotrop. Monogr. 9S: 22 (1989). Type: Ecuador, Pichincha, Reserva Forestal ENDESA, J. Jaramillo 7413 (holotype NY; isotypes MO, QCA).

distribution and habitat. Ecuador, known only from the Andean foothills. Primary forest.

24. Moquilea guatemalensis (Lundell) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472917-2

Licania guatemalensis Lundell, Wrightia 5: 39 (1974). Type: Guatemala, Izabal, between Seja and Fronteras on Petén-Guatemala road, E. Contreras 10742 (holotype LL; isotypes P, US).

distribution and habitat. Guatemala; known only from the type collection. High forest.

25. Moquilea guianensis Aubl. (Aublet 1775: 521).

Licania guianensis (Aubl.) Griseb. (Grisebach 1857: 198). Type: French Guiana, J. B. C. F. Aublet s.n. (holotype BM; phototype BR).

distribution and habitat. Guianas, Venezuela, the Amazon delta, W Amazonia, Ecuador, Peru and in Brazil. Riverine, terra firme and premontane forests.

26. Moquilea hedbergii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472919-2

Licania hedbergii Prance, Opera Bot. 121: 279 (1993). Type: Ecuador, Napo, El Chaco Cantón, Tres Cruces, between Ríos Santa Rosa and El Salado, W. Palacios 6261 (holotype K; isotypes MO, QCNE).

distribution and habitat. Ecuador; known only from the type collection. Lower montane moist forest, 1600 m.

27. Moquilea imbaimadaiensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472921-2

Licania imbaimadaiensis Prance, Kew Bull. 47: 635 (1992).Type: Guyana, Cuyuni-Mazaruni region, Imbaimadai, L. J. Gillespie 2695 (holotype K; isotype US).

distribution and habitat. Guyana; known only from the type collection. Low sclerophyllous forest adjacent to rapids on conglomerate.

28. Moquilea jaramilloi (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472922-2

Licania jaramilloi Prance, Kew Bull. 47: 249 (1992). Type: Colombia, Amazonas, Quebrada El Sol, trib. of Río Caquetá, R. Jaramillo & P. A. Palacios 7940 (holotype COL).

distribution and habitat. Colombia; known only from the type collection. River margin.

29. Moquilea kallunkiae (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472924-2

Licania kallunkiae Prance, Acta Amazon. 8 (4): 583 (1978). Type: Panama, Colón, Santa Rita Rd., S. A. Mori & J. A. Kallunki 5052 (holotype NY; isotypes BM, CAS, K, MO).

distribution and habitat. Costa Rica and Panama. Wet forest.

30. Moquilea klugii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472925-2

Licania klugii Prance, Fl. Neotrop. Monogr. 9: 47 (1972a). Type: Peru, Loreto, Río Putumayo, Flórida, G. Klug 2221 (holotype NY; isotypes BM, F, GH, K, US).

distribution and habitat. Amazonian Peru. Riverine forest.

31. Moquilea leucosepala (Griseb.) R. O. Williams (1932: 315)

Licania leucosepala Griseb. (Grisebach 1857: 198). Type: Leeward Is., Guadeloupe, without locality, E. P. Duchassaing s.n. (holotype GOET; isotypes K, P).

distribution and habitat. Antilles from Guadeloupe to Trinidad, adjacent Venezuela and Colombia. Forests.

32. Moquilea longicuspidata (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472926-2

Licania longicuspidata Prance, Kew Bull. 50: 707 (1995). Type: Ecuador, Carchi, Tulcan Cantón, Parroquia Tobar Donoso, G. Tipaz et al. 1446 (holotype MO; isotype QCNE; phototype K).

distribution and habitat. Ecuador. Premontane tropical rainforest; 650 – 1800 m.

33. Moquilea longipedicellata (Ducke) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472927-2

Licania longipedicellata Ducke, Bull. Mus. Natl. Hist. Nat., Sér. 2, 4: 725 (1932). Type: Brazil, Amazonas, Fonte Boa, rio Solimões, A. Ducke s.n. (holotype RB 23603; isotypes K, P, S, US).

distribution and habitat. Colombia, Ecuador, Peru, and the upper part of Brazilian Amazonia. Riverine forest.

34. Moquilea longipetala (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472928-2

Licania longipetala Prance, Fl. Neotrop. Monogr. 9: 62 (1972a). Type: Peru, Loreto, Río Mazan, Gamitanacocha, J. M. Schunke 288 (holotype NY; isotypes A, F, S, US).

distribution and habitat. Amazonian regions of Peru, Bolivia and Brazil. Periodically flooded forest.

35. Moquilea macrocarpa (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472929-2

Licania macrocarpa Cuatrec., Fieldiana, Bot. 27 (2): 107 (Cuatrecasas 1951). Type: Colombia, Valle, Río Cajambre, Quebrada del Corosal, J. Cuatrecasas 17740 (holotype F; isotype COL).

distribution and habitat. Panama, Colombia, Ecuador and Peru. Lowland forests to 600 m and in premontane rain forest in Panama.

36. Moquilea maranhensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472930-2

Licania maranhensis Prance, Fl. Neotrop. Monogr. 9: 58 (1972a). Type: Brazil, Maranhão, Barra do Corda, A. Lisboa s.n. (holotype NY; isotypes MG 2462, RB).

distribution and habitat. Brazil; known only from the type collection. River margin.

37. Moquilea maritima (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472931-2

Licania maritima Prance, Fl. Neotrop. Monogr. 9: 44 (1972a). Type: Colombia, Chocó, south of Curiche, J. A. Duke 9659 (holotype NY).

distribution and habitat. Colombia (Chocó). Endemic to coastal forests.

38. Moquilea megalophylla (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472932-2

Licania megalophylla Prance, Kew Bull. 46: 105 (1991). Type: Ecuador, Pichincha, Centinela, km 12 Carretera Patricia Pilar-24 de Mayo, km 47 Santo Domingo -Quevado, en la cima de las montanas de Ila, C. H. Dodson, A. Gentry, W. Palacios & J. Zaruma 14484 (holotype MO; isotype K).

distribution and habitat. Western Ecuador. Lower montane forest 600 – 900 m.

39. Moquilea minutiflora Sagot (1883: 308).

Licania minutiflora (Sagot) Fritsch (1889: 54), non Licania minutiflora Cuatrec. Type: French Guiana, Maroni, M. Mélinon s.n. (holotype P; isotypes A, BM, F, G, K, NY, R, US).

Licania rondonii Pilg. (Pilger 1923: 540). Type: Brazil, Mato Grosso, Cáceres ("São Luiz de Cáceres"), F. C. Hoehne 4602 (holotype B, lost; lectotype designated here, R; isotype SP).

Moquilea riparia Gleason (in Gleason & Smith 1934: 192); Licania riparia (Gleason) Standl. (Standley 1937: 258). Type: Brazil, Maranhão, Maracaçumé, Campo da Boa Esperança, R. L. Fróes 1961 (holotype NY; isotypes BM, F, K, MICH, MO, P, S).

distribution and habitat. Guianas, Venezuela, Bolivia and Amazonian Brazil. Non-flooded forests and gallery forests.

40. Moquilea montana (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472933-2

Licania montana Prance, Brittonia 28: 212 (1976). Type: Venezuela, Lara, vic. Laguna Negra, Loma de Los Naranjos, J. A. Steyermark, R. Smith & C. Espinoza 111541 (holotype NY; isotypes P, VEN).

distribution and habitat. Venezuela; known only from the type collection. Cloud forest, at 1300 – 1500 m.

41. Moquilea palcazuensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472934-2

Licania palcazuensis Prance, PhytoKeys 42: 2 (2014). Type: Peru, Pasco, Oxapampa, Palcazu Distr., Parque Nacional Yanachaga-Chemillén, Cerro Panjil-Ozuz, Permanent plot tree 24, 10°10'S, 75°10'W, 850 m, A. Monteagudo, A. Peña, R. Francis et al. 8250 (holotype K; isotypes AMAZ, HUT, MO, MOL, USM).

distribution and habitat. Peru; known only from the type. Primary lowland rainforest.

42. Moquilea platypus Hemsl. (Hemsley 1878: 9).

Licania platypus (Hemsl.) Fritsch (1889: 53). Type: Panama, without precise locality, H. Cuming 1272 (lectotype K, fide Prance 1972a: 55; isolectotype MO).

distribution and habitat. From southern Mexico through Central America to northern Colombia. Forest, most common near rivers, but also in forest away from rivers.

43. Moquilea pyrifolia (Griseb.) R. O. Williams (1932: 314)

Licania pyrifolia Grisebach (1860: 230). Type: Trinidad-Tobago, Trinidad, without locality, W. Purdie s.n. (holotype K).

Moquilea macrocarpa Pittier (1938: 352). Type: Venezuela, Carabobo, Valencia in Parks and Gardens, H. F. Pittier 8202 (holotype VEN; isotypes F, GH, MICH, MO, US).

distribution and habitat. Martinique to Trinidad, adjacent Venezuela and Colombia. Lowland forest and riverine forest in open places. Widely cultivated for its edible fruit.

44. Moquilea salicifolia (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472935-2

Licania salicifolia Cuatrec., Fieldiana, Bot. 27: 111 (Cuatrecasas 1951). Type: Colombia, Antióquia, Rionegro, Bro. J. Daniel 477 (holotype US; isotypes COL, F, GH, NY).

distribution and habitat. Known only from the vicinity of the type locality in Colombia. A high altitude endemic.

45. Moquilea salzmannii Hook. f. (Hooker 1867: 21)

Licania salzmannii (Hook. f.) Fritsch (1889: 49). Type: Brazil, Bahia, without precise locality, P. Salzmann s.n. (holotype K; isotypes E, G, LE, MO, P).

distribution and habitat. Brazil (Espírito Santo and Bahia). Restinga.

46. Moquilea silvatica (Glaz. ex Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472936-2

Licania silvatica Glaz. ex Prance, Fl. Neotrop. Monogr. 9: 90 (1972a). Type: Brazil, Espírito Santo, Serra de Itabapoana, A. F. M. Glaziou 10702 (holotype C; isotypes B, K, P).

Licania silvatica Glaz. (Glaziou 1906: 195), nom. nud.

distribution and habitat. Brazil; known only from the type collection. Cerrado.

47. Moquilea subarachnophylla (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472938-2

Licania subarachnophylla Cuatrec., Fieldiana, Bot. 27: 110 (Cuatrecasas 1951). Type: Colombia, Boyacá, Orocué, O. Haught 2629 (holotype F; isotypes A, COL, NY, US).

distribution and habitat. Known only from central Colombia and Venezuela. Gallery forest along the rivers of the savannas.

48. Moquilea tachirensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472940-2

Licania tachirensis Prance, Fl. Neotrop. Monogr. 9S: 30 (1989). Type: Venezuela, Táchira, Río San Buena, 10 km W of La Fundación, R. L. Liesner, A. González & R. F. Smith 9606 (holotype NY; isotypes MO, VEN).

distribution and habitat. Venezuela; known only from the type collection. Primary forested areas on sandy soil, 700 – 1000 m.

49. Moquilea tambopatensis (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472981-2

Licania tambopatensis Prance, Fl. Neotrop. Monogr. 9S: 36 (1989). Type: Peru, Madre de Dios, Tambopata Nature Reserve, laguna Coco Cocha, V. A. Funk, B. Kahn & S. Wiser 8415 (holotype NY; isotype US).

distribution and habitat. Peru and Bolivia. Along river edges.

50. Moquilea tomentosa Benth. (Bentham 1840: 215)

Licania tomentosa (Benth.) Fritsch var. latifolia Hook. f. (Hooker 1867: 21); Licania tomentosa (Benth.) Fritsch (1889: 52). Type: Brazil, Pernambuco, G. Gardner 992 (holotype K; isotypes BM, CGE, E, GH, NY, OXF, P, S, US).

Pleragina odorata Arruda ex H. Kost. (Koster 1816: 499), nom. inval.

Licania tomentosa (Benth.) Fritsch var. angustifolia Hook. f. (Hooker 1867: 21). Type: Brazil, Piauí, G. Gardner 2559 (holotype K; isotypes BM, GH, NY, OXF, P).

distribution and habitat. Northeastern, southeastern and southern Brazil. Native to the coastal restinga forest, but widely cultivated outside the region, especially in Amazonia.

51. Moquilea unguiculata (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472982-2

Licania unguiculata Prance, Fl. Neotrop. Monogr. 9: 60 (1972a). Type: Brazil, Amazonas, Rio Negro, above mouth of rio Curicuriari, A. Ducke s.n. (holotype K; isotypes NY, RB 23599).

distribution and habitat. Colombia, Peru and Brazilian Amazonia. Non-flooded forest.

52. Moquilea vasquezii (Prance) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472983-2

Licania vasquezii Prance, Kew Bull. 47: 251 (1992). Type: Peru, Loreto, Caballococha, road to Cashillo-Cocha, R. Vásquez & J. Jaramillo 12726 (holotype K; isotype MO).

distribution and habitat. Peru; known only from the type collection. Primary swamp forest.

53. Moquilea velata (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472984-2

Licania velata Cuatrec., Fieldiana, Bot. 27: 60 (Cuatrecasas 1950). Type: Colombia, Valle, Río Calima, Quebrada La Brea, J. Cuatrecasas 21179 (lectotype F, fide Prance 1972a: 88; isolectotypes COL, MAD, P).

distribution and habitat. Colombia; collected only in the Pacific coastal regions of Valle. Forest.

54. Moquilea veneralensis (Cuatrec.) Sothers & Prance comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:60472985-2

Licania veneralensis Cuatrec., Fieldiana, Bot. 27: 109 (Cuatrecasas 1951). Type: Colombia, Valle, Río Yurumanguí, Veneral, J. Cuatrecasas 15786 (holotype F; isotype COL).

distribution and habitat. Northwestern Colombia. Terra firme forest.

VIII. Licania Aublet, Hist. Pl. Guiane 1: 119, t. 45 (Aublet 1775).

Hedycrea Schreb. (Schreber 1789: 160), nom. superfl. illegit. Type: Licania incana Aubl.

Licania sect. Cymosa Prance (1967: 226). Type: Licania hypoleuca Benth.

Licania sect. Pulverulenta Prance (1967: 226). Type: Licania coriacea Benth.

Trees. Leaf lanate, pulverulent-furfuraceous, pulverulent-farinaceous, or glabrous, with or without stomatal cavities. Petiole with a pair of glands present or not. Bracteoles usually much shorter than or rarely equalling calyx, not enclosing flower buds. Inflorescences racemose panicles, panicles or spikes. Petals absent. Stamens 2 – 7 (8 – 11), rarely more than 8 (8 – 11 only in L. mollis, L. stewardii and L. majuscula), unilateral or less frequently inserted in a complete circle; filaments shorter than or equalling the calyx lobes, mostly free to base (a few species with filaments slightly connate at base) and glabrous or less often sparsely pubescent. Ovary pilose, tomentose, hirsutulous, inserted at the base of receptacle. Fruit globose to elliptic, to 10 × 6 cm, epicarp tomentose, pubescent, pulverulent, or smooth; endocarp glabrous or variously pilose within. Fig. 3A.

distribution. A genus of c. 100 species, two of which imperfectly known, distributed throughout Central and South America and in the Leeward Islands, from Mexico and Costa Rica to southeastern Brazil.

notes. The genus Licania s.s. (= core Licania) now comprises the former sections Licania, Cymosa and Pulverulenta of subgen. Licania, and the apetalous species of sect. Hymenopus of the previous classification, and three new species described since the last monograph of the family (Prance & Sothers 2003a, b). A priori here we have not circumscribed sections Cymosa and Pulverulenta, despite species appearing in subclades within the Licania clade, and these may in fact represent sections of Licania s.s., but at present we are not proposing any sections. Licania s.s. holds morphological affinities with Leptobalanus (both genera are apetalous), but differ in the number and the exsertion of stamens. In Fig. 1 Licania appears in a clade with Hirtella, and distant to Leptobalanus and Moquilea.

Key to species of Licania (key does not include L. roraimensis and L. tepuiensis as they are known from incomplete material)

• 1. Leaf undersurface glabrous

• 2. Leaves thin and membranous, the base cuneate, the venation equally prominent on both surfaces; receptacle urceolate; east-central Brazil………………………………. . . . . . . . . . . . . . . . 38. L. glazioviana

• 2. Leaves thick and coriaceous, the base usually rounded to cordate, rarely rounded-subcuneate (to cuneate in L. marleneae), the venation obscure on upper surface; receptacle conical to globose- cupuliform

• 3. Leaves rounded to obtuse at apex, rarely exceeding 9 cm in length

• 4. Flowers c. 3 mm long; petioles terete; E Brazil. . . . . . . . . . . . . . . ………………………………….55. L. littoralis

• 4. Flowers c. 2 mm long; petioles usually canaliculate

• 5. Inflorescence and flowers with brown pubescence; stamens 3; venation of leaf lower surface often papillose; stipules intrapetiolar; Venezuela, Guyana. . . . . . . . . . . . . . . ………..33. L. fanshawei

• 5. Inflorescence and flowers grey-puberulous or glabrescent; stamens 5; venation of leaf lower surface smooth-papillose; stipules adnate to extreme base of petiole

• 6. Leaf 2.5 – 6.5 cm long, the apex and base rounded; Guianas. . . . . . . . . . . . . . . ……………48. L. irwinii

• 6. Leaf 5.5 – 9.5 cm long, the apex acute, the base cuneate; Brazil (C Amazonia) . . . . . . . . . . . . . . . ……………………………………………………………………….57. L. marleneae

• 3. Leaves distinctly acuminate at apex, usually exceeding 8 cm in length

• 7. Midrib slightly impressed above; petioles canaliculate; French Guiana . . . . . . . . . . . . . . . ………………………………………………………………….26. L. cyathodes

• 7. Midrib prominulous above; petioles terete

• 8. Flowers 2.5 – 3.5 mm long, ferrugineous-brown-pubescent; stipules 4 – 7 mm long, persistent; Venezuela, Guianas, Brazil (Amazonia). . . . . . . . . . . . . . . …..76. L. polita

• 8. Flowers 1.5 mm long, grey-brown-pubescent; stipules small, caducous; Colombia, Venezuela, Brazil (Pará). . . . . . . . . . . . . . . ……………………………85. L. silvae

• 1. Leaf undersurface pubescent, either pulverulent or tomentose or lanate

• 9. Flowers in small cymules, on long slender secondary branches (peduncles) less than 0.5 mm thick and attached to primary inflorescence branches; pedicels usually 0.25 – 3 mm long, rarely absent; fruit often very small, usually not exceeding 2 cm in length

• 10. Leaf lower surface with stomatal cavities or thick, coarse, deeply cut venation, lanate-arachnoid

• 11. Bracteoles large and enclosing groups of flower buds; flowers subsessile; peduncles of cymules short and rather thick; Venezuela, Guianas. . . . . . . . . . . . . . . …………………………...30. L. densiflora

• 11. Bracteoles small and not enclosing groups of flower buds; flowers pedicellate; peduncles of cymules long and slender

• 12. Inflorescence and exterior of flowers ferrugineous-tomentose; young stems ferrugineous- tomentellous (fruit 3 – 5 mm long, stipitate, densely tomentose, in L. cuprea)

• 13. Flowers c. 2 mm long; stipules c. 5 mm long; stamens 3; Guyana. . . . . . . . . . . . . . . ………24. L. cuprea

• 13. Flowers c. 6 mm long; stipules 7 – 10 mm long; stamens 6 – 7; Brazil (Espírito Santo) . . . . . . . . . . . . . . . ………………………………………………………………………….6. L. arianeae

• 12. Inflorescence and exterior of flowers grey-puberulous or grey- or brown-tomentose; young stems glabrous or puberulous (fruit 1 – 2 cm long, not stipitate)

• 14. Midrib impressed for entire length; petioles 7 – 14 mm long; leaves oblong- lanceolate; Brazil (Amazonia). . . . . . . . . . . . . . . …………………………………….45. L. impressa

• 14. Midrib plane throughout or impressed at base only; petioles 2 – 10 mm long; leaves ovate to oblong

• 15. Leaf apex rounded; leaves orbicular-ovate, petioles 8 – 10 mm long; stamens 7; E Brazil. . . . . . . . . . . . . . . ……………………………………………………..83. L. santosii

• 15. Leaf apex acute to acuminate; leaves elliptic; petioles 2 – 7 mm long; stamens 3 – 7

• 16. Leaf lower surface with lanate pubescence obscuring venation; stamens 6 – 7; C Brazil. . . . . . . . . . . . . . . ………………………………………….29. L. dealbata

• 16. Leaf lower surface pubescent only in the mouth of stomatal cavities, the reticulations glabrous; stamens 3 – 5

• 17. Leaves ovate, broadest near base, 3 – 8.5 cm long; stipules axillary; Widespread. . . . . . . . . . . . . . . ………………………………………..44. L. hypoleuca

• 17. Leaves oblong-elliptic, broadest about middle, 5 – 13 cm long; stipules adnate to petiole base; Venezuela, Guianas, Brazil (Amazonia), Ecuador, Peru. . . . . . . . . . . . . . . …………………………...70. L. pallida

• 10. Leaf lower surface with a very fine, plane or prominulous venation, usually pulverulent-farinaceous

• 18. Exterior of flowers and branches of inflorescence entirely glabrous or glabrescent; fruit exterior often drying purple

• 19. Leaves oblong-lanceolate to oblong, the lower surface white-lanate; Brazil (Amazonia). . . . . . . . . . . . . . . ……………………………………………………………39. L. gracilipes

• 19. Leaves ovate to oblong, the lower surface sparsely grey-pulverulent-farinaceous; Venezuela, Guianas, Brazil (Amazonia), Colombia . . . . . . . . . . . . . . . ……………….73. L. parvifructa

• 18. Exterior of flowers and usually branches of inflorescence puberulous to tomentose; fruit exterior grey to brown

• 20. Leaf apex rounded, the margins revolute; inflorescence glabrous; Brazil (Pará, Bahia) . . . . . . . . . . . . . . . ……………………………………………………………27. L. cymosa

• 20. Leaf apex acuminate, the margins not revolute; inflorescence usually puberulous or tomentose

• 21. Fruit elongate-pyriform, to 2.5 cm long, the exterior with short rufous-velutinous pubescence; either inflorescence and exterior of flowers brown-tomentellous or petioles 7 – 12 mm long

• 22. Petioles 3 – 6 mm long, terete; inflorescence and flowers brown-tomentellous; leaves triangular or nearly so; W Indies. . . . . . . . . . . . . . . ……92. L. ternatensis

• 22. Petioles 7 – 12 mm long, canaliculate; inflorescence and flowers grey- puberulous; leaves oblong; Trinidad, Guianas, Amazonia . . . . . . . . . . . . . . . …………….…………………………………………………………59. L. membranacea

• 21. Fruit ovoid to pyriform, rarely exceeding 1.2 cm long; exterior of flowers grey-puberulous or glabrescent; petioles 3 – 6 mm long

• 23. Leaves 6.5 – 16 cm long; midrib and primary veins prominent on upper surface; rachis and branches of inflorescence glabrescent; Guianas, Brazil (Amapá) . . . . . . . . . . . . . . . …………………………………………….74. L. piresii

• 23. Leaves 2.5 – 10 cm long; midrib plane or prominulous on upper surface; rachis and branches of inflorescence puberulous

• 24. Inflorescence spreading, the rachis 1 – 15 mm thick; lower leaf surface pulverulent; Venezuela . . . . . . . . . . . . . . . …………………..36. L. furfuracea

• 24. Inflorescence compact, the rachis 0.5 mm thick; lower surface tomentellous; widespread . . . . . . . . . . . . . . . …………………………44. L. hypoleuca

• 9. Flowers sessile or subsessile on primary branches of inflorescence or on short (less than 0.5 mm long) thick peduncles only; pedicels usually sessile; fruit rarely less than 2 cm long

• 25. Leaf lower surface pulverulent-farinaceous

• 26. Leaf with rounded to acute apex, the margins usually revolute

• 27. Flowers 1.5 – 2 mm long; receptacle campanulate-cupuliform

• 28. Young branches and inflorescence densely tomentose; inflorescence to 4 cm long, recurved; Venezuela, Guyana . . . . . . . . . . . . . . . ………………………………………………...11. L. boyanii

• 28. Young branches and inflorescence branches glabrous; inflorescence usually exceeding 4 cm long, erect; Guyana . . . . . . . . . . . . . . . …………………………………………13. L. buxifolia

• 27. Flowers c. 3 mm long; receptacle urceolate

• 29. Leaves predominantly orbicular, occasionally oblong-elliptic, the apex rounded to retuse; Amazonia (Venezuela, Brazil) . . . . . . . . . . . . . . . ………………………68. L. orbicularis

• 29. Leaves elliptic, the apex acute

• 30. Receptacle narrowly urceolate-cylindrical; calyx lobes lanceolate; Brazil (Amazonas, Rondônia) . . . . . . . . . . . . . . . ……………………………………………..66. L. niloi

• 30. Receptacle broadly urceolate; calyx lobes deltoid; Venezuela, Guianas, Brazil (Amazonia) . . . . . . . . . . . . . . . …………………………………………………20. L. coriacea

• 26. Leaf with well-developed acumen, the margins not revolute

• 31. Flowers 3 – 4 mm long; receptacle campanulate or urceolate

• 32. Receptacle campanulate; leaves 8.5 – 10 cm broad; Ecuador . . . . . . . . . . . . . . . ……..25. L. cuyabenensis

• 32. Receptacle urceolate, leaves to 8 cm broad

• 33. Leaf apex caudate; leaves 4 – 5.5 cm long, chartaceous; exterior of receptacle red-brown-pubescent contrasting with white pubescence on interior of calyx lobes; Brazil (Rondônia) . . . . . . . . . . . . . . . ……………………………………………………………91. L. teixeirae

• 33. Leaf apex acuminate or acute; leaves 4 – 15 cm long, usually coriaceous; exterior of receptacle and calyx lobes grey-pubescent

• 34. Venation of leaf lower surface minutely reticulate, forming a network with less than 0.25 mm between reticulations; reticulation apparent because of absence of pubescence on veins; leaves thin-chartaceous; stipules usually caducous; western Amazonia (Peru, Colombia, Brazil, Venezuela) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96. L. urceolaris

• 34. Venation and reticulation coarse, with 1 – 2 mm between reticulations; pubescence obscuring much of veins; leaves thick-coriaceous; stipules persistent

• 35. Leaf apex with finely pointed, well-developed acumen; rachis of inflorescence glabrous; Costa Rica, Panama, Guianas, Brazil (Pará) . . . . . . . . . . . . . . . ………………………………………………………………….1. L. affinis

• 35. Leaf apex acute or with short blunt acumen; rachis of inflorescence pubescent; Venezuela, Guianas, Brazil (northern Amazonia) . . . . . . . . . . . . . . . ………………20. L. coriacea

• 31. Flowers 1.5 – 2 mm long; receptacle campanulate

• 36. Primary veins slightly impressed on upper surface; fruit exterior sordid-rufous-pubescent; branches of inflorescence densely tomentose to puberulous

• 37. Leaves thick-coriaceous; stipules caducous; flowers in clusters on short thick peduncles; stamens 3; Colombia (Pacific Coast) . . . . . . . . . . . . . . . ………………………37. L. glauca

• 37. Leaves thin, chartaceous-membranous; stipules persistent; flowers on primary and secondary branches of inflorescence; stamens 6 – 7; Guianas, Brazil (Amapá, Pará) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. L. davillifolia

• 36. Primary veins plane or prominent on upper surface; fruit exterior glabrous, drying yellow; branches of inflorescence glabrous to puberulous

• 38 Leaves 11 – 13 (– 18) × 4 – 8 cm; Guianas, Brazil (Amazonia) . . . . . . . . . . . . . . . ………32. L. elliptica

• 38. Leaves 4 – 10 (– 12) × 2 – 5.5 cm; Amazonia (Bolivia, Brazil, Guianas, Peru, Venezuela) …………………………………………………………………………...........15. L. canescens

• 25. Leaf lower surface densely lanate-arachnoid or with stomatal cavities, never pulverulent

• 39. Flowers 6 – 7.5 mm long, stamens often connate for ½ their length or free

• 40. Stamens connate for ½ their length; leaves prominently reticulate but without stomatal cavities; petioles eglandular; Guyana, Suriname . . . . . . . . . . . . . . . …………………………….21. L. couepiifolia

• 40. Stamens free to base; leaves with conspicuous stomatal cavities; petioles with 2 glands near base of lamina; Brazil (Amapá) . . . . . . . . . . . . . . . …………………………………………..64. L. naviculistipula

• 39. Flowers not exceeding 5.5 mm in length; stamens free almost to base

• 41. Leaf base distinctly cordate to subcordate; leaves usually ovate-orbicular

• 42. Leaves triangular-ovate, 10 – 16 cm long, membranous; Peru (Loreto) . . . . . . . . . . . . . . . …95. L. trigonioides

• 42. Leaves orbicular to ovate, 3 – 9 cm long, usually coriaceous

• 43. Young stems hispid; lower surface of leaves with hirsutulous-hispid venation, the apex with well-developed acumen; Venezuela (Amazonas) . . . . . . . . . . . . . . . ……………19. L. cordata

• 43. Young stems puberulous to tomentose; lower surface of leaves glabrous or lanate on venation, the apex acute or bluntly acuminate

• 44. Flowers 4.5 – 5.5 mm long; receptacle urceolate; stipules 5 – 6 mm long; Venezuela (Amazonas) . . . . . . . . . . . . . . . …………………………………………..35. L. foldatsii

• 44. Flowers 1.5 – 3.5 mm long; receptacle campanulate; stipules 1 – 3 mm long

• 45. Leaves submembranous, the lower surface with deeply cut venation, the pubescence occurring in cavities and hard to remove; fertile stamens 5 – 11

• 46. Stamens 8 – 11; petioles c. 5 mm long; stipules 3 – 6 mm long; Amazonian Colombia, Venezuela, Brazil . . . . . . . . . . . . . . . ………………..62. L. mollis

• 46. Stamens 5 – 6; petioles 1.5 – 3 mm long; stipules 2 – 2.5 mm long; Amazonian Venezuela, Colombia . . . . . . . . . . . . . . . ……………………41. L. hebantha

• 45. Leaves thick-coriaceous, the lower surface with shallow venation, not forming cavities, the lanate pubescence easily rubbed off; fertile stamens 3; Venezuela

• 47. Pubescence obscuring venation; stipules adnate to extreme base of petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88. L. steyermarkii

• 47. Pubescence not obscuring venation; stipules axillary . . . . . . . . . . . . . . . ………………………………………………………………………….90. L. subrotundata

• 41. Leaf base rounded to cuneate; leaves only rarely ovate-orbicular

• 48. Midrib and primary veins distinctly impressed on leaf upper surface

• 49. Leaf lower surface with well-developed stomatal cavities filled with lanate pubescence; petioles 8 – 17 mm long

• 50. Stomatal cavities conspicuous because of glabrous venation; leaves 4 – 8.5 cm long; Venezuela, Brazil (Amazonas) . . . . . . . . . . . . . . . …………………….22. L. crassivenia

• 50. Stomatal cavities less conspicuous because of puberulous venation; leaves (8 –) 10 – 25 cm long

• 51. Leaf apex acute; flowers 1.5 – 2 mm long; Ecuador . . . . . . . . . . . . . . . ……………4. L. apiknae

• 51. Leaf apex acuminate; flowers 1.5 – 5 mm long; Brazil and Guianas

• 52. Flowers c. 1.5 – 2.5 mm long; inflorescence much branched, predominantly axillary; fertile stamens 3 with 4 staminodes; petioles conspicuously 2 – 4-glandular; Suriname . . . . . . . . . . . . . . . ………………49. L. jimenezii

• 52. Flowers 3 – 5 mm long; inflorescence little branched, terminal; fertile stamens 6 – 11, staminodes absent; petiole glands inconspicuous

• 53. Receptacle conical, 4 – 5 mm long; fruit tomentellous, the stipe 2 – 6 mm long; leaf lower surface brown-lanate; Guianas, Brazil (Amazonas). . . . . . . . . . . . . . . ………………56. L. majuscula

• 53. Receptacle campanulate, 2.5 – 3 mm long; fruit pulverulent, the stipe 8 – 15 mm long; leaf lower surface white-lanate; Venezuela, Guianas, Brazil (Amazonas) . . . . . . . . . . . . . . . ………………………………2. L. alba

• 49. Leaf lower surface without stomatal cavities but often prominently reticulate; petioles 2 – 6 (– 8) mm long

• 54. Exterior of receptacle sparsely puberulous, the pubescence not completely covering the surface; leaves oblong-lanceolate; widespread . . . . . . . . . . . . . . . ……..50. L. kunthiana

• 54. Exterior of receptacle densely tomentose to tomentellous, the pubescence completely covering the surface; leaves oblong to elliptic

• 55. Leaves 2.5 – 5.5 cm long, not prominently reticulate beneath; petioles 1.5 – 3 mm long; Venezuela (Bolívar) . . . . . . . . . . . . . . . …………………………42. L. hitchcockii

• 55. Leaves 5 – 22 cm long, prominently reticulate beneath; petioles 4 – 8 mm long

• 56. Flowers c. 3.5 – 5 mm long; receptacle broadly cupuliform; leaves orbicular with rounded apex, the lower surface hirsute along venation; Guyana . . . . . . . . . . . . . . . ………………………………………………82. L. sandwithii

• 56. Flowers 2 – 3 mm long; receptacle campanulate; leaves oblong to elliptic, acute to acuminate at apex, the lower surface usually tomentellous, rarely hirsute on venation

• 57. Leaf lower surface with parallel secondary veins, giving a prominently reticulate appearance; pubescence brown; primary veins 7 – 10; Venezuela, Guianas, Brazil (Amazonia) . . . . . . . . . . . . . . . ………….…………………………………….……………….53. L. laxiflora

• 57. Leaf lower surface with diffuse secondary veins, less prominent; pubescence rufous; primary veins 5 – 6; Venezuela, Guianas . . . . . . . . . . . . . . . ……………………………………………………..81. L. rufescens

• 48. Primary veins and usually midrib plane or prominent on upper surface

• 58. Exterior of flowers and inflorescence branches grey-puberulous, the pubescence not completely covering surfaces

• 59. Receptacle campanulate; leaves without stomatal cavities; widespread…………….....………………………………………………………50. L. kunthiana

• 59. Receptacle urceolate; leaves with shallow stomatal cavities; Brazil (Rondônia)……………………………………………………...……….....9. L. bellingtonii

• 58. Exterior of flowers densely tomentellous or tomentose, the pubescence completely covering surfaces

• 60. Leaf apex rounded, obtuse or bluntly acute, or rarely short-apiculate (never acuminate)

• 61. Lanate pubescence of leaf lower surface hard to remove, covering deep stomatal cavities

• 62. Leaf veins 9 – 11; dense pubescence covering stomatal cavities receptacle urceolate; stamens 4; Guyana. . . . . . . . . . . . . . . ……………...17. L. compacta

• 62. Leaf veins 13 – 15; stomatal cavities visible; receptacle campanulate; stamens 7 – 8; Ecuador. . . . . . . . . . . . . . . …………………………..18. L. condoriensis

• 61. Lanate pubescence of leaf lower surface easily removed, revealing little protruding venation, stomatal cavities absent; receptacle usually campanulate, urceolate in L. ovalifolia and L. savannarum only

• 63. Receptacle urceolate; stipules 3 – 5 mm long, adnate to petiole well above base

• 64. Stamens 3; leaves thickly coriaceous; petioles 4 – 7 mm long; Guianas, Brazil (Amapá) . . . . . . . . . . . . . . . …………………………..69. L. ovalifolia

• 64. Stamens 9 – 11; leaves chartaceous; petioles 1.5 – 2.5 mm long; Brazil (Amazonas). . . . . . . . . . . . . . . ………………………………..87. L. stewardii

• 63. Receptacle usually campanulate, rarely urceolate; stipules 1 – 3 mm long, axillary or adnate to extreme base of petiole

• 65. Leaves ovate-orbicular, the apex often shortly apiculate

• 66. Lanate pubescence obscuring venation; stipules adnate to extreme base of petiole; Venezuela (Bolívar, Amazonas)…………..............……………....……………....88. L. steyermarkii

• 66. Lanate pubescence not obscuring venation; stipules axillary; Venezuela (Distrito Federal, Aragua) . . . . . . . . . . . . . . . ……………………………………90. L. subrotundata

• 65. Leaves oblong-lanceolate to elliptic, the apex rounded, acute or acuminate

• 67. Stipules adnate to petiole base; petioles 8 – 12 mm long; inflorescence branches very sparsely puberulous; stamens 5; Brazil (Bahia) . . . . . . . . . . . . . . . ……………………7. L. bahiensis

• 67. Stipules axillary, caducous; petioles 2 – 7 mm long; inflorescence branches densely puberulous or tomentellous; stamens 3

• 68. Leaves oblong-elliptic, sparsely lanate, the lower surface minutely reticulate; primary veins 9 – 11; petioles 4 – 7 mm long; receptacle campanulate; Colombia . . . . . . . . . . . . . . . …. ………………………14. L. caldasiana

• 68. Leaves oblong to oblong-lanceolate, the lower surface densely farinaceous-lanate with obscured venation; primary veins 5 – 8; petioles 2 – 3 mm long; receptacle campanulate-urceolate; Brazil, Venezuela (Amazonas) . . . . . . . . . . . . . . . ……………………….....84. L. savannarum

• 60. Leaf apex distinctly acuminate or sharply acute

• 69. Leaves 3 – 4 cm long, with 2 conspicuous glands at junction of petiole with upper surface of lamina; Guyana . . . . . . . . . . . . . . . …………………………61. L. microphylla

• 69. Leaves predominantly large, exceeding 5 cm long, lacking conspicuous glands

• 70. Stamens 3

• 71. Leaf lower surface with stomatal cavities

• 72. Leaf venation glabrous or glabrescent, hence conspicuous, with glands present at base of lower surface; leaves coriaceous; stipules less than 1.5 mm broad at base; Peru, Brazil, Colombian Amazonia . . . . . . . . . . . . . . . ………………….94. L. triandra

• 72. Leaf venation pubescent, lanate pubescence covering entire leaf and obscuring stomatal cavities of lower surface, lacking glands at leaf base; leaves chartaceous; stipules 2.5 mm broad at base; Brazil (Pará) . . . . . . . . . . . . . . . …………………………………93. L. tocantina

• 71. Leaf lower surface with prominent venation but no stomatal cavities, the venation pubescent and hence less conspicuous

• 73. Stipules caducous, petioles glabrous or tomentose; receptacle campanulate

• 74. Petioles tomentose when young; leaves elliptic to oblong, with well-developed acumen; primary veins 7 – 9 pairs; Venezuela, Guianas, Brazil . . . . . . . . . . . . . . . ………………...31. L. discolor

• 74. Petioles glabrous; leaves oblong-lanceolate, finely apiculate; primary veins 10 – 12 pairs; Brazil (Amazonia). . . . . . .3. L. apiculata

• 73. Stipules persistent, petioles glabrous or puberulous when young; receptacle campanulate or urceolate

• 75. Inflorescence 8 – 15 cm long, spreading, with numerous primary branches; lower leaf surface brown-lanate; receptacle campanulate; Colombia, Venezuela, Guianas, Brazil (Amazonia, Bahia) . . . . . . . . . . . . . . . ……………….60. L. micrantha

• 75. Inflorescence 5 – 6 cm long, with a few primary branches; lower leaf surface whitish-grey-pubescent; receptacle urceolate

• 76. Leaf apex with a finely pointed acumen; leaves thin- membranous; French Guiana, Brazil (Amapá). . . . . . . . . . . . . . . ……………………………....77. L. pruinosa

• 76. Leaf apex acute to bluntly acuminate; leaves coriaceous

• 77. Leaves thinly coriaceous, 4.5 – 10.5 × 2 – 5.5 cm; flowers 4 – 5 mm long; interior of receptacle pubescent on upper portion; E-central Brazil . . . . . . . . . . . . . . . ………………67. L. nitida

• 77. Leaves thickly coriaceous, 3 – 5.5 × 2 – 3 cm; flowers 2 mm long; interior of receptacle glabrous on upper portion; Brazil (NW Amazonia). . . . . . . . . . . . . . . ………………5. L. aracaensis

• 70. Stamens 4 – 8 (– 10)

• 78. Leaf lower surface with deep, extremely conspicuous stomatal cavities, the pubescence confined to cavities made obvious by almost glabrous venation

• 79. Inflorescence and flowers ferrugineous-pubescent; stipules 1 – 2 mm long × 1 – 1.5 mm broad at base, persistent, ferrugineous; leaf apex acute to bluntly acuminate; E-central Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78. L. riedelii

• 79. Inflorescence and flowers grey-puberulous; stipules 2 – 4 mm long, 0.2 – 0.5 mm broad at base, persistent or caducous, pubescent but not ferrugineous; leaf with well-developed acumen

• 80. Bracteoles persistent, lanceolate; upper surface of midrib impressed; stipules caducous; Brazil (Amazonia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12. L. bracteata

• 80. Bracteoles caducous, triangular; upper surface of midrib plane; stipules small but persistent

• 81. Inflorescence branches thick; flowers subsessile; fruit exterior ferrugineous-velutinous; Venezuela, Guianas, central and western Amazonia . . . . . . . . . . . . . . . …………..72. L. parviflora

• 81. Inflorescence branches slender; flowers distinctly pedicellate; fruit exterior reddish-brown, short-pulverulent; Venezuela, Brazil (Amazonas) . . . . . . . . . . . . . . . ……………………………………………..70. L. pallida

• 78. Leaf lower surface with poorly developed stomatal cavities or none, venation pubescent

• 82. Petioles 1.5 – 2 cm long; inflorescence rachis and branches glabrous or glabrescent; Guianas, Brazil (Pará) . . . . . . . . . . . . . . . ……79. L. robusta

• 82. Petioles to 1 cm long (to 1.5 cm in L. bahiensis and L. lamentanda); inflorescence rachis and branches usually densely tomentose or tomentellous

• 83. Inflorescence predominantly of axillary and terminal spikes, terminal inflorescences rarely little branched or with minute spikes along rachis

• 84. Leaf lower surface deeply reticulate, with poorly developed stomatal cavities

• 85. Leaves lanceolate, apex acuminate; stamens 6 – 8; Venezuela (Amazonas), Brazil (Roraima) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .52. L. lanceolata

• 85. Leaves ovate-elliptic, apex abruptly acuminate; stamens 4; Colombia, Venezuela (Aragua) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .75. L. pittieri

• 84. Leaves oblong to ovate-elliptic, the lower surface not deeply reticulate, lacking stomatal cavities

• 86. Flowers 2.5 mm long; receptacle broadly cupuliform; inflorescence puberulous; reticulate venation of leaf lower surface conspicuous; E-central Brazil . . . . . . . . . . . . . . . ………….………………………………………..86. L. spicata

• 86. Flowers 1.5 – 2 mm long; receptacle campanulate; inflorescence tomentose; venation of leaf lower surface inconspicuous except in L. nelsonii

• 87. Flowers in short, dense, minute spikes attached to long rachis; Suriname . . . . . . . . . . . . . . . ……………..89. L. stricta

• 87. Flowers solitary along rachis or in dense glomerules, but not in minute spikes

• 88. Leaf lower surface conspicuously reticulate; stipules adnate to base of petiole; receptacle rufous-pubescent; flowers densely and evenly clustered along inflorescence rachis; large tree; Brazil (Amazonas) . . . . . . . . . . . . . . . ………………...………………………………65. L. nelsonii

• 88. Leaf lower surface lanate, not conspicuously reticulate; stipules axillary; receptacle grey-brown pubescent; flowers in dense glomerules or small groups; small shrub to medium sized tree (15 m)

• 89. Leaves thin and membranous, the acumen finely pointed; flowers in dense glomerules; inflorescences largely axillary; Venezuela, Guianas, Brazil (Bahia, Pará) . . . . . . . . . . . . . . .54. L. leptostachya

• 89. Leaves thick and coriaceous, the acumen usually blunt; flowers in small groups or solitary but not glomerulate; inflorescences largely terminal; Venezuela, Guianas, Brazil (Amazonia) . . . . . . . . . . . . . . . ………………………….46. L. incana

• 83. Inflorescence of terminal and sub-terminal racemose panicles

• 90. Leaves lanceolate; low shrub or subshrub; Venezuela (Amazonas), Brazil (Roraima). . . . . .52. L. lanceolata

• 90. Leaves ovate to oblong-lanceolate; trees or tall shrubs

• 91. Stipules 3 – 10 mm long and at least 1 mm broad at base, distinctly adnate to petiole or intrapetiolar, persistent and obvious

• 92. Midrib distinctly impressed above; bracteoles 0.2 – 1.5 mm long

• 93. Inflorescence ferrugineous-pubescent; petioles canaliculate; stipules adnate to petiole well away from axil; Bolivia, Brazil (Pará) . . . . . . . . . . . . . . . . . . . . . . . . . . . .71. L. paraensis

• 93. Inflorescence grey-puberulous; petioles terete; stipules intrapetiolar or adnate to inside of extreme base of petiole

• 94. Leaves with a finely pointed acumen; primary veins 11 – 12 pairs, the lower surface glandular at base; Brazil, Colombia, Venezuela (northwestern Amazonia) . . . . . . . . . . . . . 97. L. vaupesiana

• 94. Leaves with blunt acumen; primary veins 6 – 8 pairs, the lower surface eglandular at base; E-central Brazil . . . . . . . . . . . . . . . . .7. L. bahiensis

• 92. Midrib plane or prominulous above; bracteoles 1.5 – 3 mm long

• 95 Leaves velutinous beneath; stipules fimbri-ate; Ecuador. . . . . . . . . . . 98. L. velutina

• 95. Leaves not velutinous beneath; stipules with entire margin

• 96. Leaves membranous; flowers c. 2 mm long, grey-tomentellous; inflorescence grey-puberulous; stamens 5; Brazil (Amapá) . . . . . . . . . . . . . . . ……………………………….58. L. maxima

• 96. Leaves coriaceous; flowers 3 – 3.5 mm long, brown-tomentose; inflorescence brown-tomentose; stamens 8 – 11; Amazonian Colombia, Peru, Venezuela, Brazil . . . . . . . . . . . . . . . …………………...……….62. L. mollis

• 91. Stipules usually less than (rarely exceeding) 2.5 mm long and very narrow to base, on outside of axils or adnate to extreme base of petiole, caducous or persistent, often inconspicuous

• 97. Stipules adnate to extreme base of petiole, persistent or subpersistent

• 98. Petioles 7 – 15 mm long; lamina 7 – 16 × 3 – 8 cm; stipules subpersistent

• 99. Petioles glabrous; leaves not deeply reticulate; Brazil (Amazonas) ……………………......……16. L. cidii

• 99. Petioles tomentellous even when old, leaf lower surface deeply reticulate; Brazil (Pará and central) . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. L. blackii

• 98. Petioles 2 – 6 mm long; lamina 3 – 12 × 2.2 – 5.5 cm; stipules persistent

• 100. Leaf lower surface with stomatal cavities. . . . . . . . . . . . . . . . . 72. L. parviflora

• 100. Leaf lower surface without stomatal cavities

• 101. Flowers 1.5 – 2 mm; inflorescence and flowers sparse- grey puberulous...……50. L. kunthiana

• 101. Flowers 2.5 mm; inflorescence and flowers dense-brown tomentose……….63. L. monteagudensis

• 97. Stipules axillary, caducous or persistent

• 102. Petioles 10 – 15 mm long, receptacle velutinous on exterior; Brazil (Bahia)

• 103. Leaves thickly coriaceous with recurved margins; apiculate at apex; flowers 4.5 – 5.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . .51. L. lamentanda

• 103. Leaves chartaceous, finely acuminate at apex; flowers 2 – 2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . 8. L. belemii

• 102. Petioles 2 – 10 mm long, receptacle rarely velutinous (only in L. rodriguesii)

• 104. Leaf lower surface with smooth inconspicuous reticulation, the pubescence easily removed; primary veins widely spaced, 1.2 – 2.5 cm apart; exterior of receptacle velutinous; Brazil (Amazonas) . . . . . . . . . . 80. L. rodriguesii

• 104. Leaf lower surface with deeply cut reticulation, and hence pubescence hard to remove; primary veins not more than 1 cm apart; exterior of receptacle tomentellous

• 105. Receptacle globose; upper surface of youngest leaves appressed-strigose, soon becoming glabrous; inflorescence of axillary spikes and terminal panicles; E-central Brazil . . . . . . . . . . . . . . .86. L. spicata

• 105. Receptacle campanulate; upper surface of youngest leaves glabrous; inflorescence of terminal and subterminal racemose panicles

• 106. Leaves oblong-lanceolate; midrib slightly impressed above . . . . . .47. L. indurata

• 106. Leaves ovate-elliptic to oblong; midrib plane above

• 107. Inflorescence much branched, spreading, lax; lower leaf surface rufous pubescent; Venezuela, Guianas, N Brazil.....81. L. rufescens

• 107. Inflorescence densely eroded, compact, little- branched; lower leaf surface grey-brown pubescent

• 108. Leaves finely acuminate, the acumen 8 – 12 mm long; stipules caducous; Ecuador . . . . . . . . . . . . . . .40. L. harlingii

• 108. Leaves bluntly acuminate, obtuse or with abrupt acumen 0 – 6 mm long; stipules usually persistent (caducous in L. ferreirae)

• 109. Stipules caducous; leaf apex abruptly acuminate, the acumen 4 – 6 mm long; young leaves with large glands at junction with petiole; Brazil (Amazonas)........34. L. ferreirae

• 109. Stipules persistent, 1 – 3 mm long; leaf apex obtuse, acute or bluntly acuminate; young leaves without large glands at junction with petiole

• 110. Petioles soon becoming glabrous; leaf apex bluntly acuminate; young fruit pyriform; Brazil (E-central)...43. L. hoehnei

• 110. Petioles tomentose, becoming less so with age; leaf apex obtuse to acute; young fruit cylindrical, becoming pyriform; Panama, Trinidad, Venezuela . . . . .23. L. cruegeriana

1. Licania affinis Fritsch (1889: 50). Type: Guyana, R. & R. Schomburgk 822 (holotype W; isotypes CGE, G, K, P).

Licania schomburgkiana Klotzsch in M. R. Schomb. (Schomburgk 1848: 1199), nom. nud.

distribution and habitat. Costa Rica, Panama, Guianas and N Brazil (Pará). Riverine and periodically flooded forest.

2. Licania alba (Bernoulli) Cuatrec. (Cuatrecasas 1964: 588).

Theobroma alba Bernoulli (1869: 14). Type: Guyana, ‘Essequibo and Cuyounie’, C. F. Appun 1 (holotype K).

Licania venosa Rusby (1920: 26). Type: Venezuela, Delta Amacuro, H. H. Rusby & R. W. Squires 423 (holotype NY; isotypes A, BM, F, GH, MICH, MO, S, US, VEN, Z).

Licania longifolia Benoist (1922: 252). Type: French Guiana, L. C. Richard s.n. (lectotype P, fide Prance & Sothers 2003a: 146).

distribution and habitat. Guianas and adjacent Venezuela and Brazil (Amapá, Amazonas, Pará). Primary forest.

3. Licania apiculata Prance (1972a: 151). Type: Brazil, Amazonas, Rio Cuieiras, W. A. Rodrigues & L. Coelho 4881 (holotype NY; isotype INPA).

distribution and habitat. N Brazil (Amazonas); known only from the Rio Cuieiras. Beaches and open habitats beside rivers.

4. Licania apiknae Prance (2014: 4). Type: Peru, Amazonas, Bagua Prov., Imaza Distr., Comunidad Aguaruna de Putuim, Monte Alto de Putuim, 5°00'54"S, 78°22'44"W, 500 m, C. Diaz, S. J. Kayip, & P. Atamain 7016 (holotype K; isotype MO).

distribution and habitat. Known only from Peru. Habitat unknown.

5. Licania aracaensis Prance (1976: 223). Type: Brazil, Amazonas, Serra Aracá, J. M. Pires 15027 (holotype IAN; isotypes INPA, NY, RB).

distribution and habitat. Brazil (Amazonas). Endemic to the summit of the sandstone mountain, at c. 1000 m.

6. Licania arianeae Prance (1989: 44). Type: Brazil, Espírito Santo, Linhares, Reserva Florestal CVRD, D. A. Foli 228 (holotype RB; isotypes CVRD, NY).

distribution and habitat. Brazil (Espírito Santo). Restinga and tabuleiro forests.

7. Licania bahiensis Prance (1972a: 165). Type: Brazil, Bahia, Ilhéus, H. P. Vellozo 736 (holotype R; isotype NY).

distribution and habitat. Brazil; known only from the type collection. Coastal forest.

8. Licania belemii Prance (1972a: 172). Type: Brazil, Bahia, Belmonte, R. P. Belém & Pinheiro 3211 (holotype UB; isotypes NY, RB).

distribution and habitat. E Brazil (Bahia and Espirito Santo). Common in forests.

9. Licania bellingtonii Prance (1972a: 145). Type: Brazil, Rondônia, Santa Bárbara, 15 km E of Km 117 of Porto Velho-Cuiabá Highway, G. T. Prance & J. F. Ramos 7000 (holotype NY; isotypes INPA, K, P, US).

distribution and habitat. NW Brazil (Amazonas and Rondônia). Non-flooded forest.

10. Licania blackii Prance (1972a: 167). Type: Brazil, Tocantins [Goiás], Araguaína, G. T. Prance & N. T. Silva 58989 (holotype NY; isotypes K, RB, UB).

distribution and habitat. Bolivia, Peru, and Brazil in Amazonia, and S to the borders of the planalto central. Gallery forest, and secondary forest in open places.

11. Licania boyanii Tutin (1940: 255). Type: Guyana, Bartica-Potaro road, T. G. Tutin 293 (holotype BM; isotypes K, NY, RB, U, US).

distribution and habitat. Venezuela and Guyana. Wallaba forest and forest on slopes.

12. Licania bracteata Prance (1972a: 155). Type: Brazil, Amazonas, Manaus, Estrada do Aleixo, A. Ducke 293 (holotype K; isotypes AA, F, IAN, NY, S, US).

distribution and habitat. Peru and Brazil (vicinity of Manaus and the lower rio Negro). Non-flooded forest.

13. Licania buxifolia Sandwith (1931: 369). Type: Guyana, Moraballi Creek, near Bartica, N. Y. Sandwith 327 (holotype K; isotypes NY, P, RB, U).

distribution and habitat. Guyana. Wallaba forest.

14. Licania caldasiana Cuatrec. (Cuatrecasas 1951: 106). Type: Colombia, without locality, J. C. Mutis 3868 (holotype US; isotypes F, MA).

distribution and habitat. Colombia; known only from the type collection, which lacks detailed field notes.

15. Licania canescens Benoist (1919: 514). Type: French Guiana, Maroni, M. Mélinon s.n. (lectotype P, fide Prance 1972a: 134; isolectotypes GH, K).

distribution and habitat. Guianas, Venezuela, Bolivia, Peru and widespread in Amazonian Brazil. Riverine and periodically flooded forest.

16. Licania cidii Prance (1992: 247). Type: Brazil, Amazonas, Mun. de Jutaí, São Francisco, C. A. Cid Ferreira et al. 8352 (holotype INPA; isotypes K, NY).

distribution and habitat. Peru and western Amazonian Brazil in the rio Solimões region. Forest on terra firme.

17. Licania compacta Fritsch (1889: 52). Type: Guyana, Roraima, R. & R. Schomburgk 519 (812) (holotype W; isotypes BM, BR, CGE, F, G, K, L, NY, OXF, P, W).

Licania flavicans Klotzsch in M. R. Schomb. (Schomburgk 1848: 1198), nom. nud.

distribution and habitat. Guyana (vicinity of Mt Roraima); known only from the type. Habitat unknown.

18. Licania condoriensis Prance (2013: 72). Type: Ecuador, Zamora-Chinchipe, El Pangui, Cordillera del Cóndor, plateau of Contrafuerte, Tres Patines, W of main Cóndor ridge above Jardin Botánico of EcuaCorriente Copper Company, 03°37'48"S, 78°26'50"W, 1685 m, D. Neill & W. Quizhpe 15076 (holotype K; isotypes MO, QCNE).

distribution and habitat. Known only from Ecuador, at mid-altitudes.

19. Licania cordata Prance (1972a: 136). Type: Venezuela, Amazonas, NW base of Cerro Yapacana, B. Maguire & J. J. Wurdack 34505 (holotype NY; isotypes K, P).

distribution and habitat. Venezuela (Cerro Yapacana, where it is reported to be abundant) and in Amazonian Brazil (Amazonas). Savannas.

20. Licania coriacea Benth. (Bentham 1840: 221). Type: Guyana, banks of Essequibo R., R. & R. Schomburgk 50 (holotype K; isotypes BM, BR, CGE, G, L, OXF, P, US).

Licania pallidula Standl. (Smith 1940: 286). Type: Venezuela, Bolívar, A. S. Pinkus 89 (holotype F; isotypes GH, K, NY, US).

distribution and habitat. Guianas, Venezuela and C & N Amazonia in Brazil. Non-flooded forest.

21. Licania couepiifolia Prance (1972a: 134). Type: Guyana, Shodikar Creek, Essequibo R., A. C. Smith 2839 (holotype NY; isotypes A, F, K, MAD, P, S, U, US).

distribution and habitat. Guianas. Lowland forest.

22. Licania crassivenia Spruce ex Hook. f. (Hooker 1867: 14). Type: Brazil, Amazonas, rio Uaupés, R. Spruce 2678 (lectotype K, fide Prance 1972a: 140; isolectotypes BM, BR, CGE, OXF, P, W).

distribution and habitat. Venezuela and Brazil (Amazonas). Caatinga.

23. Licania cruegeriana Urb. (Urban 1908: 352). Type: Trinidad-Tobago, Trinidad, Las Cuevas, H. Crueger 205 (2065 some sets) (holotype GOET; isotypes K, NY, TRIN).

distribution and habitat. Panama, N coastal Venezuela and Trinidad-Tobago. Forests and coastal locations.

24. Licania cuprea Sandwith (1937: 108). Type: Guyana, Demerara R., G. S. Jenman 6300 (lectotype K, fide Prance 1972a: 117).

distribution and habitat. Guyana. Wallaba forest and savanna margins. Flowering October to February.

25. Licania cuyabenensis Prance (1999: 104). Type: Ecuador, Sucumbios, Lago Agrio Cantón, Reserva Cuyabeno, Laguna Grande, W. Palacios, G. Tipaz, Rubio, Gudiño & Aulestia 9376 (holotype K; isotypes MO, QCNE).

distribution and habitat. Ecuador; known only from the type collection. Lowland primary forest on terra firme.

26. Licania cyathodes Benoist (1919: 513). Type: French Guiana, P. Sagot s.n. (lectotype P, fide Prance 1972a: 113).

distribution and habitat. French Guiana, Suriname and Venezuela. Riverine forests.

27. Licania cymosa Fritsch (1889: 47). Type: Brazil, Bahia, J. S. Blanchet 3200 (holotype W; isotypes F, G, GOET, OXF, P).

distribution and habitat. Northern and northeastern Brazil (Bahia, Pará). Littoral forest and in secondary forest.

28. Licania davillifolia Benoist (1919: 513). Type: French Guiana, Maroni, M. Mélinon s.n. (lectotype P, fide Prance 1972a: 133; isolectotypes A, BM, F, GH, K, R, US).

distribution and habitat. Guianas and in Brazil in E Amazonia (Amazonas, Amapá, Pará). Riverine forest and non-flooded forest.

29. Licania dealbata Hook. f. (Hooker 1867: 14). Type: Brazil, Bahia, Rio Preto, G. Gardner 2836 (holotype K; isotypes BM, CGE).

distribution and habitat. Planalto of Central Brazil and in the northeast. Cerrados.

30. Licania densiflora Kleinhoonte (in Pulle 1925: 383). Type: Suriname, Jan Passie, B. W. (Boschwezen) 5346 (lectotype U, fide Prance 1972a: 117).

Licania kanukuensis Standl. (in Smith 1939: 182). Type: Guyana, NW slopes of Kanuku Mts, Moku-Moku Creek, A. C. Smith 3420 (holotype F; isotypes A, IAN, K, LE, MO, NY, P, U, US).

distribution and habitat. Guianas and adjacent Venezuela and Brazil. Primary forest on high ground and slopes.

31. Licania discolor Pilg. (Pilger 1914: 137). Type: Brazil, Roraima, Surumu, Serra de Mairary, E. Ule 8393 (holotype B, destroyed; lectotype K, fide Prance 1972a: 151; isolectotypes L, LA, MG).

distribution and habitat. Guianas and adjacent Venezuela and Brazil. Forest on high ground, especially on slopes.

32. Licania elliptica Standl. (Standley 1937: 255). Type: Brazil, Amazonas, near mouth of rio Embira, trib. of rio Tarauacá, B. A. Krukoff 5014 (holotype F; isotypes A, BM, K, LE, M, MICH, MO, NY, S, U, US).

Licania paniculata Fanshawe & Maguire (Maguire 1948a: 323). Type: Suriname: Toekoemoetoe Creek, B. Maguire 24068 (holotype NY; isotypes A, BR, F, K, US).

distribution and habitat. Guianas and Amazonia (Peru and Brazil). Primary forest.

33. Licania fanshawei Prance (1972a: 112). Type: Venezuela, Delta Amacuro, Río Cuyubini, Cerro La Paloma, J. A. Steyermark 87557 (holotype NY).

distribution and habitat. Guianas and Venezuela, in the Orinoco delta. Primary forest and in the delta area.

34. Licania ferreirae Prance (1999: 104). Type: Brazil, Amazonas, Município de Alavarães, road Alvarães -Vila Nogueira, C. A. Cid Ferreira 8442 (holotype INPA; isotypes CAS, K, MO, NY).

distribution and habitat. NW Brazil; known only from the type collection. Terra firme forest.

35. Licania foldatsii Prance (1972a: 136). Type: Venezuela, Amazonas, Río Atabapo, Santa Cruz, E. Foldats 3857 (holotype VEN 49613; isotype NY).

distribution and habitat. Venezuela (Upper Orinoco and Río Negro). Savannas and river margins.

36. Licania furfuracea Prance (1976: 221). Type: Venezuela, Bolívar, El Dorado-Santa Elena Rd, km 251 – 253, J. A. Steyermark 111390 (holotype NY; isotype VEN).

distribution and habitat. Venezuela; known only from the type collection. Tall forest, beside a small stream.

37. Licania glauca Cuatrec. (Cuatrecasas 1951: 109). Type: Colombia, Valle, highway Buenaventura to Cali, E. P. Killip & J. Cuatrecasas 39000 (holotype F; isotypes COL, K, MO, NY, S, US).

distribution and habitat. Colombia and Ecuador. Pacific coastal forest.

38. Licania glazioviana Warm. (Warming 1874: 68). Type: Brazil, Rio de Janeiro, Rio de Janeiro, Tijuca, A. F. M. Glaziou 2561 (holotype C; isotypes BR, K, P).

distribution and habitat. Southeastern Brazil. Forests and possibly coastal restingas.

39. Licania gracilipes Taub. (Taubert 1892: 8). Licania gracilis Taub., loc. cit., nom. nud. Type: Brazil, without locality, A. F. M.Glaziou 13800 (holotype B, destroyed; lectotype K, fide Prance & Sothers 2003a: 129; isolectotypes C, P).

Licania duckei Maguire (1948c: 29). Type: Brazil, Amazonas, A. Ducke 248 (holotype NY; isotypes K, R, RB).

distribution and habitat. Amazonian Brazil, Venezuela and Colombia. Non-flooded forest.

40. Licania harlingii Prance (1979a: 9). Type: Ecuador, Napo, c. 6 km S of Puerto Napo, H. Lugo S. 1054 (holotype GB; isotype NY).

distribution and habitat. Western Amazonia: Bolivia, Colombia, Ecuador, Peru and Brazil (Acre). Terra firme forest.

41. Licania hebantha Mart. ex Hook. f.(Hooker 1867: 17). Type: Colombia, Amazonas, Monte Araracoara, C. F. P. Martius s.n. (holotype M).

distribution and habitat. Amazonian Colombia and Brazil (Amazonas). Savannas.

note. A leaf fragment at K was erroneously cited as an isotype of this species (Prance 1972a; Prance & Sothers 2003a), but is most probably a fragment from a collection by Wied-Neuwied. We are unable to identify to which species the fragment belongs.

42. Licania hitchcockii Maguire (in Maguire & Wurdack 1957: 477). Type: Venezuela, Bolívar, Uaipan Tepui, K. D. Phelps & A. S. Hitchcock 407 (holotype NY; isotype VEN).

distribution and habitat. Venezuela; known only from two inadequate collections. Forested slopes.

43. Licania hoehnei Pilg. (Pilger 1923: 541). Type: Brazil, São Paulo, São Paulo, Butantã, F. C. Hoehne 1494 (syntype B, destroyed; lectotype SP, fide Prance 1972a: 171; isotypes A, SPF, US).

distribution and habitat. Bolivia and east-central Brazil (Mato Grosso, Bahia, Espirito Santo, Minas Gerais, Rio de Janeiro, São Paulo). Forests.

44. Licania hypoleuca Benth. (Bentham 1844: 91). Type: Panama, R. B. Hinds s.n. (holotype K; isotype LE).

44a. Licania hypoleuca Benth. var. hypoleuca

Licania microcarpa Hook. f. (Hooker 1867: 15). Type: Venezuela, San Carlos, R. Spruce 3696 (lectotype K, fide Prance 1972a: 126; isolectotypes BM, BR, C, CGE, F, GH, GOET, LD, LE, NY, OXF, P, RB, TCD, W).

Licania grisea Kleinhoonte (in Pulle 1925: 382). Type: Suriname, Sectie O, B. W. (Boschwezen) 3392 (lectotype U, fide Prance, 1972a: 126; isotypes A, MO).

Licania parvifolia Pittier (1938: 351), nom. illegit. non L. parvifolia Huber (1909). Type: Venezuela, Bolívar, Ciudad Bolívar, E. G. Holt & W. Gehringer 69 (holotype VEN; isotypes F, NY, US).

Licania parviflora Benth. var. conduplicata Maguire (1952: 253). Type: Venezuela, Bolívar, J. A. Steyermark 60277 (holotype NY; isotypes F, VEN).

distribution and habitat. A wide ranging variety from Southern Mexico to Bolivia, the Guianas, Venezuela, Colombia, Peru and scattered localities in NE and Amazonian Brazil. Sandy soils in forest and savannas.

44b. Licania hypoleuca var. foveolata Prance (1972a: 127). Type: Brazil, Amazonas, rio Curicuriari, R. E. Schultes & López 9719 (holotype US; isotype IAN).

distribution and habitat. Brazil (Amazonas). Forest.

45. Licania impressa Prance (1972a: 118). Type: Brazil, Amazonas, Manaus, Colônia dos Franceses, A. Ducke s.n. (holotype K; isotype RB 25032).

distribution and habitat. Eastern and central part of Brazilian Amazonia. Non-flooded forest.

46. Licania incana Aubl. (Aublet 1775: 119).

Hedycrea incana (Aubl.) J. F. Gmel. (Gmelin 1796: 428); Chrysobalanus incanus (Aubl.) M. Gómez (1887: 39). Type: French Guiana, J. B. C. F. Aublet s.n. (holotype BM; isotype P-Rousseau; phototype BR).

Licania crassifolia Benth. (Bentham 1840: 221); Licania leptostachya Benth. var. crassifolia (Benth.) Benoist (1919: 512). Type: Guyana, R. & R. Schomburgk 388 (381 some sets) (holotype K; isotypes BM, CGE, L, NY, OXF, P, TCD, US, W).

distribution and habitat. Venezuela, Guianas, N and NE Brazil. Savannas, savanna margins and savanna forests.

47. Licania indurata Pilg. (Pilger 1923: 542). Type: Brazil, São Paulo, Santo André, Paranapiacaba, Alto da Serra, E. Schwebel 81 (holotype B, destroyed; lectotype R, fide Prance 1972a: 170; isotypes NY, SP, SPF; phototypes F, GH, US).

distribution and habitat. SE Brazil (Rio Janeiro and São Paulo).

48. Licania irwinii Prance (1972a: 113). Type: Suriname, Wilhelmina Mts, 3 km SSE of Juliana Top, H. S. Irwin, G. T. Prance, T. Soderstrom & N. Holmgren 55002 (holotype NY; isotypes K, P).

distribution and habitat. Guianas, Venezuela and Brazil (Amazonas). Forested slopes.

49. Licania jimenezii Prance (1972b: 7). Type: Suriname, Nickerie Distr., Falawatra, H. Jiménez-Sáa 1549 (holotype NY; isotypes LBB, P).

distribution and habitat. Known only from Guyana and Suriname. Rainforests.

50. Licania kunthiana Hook. f. (Hooker 1867: 16). Type: Guyana, Pirara, R. & R. Schomburgk 728 (lectotype K, fide Prance 1972a: 145; isolectotypes BM, BR, CGE, GH, L, NY, OXF, P, US, W).

Licania hypargyrea Malme (1930: 12). Type: Brazil, Mato Grosso, Chapada dos Guimarães (Santa Anna da Chapada), G. O. A.Malme 2378 (holotype S; isotypes LD, R).

Licania parviflora Benth. var. submembranacea Maguire (1952: 254). Type: Venezuela, Monagas, Quebrada Pajarral, Río Caripe, NE of Alto Aguacate, J. A. Steyermark 62173 (holotype NY; isotypes F, VEN).

distribution and habitat. A wide ranging species from Costa Rica to Bolivia, the Guianas, Venezuela, Peru and throughout Brazil. Non-flooded forest, riverine forest, cerrados, or secondary forest.

51. Licania lamentanda Prance (1989: 51). Type: Brazil, Bahia, Mun. de Ilhéus, 4 km N of Olivença, S. A. Mori, B. M. Boom & A. M. de Carvalho 13673 (holotype CEPEC; isotypes K, NY).

distribution and habitat. Brazil (Bahia). Restinga on sandy soil, 25 m altitude, growing with many individuals of piaçaba palm (Attalea funifera Mart. ex Spreng.).

52. Licania lanceolata Prance (1972a: 158). Type: Venezuela, Amazonas, Río Guainía, Sabana El Venado, B. Maguire & J. J. Wurdack 35568 (holotype NY).

distribution and habitat. Venezuela (Upper Orinoco region) and N Brazil. Savannas.

53. Licania laxiflora Fritsch (1889: 46). Type: Guyana, R. & R. Schomburgk 976 (holotype W; isotypes K, P).

Licania gracilis Kleinhoonte (Pulle 1925: 382). Type: Suriname, Gonini R., B. W. (Boschwezen) 3791 (holotype U; isotypes K, NY).

Licania macrophylla Klotzsch (in Schomburgk 1848: 1198), nom. nud.

distribution and habitat. Guianas and Amazonian Brazil and Venezuela. Non-flooded forest and clump wallaba forest.

54. Licania leptostachya Benth. (Bentham 1840: 220). Type: Guyana, bank of Rupununi R., R. & R. Schomburgk 111 (holotype K; isotypes CGE, F, G. L, LE, TCD, NY, P, US, W).

Licania incana var. axilliflora Sagot (1883: 305); Licania leptostachya var. axilliflora (Sagot) Fritsch (1889: 46); Licania axilliflora (Sagot) Hochr. (Hochreutiner 1910: 273). Type: Guyana, Roraima, R. & R. Schomburgk 868 (holotype NY; isotypes BM, K, IAN, P, W).

distribution and habitat. Guianas, Venezuela and northern and northeastern Brazil. Common along river margins and also found in Atlantic coastal Brazil.

55. Licania littoralis Warm. (Warming 1874: 67). Type: Brazil, Rio de Janeiro, Rio de Janeiro, Restinga da Tijuca, A. F. M. Glaziou 6168 (holotype C; isotypes F, K, P, R).

55a. Licania littoralis Warm. var. littoralis

distribution and habitat. E Brazil, from Paraíba to Rio de Janeiro. Confined to coastal restingas and coastal forests.

55b. Licania littoralis var. cuneata Kuhlm. (Kuhlmann 1940: 77). Type: Brazil, Espírito Santo, Linhares, Rio Doce, Lagoa do Durão, J. G. Kuhlmann 208 (holotype RB; isotype NY).

distribution and habitat. Brazil; known only from the type collection. In restinga vegetation.

56. Licania majuscula Sagot (1883: 306). Type: French Guiana, Maroní R., M. Mélinon s.n. (holotype P; isotypes BM, GH, K, NY, P, US).

Licania hostmannii Fritsch (1889: 42). Type: Suriname, W. R. Hostmann & A. Kappler 1250 (holotype W; isotypes BM, C, CGE, F, K, LE, NY, P, S, U).

distribution and habitat. Guianas and Amazonian Brazil. Upland forest and high riverine forest.

57. Licania marleneae Prance (1976: 218). Type: Brazil, Amazonas, Manaus-Porto Velho Hwy., between Castanho and Tupana rivers , M. F. Silva 873 (holotype INPA; isotype NY).

distribution and habitat. Brazil; known only from the type. Rainforest on terra firme.

58. Licania maxima Prance (1972a: 165). Type: Brazil, Amapá, Oiapoque, road between Oiapoque and Clevelândia, B. Maguire, J. M. Pires & C. Maguire 47081 (holotype NY; isotypes IAN, MG, P).

distribution and habitat. Brazil; known only from the type collection. Riverine forest.

59. Licania membranacea Sagot ex Laness. (Lanessan 1886: 130).

Licania galibica Benoist (1919: 515). Type: French Guiana, Karouary, P. Sagot 1081 (holotype P; isotypes BM, BR, GOET, K, NY, S, W).

distribution and habitat. Trinidad, the Guianas, Venezuela and Brazil. Forested slopes and non-flooded forest.

60. Licania micrantha Miq. (Miquel 1851: 20). Type: Suriname, W. R. Hostmann 1257 (holotype U; isotypes C, F, GH, K, LE, NY, P, W).

60a. Licania micrantha Miq. subsp. micrantha

Licania helvola Spruce ex Hook. f. (Hooker 1867: 18), nom. nud.

Licania anisophylla Standl. (Standley 1937: 253). Type: Brazil, Amazonas, B. A. Krukoff 8451 (holotype NY; isotypes A, BM, BR, F, K, MO, NY, P, S, US).

distribution and habitat. Widespread from Costa Rica to the Guianas, Venezuela, Colombia, Peru, Amazonian Brazil and Bolivia. Its known range has been recently extended west of the Andes in Colombia (Chocó and Valle) and in Brazil (Bahia). Non-flooded forests and Atlantic coastal forests.

60b. Licania micrantha subsp. atabapoensis Prance (1995: 717). Type: Venezuela, Amazonas, Atabapo Dept., Cabeceras del Río Yudi, F. Delgado 1799 (holotype PORT; isotype K).

distribution and habitat. Venezuela; known from the type collection. Forest on terra firme.

61. Licania microphylla Fanshawe & Maguire (Maguire 1948a: 322). Type: Guyana, Potaro R., D. B. Fanshawe 1078 (F. D. 3814) (holotype NY; isotype K).

distribution and habitat. Guyana; known only from the type collection. Forest.

62. Licania mollis Benth. (Bentham 1840: 219). Type: Brazil, Amazonas, R. & R. Schomburgk 910 (holotype K; isotypes BM, CGE, F, G, GH, L, NY, OXF, P, TCD, US, W).

Licania lucida J. F. Macbr. (Macbride 1934: 369). Type: Peru, Loreto, Mishuyaru near Iquitos, G. Klug 462 (holotype F; isotypes NY, US).

distribution and habitat. Amazonian Colombia, Peru, Venezuela and Brazil (Amazonas and Roraima). River banks and river islands and in riverine forest on sandy soil.

63. Licania monteagudensis Prance (2014: 6). Type: Peru, Pasco, Distr. Palcazu, Parque Nacional Yanachaga-Chemillén, Estación Biologica Paujil, 10°43'S, 74°54'W, 800 m, A. Monteagudo, G. Ortiz & R. Francis 5164 (holotype K; isotypes AMAZ, MO, MOL, USM).

distribution and habitat. Peru. Primary rainforest.

64. Licania naviculistipula Prance (1972a: 398). Type: Brazil, Espírito Santo, Linhares, Vale do Rio Doce, R. P. Belém 3814 (holotype NY; isotypes FHO, SP, UB).

distribution and habitat. E & SE Brazil (Bahia, Espírito Santo). Margin of secondary forest.

65. Licania nelsonii Prance (1989: 53). Type: Brazil, Amazonas, 3 km S of Serra Aracá, W. A. Rodrigues et al. 10501 (holotype INPA; isotypes NY, RB).

distribution and habitat. NW Brazil (Amazonas). Igapó forest in sandy soil.

66. Licania niloi Prance (1972a: 129). Type: Brazil, Rondônia, Porto Velho, N. T. Silva 392 (holotype NY; isotypes IAN, K).

distribution and habitat. Bolivia and N Brazil (Amazonas, Rondônia). Primary forest.

67. Licania nitida Hook. f. (Hooker 1867: 17).

Licania coriacea Benth. var. nitida (Hook. f.) Fritsch (1889: 49). Type: Brazil, Bahia, Jacobina, J. S. Blanchet 3290 (holotype BM; isotypes BR, F, G, K, LE, NY, P, W).

distribution and habitat. SE and C Brazil. Cerrados.

68. Licania orbicularis Spruce ex Hook. f. (Hooker 1867: 13). Type: Venezuela, Amazonas, Río Guainía, above mouth of Río Casiquiare, R. Spruce 3760 (lectotype K, fide Prance 1972a: 129; isolectotypes BM, BR, CGE, NY, P, TCD, W).

distribution and habitat. Amazonian Venezuela and adjacent Brazil (Amazonas). Riverine forest and savanna margins.

69. Licania ovalifolia Kleinhoonte (Pulle 1933: 180). Type: Suriname, Brownsberg, B. W. (Boschwezen) 6457 (holotype U; isotypes IAN, K, NY).

Licania stahelii Kleinhoonte (Pulle 1933: 181). Type: Suriname, Brownsberg, B. W. (Boschwezen) 6818 (holotype U; isotypes IAN, K, NY).

distribution and habitat. Guianas and Brazil (Amapá). Primary forest.

70. Licania pallida Spruce ex Sagot (1883: 306).

Licania parviflora var. pallida Hook. f. (Hooker 1867: 18). Type: Brazil, Amazonas, Managuiri, R. Spruce 1576 (lectotype P, fide Prance 1972a: 119; isolectotypes BM, BR, C, CGE, E, F, K, LE, M, NY, OXF, TCD ).

Licania pallida Spruce ex Hook. f. (Hooker 1867: 18), nom. nud., in synon.

distribution and habitat. Guianas and Venezuela to Brazilian Amazonia, Peru and Ecuador. Non-flooded forest.

71. Licania paraensis Prance (1972a: 163). Type: Brazil, Pará, Santarém, Piquiatuba, A. Ducke s.n. (holotype K; isotypes NY, RB 8818).

distribution and habitat. Amazonian Brazil, Peru and Bolivia. Non-flooded forest.

72. Licania parviflora Benth. (Bentham 1840: 221).

Licania parviflora var. subfalcata Spruce ex Hook. f. (Hooker 1867: 18). Type: Brazil, rio Uaupés, near Panuré, R. Spruce 2885 (holotype K; isotypes F, G, NY, RB).

Licania parviflora f. brevifolia Fritsch (1889: 51). Type: Brazil, Amazonas, rio Negro, R. & R. Schomburgk 977 (holotype K; isotypes BM, CGE, F, G, L, OXF, P, W).

Licania parviflora f. longifolia Fritsch (1889: 51). Type: Brazil, Amazonas, Tefé, E. Poeppig 2770 (holotype not traced).

distribution and habitat. Guianas, Venezuela, and Amazonia in Colombia, Peru, Brazil. Periodically flooded and riverine forest.

note. The type specimen of Licania parviflora f. longifolia, Poeppig 2770, was cited by Fritsch (1889: 51) as pro parte, with the type of Leptobalanus egensis (synonym of Leptobalanus octandrus subsp. pallidus) having the same number. We have only seen the part of the specimen which refers to Leptobalanus egensis.

73. Licania parvifructa Fanshawe & Maguire (Maguire 1948b: 374). Type: Guyana, Berbice-Rupununi Cattle Trail, A. A. Abraham 258 (lectotype NY, fide Prance 1972a: 121; isolectotypes K, U).

distribution and habitat. Guianas, Venezuela, Colombia and Amazonian Brazil. Primary forests.

74. Licania piresii Prance (1972a: 124). Type: Brazil, Amapá, rio Araguari, junction with rio Mururé, J. M. Pires, W. A. Rodrigues & G. C. Irvine 50435 (holotype NY).

distribution and habitat. Guianas and Brazil (Amapá). Non-flooded forest.

75. Licania pittieri Prance (1992: 249). Type: Venezuela, Aragua, P. N. Henri Pittier, A. Cardozo et al. 1397 (holotype MY; isotype K).

distribution and habitat. Venezuela (coastal Cordillera) and Colombia. Cloud forest at 1,100 – 2,200 m.

76. Licania polita Spruce ex Hook. f. (Hooker 1867: 17). Type: Brazil, Amazonas, rio Uaupés, near Panuré, R. Spruce 2676 (lectotype K, fide Prance 1972a: 115; isolectotypes BM, BR, C, CGE, E, F, GH, GOET, LD, LE, NY, OXF, P, RB, W).

Licania polita f. angustifolia Fritsch (1889: 42). Type: Brazil, Amazonas, rio Uaupés, Panuré, R. Spruce 2762 (lectotype K, fide Prance & Sothers 2003a: 124; isolectotypes BM, BR, C, F, GOET, LD, LE, OXF, P).

Licania poeppigii Fritsch (1889: 49). Type: Brazil, Amazonas, Tefé, E. Poeppig 2785 (holotype W; isotypes BR, G, GOET).

Licania laurifolia Huber (1909: 365). Type: Brazil, Pará, Oriximiná, rio Cuminá-mirim, lower rio Trombetas, A. Ducke s.n. (holotype MG 7958; isotype BM).

Licania laxa Fanshawe & Maguire (Maguire 1948a: 321). Type: Guyana, Essequibo R., Kuriki Rapids, D. B. Fanshawe 1665 (F.D. 4401) (holotype NY; isotype K).

distribution and habitat. Guianas, Venezuela and Amazonian Brazil. Non-flooded and periodically flooded forest.

77. Licania pruinosa Benoist (1919: 516). Type: French Guiana, Cayenne, L. C. Richard s.n. (holotype P).

distribution and habitat. French Guiana and adjacent Brazil. River and island margins.

78. Licania riedelii Prance (1972a: 155). Type: Brazil, Minas Gerais, Catas Altas, Serra do Caraça, L. Riedel 518 (holotype NY; isotype US).

distribution and habitat. SE Brazil (Minas Gerais and Rio de Janeiro). Forested regions.

79. Licania robusta Sagot (1883: 306). Type: French Guiana, Maroní R., M. Mélinon s.n. (1862) (holotype P; isotypes A, BM, F, GH, K, NY, R, US).

Licania pachystachya Kleinhoonte (Pulle 1925: 384). Type: Suriname: Sectie O, B. W. (Boschwezen) 2729 (holotype U; isotype P).

distribution and habitat. Guianas and Brazil (Pará). Primary forest.

80. Licania rodriguesii Prance (1972a: 168). Type: Brazil, Pará, Breves, rio Jaburuzinho, A. Ducke s.n. (holotype K; isotype RB 18812).

distribution and habitat. French Guiana and Brazilian Amazonia (Amazonas, Pará). Non-flooded forest.

81. Licania rufescens Klotzsch ex Fritsch (1889: 52). Type: Guyana, Roraima, R. & R. Schomburgk 601 (935B) (holotype W; isotypes BR, CGE, K, NY, P).

Licania rufescens Klotzsch in M. R. Schomb. (Schomburgk 1848: 1103), nom. nud.

distribution and habitat. Guianas and adjacent Venezuela and Brazil (Pará). Primary forest on slopes and high ground.

82. Licania sandwithii Prance (1972a: 142). Type: Guyana, Upper Mazaruni Distr., Tuibarodai trail, R. Boyan 92 (F.D. 7916) (holotype NY; isotype K).

distribution and habitat. Guyana and Central Amazonian Brazil (Amazonas). Terra firme forests.

83. Licania santosii Prance (1979b: 28). Type: Brazil, Bahia, Itacaré, J. Almeida & T. S. dos Santos 150 (holotype CEPEC; isotypes AAU, FHO, MO, NY, U, US).

distribution and habitat. Known only from Brazil (Bahia). Coastal region.

84. Licania savannarum Prance (1972a: 149). Type: Venezuela, Amazonas, Río Pacimoni, 50 km above mouth, B. Maguire, J. J. Wurdack & G. S. Bunting 37584 (holotype NY; isotype K).

distribution and habitat. Venezuela (Upper Río Orinoco region) and Brazil (Amazonas; Rio Negro region). Savannas or open places by river margins.

85. Licania silvae Prance (1972a: 115). Type: Brazil, Tocantins, Cariri do Tocantins, Belém-Brasília road, 30 km south of Gurupi, G. T. Prance & N. T. Silva 58968 (holotype NY; isotypes K, UB).

distribution and habitat. Amazonian Brazil, Colombia, Peru, Guianas and Venezuela. Non-flooded forest.

86. Licania spicata Hook. f. (Hooker 1867: 16).

Moquilea organensis Miers (1879: 374); Licania organensis (Miers) Fritsch (1889: 59). Type: Brazil, Rio de Janeiro, Organ Mts, J. Miers 4095 (holotype K; isotypes BM, P).

distribution and habitat. C and SE Brazil. Forested regions on hills.

87. Licania stewardii Prance (1976: 223). Type: Brazil, Amazonas, Manaus-Caracaraí Rd, km 130, W. C. Steward et al. P20251 (holotype INPA; isotypes FHO, K, MO, NY, P, US).

distribution and habitat. Colombia and northern Brazil in Amazonas and Roraima. Low campina forest around a sandstone rock outcrop in an area where the original forest has now been destroyed. Common around the vicinity of the type location.

88. Licania steyermarkii Maguire (1952: 254). Type: Venezuela, Bolívar, Río Karuai, base of Sororopan Tepui, 1220 m, J. A. Steyermark 60768 (holotype NY; isotypes F, MO, VEN).

distribution and habitat. Venezuela and Colombia. Forests.

89. Licania stricta Kleinhoonte (Pulle 1925: 380). Type: Suriname, beside the Maratakka River, B. W. (Boschwezen) 3463 (holotype U).

distribution and habitat. Suriname; known only from the type collection. River margin.

90. Licania subrotundata Maguire (1952: 255). Type: Venezuela, Distrito Federal, Cordillera del Avila, between Pico del Galipán and Boca del Tigre, J. A. Steyermark 56917 (holotype NY; isotypes F, VEN).

distribution and habitat. Venezuela (Coastal Cordillera). Confined to the highland cloud forest.

91. Licania teixeirae Prance (1989: 47). Type: Brazil, Rondônia, Itapuã do Oeste, Mineração Santa Bárbara, L. O. A.Teixeira 728 (holotype INPA; isotypes FHO, K, NY).

distribution and habitat. Brazil; known only from the type collection. Forest.

92. Licania ternatensis Hook. f. ex Duss (1896/1897: 259). Type: Leeward Is., Guadeloupe, Houëlmont, Vieux-Habitants, La Bouillante, Trois Rivières, A. Duss 2868 (lectotype NY, fide Prance & Sothers 2003a: 130; isolectotype US).

Licania ternatensis Hook. f. (Hooker 1893: 251), nom. nud.

distribution and habitat. Lesser Antilles from Guadeloupe to Grenada. Forests and forested slopes.

93. Licania tocantina Prance (1983: 28). Type: Brazil, Pará, Breu Branco, rio Tocantins, Tucuruí, M. G. Silva & R. Bahia 3508 (holotype MG; isotypes INPA, NY, RB).

distribution and habitat. N Brazil; known only from near the type locality. Terra firme forest.

94. Licania triandra Mart. ex Hook. f. (Hooker 1867: 18).

Licania micrantha Miq. var. triandra (Mart. ex Hook. f.) Fritsch (1889: 50). Type: Brazil, Amazonas, Panuré, rio Uaupés, R. Spruce 2490 (2409 in some sets) (lectotype K, fide Prance 1972a: 140; isolectotypes BR, C, CGE, GH, GOET, LD, LE, NY, P, W).

Licania pulchravenia Killip & Cuatrec. (Cuatrecasas 1951: 106). Type: Colombia, Vaupés, J. Cuatrecasas 7054 (holotype COL; isotype F (fragments)).

distribution and habitat. Amazonia: Venezuela, Colombia, Peru and Brazil (Acre, Amazonas). Non-flooded forest.

95. Licania trigonioides J. F. Macbr. (Macbride 1934: 368). Type: Peru, Loreto, near Mishuyacu, Iquitos, G. Klug 449 (holotype F; isotypes NY, US).

distribution and habitat. Peru; known only from the type. Forest.

96. Licania urceolaris Hook. f. (Hooker 1867: 15). Type: Brazil, Amazonas, rio Uaupés, R. Spruce 2422 (lectotype K, fide Prance 1972a: 131; isolectotypes BM, BR, C, CGE, E, GH, GOET, LD, LE, NY, OXF, P, RB, W).

distribution and habitat. Western Amazonia in Venezuela, Colombia, Peru, Brazil (Acre, Amazonas). Non-flooded forest.

97. Licania vaupesiana Killip & Cuatrec. (Cuatrecasas 1951: 105). Type: Colombia, Vaupés, Río Cuduyarí, tributary of Río Vaupés, J. Cuatrecasas 6837 (holotype COL; isotype fragments F, US).

Licania parviflora var. grandifolia Hook. f. (Hooker 1867: 19). Type: Colombia, Amazonas, C. F. P. Martius s.n. (holotype M).

distribution and habitat. Venezuela, Amazonian Colombia and Brazil (Amazonas). Riverine forest.

98. Licania velutina Prance (1992: 638). Type: Ecuador, Pastaza, Pastaza Cantón, 20 km S of Curaray, V. Zak & S. Espinoza 5052 (holotype K; isotypes MO, QCNE).

distribution and habitat. Ecuador; known only by two collections from a single locality. Tropical moist forest.

Imperfectly known species

1. Licania roraimensis Standl. (Smith 1940: 286). Type: Guyana, A. S. Pinkus 61 (holotype F; isotypes GH, K, NY, S, US).

distribution and habitat. Known only from the type collection, which is a very poor specimen with young fruit. No habitat information given. This is closely related to Licania micrantha and may be conspecific with it.

2. Licania tepuiensis Prance (1972a: 174). Type: Venezuela, Bolívar, Ilu-Tepui, Gran Sabana, B. Maguire 33389 (holotype NY).

distribution and habitat. Known only from the type collection; flowers unknown. Low open woodland at edge of mesa, 4500 ft. [c. 1350 m] elevation.

Appendix 1

ᅟ List of taxa used in the molecular analyses with an indication of markers obtained for each taxon. All DNA extracted from silica-gel dried leaf material, except those marked with an *, which were extracted from herbarium specimens at the Royal Botanic Gardens, Kew (K). Taxa in bold represent additions to our analyses that were not included in Sothers 2010 or Sothers et al. 2014. For Licania s.l., species names from the previous classification are used.