A Taxonomic Update of Neotropical Pradosia (Sapotaceae, Chrysophylloideae) Author(s): Mário H. Terra-Araujo, Aparecida D. de Faria, and Ulf Swenson Source: Systematic Botany, 41(3):634-650. Published By: The American Society of Plant Taxonomists URL: http://www.bioone.org/doi/full/10.1600/036364416X692389 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Systematic Botany (2016), 41(3): pp. 634–650 © Copyright 2016 by the American Society of Plant Taxonomists DOI 10.1600/036364416X692389 Date of publication August 26, 2016 A Taxonomic Update of Neotropical Pradosia (Sapotaceae, Chrysophylloideae) Mário H. Terra-Araujo,1,2,4 Aparecida D. de Faria,3 and Ulf Swenson2 1 Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Botânica (PPG-BOT), Av. André Araújo 2.936, 69067-375, Manaus, Amazonas, Brazil. 2 Department of Botany, Swedish Museum of Natural History, Box 50007, 104 05 Stockholm, Sweden. 3 Universidade Estadual de Londrina, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Box 6001, 86051-980, Londrina, Paraná, Brazil 4 Author for correspondence (araujo.mht@gmail.com) Communicating Editor: Chrissen E. C. Gemmill Abstract—We provide a systematic update of Pradosia (Sapotaceae, Chrysophylloideae), including overall morphology, a key to all species, comprehensive morphological descriptions, geographic distributions, and important characteristics for each species. Phylogenetic analyses based on molecular data demonstrated that the genus is monophyletic and includes three main clades. Twenty-three species of Pradosia are accepted, which are mostly distributed in lowland rainforests on either white-sand or clayish soils in tropical South America. A rotate corolla with a short tube, lack of staminodes, a drupaceous fruit with plano-convex cotyledons, an exserted radicle below the cotyledons, and the absence of endosperm are diagnostic for the genus. Two names are reduced into synonymy, viz. Pradosia atroviolacea Ducke, syn. of P. grisebachii (Pierre) T. D. Penn., and Pradosia verrucosa Ducke, syn. of P. glaziovii (Pierre) T. D. Penn. The affinity of P. argentea (Kunth) T. D. Penn., a species known only from the type collection, remains uncertain and for now excluded from the genus. Keywords—Amazon, Atlantic forest, extinct species, phylogeny Pradosia Liais (Sapotaceae, Chrysophylloideae) is a genus of 23 species of trees and shrubs that are mainly distributed throughout South America across a wide variety of habitats, including savannahs, evergreen and deciduous forests in the Amazon, the Brazilian Atlantic coast, and the Andes (Pennington 1991; Alves-Araújo and Alves 2012; Terra-Araujo et al. 2012, 2013). The Amazon region and the Brazilian Atlantic coast are major centers of diversity for the genus with 16 species (Pennington 2006; Terra-Araujo et al. 2012, 2013). Many of the species were previously known only from the type specimen (see Pennington 1990, 1991), or were described based on poor-quality collections, lacking fruits and/or flowers. This resulted in many species assessed to be either extinct or endangered (IUCN 2015). However, recent fieldwork conducted in these areas has located some of the presumed extinct or endangered species and revealed them being more abundant than previously reported (Terra-Araujo et al. 2015). The latest taxonomic revision of Pradosia was provided by Pennington (1990), but over the past twenty-five years it has become outdated, with newly described species and an accumulation of new material that has extended the geographical distribution of many species (Alves-Araújo and Alves 2012; Terra-Araujo et al. 2012, 2013). Recent phylogenetic analyses have demonstrated that Pradosia is monophyletic, provided that the African species Pradosia spinosa Ewango & Breteler (Ewango and Breteler 2001) is excluded from the genus (Anderberg and Swenson 2003; Swenson and Anderberg 2005; Swenson et al. 2008). Monophyly of Pradosia was initially inferred based on three species only, and a single morphological character, the drupaceous fruit, appears to be the only synapomorphy for the genus (Swenson et al. 2008; Terra-Araujo et al. 2012, 2013). A more detailed study, including 18 of the 23 species in Pennington’s revision, plus three recently described species (Alves-Araújo and Alves 2012; Terra-Araujo et al. 2012, 2013), was recently undertaken (Terra-Araujo et al. 2015). Results confirmed that Pradosia is monophyletic and seems to have originated in the Middle Eocene at 47.5 Ma somewhere in the rainforests of South America, likely in the Amazon basin, and may have reached the Brazilian Atlantic coast for the first time around 34.4 Ma. Historical Review Liais (1872) described Pradosia based on a single species (Lucuma glycyphloea (Casar.) Mart. & Eichler) published in Martius’s Flora Brasiliensis (Miquel 1863; pp. 82–83, Table 25). The generic name was instated in honor of “Visconde de Prados,” a person with political prominence who lived in Rio de Janeiro and Minas Gerais, Brazil, in the 19th century. Diagnostic features for Pradosia as circumscribed at the time included opposite leaves, absence of staminodes, fruits with a soft pericarp, and lack of endosperm. These characters were not attributes of either Chrysophyllum L. or Lucuma Molina, so Liais (1872) instated his genus Pradosia. Fifty years later, Ducke (1922) noticed in northeastern Amazonia that some trees with opposite or subopposite leaves and folded stamens (at least in bud), formed a distinct group within Sapotaceae, and named these Glycoxylon Ducke. In addition to the leaf arrangement, the inner bark of these species also shared the presence of a sweet, afterward astringent taste. Glycoxylon refers to the inner bark with a sweet taste, which is called “casca doce” or “pau doce” in northern Brazil. Ducke (1922) based the genus on Chrysophyllum inophyllum Mart. ex Miq. from Manaus in central Amazon and subsequently added two species discovered in the State of Pará (Glycoxylon huberi Ducke and G. pedicellatum Ducke). An additional species (Ducke 1925; G. praealtum Ducke), with alternate or subverticillate leaves, was also described from Pará. At that time, opposite leaves, folded stamens, and the sweet taste of the bark were no longer valuable generic characters. Ducke himself recognized the close relationship between his new genus Glycoxylon and the south Brazilian species Pradosia lactescens (Vell.) Radlk. and P. kuhlmannii Toledo. 634 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA Eyma (1936), after a detailed review, proposed the suppression of Glycoxylon and placed Pradosia within Pouteria Aubl. Cronquist (1946a, 1946b), on the other hand, in his revision of several American groups of Sapotaceae, maintained Pradosia and proposed new species and combinations in the genus. In a similar manner, Ducke (1942) also recognized Pradosia and provided a synopsis of the genus, and later an identification key together with a new species (Ducke 1953). In contrast, Pradosia and Glycoxylon were still considered as distinct groups by Aubréville (1964) in his Malacanthées, recognizing the genera based on leaf arrangement: opposite in Glycoxylon and alternate in Pradosia. One year later, Baehni (1965) united Pradosia with Chrysophyllum. Over the years, some authors have accepted Pradosia at the generic level, but retained some species in Chrysophyllum, Pouteria, Pometia Vell. (an illegitimate homonyn of Pometia J. R. Forst & G. Forst, Sapindaceae; renamed Neopometia by Aubréville), or even Lucuma (Aubréville 1961). Species of Pradosia have further been placed in Chrysophyllum, Ecclinusa Mart., Diploon Cronquist, Micropholis (Griseb.) Pierre, and Pouteria, demonstrating the difficulties there are to find the natural limits of genera in neotropical Chrysophylloideae (Swenson et al. 2008). Pennington (1990), in his treatment of neotropical Sapotaceae, united Glycoxylon with Pradosia and recognized a total of 23 species. He distinguished the genus from other neotropical genera by the rotate corolla with a short tube, lack of staminodes, a stipitate attachment between the anther and its filament, and a drupe (a drupaceous fruit). However, it is now clear that Pradosia including Glycoxylon forms a strongly supported group (Terra-Araujo et al. 2015). We here present a taxonomic update of Pradosia that incorporates novel molecular phylogenetic data, field observations, and the examination of herbarium collections, many of which were not available to Pennington. The taxonomic changes herein proposed include synonymy of two species names (i.e. Pradosia atroviolacea Ducke and Pradosia verrucosa Ducke), a review of the species geographical distribution, and a new diagnostic key followed by a short morphological description for each recognized species, as well as notes regarding species circumscription and field tips. Materials and Methods Pradosia was revised with full nomenclatural account in Flora Neotropica (Pennington 1990). Since few new nomenclatural changes are necessary here, nomenclature is limited for simplicity to the current accepted names and basionyms. This taxonomic update includes an identification key to species and their distributions, which are based on fieldwork conducted throughout the Neotropics by one of us (MHTA), bibliographic review, and examination of specimens deposited in Brazil (ALCB, CEPEC, CVRD, HRB, IAN, INPA, IPA, MBML, MG, PEUFR, RB, SP and UEFS), USA (CA, NY, MO, PH and US), and in Europe (K, P and S) (abbreviations follow Holmgren et al. 1990). A complete list of the examined specimens is provided in Supplemental Appendix S1. Morphological data were obtained from herbarium specimens and from fresh material collected from 2008 and onwards. Morphological features such as habit, bark patterns, flowers, fruits, and seeds were also studied in the field. The terminology used follows Harris and Harris (2001). A database with species georeferences was created from label data from 275 specimens to generate distribution maps in QGIS program (QGIS Development Team 2014). The Pradosia phylogeny used here is a simplified version including 21 of the 23 recognized species reported by Terra-Araujo et al. (2015). Here, we adopted the general lineage-based concept (GLC) (de Queiroz 1998) in delimiting species. Under GLC, a species is formed by 635 a separately evolving metapopulation lineage, where reproductive isolation, reciprocal monophyly or ecological divergence is reached independently at different times along the evolutionary history (de Queiroz 2007). Thus, a lineage does not have to be necessarily monophyletic, morphologically distinct, or reproductively isolated to be recognized as a species. In practice, any property that provides evidence of lineage separation is relevant to infer the boundaries and number of species. For this taxonomic treatment we considered ecology, species geographical distribution, phylogenetic relationships, and morphology important for the delimitation of the species. Two species here recognized, P. beardii (Monach.) T. D. Penn. and P. huberi (Ducke) Ducke, have not been tested phylogenetically because no samples for molecular data have been available. Results Habit — As in other neotropical Sapotaceae, Pradosia comprises woody taxa ranging in habit from shrubs such as Pradosia schomburgkiana (A. DC.) Cronquist, understory trees around 10–15 m (most of the species), to canopy trees up to 35 m tall like P. decipiens Ducke, P. cochlearia (Lecomte) T. D. Penn., P. glaziovii (Pierre) T. D. Penn., and P. kuhlmannii (Figs. 1A–B). Most tree species are buttressed, usually about 1 m high, but occasionally up to 2 m (Fig. 1C). Only P. brevipes (Pierre) T. D. Penn. has a specialized habit with most of the plant being below the ground (geoxylic subshrub), with only a few branches emerging above the ground. Leaves — The species have entire, alternate (Fig. 1D), subverticillate, verticillate (Fig. 1E), or opposite leaves (Fig. 1F); the latter condition is found in Pradosia beardii, P. huberi and P. schomburgkiana. The petioles are flat above (semiterete) or canaliculate, rarely carrying paired scales attached midway up the petiole as in P. grisebachii (Pierre) T. D. Penn. and P. lahoziana Terra-Araujo (Fig. 1G). The midvein may be deeply sunken or less frequently flat or raised on the upper leaf surface. The leaf venation of the majority of species is eucamptodromous, but eucampto-brochidodromous venation occurs when secondaries are joined by loops in the upper half of the blade but not in the lower half (Ellis et al. 2009). The secondary veins frequently are arcuate or sometimes straight. Intersecondaries are unusual, but if present they extend from the midvein and disappear or integrate into a higher venation pattern as in P. kuhlmannii, P. mutisii, and P. schomburgkiana (Fig. 1H). The tertiaries, if visible, are usually oblique-reticulate (Fig. 1I) or oblique (Fig. 1J). A horizontal-reticulate pattern (Fig. 1H) is a less common condition within the genus and only found in P. kuhlmannii, P. mutisii, and P. schomburgkiana. The leaf blade surfaces range from glabrous to sparsely pubescent (Fig. 1K), or densely covered in a reddish-brown indument as in Pradosia beardii (Fig. 1L). Trunk and Inner Bark — In addition to leaf and flower characters, the combination of some bark features are very useful as field characters at the species or subclade level. For example, a smooth bark of gray-brown or yellowish-gray color, usually bearing discolored scars (red or brown) are typical for Pradosia cochlearia, P. decipiens, P. glaziovii, P. kuhlmannii, P. schomburgkiana, P. subverticillata, P. surinamensis, and P. verticillata (Figs. 1M–T). In additional, a bark with sweet taste is found in P. cochlearia, P. kuhlmannii, P. schomburgkiana, P. subverticillata, and P. surinamensis, a character not present in any other genera of neotropical Sapotaceae. On the other hand, some species are characterized by a gray or brown colored bark with rather deep vertical fissures like in P. granulosa and P. longipedicellata (Figs. 1U–V). Most exude white latex 636 SYSTEMATIC BOTANY [Volume 41 Fig. 1. Habitat, leaf venation and bark variation in Pradosia (Sapotaceae, Chrysophylloideae). A–B. Habit. A. P. granulosa, an understory tree growing in rainforests of the Amazon region. B. P. kuhlmannii, a tall canopy tree from the Brazilian Atlantic coast. C. Buttresses to 1.5 m high in P. cochlearia. D–F. Leaf arrangement. D. Alternate in P. granulosa. E. Verticillate in P. verticillata. F. Opposite in P. schomburgkiana. G. Scales on the petiole of P. lahoziana. H–L. Lower leaf surfaces showing the midvein, secondaries or higher leaf venation and indument. H. P. schomburgkiana. I. P. longipedicellata. J. P. granulosa. K. P. cochlearia. L. P. beardii. M–V. Bark patterns and slash of ten species. The scale bar in H–L corresponds to 1 cm. Photos: M. H. TerraAraujo (A–E, G–V), A.Vicentini (F). 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA and the color of the inner bark ranges from light orange or yellowish in P. decipiens, P. glaziovii, P. granulosa, P. lactescens, P. lahoziana, P. ptychandra, P. subverticillata, and P. verticillata to pinkish or reddish in P. cochlearia, P. kuhlmannii, P. longipedicellata, P. montana, and P. schomburgkiana. Flowers — In Sapotaceae flowers are borne in fascicles and in Pradosia the fascicles usually appear below the apical clusters of leaves, rendering the majority of the species ramiflorous. Flowers of some species, viz. Pradosia lactescens, P. lahoziana, and P. ptychandra (Eyma) T. D. Penn., are borne on the trunk, making them cauliflorous (Fig. 1Q). One species, P. longipedicellata, has a terminal inflorescence (Fig. 2A). Flowers are bisexual and frequently 5-merous; the corolla is rotate with a short tube and spreading to flat corolla lobes. The color varies, ranging from white, green, and reddish to deep wine red (Fig. 2A–F). The stamens are exserted and equal the number of the corolla lobes. They are further usually attached in the corolla tube orifice (seldom just below the tube orifice). The filaments are long, geniculate in bud, becoming straight in open flowers, and are attached to the glabrous anthers by a narrow stipe (Fig. 2C). Staminodes are lacking, a trait considered diagnostic for the genus (Liais 1872; Ducke 1953; Pennington 1990; Swenson et al. 2008). The ovary is 5(–6)-locular and pubescent, in contrast to the glabrous style that is simple, without any visible stigmatic areas (cf. Planchonella; Swenson et al. 2007). Fruits — The fruits usually have an obovoid or ellipsoid form, less frequently ovoid, and normally with a smooth surface (Figs. 2G–I). Only Pradosia glaziovii (Pierre) T. D. Penn. and P. granulosa Pires & T. D. Penn. have fruits with small spicules covering the surface, giving the fruit a muricate aspect (Fig. 2J). The mesocarp is soft and most of the species are edible, having a sweet taste. In Pradosia, fruits have been characterized as a drupe with a thin cartilaginous endocarp (Pennington 1990, 1991; Swenson et al. 2008). However, it is better characterized as a drupaceous fruit, because the endocarp is rather jelly-like, partly transparent, never hardened (Terra-Araujo et al. 2012, 2013; Figs. 2K–N). Fruits are further considered as one-seeded, but more than one seed have been observed in P. cochlearia, P. granulosa, P. longipedicellata and P. restingae Terra-Araujo. The seeds are a little laterally compressed with a smooth, shinning testa, and with a seed scar as long as the seed itself; the cotyledons are plano-convex, the radicle is exserted below the cotyledons and there is no endosperm. Phylogenetic Relationships — A recent phylogenetic study of Pradosia was recently undertaken (Terra-Araujo et al. 2015). The results of this study support Pradosia as monophyletic and indicate P. longipedicellata as the sister to all other species of the genus (Fig. 3). According to this study, the species are grouped in three clades, which were strongly supported and are morphologically, geographically, and ecologically coherent. The Montane clade includes three species that have narrow distributions in the northeast corner of Colombia, Venezuela, and Ecuador (Fig. 4A). All three species are trees having an inner bark lacking a sweet taste, leaves with sunken midvein, fascicles of flowers borne along the branches (ramiflorous) of old wood below the leaves without any adjacent leaf scars (axillary in P. colombiana), and greenish flowers. The Sweetbark clade includes four species that are almost exclusively distributed in the Amazon region and have leaves with flat or raised midvein, fascicles of flowers below the leaves, 637 and greenish flowers with small corollas. The Red-flowered clade includes species from the Amazon region (eight species), the Brazilian Atlantic coast (four species), the savannah of central Brazil, and the Chocó region on the Pacific coast in Colombia. Within this clade the majority of species have non-sweet bark, leaves with sunken midvein, reddish or wine red flowers, and larger (> 4 mm long) corolla. This latter clade further includes plants with flowers borne on the trunk, scales attached at the middle of the petiole, and fruits with muricate surface, features not found in any other clade of Pradosia. Distribution and Habitat — Pradosia occurs throughout South America, although Pradosia grisebachii extends to Costa Rica and Panama. The highest diversity of Pradosia is found in the Amazon region (10 species), followed by the Brazilian Atlantic coast with six species (Fig. 4). Most of the species grow in lowland rainforest on either white-sand or clayish soils. Nine species are found in forests on white-sand soil, locally known as campina, campinarana, and restinga. Such patches of forests are nutrient-poor, have low pH and low water retention, resulting in completely different ecological conditions (Prance and Schubart 1978; Anderson 1981; Prance 1996). One species has been recorded in permanent flooded-forests, or swampy land in northeast Amazon (P. huberi). Only P. brevipes occurs in savannas ranging from 300–1000 m altitude in central Brazil. Three species, P. colombiana (Standl.) T. D. Penn. ex T. J. Ayres & Boufford, P. montana T. D. Penn., and P. mutisii Cronquist are found in dry montane forest up to 1200 m altitude in north and northwest South America. Conservation assessments — During the past years have Red List assessments for conservation using IUCN (2015) guidelines and criteria have usually accompanied each newly described or revised species. Assessments are possibly the most authoritative list to conservation of species. By using the IUCN criteria, species are categorized based on multiple sources of evidence, including population size, population decline, range area, and range decline. For the past century, many species of Pradosia have been, and are still, poorly documented. In the light of the IUCN assessments, this resulted in three extinct species, viz. P. argentea (Kunth) T. D. Penn., P. mutisii and P. glaziovii, where others were classified as Critically Endangered or Vulnerable (IUCN 2015). Recent fieldwork in the Amazon and the Atlantic forest have revealed new populations of almost all species. We here report previously unknown populations of P. cochlearia, P. decipiens, P. glaziovii, P. kuhlmannii, P. lahoziana, P. mutisii and P. subverticillata, which extend their distributions and point out that some of the presumed extinct or endangered species are more common than previously reported. For this reason, we refrain from red list assessments because with the current knowledge, most species would in fact be classified as data deficient (DD) and such assessments are not meaningful. Taxonomic Treatment Pradosia Liais, Clim. Géol. Géogr. Bot. Brésil 614. 1872.— TYPE: BRAZIL, Pradosia glycyphloea (Casar.) Liais. (= Pradosia lactescens (Vell.) Radlk.) Tall canopy trees, treelets, or rarely geoxylic shrubs; buttresses short or lacking, less frequently up to 2 m tall. 638 SYSTEMATIC BOTANY [Volume 41 Fig. 2. Flowers and fruits of Pradosia (Sapotaceae, Chrysophylloideae). A–F. Flowers. A. P. longipedicellata. B. P. schomburgkiana. C. P. montana. D. P. restingae. E. P. lactescens. F. P. granulosa. G–J. Fruit. G. P. restingae. H. P. lactescens. I. P. surinamensis. J. P. granulosa. K–N. Transection of fruits showing the half-transparent cartilaginous endocarp. K. P. cochlearia. L. P. granulosa. M. P. restingae. N. P. surinamensis. Photos: A. Alves-Araújo (D, G, M), R. I. Barbosa (I, N), J. Jardim (A), M. H. Terra-Araujo (E, F, H, J, K, L), A. Vicentini (B). 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA 639 Fig. 3. Phylogeny of Pradosia (Sapotaceae, Chrysophylloideae) obtained from Bayesian analyses using three nuclear molecular markers (ITS, ETS and RPB2) by Terra-Araujo et al. (2015) with herein adopted nomenclature. Posterior probabilities (PP) are indicated as black (PP > 0.95) and gray (0.85 < PP < 0.95) branches. Bark gray-brown, or yellowish-gray colored, smooth or rough, shallowly fissured or scaling in thin asymmetric plates leaving reddish, orange, or dark marks; slash of trunk light orange, yellowish, pinkish or reddish; inner bark non-sweet or sweet. Leaves simple, entire, sometimes revolute, usually alternate or subverticillate, less frequently verticillate or opposite, coriaceous or chartaceous, clustered at branch tips or scattered along branches; upper 640 SYSTEMATIC BOTANY [Volume 41 Fig. 4. Known geographic distribution to the species of Pradosia (Sapotaceae, Chrysophylloideae). A. Sister species and Montane clade. B. Sweet-bark clade. C. Clade “a” from Red-flowered clade. D. Clade “b” from Red-flowered clade. * = Species not included in the phylogeny of the genus. surface glabrous or with residual indument at the midvein and secondary veins, lower surface glabrous, pubescent, puberulous or tomentulose; trichomes malpighiaceous, Y-shaped, usually brownish, less frequently whitish, golden or gray; midvein usually sunken, less frequently flat or raised; secondary venation eucamptodromous, eucamptobrochidodromous or rarely brochidodromous, parallel or slightly divergent, flat, raised or impressed above; intersecondaries sometimes present, but rarely well-developed; tertiaries oblique, horizontal, reticulate, or a combination of these patterns; higher venation usually areolate; petiole semiterete or canaliculate, pubescent, glabrous or with some residual indument, sometimes with small scales, 0.5– 2.0 mm long, attached midway on the petiole. Flowers bisexual, clustered in fascicles, usually below the leaves (ramiflorous), sometimes on the trunk (cauliflorous), rarely clustered at the apices of short branches (terminal) or axillary; pedicel glabrous, usually tomentulose. Sepals are in a single whorl, 5(–6), usually ovate, with a rounded apex, glabrous, tomentose or tomentulose outside, glabrous inside, and persistent in fruit. Corolla is rotate with 5(–6) corolla lobes; corolla tube always shorter than the lobes, wine red, reddish, greenish or less frequently white, glabrous, sericeous or tomentulose outside, glabrous inside. Staminodes absent. Disk absent. Stamens opposite and equal the corolla lobes, glabrous; filaments geniculate in bud, slightly exserted in open flowers; anthers glabrous, cream or white. Ovary with 5(–6) locules, frequently puberulous or strigose; style simple, glabrous. Fruit drupaceous, ellipsoid, obovoid or ovoid, greenish, yellow or orange when mature, smooth, lenticellate or muricate, glabrous, puberulous or tomentulose, usually one-seeded; mesocarp soft and endocarp cartilaginous. Seeds usually laterally compressed; testa smooth, brown, shinning; seed scar adaxial, 100% of seed length, 10–25% of seed circumference; embryo with plano-convex cotyledons, exserted radicle, and no endosperm. Identification tips — The genus is very easy to recognize by its rotate corolla with a short tube, lack of staminodes and drupaceous fruits, unlike any other Sapotaceae in South America. Most species are distinguished by some vegetative features such as the taste of the inner bark, leaf arrangement, venation, leaf surface, and presence or absence of appendages on the petiole. If fertile specimens are present, useful characters include the position of the fascicles, the size and color of the corolla, and the fruit surface. Even the ecology is usually species specific. A combination of these characters usually leads to the right species as exemplified in Figs. 1 and 2. 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA 641 Key to the Species of PRADOSIA 1. 1. Geoxylic subshrub in savannahs from central Brazil to Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. brevipes Tree, treelets or shrubs in evergreen and deciduous forests, not in savannahs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Upper leaf surface with a flat or raised midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Secondary venation brochidodromous; intersecondaries present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaves alternate, chartaceous; intersecondaries present, short. On clayish soils in the Brazilian Atlantic coast . . . . . . . . . . P. kuhlmannii 4. Leaves opposite, coriaceous; intersecondaries long, extending towards the margin. In campina or campinarana forests throughout Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. schomburgkiana 3. Secondary venation eucamptodromous; intersecondaries absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Petiole semiterete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Secondary veins divergent. Lower leaf surface glabrous; ramiflorous; pedicel 1.0–4.0 mm long; corolla greenish. In swampy lands or flooded forest along the northeastern Amazon coast (estuary areas) . . . . . . . . . . . P. huberi 6. Secondary veins parallel. Lower leaf surface covered by whitish or ferruginous trichomes; inflorescence terminal; pedicels 18–30 mm long; corolla white. In restinga forested in southern Bahia . . . . . . . . P. longipedicellata 5. Petiole slightly canaliculate or canaliculate in the upper half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Inner bark with sweet taste; leaves chartaceous, glabrous or appressed brown-puberulous on the lower surface; petiole 0.7–1.6 cm long; pedicel 0.5–4.0 mm long; corolla 1.7–3.0 mm long; fruits 3.5–5.0 cm long . . . . . . . . P. cochlearia 7. Inner bark without a sweet taste; leaves coriaceous, glabrous; petiole 1.7–2.6 cm long; pedicel 5.0–13.0 mm long; corolla 4.5–5.2 mm long; fruits 3.0–3.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. decipiens 2. Upper leaf surface with a sunken midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Petiole with scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Ramiflorous; pedicel 1.0–3.0 mm long; corolla 4.0–5.0 mm long; fruits 3.5–5.0 cm long. Known from western Amazonia to Panama and Costa Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grisebachii 9. Cauliflorous; pedicel 7.0–9.0 mm long; corolla 6.4–9.5 mm long; fruits 2.5–3.0 cm long. Known from central Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lahoziana 8. Petiole without scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Lower leaf surface sparsely gray-puberulous, glabrous or glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Intersecondaries present and extending towards the margin. In dry montane forests of Colombia, Ecuador, and Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mutisii 11. Intersecondaries absent or short, not extending towards the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Fascicles mostly cauliflorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Pedicel 4.0–6.0 mm long; corolla usually glabrous. In rainforests on clayish soils in the Brazilian Atlantic coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lactescens 13. Pedicel 7.0–10.5 mm long; corolla sparsely tomentulose outside. In non-flooded forests on clayish soils in central and eastern Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ptychandra 12. Fascicles ramiflorous or axillary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Corolla ≥ 4.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15. Petiole 0.6–0.8 cm long. In forested and open vegetation on white-sand soils in northeast Atlantic coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. restingae 15. Petiole ≥ 0.9 cm long. In rainforests on sandy or clayish soils throughout Amazonia . . . . . . . . . . . . . . . . . . . . . 16 16. Inner bark without a sweet taste; lower leaf surface with indument at the midvein and secondary veins; petiole 0.9–2.0 cm long, canaliculate in the upper half; flowers wine red; fruit muricate. In rainforest on sandy soils in northeastern Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. granulosa 16. Inner bark with sweet taste; lower leaf surfaces glabrous; petiole 2.0–3.2 cm long, canaliculate; flowers greenish; fruit smooth. In forest on clayish or white-sandy soils in central and northwestern Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subverticillata 14. Corolla ≤ 4.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. Lower leaf surface sparsely gray-puberulous; fascicle axillary; pedicel 5.0–12 mm long. In deciduous forests in Colombia and Venezuela . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. colombiana 17. Lower leaf surface glabrous or glabrescent; fascicle ramiflorous; pedicel ≤ 2.5 mm long. In rainforest of Amazon region and northern South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18. Pedicel 1.5–2.5 mm long. Fruit 5.0–7.0 cm long, densely lenticellate, and glabrous. In wet forests of northern Colombia and north and northwest Venezuela . . . . . . . . . . . . . . . . P. caracasana 18. Pedicel 0.5–2.0 mm long. Fruit 2.0–4.0 cm long, not lenticellate, glabrous or puberulous. In rainforest on sandy or clayish soils in northeastern Amazonia, Brazil and Guyana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. surinamensis 10. Lower leaf surface densely tomentulose, puberulous or with erect two-branched pale-brownish trichomes . . . . . . . . . . . . . . . . . . . . 19 19. Corolla ≥ 5mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Leaves 20–40 cm long; petiole 2.0–2.6 cm long; pedicel 5.0–10 mm long; corolla ca. 7.5 mm long, greenish; fruit smooth. In wet lowland forest of Chocó region in Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cuatrecasasii 20. Leaves 9–24 cm long; petiole 0.8–2.2 cm long; pedicels ca. 1 mm long; corolla 5.0–6.5 mm long, reddish; fruit muricate. In rainforests on clayish soils throughout the Brazilian Atlantic coast . . . . . . . . . . . . . . . P. glaziovii 19. Corolla < 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21. Lower leaf surface with indument of erect two-branched pale-brownish trichomes; pedicel 2.0–3.0 mm long; corolla glabrous or sparsely tomentulose outside, especially the corolla tube. In deciduous coastal forests of Ecuador and Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. montana 21. Lower leaf surface densely reddish-brown tomentulose or brown-pubescent; pedicel 1.0–1.7 mm long; corolla sericeous or tomentulose outside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Leaves opposite or subverticillate, lower surface densely reddish-brown tomentulose; tertiaries obscure; corolla 2.0–3.4 mm long, greenish. Fruit 1.2–1.6 cm, brown-tomentulose. In scrublands or tall forests, usually on sandy soils of western corner of Guiana Shield . . . . . . . . . . . . . . . . P. beardii 22. Leaves verticillate, lower surface brown-pubescent; tertiaries visible; corolla 4.0–4.6 mm long, reddish. Fruit 3.5–4.5 cm long, glabrous. In non-flooded forests on clayish soils of central and eastern Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. verticillata 642 SYSTEMATIC BOTANY Pradosia beardii (Monach.) T. D. Penn., Fl. Neotrop. Monogr. 52: 665. 1990. ≡ Chrysophyllum beardii Monach., Phytologia 3: 159. 1949.—TYPE: TRINIDAD. Long Stretch Reserve, alongside main road near Turure junction, 23 Aug 1949 (fl), R. S. Ayliffe s.n. [Monachino 526] (holotype: NY 00273412!; isotype: U 0006709!). Shrub 1.5–3.0 m tall, sometimes tree up to 20 m; buttresses short; bark smooth, reddish, scaling in thin irregular plates and leaving deep marks; slash of trunk unknown. Leaves loosely clustered at branch tips, opposite or subverticillate, 5–16 × 2–6 cm long, oblanceolate or obovate, coriaceous, with a revolute margin; upper surface glabrous or with residual indument at the midvein and secondary veins, lower surface densely reddish-brown tomentulose; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, impressed above; intersecondaries absent; tertiaries oblique to horizontal, obscure; petiole 0.5–1.3 cm long, slightly canaliculate, pubescent, without scales. Flowers 3– many in each fascicle, ramiflorous; pedicel 1.0–1.5 mm long, tomentose. Sepals are tomentose outside. Corolla is 2.0– 3.4 mm long, greenish, tomentulose outside. Fruit 1.2–1.6 cm long, ellipsoid or obovoid, smooth, and brown-tomentulose. Distribution and habitat — This species is found in the western corner of Guiana Shield (Fig. 4C). It occurs in scrublands or tall forests, usually on sandy soils associated with periodic flooding, although it is also found in humid forest up to 1000 m alt. in Venezuela. Recognition — Pradosia beardii is easily distinguished from all other Pradosia species by its dense reddish-brown pubescence covering the pedicel and the lower leaf surface (Fig. 1L). Pradosia brevipes (Pierre) T. D. Penn., Fl. Neotrop. Monogr. 52: 641. 1990. ≡ Ecclinusa brevipes Pierre, Not. Bot. Sapot. 57. 1891.—TYPE: BRAZIL. Prov. Paraná. São Bento (Rincao das Pedras), prope Castro, 8 Jun 1880, C. A. W. Schwacke 2894 (holotype: GOET 010954!; isotype: P 00649435 [fragment]!) Geoxylic shrub to 30–100 cm tall. Leaves alternate, 8–23 × 2.5–8.5 cm long, oblanceolate, coriaceous, upper surface glabrous or with some residual indument at the midvein and secondary veins, lower surface usually pubescent; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, impressed above; intersecondaries absent; tertiaries oblique or oblique-reticulate; petiole 0.6–1.8 cm long, canaliculate, pubescent, without scales. Flowers 3–10 in each fascicle, ramiflorous; pedicel 5–30 mm, tomentulose. Sepals are tomentulose outside. Corolla is 4.2–7.3 mm long, reddish, sparsely sericeous outside. Fruit 2.5–4.0 cm long, ellipsoid, yellowish, smooth and puberulous. Distribution and habitat — Widespread from central Brazil, south into Paraná State and eastern Paraguay in scrublands (Cerrado), usually on sandy soils, from 300–1000 m alt. (Fig. 4D). Recognition — Pradosia brevipes is closely related to P. glaziovii and P. granulosa, species with similar leaf shape and reddish flowers, but they cannot be confused since P. brevipes is the only geoxylic species in the genus, not forming treelets or trees, and is restricted to savannahs, not occurring in wet evergreen forests. If one confronts a fruiting specimen where the habit is unclear, fruits of P. brevipes are smooth, not muricate as in other members with similar leaf shape. [Volume 41 Pradosia caracasana (Pittier) T. D. Penn., Fl. Neotrop. Monogr. 52: 668. 1990. ≡ Oxythece caracasana Pittier, Arb. Arbust. Venez. 1: 11. 1921.—TYPE: VENEZUELA. Lower Cotiza, near Caracas, 800–1200 m alt., [28] Jul 1918 (fl), H. Pittier 7955 (holotype: US 00113132!; isotypes: BKL 00002344!, F 0072112F and F 0072113F, GH, MICH 1210173!, MO 1991133, PH 00018154 and PH 00018155, VEN). Tree up to 10 m, but can reach 25 m tall; buttresses short. Bark smooth, reddish-brown, shallowly fissured, sloughing off in thin plates and leaving deep whitish marks; slash unknown; inner bark non-sweet tasting. Leaves subverticillate or alternate, 8–23 × 4–10 cm, elliptic or obovate, chartaceous, glabrous on both sides; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, slightly impressed above; intersecondaries absent; tertiaries oblique-reticulate; petiole 1.0–2.7 cm long, canaliculate, glabrous or with residual indument, without scales. Flowers 3–20 in each fascicle, ramiflorous; pedicel 1.5–2.5 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 3.5– 4.0 mm long, greenish, sericeous outside. Fruit 5.0–7.0 cm long, ellipsoid, greenish-brown, smooth, densely lenticellate, and glabrous. Distribution and habitat — Pradosia caracasana is known from northern Venezuela where it occurs in wet lowland to montane forests, from 80–900 m alt. (Fig. 4A). Recognition — Pradosia caracasana, P. colombiana, and P. montana form a strongly supported group (Fig. 3). Pradosia caracasana and P. colombiana are sister species and are sometimes confused morphologically, especially in Zulia, northwestern Venezuela, where these two species meet (Pennington 1990). However, P. caracasana differs from P. colombiana by glabrous leaves (vs. pubescent), ramiflorous fascicles (vs. axillary), flowers with 1.5–2.5 mm long pedicels (vs. 5– 12 mm long), and densely lenticellate fruits (vs. smooth). They also occur in different types of forests; P. caracasana occurs in wet evergreen forests, while P. colombiana occurs in deciduous forests. Pradosia cochlearia (Lecomte) T. D. Penn., Fl. Neotrop. Monogr. 52: 655. 1990. ≡ Chrysophyllum cochlearium Lecomte, Notul. Syst. (Paris) 4: 63. 1923.—TYPE: FRENCH GUIANA. Charvein, 13 Nov 1913 (fl), R. Benoist 224 (holotype: P 00649438!; isotypes: F 0071930F [fragment]!, P 00649439!). Tree up to 35 m tall; buttresses up to 2 m tall (Fig. 1C); bark grayish, smooth, scaling and leaving orange-brown patches; slash of trunk pinkish (Fig. 1M); inner bark sweettasting. Leaves alternate, less frequently opposite or verticillate, oblanceolate or obovate, 5–11(–15) × 2.5–4.5 cm, chartaceous, glabrous or appressed brown-puberulous on the lower surface (Fig. 1K); midvein raised on the upper surface; secondary venation eucamptodromous, parallel, raised above; intersecondaries sometimes present, short; tertiaries oblique to horizontal, sometimes obscure; petiole 0.7–1.6 cm long, slightly canaliculate, glabrous or appressed brownpuberulous, without scales. Flowers 4–10 in each fascicle, ramiflorous; pedicel 0.5–4.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 1.7–3.0 mm long, greenish, sparsely sericeous outside. Fruit 3.5–5.0 cm long, ellipsoid or obovoid, yellow-greenish, smooth, glabrous or glabrescent (Fig. 2K). 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA Distribution and Habitat — Widespread, extending from eastern to central Amazon in Brazil and French Guiana (Fig. 4B). It usually grows in tall forests, non-flooded systems on sandy soils, although some plants have been collected in tall forests on clayish soils near Manaus, central Amazon. Additional specimens have been reported for Caquetá (Colombia), and from Loreto in Peru, in upland forest on white-sandy soils. However, we have not been able to examine voucher material from these areas. Note — Raymond Benoist (1881–1970), a French botanist, was sent on an official botanical mission to French Guiana in 1913–1914. His collection Benoist 224 was later used by Lecomte (1923) to describe Chrysophyllum cochlearium, a species transferred to Pradosia by Pennington (1990). Pennington stated that the holotype is present in the Paris herbarium. In Paris, two sheets of Benoist 224 are present, P 00649438 and P 00649439, of which the first mentioned has pencil drawings of a flower, a corolla, and an ovary that concur with the diagnosis (Lecomte 1923). We therefore, in accordance with Article 9 of the International Code of Nomenclature (McNeill et al. 2012), believe that Benoist 224 (P 00649438) constitutes the holotype. Recognition — Pradosia cochlearia is similar in morphology to P. huberi and P. schomburgkiana among the other Amazonian species with sweet bark. It differs from P. huberi in that the secondary veins are parallel (vs. divergent), the tiny, 1.7– 3.0 mm long corolla (vs. 4.5–5.5 mm long) (see Pennington 1990), and that it grows in non-flooded forests, usually on sandy soils (not swampy lands or flooded forests). It is further distinguished from P. schomburgkiana in its eucamptodromous venation and by the absence or much shorter intersecondaries (vs. brochidodromous venation and long intersecondaries extending towards the margin). Pennington (1990) recognized two subspecies, P. cochlearia subsp. cochlearia and P. cochlearia subsp. praealta (Ducke) T. D. Penn., based on the presence or absence of a sericeous indument on the lower leaf surface plus whether the tertiaries are visible or obscure. We have examined the morphology of samples covering the entire geographic distribution of these subspecies and could not find morphological coherence supporting subspecies in P. cochlearia. Pradosia colombiana (Standl.) T. D. Penn. ex T. J. Ayers & Boufford, Brittonia 40: 428. 1988. ≡ Sideroxylon colombianum Standl., Trop. Woods 22: 13. 1930.—TYPE: COLOMBIA. Region of Santa Marta, 250 ft alt., Aug 1898–1901 (fl), H. H. Smith 456 (holotype: F 0072232F!; isotypes: A 00589962 and A 00589963, BM 000952570!, BR 0000005415045!, COL 000003939!, CM 1674, E 00259453!, F 0072232F, G 00439604!, MICH 1104683!, MO 345912!, MPU 019068!, NY 00296951!, PH 00024050, S 05–10509!, U 0006711!, US 00113305!). Tree up to 8 m, but can reach 25 m tall; buttresses short; bark red-brown, smooth, with irregular grayish spots along the trunk; slash of trunk unknown; inner bark non-sweet tasting. Leaves alternate, elliptic, 8–25 × 3–10 cm, chartaceous, upper surface glabrous or with some residual indument, lower surface sparsely gray-puberulous; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, not impressed above; intersecondaries sometimes present, short; tertiaries oblique or horizontal; petiole 1.7– 6.5 cm long, slightly canaliculate, glabrescent, without scales. 643 Flowers 5–20 in each fascicle, axillary; pedicel 5.0–12 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 3.0–4.0 mm long, greenish, sparsely sericeous outside. Fruit 3.0–4.0 cm long, ellipsoid, gray-greenish, not lenticellate, and puberulous. Distribution and habitat — Pradosia colombiana occurs in the north corner of Colombia, along the Atlantic coast, into northwestern Venezuela, in deciduous forests from sea level to 400 m alt. (Fig. 4A). Recognition — This species is easy to recognize because it is the only congener growing in deciduous forest with axillary flowers and smooth fruits. Pradosia cuatrecasasii (Aubrév.) T. D. Penn., Fl. Neotrop. Monogr. 52: 663. 1990. ≡ Ecclinusa cuatrecasasii Aubrév., Adansonia 7: 144. 1967.—TYPE: COLOMBIA. Departamento del Valle, Costa del Pacífico, Río Naya, Puerto Merizalde, bosque 5–20 m alt., 20 Feb 1943 (fl), J. Cuatrecasas 13988 (holotype: US 00153804! and US 00153803!; isotypes: COL 000003925, F 0071965F! and F 0071966F, P 00649441!). Large buttressed tree; bark grayish, scaling; slash of trunk unknown; inner bark sweet-tasting. Leaves alternate, 20– 40 × 8–14 cm, oblanceolate, coriaceous, upper surface glabrous or with some residual indument, lower surface densely brown-pubescent; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, impressed above; intersecondaries absent; tertiaries oblique, impressed above; petiole 2.0–2.6 cm long, canaliculate, pubescent, without scales. Flowers 5–14 in each fascicle, born on small burls, ramiflorous; pedicel 5.0–10 mm long, tomentulose. Sepals are tomentulose outside. Corolla ca. 7.5 mm long, greenish, sericeous outside. Fruit 5.5–6.5 cm long, ellipsoid, green-brownish, smooth, and glabrescent. Distribution and Habitat — This species is only known from wet lowland forests at low altitudes along the Pacific coast of Colombia (Chocó) (Fig. 4C). Note — The holotype Cuatrecasas 13988 is deposited in the Smithsonian herbarium US (Aubréville 1967), not in Paris as stated by Pennington (1990). Further, it is mounted in two parts, one with leaves and one with branches carrying flowers, fully in accordance with article 8.3 of the International Code of Nomenclature (McNeill et al. 2012). Recognition — Pradosia cuatrecasasii is readily recognized by the large, strongly coriaceous leaves, with sunken midvein, conspicuous secondary veins, and densely brown-pubescent lower surface. There is no other species of Pradosia in west Colombia with which it can be confused. Pradosia decipiens Ducke, Trop. Woods 71: 17. 1942.— TYPE: BRAZIL. Amazonas: Manaus, Cachoeira baixa do Tarumã, mata da t. f. baixa, 2 Dec 1932 (fl), A. Ducke 385 [RB24860] (holotype: RB 00379516!; isotypes: G 00439603, K 000640454! and K 000640455!, R 000054689, RB 00635940! and RB 00635945!, WIS 00000902MAD). Tree up to 20 m tall; buttresses up to 0.5 m tall; bark grayish, smooth, scaling in asymmetric thin plates leaving orange marks; slash of trunk orange (Fig. 1N); inner bark non-sweet tasting. Leaves alternate, 8–19 × 4–8 cm, oblanceolate, coriaceous, glabrous; midvein raised on the upper surface; secondary venation eucamptodromous, parallel, impressed above; intersecondaries absent; tertiaries oblique, obscure; petiole 1.7–2.6 cm long, canaliculate in the upper half, 644 SYSTEMATIC BOTANY glabrous, without scales. Flowers 5–12 in each fascicle, ramiflorous; pedicel 5.0–13.0 mm long, glabrous. Sepals are glabrous outside. Corolla is 4.5–5.2 mm long, greenish, glabrous. Fruit 3.0–3.5 cm long, ellipsoid, yellowish, smooth, and glabrous. Distribution and habitat — Two disjunct populations of Pradosia decipiens are known: near Manaus in central Amazon, where it has been collected in forest on whitesandy soil in the Reserva Florestal Adolpho Ducke and in Solano, Caquetá, southern Colombia (Fig. 4B). This species usually grows in lowland non-flooded forests or occasionally flooded forests on nutrient-poor white-sandy soils (campina and campinarana). Recognition—Pradosia decipiens, like P. cochlearia, P. kuhlmanni and P. schomburgkiana, belongs to the Sweet-bark clade and has similar bark structure, leaves with flat or raised midvein on the upper surface, and green flowers, but P. decipiens differs from these congeners in that the inner bark is not sweet in taste. In addition, the tertiary venation is weak and not prominent and the corolla is slightly longer, 4.5–5.2 mm instead of less than 4 mm long. Pradosia glaziovii (Pierre) T. D. Penn., Fl. Neotrop. Monogr. 52: 643. 1990. ≡ Ecclinusa glaziovii Pierre, Not. Bot. Sapot. 56. 1891.—TYPE: BRAZIL. Rio de Janeiro: Itaborahy, 12 Sep 1875 (fl, fr), A. Glaziou 8229 (lectotype chosen by Pennington [1990: 643]: P 00649442!; isolectotype: BR 0000005415342 and BR0000005415014, F 0BN004272 [photo negative image]!, G 00439602!, K 000518106!, P 00649443!). Pradosia verrucosa Ducke, Bol. Tecn. Inst. Agron. N. 28: 27. 1953.—TYPE: BRAZIL. Pernambuco: Recife, Estrada da Aldeia, Km 28, mata alta, 30 Apr 1952 (fl, fr), A. Ducke & A. Lima 80 (holotype: IAN!; isotype: US 00037033!). syn. nov. Tree up to 30 m tall; short buttresses; bark gray-brown, smooth; slash of trunk light orange; inner bark non-sweet tasting (Fig. 1O). Leaves clustered at branch tips, alternate or verticillate, 9–24 × 4–11 cm, oblanceolate or obovate, coriaceous, upper surface glabrous or midvein pubescent, lower surface puberulous; midvein sunken on upper surface; secondary venation eucamptodromous, parallel, slightly impressed above; intersecondaries absent; tertiaries oblique; petiole 0.8–2.2 cm long, canaliculate in upper half, pubescent, without scales. Flowers 2–10 in each fascicle, ramiflorous; pedicel ca. 1 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 5.0–6.5 mm long, reddish, sericeous outside. Fruit 3.5–5.0 cm long, ovoid to ellipsoid, yellowish, muricate, and brown-tomentulose. Distribution and Habitat — Pradosia glaziovii is found throughout the Brazilian Atlantic coast, occurring from Pernambuco in the north to the state of Espírito Santo in the south, occurring in wet lowland tall forests on clayish soils (Fig. 4D). Recognition — Pennington (1990) suggested that Pradosia glaziovii is closely related to P. brevipes, which has been confirmed by our phylogenetic analysis (Terra-Araujo et al. 2015). These two species have similar leaves, but readily recognized in that P. glaziovii is a tall canopy tree confined to wet coastal forests whereas P. brevipes is a geoxylic shrub from the savannah in central Brazil. However, Pennington overlooked the similarities between P. glaziovii and P. verrucosa, [Volume 41 two species of Pradosia with muricate fruits, restricted to the Atlantic coast, and occurring in wet forests on clayish soils. The name glaziovii was proposed in honor of A. Glaziou whereas verrucosa probably refers to the muricate fruit. Because P. glaziovii is the older name it has authority over P. verrucosa. Pradosia glaziovii is mainly characterized by being a tall canopy tree with coriaceous leaves and with appressed indument on the lower surface, short pedicel, reddish flowers, muricate fruit surface, and by its occurrence in wet forest on clayish soils. Pradosia granulosa Pires & T. D. Penn., Fl. Neotrop. Monogr. 52: 645. 1990.—TYPE: BRAZIL. Pará: Serra dos Carajás, AMZA camp 4-Alfa, ca. 25 km by road northwest of Rio Itacaiúmas, edge of forest around abandoned mining camp, 5°46′S, 50°36′W, ca. 225 m alt., 6 Jun 1982 (fl), C. R. Sperling et al. 5923 (holotype: K 001096525!; isotypes: F V0072206F!, MG!, NY 01168534!, US 00516779!). Treelet 4–6 m tall; unbuttressed; bark gray-brownish, rough, lenticellate, shallowly fissured; slash of trunk light yellow (Fig. 1U); inner bark non-sweet tasting. Leaves subverticillate, 8–22 × 5–8 cm, oblanceolate or obovate, coriaceous, upper surface glabrous, lower surface glabrous or indument if present, restrict to the midvein or secondary veins; midvein sunken on upper surface; secondary venation eucamptodromous, parallel, straight or slightly arcuate, slightly impressed above; intersecondaries absent; tertiaries oblique (Fig. 1J); petiole 0.9–2.0 cm long, canaliculate in the upper half, tomentulose or glabrescent, without scales. Flowers 10– many in each fascicle, ramiflorous or axillary; pedicel 1.6– 3.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 4.5–5.5 mm long, wine red, sericeous outside (Fig. 2F). Fruit 3.0–4.5 cm long, ellipsoid, yellow-orange, muricate, and brown-tomentulose (Fig. 2J, 2L). Distribution and habitat — Pradosia granulosa is known from northeastern Amazon, occurring in the states of Goiás, Maranhão and Pará in Brazil (Fig. 4D). It grows in open lowland forests on sandy soils (campina forest) or occasionally on clay, from sea level to 250 m altitude. Recognition — This species is easily distinguished from all other Amazonian species by its muricate fruits. Pradosia granulosa is sister of P. brevipes and P. glaziovii in the Red-flowered clade (Fig. 3), and apart from the morphology, also has other ecological preferences, growing in campina forests or occasionally on clay, not in Cerrado as P. brevipes or in wet lowland tall forests on clayish soils as P. glaziovii. Pradosia grisebachii (Pierre) T. D. Penn., Fl. Neotrop. Monogr. 52: 655. 1990. ≡ Ecclinusa grisebachii Pierre, Not. Bot. Sapot. 57. 1891.—TYPE: TRINIDAD AND TOBAGO. Trinidad: Versimmiliter ex insula Trinitatis et ex Cruegeri colectione, Aug 1860 (fl), H. Crüger 157 (holotype: GOET 010955!; isotypes: GOET 010956 [fragment]!, K 000640458!, NY 0273462 [fragment]!, P 00649446!). Pradosia atroviolacea Ducke, Trop. Woods 90: 25. 1947.–– TYPE: COLOMBIA. Amazonas: Leticia, mata de t. firme, baixa ao Nordeste, 3 Nov 1945 (fl), A. Ducke 1800 (lectotype chosen by Pennington [1990: 653]: RB 00380742!; isolectotypes: A 00460602! CAS 214950!, F V0072205F!, GH 00075855!, IAN!, K 000640438! and K 000640439!, MG!, NY 00273667! and NY 00273668!, 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA R 000075072!, RB 00635954!, US 00113358!, US 00323586!, and US 00930768!). syn. nov. Tree up to 15 m tall; buttressed; bark grayish, smooth, sloughing off in thin plates and leaving dark marks; slash of trunk unknown; inner bark non-sweet tasting. Leaves alternate or subverticillate, 8–22 × 4–8 cm, oblanceolate or obovate, chartaceous to coriaceous, upper surface glabrous, lower surface glabrous or puberulous; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, slightly impressed above; intersecondaries absent; tertiaries oblique to horizontal; petiole 0.8–2.0 cm long, canaliculate in the upper half, glabrous; scales present, paired, 1.0–2.0 mm long, fixed at the midway of the petiole. Flowers 2–many in each fascicle, ramiflorous; pedicel 1.0– 3.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 4.0–5.0 mm long, reddish, sparsely sericeous outside. Fruit 3.5–5.0 cm long, ellipsoid, yellowish, smooth, and sparsely puberulous. Distribution and habitat — Widespread species from southwestern Amazon, Acre, Brazil to northwestern Amazon in Colombia, reaching the north of Venezuela and Costa Rica (Fig. 4C). Pradosia grisebachii grows in lowland non-flooded forests on clayish soils from 100–700 m alt. Note — Hermman Crüger collection was used by Pierre (1891) to describe Ecclinusa grisebachii, a species later transferred to Pradosia by Pennington (1990). In the herbarium of the Universität Göttingen (GOET), there are two specimens, GOET 010955 and GOET 010956, of which only the first is a good sample bearing leaves and flowers while the second one has only fragments of leaves, flowers, and a pencil drawing of a flower, which agree with the diagnosis of Pierre. In accordance with article 9 of the International Code of Nomenclature (McNeill et al. 2012), we believe that Crüger 157 (GOET 010955) constitutes the holotype. Recognition — We unite here Pradosia grisebachii and P. atroviolacea based on morphological similarities. These species are the only Pradosia having scales on the petiole combined with ramiflorous fascicles. They have been distinguished by the presence or absence of trichomes on the lower leaf surface, petiole length, and the number of flowers in their fascicles (Pennington 1990). In general, P. grisebachii is known from Trinidad and Venezuela, whereas P. atroviolacea occurs in western Amazonia to Costa Rica and Panama. Studies of the types and many new collections covering the geographic range of both species cannot reveal any clear morphological discontinuities between them. The abovementioned morphological characters used by Pennington (1990) to distinguish them do not clearly separate the material in two distinct morphological groups. We consider it therefore appropriate to unite P. atroviolacea with P. grisebachii. Pradosia huberi (Ducke) Ducke, Trop. Woods 71: 16. 1942. ≡ Glycoxylon huberi Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 253. 1922.–– TYPE: BRAZIL. Pará: Furo Macujubim, Ilhas de Breves, mata alagada, 16 Nov 1923 (fl, fr), A. Ducke [RB3782] (lectotype chosen by Pennington [1990: 657]: RB 00560333!; isolectotypes: G 00439601!, IAN!, K 000640447!, MG!, P 00649447!, S 05–10587!, U 0006712!). Tree up to 30 m tall; large buttresses to 2 m tall; bark smooth, grayish; slash of trunk unknown; inner bark sweettasting. Leaves opposite, 5–15 × 3–4 cm, elliptic or obovate, 645 chartaceous, glabrous; midvein flat and slightly raised on upper surface; secondary venation eucamptodromous, divergent and slightly raised above; intersecondaries absent; tertiaries oblique-reticulate; petiole 0.5–1.5 cm long, semiterete, glabrous, without scales. Flowers 2–5 in each fascicle, ramiflorous; pedicel 1.0–4.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 4.5–5.5 mm long, greenish, glabrous outside. Fruit 2.5–4.5 cm long, obovoid to ellipsoid, greenish, smooth, glabrous. Distribution and habitat — Pradosia huberi occurs in northeastern Amazon, in the states of Amapá and Pará in Brazil and nearby Cayenne in French Guiana (Fig. 4B). In spite of the low number of recent collections this species is believed to be a characteristic member of swampy lands or flooded forests along the coast (estuarine areas) and associated with riverine systems (igarapé or furo) (Ducke 1953). Recognition — Pradosia huberi is similar in morphology to P. cochlearia, but is readily recognized on its semiterete petiole (vs. slightly canaliculate), divergent secondary veins (vs. parallel), larger corolla (vs. 1.7–3.0 mm) and that it occur in swampy or flooded forests (vs. non-flooded forests, usually on sandy soils). Pradosia kuhlmannii Toledo, Arq. Bot. Estado São Paulo 2: 29. 1946.—TYPE: BRAZIL. Rio de Janeiro: Mata do morro do Jardim Botânico, 22 May 1926 (fl, fr), D. Constantino & P. Occhioni [RB22231] (lectotype chosen by Pennington [1990: 651]: RB 00560363!; isolectotypes: IAN!, NY 00375787!, P 00649448!, RB 00560364!, S 05– 10534!, U 0006713!, US 00323588!). Tree up to 15 m tall; buttresses well developed; bark smooth, grayish, scaling in thin irregular plates leaving deep orange to brown patches; slash of trunk reddish; inner bark sweet-tasting (Fig. 1P). Leaves alternate, 4–11 × (1.4–)2–4 cm, elliptic or oblanceolate, chartaceous, glabrous; midvein flat on the upper surface; secondary venation brochidodromous, parallel, not impressed above; intersecondaries present, short; tertiaries horizontal-reticulate; petiole 0.8–2.0 cm long, semiterete, glabrous, without scales. Flowers 3–6 in each fascicle, ramiflorous or axillary; pedicel 0.5–1.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 3.0– 4.0 mm long, greenish, glabrous outside. Fruit 2.5–3.5 cm long, ellipsoid, greenish, smooth, and glabrous. Distribution and habitat — Pradosia kuhlmannii occurs along the Brazilian Atlantic coast, from Pernambuco in the north to Rio de Janeiro State in the south, and always in wet lowland tall forests on clayish soils (Fig. 4B). Recognition — Sterile specimens of Pradosia kuhlmannii are easy to recognize because of its sweet tasting bark, semiterete petioles, brochidodromous venation, flat midvein, and horizontal tertiaries. Other members of the Sweet-bark clade have similar character combinations, but occur in rainforests on sandy soils in the Amazon. On the other hand, P. kuhlmannii grows on clayish soils in wet rainforests and is sympatric with P. lactescens, a species with which it has been confused (see Kuhlmann 1930; Toledo 1946). However, P. lactescens differs in an astringent inner bark, eucamptodromous leaf venation, canaliculate petiole, cauliflorous fascicles, and larger and wine-reddish flowers. Pradosia lactescens (Vell.) Radlk., Sitzungsber. Math.Phys. Cl. Königl. Bayer. Akad. Wiss. München 18: 407. 1888. ≡ Pometia lactescens Vell., Fl. Flum. 81, 2: Table 87. 646 SYSTEMATIC BOTANY 1825.—TYPE: BRAZIL: J. M. Vellozo s.n. (lectotype chosen by Pennington [1990: 650]: P 00649449!). Tree up to 15 m tall; buttresses poorly developed; bark reddish-brown, scaling in thin plates; slash of trunk orange; inner bark non-sweet tasting (Fig. 1Q). Leaves alternate, 5–20 × 2–6 cm, elliptic to obovate, chartaceous, glabrous; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, slightly sunken above; intersecondaries absent; tertiaries oblique; petiole 0.6–1.8 cm long, canaliculate, glabrous or with residual appressed indument, without scales. Flowers usually many in each fascicle, cauliflorous; pedicel 4.0–6.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 4.8–6.5 mm long, wine red, usually glabrous (Fig. 2E). Fruit 3.0–4.5 cm long, obovoid to ellipsoid, yellow-orange, smooth, and glabrous (Fig. 2H). Distribution and habitat — Pradosia lactescens is a very common and widespread species along the whole range of the Brazilian Atlantic coast rainforest, from Pernambuco in the north to Paraná State in the south. It grows in wet lowland tall forests on clayish soils from sea level to 980 m altitude (Fig. 4D). Recognition — Pradosia lactescens is in overall morphology most similar to P. ptychandra. These two species are cauliflorous, have similar leaf shapes, and wine red flowers. However, they are not closely related and are best distinguished by flower features (e.g. pedicel and corolla length), which are larger in P. ptychandra. Moreover, they are allopatric: P. lactescens is restricted to the Brazilian Atlantic coast, while P. ptychandra occurs in Guyana, Suriname, and French Guyana. On the other hand, P. lactescens has been historically confused with P. kuhlmannii, a species also restricted to the Brazilian Atlantic coast region. In part, the confusion came from a swap of samples committed by Casaretto (Toledo 1946). The diagnosis was based on two different samples: leaves of Pradosia lactescens and bark of P. kuhlmannii, a mistake highlighted by Toledo (1946) and which was dealt with subsequently with new descriptions. In the southern coast of the state of Rio Grande do Norte in Brazil, P. lactescens could be confounded with P. restingae. Pradosia lactescens is best recognized in having decurrent leaf bases, being cauliflorous with 4.0–6.0 mm pedicellate, wine reddish flowers, and growing in forests on clayish soils. In contrast, P. restingae has cuneate leaves, being ramiflorous with subsessile, greenish flowers, and is only known from white-sand forests (Terra-Araujo et al. 2013). Pradosia lahoziana Terra-Araujo, Brittonia, 64: 142. 2012.—TYPE: BRAZIL. Amazonas: Manaus, Estrada do Aleixo, grounds of Companhia das Plantações, forest on terra firme [03°05′ S, 59°55′ W, 50–94 m], 30 Aug 1973 (fl), G. T. Prance 18763 (holotype: INPA!; isotypes: MG!, MO!, NY 01171401!). Tree up to 15 m tall; unbuttressed; bark rough, lenticellate, grayish-brown; slash of trunk light orange; inner bark nonsweet tasting. Leaves alternate or subverticillate, 9–20 × 3– 7 cm long, oblanceolate or elliptic, chartaceous, upper surface glabrous, lower surface sparsely tomentulose, at least on the venation (visible with a hand-lens); secondary venation eucamptodromous; midvein sunken on the upper surface; secondary veins parallel, slightly raised or flat above, usually tomentulose, glabrescent; intersecondaries absent; [Volume 41 tertiaries horizontal or oblique; petiole 1.6–2.6 cm long, canaliculate, tomentulose; scales present, attached below the blade to the middle of the petiole, 1.0–2.0 mm long, paired (Fig. 1G). Flowers 5–10 in each fascicle, cauliflorous; pedicel 7.0–9.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 6.4–9.5 mm long, wine red, glabrous outside. Fruit 2.5–3.0 cm long, obovoid, yellowish, smooth, and tomentulose. Distribution and habitat — Pradosia lahoziana is known from central Amazon, where it occurs in forests north of Manaus, in the permanent plots of the Dinâmica Biológica de Fragmentos Florestais Project, Dimona Reserve and from the Estação Experimental de Sivicultura Tropical-ZF2, both from the Instituto Nacional de Pesquisas da Amazônia (Fig. 4C). It grows in rainforest on sandy (e.g. campina and campinarana forests) or occasionally clayish soils from 50–125 m alt. Recognition — This species has a unique combination of characters in the genus: having scales on the petiole below the blade and flowers on the stem (cauliflorous). Similar scales are present in Pradosia grisebachii, a ramiflorous species. Other cauliflorous species such as P. ptychandra and P. lactescens lack these scales. Pradosia longipedicellata Alves-Araújo & M. Alves, Brittonia 64: 24. 2012.—TYPE: BRAZIL. Bahia: Una, Reserva Biológica do Mico-Leão (IBAMA), BA-001 Km 46, 15°09′S, 39°05′W, 9 Mar 1993 (fl, fr), J. G. Jardim et al. 92 (holotype: CEPEC!; isotype: NY 00585395!). Tree up to10 m, but can reach 20 m tall; unbuttressed or occasionally with buttresses to 0.5 m tall; bark grayish, rough, lenticellate, shallowly fissured, scaling; slash of trunk pinkish; inner bark non-sweet tasting (Fig. 1V). Leaves clustered, alternate, 6–17 × 3–7 cm, ovate to elliptic, chartaceous, both leaf surfaces covered by whitish to ferruginous trichomes, glabrescent; secondary venation eucamptodromous; midvein flat; secondary vein parallel, flat or slightly raised; intersecondaries absent; tertiaries oblique-reticulate (Fig. 1I); petiole 0.4–1.6 cm, semiterete, with whitish to ferruginous trichomes, without scales. Flowers 2–8 in each fascicle, terminal; fascicles in clusters, umbelliform; pedicel 18–30 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 7.0–8.0 mm long, white, glabrous (Fig. 2A). Fruit 3.0–4.0 cm long, ellipsoid, smooth, and brown-puberulous. Distribution and habitat—Pradosia longipedicellata is restricted to the coastal region of southern Bahia, Brazil, and is found in white-sandy restinga forests, from sea level to around 60 m altitude (Fig. 4A). Recognition — Sterile specimens of Pradosia longipedicellata are easily distinguished from all other Brazilian Atlantic coast species by its leaves covered by whitish to ferruginous trichomes and flat midvein. If fertile specimens are available, it is readily distinguished on dense, terminal inflorescences with long pedicels (18–30 mm) and white flowers. Pradosia montana T. D. Penn., Fl. Neotrop. Monogr. 52: 661. 1990.—TYPE: ECUADOR. Province El Oro, ca. 60 km SE of Arenillas on road to Loja, 13 Nov 1982 (fl), T. D. Pennington & G. Tenorio 10719 (holotype: K 000640437!; isotypes: MO 1991078, QCA 100610!, QCNE 735!). Tree up to 8 m, but can reach 25 m tall; unbuttressed or with slight buttress to 0.5 m tall; bark light gray, lenticellate, 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA rough, fissured, sloughing off in small plates; inner bark non-sweet tasting; slash of trunk pinkish. Leaves alternate or subverticillate, 5–15 × 2–7 cm, obovate or elliptic, coriaceous, upper surface sparse pubescent on midvein and secondary veins, lower leaf surface with indument of erect twobranched pale-brownish trichomes; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, slightly impressed above; intersecondaries absent; tertiaries oblique to horizontal; petiole 0.5–1.3 cm long, slightly canaliculate, tomentulose, without scales. Flowers 5–10 in each fascicle, ramiflorous; pedicel 2.0–3.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 3.0–4.0 mm long, greenish, glabrous or sparsely tomentulose outside, especially the corolla tube (Fig. 2C). Fruit 1.5–2.5 cm long, obovoid or ellipsoid, smooth, and densely brown-pubescent. Distribution and habitat — This species is restricted to western Ecuador and northwest corner of Peru, being considered a common component of tropical deciduous forests from sea level to 720 m altitude (Fig. 4A). Recognition — Pradosia montana is readily distinguished from all other species of Pradosia by the soft pubescence on the lower leaf surface. There are two other species associated with deciduous forests, P. colombiana and P. mutisii, but they have glabrous or glabrescent leaves. Pradosia mutisii Cronquist, Bull. Torrey Bot. Club 73: 470. 1946.—TYPE: COLOMBIA. (fl), J. C. Mutis 4004 (holotype: US 00037017!; isotype G 00439600 [fragment]!). Tree up to 8 m, but can reach 30 m tall; buttress up to 0.5 m tall; bark dark brown; slash of trunk unknown. Leaves clustered, alternate, 5–14 × (2.5–)3–6 cm, obovate or elliptic, chartaceous, upper surface glabrous, lower surface glabrous or with sparse short whitish trichomes; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, not impressed above; intersecondaries present and extending towards the margin; tertiaries horizontalreticulate; petiole 1.1–2.2 cm long, canaliculate, glabrous, without scales. Flowers 2–10 in each fascicle, ramiflorous; pedicel ca. 1 mm long, glabrescent. Sepals are glabrescent outside. Corolla is 2.5–3.0 mm long, greenish, glabrescent outside. Fruit ca. 2 cm long, ellipsoid, greenish, smooth, and glabrous. Distribution and habitat — Pradosia mutisii occurs from northwestern Colombia to northwest Peru. It is a rare species found in tropical deciduous forest from 180–1200 m alt. (Fig. 4A). Recognition — This species can be mistaken for P. montana, and their ranges overlap in the northwest corner of Peru. However, P. mutisii is easily distinguished from P. montana by its glabrous leaves (vs. pubescent), presence of long intersecondaries veins (vs. absent), and glabrous fruits (vs. pubescent). Pradosia ptychandra (Eyma) T. D. Penn., Fl. Neotrop. Monogr. 52: 648. 1990. ≡ Pouteria ptychandra Eyma, Recueil Trav. Bot. Néerl. 33: 189. 1936.—TYPE: SURINAM. Wilhelminagebergte, Eindkamp Lucie rivier, 10 Apr 1926 (fl), G. Stahel [B. W. 6943] (holotype: U 0006715!; isotypes: G 00439599 [fragment], K 000640453 [fragment]!, U 0006716!). Tree up to 13 m, but can reach 25 m tall; unbuttressed or with poorly developed buttress to ca. 30 cm tall; bark 647 smooth, grayish-brown, scaling in thin plates and leaving pock-like depressions; slash of trunk cream with orange streaks; inner bark non-sweet tasting. Leaves clustered, alternate, 7–22 × 2–8 cm long, elliptic, oblanceolate or obovate, glabrous; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, slightly sunken above; intersecondaries absent; tertiaries oblique; petiole 0.6–2.2 cm long, canaliculate, glabrous or with appressed indument, without scales. Flowers usually many in each fascicle, cauliflorous, less frequently ramiflorous; pedicel 7.0–10.5 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 5.0–5.5 mm long, wine red, sparsely tomentulose outside. Fruit 2.5–4.0 cm long, ovoid to ellipsoid, yellow or orange, smooth, and glabrous. Distribution and habitat — This species has a disjunct distribution, in central Amazon in the surroundings of Manaus, and also known from several collections in French Guiana, Guyana and Surinam (Fig. 4C). Pradosia ptychandra grows in non-flooded forests on clayish soils, from 75–770 m alt. Recognition — Pradosia ptychandra is most similar to P. lasctescens, a species of the Atlantic coast, and are best separated on flower features (see under P. lactescens for further discussion). It could also be confused with P. lahoziana, a closely related species, also having wine-red flowers borne along the trunk, but P. ptychandra lacks scales on the petiole (present in P. lahoziana). Pradosia restingae Terra-Araujo, Nord. J. Bot. 31: 437. 2013.—TYPE: BRAZIL. Rio Grande do Norte: Pipa, estrada para o Santuário Ecológico, 06°13′43.7″S, 35°03′ 31.7″W, 55 m alt., 26 Sep 2011 (fl, fr), A. Alves-Araújo et al. 1373 (holotype: UFP!, isotypes: INPA!, RB, S!, UFRN!). Tree about 2–6 m, sometimes up to 9 m tall; slash of trunk unknown. Leaves alternate or subverticillate, (7–)8–13 × 3– 4 cm, obovate or elliptic, chartaceous, glabrous; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, slightly sunken above; intersecondaries absent; tertiaries, visible with lens are oblique or horizontal; petiole 0.6–0.8 cm long, canaliculate, tomentulose, without scales. Flowers 2–10 in each fascicle, ramiflorous, usually not on the trunk; pedicel 0.8–1.6 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 4.8–5.5 mm long, greenish with a flesh colored ring, tomentulose outside, especially the corolla tube and central part of the lobes (Fig. 2D). Fruit 2.5–3.7 cm long, ovoid, yellowish, smooth, and glabrous (Figs. 2G, 2M). Distribution and habitat — Pradosia restingae is only known from the southern coast of the state of Rio Grande do Norte in Brazil, where it is found in both forested (restinga alta) and open vegetation on white-sand soils (dunas), from sea level to 55 m altitude (Fig. 4D). Recognition — This species can be confused with P. lactescens; see under P. lactescens for discussion. Pradosia schomburgkiana (A. DC.) Cronquist, Bull. Torrey Bot. Club 73: 311. 1946. ≡ Chrysophyllum schomburgkianum A. DC. in Candolle A. L. P. P. de, Prodr. 8: 157. 1844.— TYPE: GUYANA, 1838 (fl), M. R. Schomburgk 505 (lectotype chosen by Pennington [1990: 659]: G 00139569!; isolectotypes: BM 000952571!, BR 0000005415137!, E 00259450!, F V0071946F!, G 00439595 and G 00439596, 648 SYSTEMATIC BOTANY K 000640445! and K 000640446!, NY 902225!, OXF, P 00649452!, US 00113129!). Shrub or small tree up to 4 m, but can reach 20 m tall; unbuttressed or occasionally with large buttresses to ca. 1 m tall; bark grayish, smooth, scaling and leaving orange-brown patches; slash of trunk reddish; inner bark sweet-tasting (Fig. 1R). Leaves opposite (Fig. 1F), 4–18 × 2–7 cm, oval (broadly elliptic) to obovate, coriaceous, usually glabrous on both sides, sometimes sparsely ferruginous tomentulose below; midvein flat; secondary venation brochidodromous, parallel, flat or slightly raised on upper leaf surface; intersecondaries long, extending towards the margin; tertiaries horizontal-reticulate; petiole 0.4–1.5 cm long, semiterete, glabrous or with appressed indument, without scales. Flowers 5–many in each fascicle, ramiflorous; pedicel 2–10 mm long, tomentulose to glabrous. Sepals are tomentulose to glabrous outside. Corolla is 2.0–4.0 mm long, greenish, sparsely tomentulose outside, especially the corolla tube (Fig. 2B). Fruit 1.0–2.0 cm long, obovoid to ellipsoid, greenish, smooth, glabrous or brown-puberulous. Distribution and habitat—Pradosia schomburgkiana is widely distributed in the Amazon region, being found in campina, campinarana or restinga forests, from sea level in Pará State up to 1200 m altitude in Guiana Shield (Fig. 4B). Recognition — A sweet-tasting bark combined with an opposite leaf arrangement and a brochidodromous leaf venation readily distinguish Pradosia schomburgkiana from all other congeners. It furthers have long intersecondaries that extend towards the margin, a rather unusual characters for Pradosia. Pennington (1990) recognized two subspecies of Pradosia schomburgkiana (subsp. schomburgkiana and subsp. sericea), which are sympatric in the Serra do Aracá region. The subspecies are distinguished by the presence and absence, respectively, of a dense indument on the lower leaf surface. Both subspecies were included in a recent phylogenetic analysis using molecular data, but there was no phylogenetic signal (Terra-Araujo 2013) supporting subspecies, or correlation between morphological variation, habitat or geography. Furthermore, both subspecies have the same ecological preferences and grow in campina and campinarana forests. Species of Sapotaceae in the Neotropics are frequently poorly collected and their full morphological variation and distribution range is therefore unclear. We propose that the taxon limits within P. schomburgkiana requires a closer examination to better understand whether it contains more than one species. Pradosia subverticillata Ducke, Trop. Woods 71: 13. 1942.—TYPE: BRAZIL. Amazonas: Manaus, Rio Tarumã, cachoeira baixa, mata de t. f. em solo arenoso, 27 Jun 1941 (fl), A. Ducke 812 (lectotype chosen by Pennington [1990: 646]: RB 00544076!; isolectotypes: B 10 0413399!, F 0072208F!, IAN!, K 000640456!, MG!, MO 1991009!, NY 00273673! and NY 01168598!, R 000075081!, RB 00642343!, US 00113361! and US 00323587!). Tree up to 4 m, but can reach 20 m tall; bark grayish, smooth, scaling and leaving irregular brown deep patches; slash of trunk yellowish; inner bark sweet-tasting (Fig. 1S). Leaves clustered at branch tips, alternate or subverticillate, 10–21 × 3–8 cm, oblanceolate or obovate, coriaceous, both surfaces glabrous; midvein sunken on the upper surface; sec- [Volume 41 ondary venation eucamptodromous, parallel, impressed above; intersecondaries absent; tertiaries oblique, obscure; petiole 2.0–3.2 cm long, canaliculate, tomentulose, without scales. Flowers usually many in each fascicle, usually ramiflorous but sometimes axillary; pedicel 4.0–7.0 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 5.5– 6.5 mm long, greenish, sparsely sericeous outside. Fruit 3.5– 4.0 cm long, ellipsoid, smooth, and glabrescent. Distribution and habitat — Pradosia subverticillata occurs from the lower Rio Negro basin near Manaus to upper Rio Negro around Barcelos and Issana rivers in the Upper Rio Negro region (Fig. 4C). Its distribution further seems to extend to southern Colombia, in La Uribe, Meta (M. Gaitán 155, COAH). However, we have not been able to examine voucher material from this area. In Brazil, along Rio Negro, it grows in campina and campinarana forests or occasionally on lowland non-flooded forests on clayish soils near Manaus, in central Amazon. Recognition — If Pradosia subverticillata is compared with other species in the Amazon region, its morphology is most similar to P. surinamensis. These two species have a sweettasting inner bark, glabrous leaves, sunken midvein above, and greenish flowers, and they grow in forests on sandy soils. However, P. subverticillata differ from P. surinamensis by the secondaries veins deeply impressed above and the larger flowers, 5.5–6.5 mm long, in contrast to 2.0–3.0 mm in P. surinamensis. The grayish, scaling, and sweet-tasting bark, however, increase the risk to confound P. subverticillata with P. cochlearia and P. schomburgkiana, two species of the Sweetbark clade that also grow on sandy soils. The midvein is the best vegetative character to separate these species, being sunken in P. subverticillata and raised in P. cochlearia and P. schomburgkiana. Pradosia surinamensis (Eyma) T. D. Penn., Fl. Neotrop. Monogr. 52: 652. 1990. ≡ Pouteria surinamensis Eyma, Recueil Trav. Bot. Néerl. 33:189. 1936.—TYPE: SURINAM. Boven-Suriname Rivier bij Goddo, 23 Jan 1926 (fl), G. Stahel 39 (holotype: U 0006719!; isotypes: G 00439591, K 000640450!, MO 345916!, RB 00544069!, U 0006720! and U 0006721!). Tree up to 15 m, but can reach 30 m tall; buttressed 0.30– 1.0 m tall; bark grayish, smooth, scaling and leaving orange patches; slash of trunk light orange; inner bark sweet-tasting. Leaves spaced, loosely clustered, alternate, 4–16 × 2–6 cm, elliptic or obovate, coriaceous, upper surface glabrous, lower surface glabrous or glabrescent; midvein sunken on the upper surface; secondary venation eucamptodromous, slightly impressed above; intersecondaries absent; tertiaries obliquereticulate, obscure; petiole 0.5–1.9 cm long, canaliculate, tomentulose, without scales. Flowers 1–4 in each fascicle, ramiflorous; pedicel 0.5–2.0 mm long, tomentulose. Sepals are sparsely tomentulose outside. Corolla is 2.0–3.0 mm long, greenish, with sparsely tomentulose tube. Fruit 2.0–4.0 cm long, obovoid to ellipsoid, yellowish, smooth, not lenticellate, glabrous or puberulous (Fig. 2I, 2N). Distribution and habitat — This species is known from northeastern Amazonia, in Roraima and Pará State, Brazil, Guyana and Surinam where it is found in lowland non-flooded forest from 60–600 m. alt. (Fig. 4D). In Brazil, it has been collected usually on sandy soils (campinarana forest). 2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA Recognition — Pradosia surinamensis belongs to the Redflowered clade and is morphologically similar to P. subverticillata. Its relationships with P. subverticillata are discussed under the latter. Pradosia verticillata Ducke. Trop. Woods 71: 12. 1942.— TYPE: BRAZIL. Amazonas: Manaus, Estação do Aleixo além do V. Municipal, restos de mata de terra firme, argilosa, 14 Oct 1941 (fl, fr), A. Ducke 811 (lectotype chosen by Pennington [1990: 664]: RB 00379868!; isolectotypes: F 0072209F!, IAN!, K 000640440!, MG!, MO 1991017, NY 01365138!, R 000075080!, RB 00635941! and RB 00635944!, US 00113362!, US 00930762! and US 00930763!). Tree up to 15 m, but can reach 30 m tall; buttresses up to 0.5 m tall; bark greenish, smooth, scaling and leaving brown deep patches; slash of trunk orange; inner bark non-sweet tasting (Fig. 1T). Leaves clustered at branch tips, verticillate (Fig. 1E), 13–31 × 4–11 cm, oblanceolate, coriaceous, upper surface glabrous or with some residual indument, lower surface brown-pubescent, denser on the venation; midvein sunken on the upper surface; secondary venation eucamptodromous, parallel, impressed above; intersecondaries absent; tertiaries oblique; petiole 1.0–4.1 cm long, canaliculate, pubescent, without scales. Flowers 5–10 in each fascicle, ramiflorous; pedicel 1.0–1.7 mm long, tomentulose. Sepals are tomentulose outside. Corolla is 4.0–4.6 mm long, reddish, sericeous outside. Fruit 3.5–4.5 cm long, obovoid, smooth, and glabrous. Distribution and habitat — Pradosia verticillata is presently known as disjunct, with one population in the lower Rio Negro basin near of Manaus in Brazil, and one in La Fumée Mountain, French Guiana (Fig. 4D). It grows in lowland non-flooded forest on clayish soils, from 59– 400 m alt. Recognition — A very distinct species from all other congeners because of the verticillate clusters of leaves near the branch apices, an impressed venation, and a dark brown indument on the lower leaf surface. Doubtful Taxon— Pradosia argentea (Kunth) T. D. Penn., Fl. Neotrop. Monogr. 52: 660. 1990. ≡ Nycterisition argenteum Kunth in Humbolt, Bonpland & Kunth. Nov. Gen. Sp. 3: 238. 1819.—TYPE: PERU. [Crescit prope Jaen de Bracamoros (Regno NovoGranatensi), 320 m alt.] (fl), A. J. A. Bonpland & F. W. H. A. von Humboldt s.n. (holotype: P 00670922!; isotype: G 00439608 [fragment]!). Pradosia argenta is only known from the type collection, dating back to 1819 in Peru, having flowers but no fruits. Since it has never been relocated, P. argentea was treated as extinct by the Red List in Peru (IUCN 2015). Pennington (1990) in his monograph placed it in Pradosia because of 5-merous, rotate flowers with long-exserted stamens, and lack of staminodes. It was further considered closely related to P. schomburgkiana due to a brochidodromous venation with long intersecondaries, but readily distinct from the latter in having alternate leaves with a fine silvery indument. The overall flower morphology is coherent with Pradosia, but since fruits and molecular data are unknown, it remains unclear if P. argentea is a member of the genus. 649 Acknowledgments. This study was part of the Ph.D. thesis of M. H. Terra-Araujo. The directors at the herbaria ALCB, CA, CEPEC, CVRD, HRB, IAN, INPA, IPA, MBML, MG, NY, PEUFR, PH, RB, S, SP, UEFS and US are acknowledged for permission to examine the herbarium material. We thank Alberto Vicentini, Ana Andrade, Anderson Alves-Araújo, Eduardo Prata, Fernanda A. Carvalho, Flávio Costa, Flávio Obermüller, Jefferson Carvalho-Sobrinho, José Ribamar, Jomar Jardim, Nallarett Dávila, Nory Daniel, Rafaela Forzza, Ricardo Perdiz, Saci and Xavier Cornejo for their help in the field. This research was financed through a scholarship to the first author by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, 143693/ 2008-5) and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, BEX 6161/ 11-1). Literature Cited Alves-Araújo, A. and M. Alves. 2012. Two new species and a new combination of Neotropical Sapotaceae. Brittonia 64: 23–29. Anderberg, A. A. and U. Swenson. 2003. Evolutionary lineages in Sapotaceae (Ericales): A cladistic analysis based on ndhF sequence data. International Journal of Plant Sciences 164: 763–773. Anderson, A. B. 1981. White-sand vegetation of Brazilian Amazonia. Biotropica 13: 199–210. Aubréville, A. 1961. Notes sur les Sapotacées Africanes et SudAméricaines. Adansonia 1: 24–26. Aubréville, A. 1964. Les Sapotace es taxonomie et phytogeographie. Adansonia 1: 1–157. Aubréville, A. 1967. Sapotacées nouvelles de la cote Colombienne du Pacifique. Adansonia 7: 141–148. Baehni, C. 1965. Mémoires sur les Sapotacées. 3. Inventairedes genres. Boissiera 11: 1–262. Cronquist, A. 1946a. Studies in the Sapotaceae V. The South American species of Chrysophyllum. The Journal of the Torrey Botanical Society 73: 286–311. Cronquist, A. 1946b. Studies in Sapotaceae VI. The Journal of the Torrey Botanical Society 73: 465–471. de Queiroz, K. 1998. The general lineage concept of species, species criteria, and the process of speciation: A conceptual unification and terminological recommendations. Pp. 57–75 in Endless forms: Species and speciation, eds. D. J. Howard and S. H. Berlocher. Oxford, UK: Oxford University Press. de Queiroz, K. 2007. Species concepts and species delimitation. Systematic Biology 56: 879–886. Ducke, A. 1922. Plantes nouvelles ou peu connues de la région amazoniènne, Sapotaceae. Archivos do Jardim Botânico do Rio de Janeiro 3: 230–238. Ducke, A. 1925. Plantes nouvelles ou peu connues de la région amazonienne III. Archivos do Jardim Botânico do Rio de Janeiro 4: 158–166. Ducke, A. 1942. New and noteworthy Sapotaceae of Brazilian Amazonia. Tropical Woods 71: 7-25. Ducke, A. 1953. As espécie brasileiras do gênero Pradosia Liais (Fam. Sapotaceae). Boletim Técnico do Instituto Agronômico do Norte 28: 21-34. Ellis, B., D. C. Daly, L. J. Hickey, K. R. Johnson, J. D. Mitchell, P. Wilf, and S. L. Wing. 2009. Manual of leaf architecture. Ithaca, New York: The New York Botanical Garden Press and Comstock Publishing Associates. Eyma, P. J. 1936. Notes on Guiana Sapotaceae. Recueil des Travaux Botaniques Néerlandais 33: 156–210. Ewango, C. E. N. and F. J. Breteler. 2001. Présence du genre Pradosia (Sapotaceae) en Afrique: description d’une nouvelle espèce, P. spinosa. Adansonia 23: 147–150. Harris, J. G. and M. W. Harris. 2001. Plant identification terminology: An illustrated glossary. Spring Lake, Utah: Spring Lake Publishing. Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum. Part I: The herbaria of the world. Eighth edition. New York: New York Botanical Garden Press. IUCN. 2015. IUCN red list of threatened species. Version 2015–3. <www .iucnredlist.org>. Downloaded on 9 September 2015. Kuhlmann, J. G. 1930. Contribuição para o conhecimento de algumas novas espécies da região amazônica e uma do Rio de Janeiro, bem como algumas notas sobre espécies já conhecidas. Arquivos do Jardim Botânico do Rio de Janeiro 5: 201–211. Lecomte, H. 1923. Quelques espèces nouvelles de la famille des Sapotacées. Notulae Systematicae 4: 62–64. (Paris). 650 SYSTEMATIC BOTANY Liais, E. 1872. Climats, géologie, faune et géographie botanique du Brésil. Paris: Garnier Frères. McNeill, J., F. R. Barrie, W. R. Buck, V. Demoulin, W. Greuter, D. L. Hawksworth, P. S. Herendeen, S. Knapp, K. Marhold, J. Prado, W. F. Prud’Homme van Reine, G. F. Smith, J. H. Wiersema, and N. J. Turland (eds.). 2012. International Code of Nomenclature for algae, fungi, and plants (Melbourne Code): Adopted by the Eighteenth International Botanical Congress Melbourne, Australia, July 2011. Regnum Vegetabile 154. Königstein: Koeltz Scientific Books. Miquel, F. A. W. 1863. Sapotae. Pp. 38–117 in Flora brasiliensis vol. 7, eds C. F. P. Martius and A. W. Eichler. Leipzig & München: Frid. Fleischer. Pennington, T. D. 1990. Flora Neotropica Monograph 52: Sapotaceae. New York: New York Botanical Garden. Pennington, T. D. 1991. The genera of Sapotaceae. Kew: Royal Botanic Garden. Pennington, T. D. 2006. Flora da Reserva Ducke, Amazonas, Brasil: Sapotaceae. Rodriguésia 57: 251–366. Prance, G. T. 1996. Islands in Amazonia. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 351: 823–833. Pierre, L. 1891. Notes botaniques, Sapotacées .II. Paris: Paul Klincksieck. Prance, G. T. and H. O. R. Schubart. 1978. Notes on the vegetation of Amazonia I. A preliminary note on the origin of the open white sand campinas of the Lower Rio Negro. Brittonia 30: 60–63. QGIS Development Team. 2014. QGIS geographic information system. Open Source Geospatial Foundation Project. < http://qgis.osgeo.org >. Accessed October 2014. [Volume 41 Swenson, U. and A. A. Anderberg. 2005. Phylogeny, character evolution and classification of Sapotaceae (Ericales). Cladistics 21: 101–130. Swenson, U., J. Munzinger, and I. V. Bartish. 2007. Molecular phylogeny of Planchonella (Sapotaceae) and eight new species from New Caledonia. Taxon 56: 329–354. Swenson, U., J. E. Richardson, and I. V. Bartish. 2008. Multi-gene phylogeny of the pantropical subfamily Chrysophylloideae (Sapotaceae): Evidence of generic polyphyly and extensive morphological homoplasy. Cladistics 24: 1006–1031. Terra-Araujo, M. H., A. D. Faria, and A. Vicentini. 2012. A new species of Pradosia (Sapotaceae) from Central Amazonia. Brittonia 64: 139–142. Terra-Araujo, M. H. 2013. Phylogeny, biogeography and taxonomy of Pradosia (Sapotaceae, Chrysophylloideae). Ph.D. dissertation. Manaus, Amazonas: Instituto Nacional de Pesquisas da Amazônia. Terra-Araujo, M. H., A. D. Faria, A. Alves-Araújo, and M. Alves. 2013. A new species of Pradosia from the Atlantic forest, Brazil. Nordic Journal of Botany 31: 437–441. Terra-Araujo, M. H., A. D. Faria, A. Vicentini, S. Nylinder, and U. Swenson. 2015. Species tree phylogeny and biogeography of the Neotropical genus Pradosia (Sapotaceae, Chrysophylloideae). Molecular Phylogenetics and Evolution 87: 1–13. Toledo, J. F. 1946. Observações criticas sôbre nomes de algumas plantas Brasileiras. Arquivos Botânicos do Estado de São Paulo 2: 29–31.