Clusiaceae (Calophyllaceae, Clusiaceae. and Hypericaceae)

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Clusiaceae s.l. (Calophyllaceae, Clusiaceae s.s. and Hypericaceae) in the Viruá

National Park, Roraima, Brazil
Article in Phytotaxa · November 2017

DOI: 10.11646/phytotaxa.329.1.1
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Phytotaxa 329 (1): 001–027 ISSN 1179-3155 (print edition)
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Article PHYTOTAXA
Copyright © 2017 Magnolia Press ISSN 1179-3163 (online edition)
https://doi.org/10.11646/phytotaxa.329.1.1
Clusiaceae s.l. (Calophyllaceae, Clusiaceae s.s. and Hypericaceae) in the Viruá

National Park, Roraima, Brazil
FERNANDA NUNES CABRAL1,2, VOLKER BITTRICH3 & MICHAEL JOHN GILBERT HOPKINS4
1
Museu da Amazônia, MUSA, Avenida Margarida, s.n., Cidade de Deus, 69099-415, Manaus, Amazonas, Brazil.
E-mail: nandancb@gmail.com
2
Universidade do Estado do Amazonas, Pós-Graduação em Biotecnologia e Recursos Naturais, Av. Djalma Batista, 3586, Flores, Man-
aus, Amazonas, Brazil.
3
R. Mario de Nucci, 500, 13083-290 Campinas, São Paulo, Brazil.
4
Instituto Nacional de Pesquisas da Amazônia, Pós-Graduação em Botânica, Av. André Araújo, 2936, Petrópolis, 69060–001, Manaus,
Amazonas, Brazil.
Abstract
We report a floristic survey of the species of Clusiaceae s.l. (Calophyllaceae, Clusiaceae s.s., and Hypericaceae) in the Viruá
National Park (Roraima, Brazil). The fieldwork was conducted between 2009 and 2015 and covered the different vegetation
types found in the park. Calophyllaceae is represented by two genera and three species, Clusiaceae s.s. by five genera and 12
species, and Hypericaceae by one genus and five species. Three species are new records for the state of Roraima. We present
an identification key, updated taxonomic descriptions, and photographic illustrations for each species. Two lectotypifications
are proposed for Clusia. A new combination in Garcinia is also proposed.
Keywords: Amazon, biodiversity, Guttiferae, taxonomy
Resumo
Relatamos um levantamento florístico das espécies de Clusiaceae s.l. (Calophyllaceae, Clusiaceae s.s. e Hypericaceae) no
Parque Nacional do Viruá (Roraima, Brasil). O trabalho de campo foi realizado entre 2009 e 2015, cobrindo os diferentes
tipos de vegetação encontrados no parque. Calophyllaceae é representada por dois gêneros e três espécies, Clusiaceae s.s. por
cinco gêneros e 12 espécies e Hypericaceae por um gênero e cinco espécies. Três espécies são novos registros para o estado
de Roraima. Apresentamos uma chave de identificação, descrições taxonômicas atualizadas e ilustrações fotográficas para
cada espécie. Duas lectotipificações são propostas para Clusia. Uma nova combinação em Garcinia também é proposta.
Palavras-chave: Amazônia, biodiversidade, Guttiferae, taxonomia
Introduction
The clusioid clade belongs to the order Malpighiales and is represented by five families: Bonnetiaceae, Calophyllaceae,
Clusiaceae s.s., Hypericaceae and Podostemaceae (APG IV 2016, Ruhfel et al. 2011, Ruhfel et al. 2013, Wurdack &
Davis 2009, Xi et al. 2012). Calophyllaceae, Clusiaceae s.s. and Hypericaceae were previously known as Clusiaceae
or Guttiferae (Cronquist 1981, Mabberley 2008), but the latest molecular studies showed that the family in this
circumscription was not monophyletic (APG IV 2016, Ruhfel et al. 2011, Ruhfel et al. 2013, Wurdack & Davis 2009,
Xi et al. 2012). Calophyllaceae, Clusiaceae s.s. and Hypericaceae are Pantropical and include 36 genera and about
1,960 species (Stevens 2001, onwards). In Brazil, they are present in all phytogeographical domains (Amazon, Atlantic
Forest, Caatinga, Cerrado, Pampa and Pantanal), with a total of 20 genera and 267 species (BFG 2015). In the state of
Roraima the three families are represented by 13 genera and 46 species in several phytophysiognomies, such as terra-
firme forest, wetlands (igapó and várzea forests) and white-sand vegetation (BFG 2015).
The Viruá National Park (VNP) is located in the central-south state of Roraima (northern Brazil). The VNP
is an important conservation area for the state and the Brazilian Amazon Biome. The park hosts a wide variety of
Accepted by Carlos Lehnebach: 19 Oct. 2017; published: 24 Nov. 2017

environments and vegetation types (Schaefer et al. 2009) and its diversity has been the subject of several ecological and
taxonomic studies (Costa et al. 2016a, 2016b, Dávila 2011, ICMBio 2014, Melo et al. 2014, Mota et al. 2015, Pessoa et
al. 2015), most of them conducted as part of a PNADB program (Programa Nacional de Apoio e Desenvolvimento da
Botânica/CAPES). According to data reported by ICMBio (2014), Clusiaceae s.l. figures among the 10 most species-
rich families in VNP. It needs to be emphasized, however, that the species listed for these families in the VNP are
mostly based on field observations, for which no herbarium vouchers exist.
The aim of this work is to report on the diversity of the families Calophyllaceae, Clusiaceae s.s. and Hypericaceae,
collectively known as Clusiaceae s.l., occurring in the Viruá National Park. We provide a key to species, updated
taxonomic descriptions, photographs, and comments on the distribution, ecology, nomenclature and taxonomy of all
species.
Material and methods
Study area
The VNP is located in Caracaraí, Roraima, (1º28’N, 61ºW), in northern Brazil (Fig. 1). It is managed by the Instituto
Chico Mendes de Conservação da Biodiversidade (ICMBio) and has an area of 241,948 hectares. The climate is
equatorial (hot and humid) with a short dry season; the driest months between September and January. The park
presents a wide variety of environments, forming a mosaic of different vegetation types (Schaefer et al. 2009). This
mosaic is formed mostly by white-sand vegetation locally known as “campinarana”, various formations of tropical
rain forest, alluvial tropical rain forest, swamps (“buritizais”) dominated by Mauritia flexuosa Linnaeus (1782: 454)
and grassy areas (Veloso et al. 1991).
Field work
We conducted ten field trips between November 2009 and October 2012 and one trip in January 2015. The specimens
were collected and photographed in situ. We also collected flowers and fruits and these were fixed in 70% ethanol.
Both dried voucher specimens and spirits collections were deposited at the INPA, UFRR, UEC and UFP herbaria.
Herbarium acronyms according to Thiers (2017, continuously updated).
Taxonomic and morphological descriptions

The material collected at VNP was used to update morphological descriptions, obtain phenological data and construct
the identification keys. Herbarium specimens from INPA, UEC and UFP and related literature (Abdul-Salim 2002,
Bittrich 2008, Bittrich & Amaral 1996, Carvalho et al. 2003, Díaz 2013, Pipoly & Gustafsson 2003, Pipoly et al.
2003, Reichardt 1878, Van den Berg 1974) were used to complement our data when necessary. We followed the
terminology used by Endress (2010), Harris & Harris (2001), Hewson (1990), Hickey & King (2000) and Stevens
(2001, onwards).
Results
Altogether, the Clusiaceae s.l. are represented by 20 species in the VNP. Calophyllaceae is represented by two genera,
Calophyllum (one species) and Caraipa (two species). Clusiaceae s.s. is represented by five genera, Clusia (seven
species), Garcinia (one species), Platonia (one species), Symphonia (one species) and Tovomita (two species).
Hypericaceae is represented by a single genus, i.e. Vismia comprising five species. Garcinia albuquerquei, a rare
species, previously considered endemic to the Middle and Upper Rio Negro region, was collected only twice in VNP,
in white-sand vegetation. Further, two species of Caraipa (C. caespitosa and C. savannarum) and two of Clusia (C.
lopezii and C. nitida) were only found in white-sand vegetation, where they were locally abundant.
• Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

FIGURE 1. A. Map of Viruá National Park.
Key to the species of Clusiaceae s.l. in the VNP

1. Leaves alternate...................................................................................................................................................................................2
– Leaves opposite...................................................................................................................................................................................3
2. Shrubs up to 70 cm tall; leaf base cordate; secondary veins 11–19 pairs; flowers 4.0–4.5 cm wide; calyx light brown; pedicel and
peduncle tomentose; fruit 1.7–2.6 cm long........................................................................................................... Caraipa caespitosa
– Small trees or trees up to 3 m; leaf base acute to obtuse; secondary veins 5–12 pairs; flowers 1–1.2 cm wide; calyx green; pedicel
and peduncle glabrous; fruit 0.7–1.1 cm long.....................................................................................................Caraipa savannarum
3. Leaves with secondary veins extremely dense (20–36 veins/cm); ovary 1-locular, 1-ovule........................ Calophyllum brasiliense
– Leaves with secondary veins sparse (5–76 pairs of veins/leaf); ovary multilocular, 1 to many ovules..............................................4
4. Lateral branches with long basal internode; mature bud enclosed by outer sepals; endocarp bright red...........................................5
– Lateral branches with short basal internode; mature bud not enclosed by outer sepals; endocarp brown to cream...........................6
5. Internodes short, distal ones up to 5.5 cm long; midvein prominent on the abaxial surface and impressed or flat on the adaxial
surface; petiole 21–44 mm long........................................................................................................................... Tovomita spruceana
– Internodes long, distal ones up to 10 cm long; midvein prominent on both surfaces; petiole 7–20 mm long......Tovomita umbellata
Clusiaceae s.l. in the Viruá National Park Phytotaxa 329 (1) © 2017 Magnolia Press •
6. Leaves sessile, leaf blade with large basal adaxial margined pits; fruit a coriaceous berry............................. Garcinia albuquerquei
– Leaves petiolated, leaf blade with small basal adaxial margined pits; fruit a capsule or a berry........................................................7
7. Flowers bisexual; petal aestivation contorted; fruit a berry................................................................................................................8
– Flowers unisexual; petal aestivation quincuncial; fruit a capsule.....................................................................................................14
8. Shrubs or small trees; exudate brightly colored drying reddish on exposure; lamina tomentose on most species with conspicuous
dark glands; secondary veins 8–31 pairs, distant 3.8–20 mm.............................................................................................................9
– Trees; exudate yellow or cream not drying reddish on exposure; lamina glabrous without conspicuous dark glands; secondary veins
34–76 pairs, distant 1–2 mm.............................................................................................................................................................13
9. Leaf blade glabrous on both surfaces, 5.9–9.0 cm long; calyx yellowish green....................................................Vismia cayennensis
– Leaf blade adaxially glabrous, abaxially rusty to brown tomentose, 5.9–42 cm long; calyx whitish, green or brown....................10
10. Leaf blade 10–42 cm long, base cordate to rounded; secondary veins 12–31 pairs.............................................Vismia macrophylla
– Leaf blade 5.9–15.8 cm long, base acute, cuneate, obtuse to rounded; secondary veins 8–15 pairs................................................11
11. Fascicles of stamens persistent in fruit................................................................................................................................. Vismia sp.
– Fascicles of stamens deciduous in fruit.............................................................................................................................................12
12. Petals 12–15 mm long, deciduous; sepals rigidly coriaceous; fruit 1.0–1.7 cm long.............................................. Vismia japurensis
– Petals 8.0–10 mm long, persistent; sepals fleshy; fruit 0.8–0.9 cm long.................................................................... Vismia laxiflora
13. Leaf blades 5.1–10 × 2.4–3.7 cm; flowers axillary cymes; stamens forming a staminal tube surrounding the gynoecium, 3 stamens
each per fascicle; petals scarlet, shorter than 15 mm long............................................................................... Symphonia globulifera
– Leaf blades 6.9–11.7 × 3.7–7.8 cm; flowers solitary; stamens not forming a staminal tube, ca. 100 stamens per fascicle; petals
peach colored, longer than 35 mm............................................................................................................................ Platonia insignis
14. Tertiary veins conspicuous; lamina 16–24 cm long; staminate flowers with pistillode and pistillate flowers with 5 stigmas.............
.............................................................................................................................................................................. Clusia candelabrum
– Tertiary veins inconspicuous; lamina 3.5–17 cm long; staminate flowers without pistillode and pistillate flowers with more than 5
stigmas...............................................................................................................................................................................................15
15. Exudate channels in the lamina inconspicuous.................................................................................................................................16
– Exudate channels in lamina conspicuous, at least on one surface.....................................................................................................17
16. Lamina (5–)5.6–12(–13) long; bracts early deciduous; corolla inside white with red center; petals 9–13 mm long; number of
stigmas 4–5(–6)................................................................................................................................................... Clusia renggerioides
– Lamina 13–25 cm long; bracts persistent; corolla inside dark red to vinaceous; petals 40–60 mm long; number of stigmas 15–21..
.......................................................................................................................................................................................Clusia insignis
17. Corolla persistent in fruit, white-pinkish at apex; stigmas divergent; anthers annular, opening by a transversal slit.....Clusia lopezii
– Corolla deciduous in fruit, white-reddish, with pink or dark purple center; stigmas parallel; anthers not annular, opening with
apical, transverse or longitudinal slits...............................................................................................................................................18
18. Corolla internally white-red, with pink or white center; staminodes 5–6 mm long; stamens free; fruit 2.2–4.0 cm long, 6–8 seeds
per locule....................................................................................................................................................................Clusia nemorosa
– Corolla internally white-red, with pink or dark purple center or colored for most of its surface; staminodes 1.3–1.9 mm long;
stamens densely grouped in a flat circular or obtusely 5–angular disk; fruit 1–2 cm long, 1–5 seeds per locule............................19
19. Exudate channels on abaxial surface of leaf blades 1.2–3.0 mm distant; corolla white, violet-red or red, bracts and sepals white-
pinkish; staminate flower with androecium in a flat circular disk with 8–9 mm wide, anthers present on lateral side of the disk......
...............................................................................................................................................................................Clusia microstemon
– Exudate channels on abaxial surface of leaf blades 0.3–0.7 mm distant; corolla dark or blood red, bracts and sepals green; staminate
flower with androecium in a flat circular disk-like with 3–6 mm wide, anthers absent on the lateral sides of the disk.......................
..........................................................................................................................................................................................Clusia nitida
Taxonomic Treatment
Calophyllaceae J. Agardh
Type: Calophyllum Linnaeus (1753: 513).
Trees or shrubs, rarely lianas. Exudate cream, white or translucid. Leaves simple, opposite or alternate (distichous
or spiral), usually coriaceous or fleshy, stipules lacking, margin entire, with secretory cavities or channels, colleters
lacking or present. Cymes or racemes, usually terminal, rarely solitary flowers. Flowers bisexual or rarely unisexual,
actinomorphic, white or cream, 4–5-merous, sepals (2–)4–6(–17)-merous, imbricate; petals (2–)4–5(–12)-merous,
free, imbricate or convolute; stamens numerous, often with complex or simple apical oil-glands; ovary superior, 2–5
carpels, placentation axillary, basal or parietal, style usually long; ovules vary from 1 to many per carpel. Berries or
capsules. Seeds1-many. Embryo with cotyledons huge or of moderate size.
Calophyllaceae are composed of 14 genera and about 460 neo- and paleotropical species (Stevens 2001, onwards,
Wurdack & Davis 2009) and it can be divided in two tribes: Calophylleae (pantropical) and Endodesmieae (restricted
to tropical Africa) (Stevens 2001, onwards, Ruhel et al. 2011). In Brazil, the family is represented by 83 species (BFG
2015).

1 Calophyllum Linnaeus (1753: 513).
Type: Calophyllum calaba Linnaeus (1753: 514).
Trees or treelets; exudate yellow or cream. Leaves opposite, decussate, secondary veins densely parallel. Racemes,
axillary, bracts deciduous, bracteoles absent. Flowers bisexual or unisexual (plants usually dioecious); tepals 0–8,
opposite or imbricate, white. Stamens numerous. Ovary 1-locular, 1 basal ovule, style 1 and usually long, stigma
expanded. Berries, coriaceous, usually globular. Seed 1, testa woody.
Pantropical; with ca. 190 species (Díaz 2013, Stevens 2001, onwards) and four species in Brazil (BFG 2015).
1.1 Calophyllum brasiliense Cambessèdes (1825: 320) (Fig. 2A–C)

Type:—BRAZIL. Espírito Santo, s.d., fl., Saint-Hilaire 330 (holotype: P, barcode 00761176).
= Calophyllum angulare A.C.Sm (1933: 379)
= Calophyllum ellipticum Rusby (1927: 303).
= Calophyllum lucidum Bentham (1843: 370).
= Calophyllum mariae Planch. & Triana (1861: 251).
= Calophyllum piaroanum An. Castilho & C. Gil (1991: 41).
Trees or small trees up to 25 m. Exudate yellow, cream or white. Leaf blade 4.2–17.7 × 2.2–6.8 cm, coriaceous
to subcoriaceous, elliptical, oblong–elliptic to oboval, apex obtuse, rounded to retuse, cuneate, base acute; exudate
channels inconspicuous; secondary veins 20–36 veins/cm, conspicuous on both surfaces; petiole 4.8–17.2 mm long.
Inflorescence 3–5-flowered. Flowers bisexual, ca. 10 mm long, pedicel 2–10 mm long. Outer tepals 2.7–4.3 × 1.7–2.6
mm. Inner tepals 2–4(–5), 4–6.3 × 1.8–4.3 mm, recurved. Stamens 11–28 × 2.3–3.6 mm long. Ovary ca. 4 mm across,
style ca. 2 mm long, stigma 0.5–2 mm wide, peltate. Fruit 0.9–2.5 × 0.6–2.1 cm, ellipsoid or ovoid, apex obtuse. Seeds
1.1–1.5 × 1.1–1.3 cm, ellipsoid or ovoid.
Phenology:—Flowering September to November; fruiting October to January.
Distribution and habitat:—Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Panama, Peru, Suriname
and Venezuela. In Brazil, it occurs in the north region (states of Acre, Amazonas, Pará, Rondônia and Roraima), in the
northeast (states of Bahia, Maranhão and Pernambuco), in the central-west region (Distrito Federal, states of Goiás,
Mato Grosso and Mato Grosso do Sul), in the southeast region (states of Espírito Santo, Minas Gerais, Rio de Janeiro
and São Paulo), and in the south region (states of Paraná and Santa Catarina). In VNP, in white-sand vegetation and
flooded areas.
Specimens examined:—BRAZIL. Roraima: Caracaraí, Viruá National Park, elevation 62 m, 1º28’06’’N,
61º00’32’’W, 24 November 2009, fl., Cabral et al. 269 (INPA); Ibid, 20 January 2011, fr., Cabral et al. 362 (INPA).
Additional specimens:—BRAZIL. Minas Gerais: Diamantina, Biri-Biri, 25 February 1975, fl., Hatschbach et
al. 36489 (INPA). Amazonas: Manaus-Itacoatiara, Reserva Florestal Ducke, km 26, 02º53’S, 59º58’W, 04 December
1993, fr., Vicentini et al. 381 (INPA); Ibid., 15 August 1997, fl., Hopkins et al. 1632 (INPA).
Taxonomic notes:—Calophyllum is easily recognizable by its decussate leaves and parallel and extremely dense
secondary veins in combination with the presence of exudate (Díaz 2013). However, species delimitation is difficult
because of infraspecific variation, such as the number of tepals which can vary widely. The specimens we collected fit
the description and the taxonomic key prepared by Díaz (2013).
2 Caraipa Aublet (1775: 561).

Type: Caraipa parvifolia Aublet (1775: 561).
Trees, shrubs or small trees; exudate present, scarce, resiniferous and viscose. Leaves alternate, distichous or spiral;
secondary veins conspicuous. Panicules, racemes or rarely cymes. Flowers bisexual, fragrant, bracts and bracteoles
persistent or deciduous; sepals 5, greenish, petals 5, yellow to whitish, contorted. Stamens numerous, with a conspicuous
cup-shaped oil-gland on the connective that secretes a fragrant oil. Ovary 3-locular, 2–3 ovules per locule, style 1, 3-
lobed at apex, stigmas expanded. Capsules, woody, dehiscence septifragal, exocarp may separate from the endocarp.
Seeds 1–3, winged.
Neotropics; with ca. 35 species (Cabral et al. 2016a, 2016b, Kubitzki 1978, Kubitzki & Holst 1998) and 24
species in Brazil (BFG 2015).
FIGURE 2. Calophyllum brasiliense (A. Habit; B. Flower; C. Fruit), Caraipa caespitosa (D. Habit; E. Flower; F. Fruit) and Caraipa
savannarum (G. Flower; H. Fruit).
2.1 Caraipa caespitosa F.N. Cabral in Cabral et al. (2016b: 248) (Fig. 2D–F)
Type:—BRAZIL. Roraima: Caracaraí, Parque Nacional do Viruá, igarapé Anauá, 23 March 2008, fr., Ferreira et al. 13082 (holotype:
INPA!, barcode 253111).
Shrubs, up to 70 cm tall, with numerous (20–40) sparsely branched stems growing from a mound. Exudate cream
or translucid. Leaves distichous, leaf blade 4–7.5 × 1.8–4.5 cm, coriaceous, elliptical to oval, apex rounded to acute
or obtuse, base cordate to rounded, adaxial surface glabrous, abaxial surface with minute sessile stellate trichomes
or glabrous, with bulliform cells; secondary veins 11–22 pairs, distant 4–6 mm; petiole 2.4–7.6(–10.5) mm long.

Panicule, (1–)3–14-flowered, peduncle and pedicels tomentose, pedicel 3–13 mm long. Sepal lobes 2–4 × 2–4 mm.
Petals 14–27 × 10–14 mm. Stamens 3–10 mm long, gland at the apex of the thecae. Ovary ca. 2 mm across, style ca. 4
mm long. Fruit 17–26 × 14–19 mm, ovoid to pyramidal, rugulose to smooth, densely covered with sessile and stipitate
stellate hairs, exocarp separating from endocarp. Seeds ca. 10 × 8 mm.
Phenology:—Flowering September to December; fruiting January to March.
Distribution and habitat:—Brazil and Venezuela. In Brazil, it occurs in the north region (states of Amazonas and
Roraima). Caraipa caespitosa occurs on white-sand vegetation.
60º59’13’’W, 3 December 2009, fl., Cabral et al. 101 (INPA); Ibid., 1º23’39’’N, 61º13’25’’W, 22 January 2010, fr.,
Cabral et al. 167 (INPA); Ibid., 14 September 2010, fl., Cabral et al. 270 (INPA); Ibid., 12 October 2010, fl., Cabral
et al. 294 (INPA).
Additional specimens:—BRAZIL. Amazonas: Final da Perimetral 1—Norte da Perimetral Norte (RR210), 25
March 1999, fr., Vicentini 1416 (MO!).
Taxonomic notes:—Caraipa caespitosa is morphologically similar to C. grandifolia and C. longipedicellata.
Caraipa caespitosa can be easily distinguished by its habit, which is small shrub with numerous sparsely branched
stems growing from a mound, versus trees in C. grandifolia and C. longipedicellata. The species can also be identified
by leaf size, number of secondary veins, and inflorescence size, see Cabral et al. (2016b) for more details.
2.2 Caraipa savannarum Kubitzki (1978: 104) (Fig. 2G–H)

Type:—BRAZIL. Amazonas: Rio Curicuriary affl. Rio Negro, 23 November 1936, fr., Ducke RB 35414 (holotype: M, barcode 0112098,
isotype: NY, barcode 00067011).
Trees or treelets up to 3 m. Exudate sticky and translucid. Leaves spirally-alternate; leaf blade 3.3–6.4(–7.7) × (1.6–)
2.1–4.8(–5.8) cm, chartaceous, ovate or orbicular, apex rounded or acuminate, base acute or obtuse, both surfaces
glabrous, without bulliform cells; secondary veins 5–12 pairs, distant 3–7.4 mm; petiole 3.2–6.1 mm long. Panicule,
2–6-flowered, peduncle and pedicels pubescent, pedicel 1–6 mm long. Sepal lobes ca. 1.2 × 1.3 mm. Petals 5.3–6.3
× 2.2–4.5 mm. Stamens 2–6 mm long, gland between the thecae. Ovary ca. 1.5 mm across, style 1.5–2.0 mm long.
Fruit 0.8–1.1 × 0.8–1.1 cm, pyramidal to ovoid, asymmetrical, minutely rugulose, with sparse sessile stellate hairs or
glabrous, exocarp separating from endocarp. Seeds 7–8 mm long.
Phenology:—Flowering September to January; fruiting September to April.
Distribution and habitat:—Brazil, Guyana and Venezuela. In Brazil, it occurs in the north region (states of
Amazonas, Rondônia and Roraima). In VNP, it was found growing on white-sand vegetation.
60º58’10’’W, 25 November 2009, fr., Cabral et al. 17 (INPA); Ibid., elevation 57 m, 1º21’34’’N, 61º13’57’’W, 22
January 2010, fr., Cabral et al. 164 (INPA); Ibid., elevation 62 m, 1º30’32’’N, 61º37’48’’W, 23 January 2010, fl.,
Cabral et al. 178 (INPA); Ibid., 16 September 2010, fr., Cabral et al. 285 (INPA); Ibid., 22 October 2010, fl., Cabral et
al. 313 (INPA); Ibid., 25 October 2010, fr., Cabral et al. 331 (INPA); Ibid., elevation 55 m, 1o26’28”N, 60o58’45”W, 31
January 2010, fl., Dávila et al. 6025 (INPA); Ibid., elevation 57 m, 1o14’49”N, 60o57’59”W, 4 March 2010, fr., Dávila
et al. 6106 (INPA).
Additional specimens:—BRAZIL. Amazonas: Bacia do Rio Negro, 8 November 1971, fr., Prance et al. 16211
(INPA); Rio Univini, 23 April 1974, fr., Pires et al. 14159 (INPA). Pará: Região dos Tiriós, Rio Paru do Oeste, 17
March 1962, fr., Fittkau et al. 39 (INPA).
Taxonomic notes:—Caraipa savannarum can be distinguished by its spiral, glabrous, ovate or orbicular leaves,
and its small fruits, which are sparsely covered with sessile stellate hairs, with concave valves and usually with
longitudinal black lines on the surface.
Clusiaceae Lindley
Type: Clusia Linnaeus (1753: 509).
Trees, shrubs or herbs, terrestrial, hemi-epiphytic or lianas. Exudate white, cream, yellow or orange. Leaves simple,
opposite, coriaceous or cress, stipules lacking, margin entire, with secretory cavities, colleters present or lacking,
petiole base sometimes with an intrapetiolar cavity, sometimes perulae present, leaving scars on branches after falling;
leaf blades with linear exudate channels and/or glandular dots. Cymes, or solitary flowers, terminal or axillary. Flowers
bisexual or unisexual (usually dioecious), actinomorphic, sepals (2–)4–6(–17), free, imbricate; petals (2–)4–6(–14),
white, yellow or pink, imbricate or convolute; stamens 4 to numerous, sometimes grouped in fascicles or whorls,
or fused into a synandria, staminodes similar to stamens or connate in resiniferous rings or in a central mass; ovary
superior, (2–)3–5(–21) carpels, placentation axillar, ovules vary from 1 to many per carpel, style single or in equal
number to the carpels, stigma 1 per locule. Capsules or berries, fleshy. Seeds (1–)2 to many per locule, sometimes
arillate or surrounded by an edible pulp, cotyledons minute.
Clusiaceae s.s. are composed of 14 genera and about 800 neo- and paleotropical species. It has been divided into
three tribes: Clusieae (neotropical), Garcinieae (pantropical) and Symphonieae (pantropical) (Stevens 2001, onwards,
Ruhfel et al. 2011). In Brazil, the family is represented by 131 species (BFG 2015).
3 Clusia Linnaeus (1753: 509).

Type: Clusia major Linnaeus (1753: 509).
Shrubs, trees, small trees or hemi-epiphytes. Exudate white, cream, yellow, translucid or yellowish-brown, usually
abundant. Leaves opposite; exudate channels visible in most species, colleters present. Cymes or solitary flowers.
Flowers usually unisexual (mostly dioecious); with epicalyx bracts, sepals 2–numerous, petals 4–10, quincuncial;
floral resin secreted by staminodes and/or stamens. Staminate flowers with 4-numerous stamens, staminodes and
pistillode present or absent. Pistillate flowers with staminodes shorter than stamens, stigmas sessile or on short styles,
equal to the number of locules of the ovary. Ovary with 4-numerous locules, with 1 to usually many ovules per locule.
Capsule, carnose-coriaceous, dehiscence septifragal, stigma persistent, calyx persistent in most species. Seeds usually
more than one per locule, with reddish, orange or yellowish aril.
Neotropics and subtropics; with ca. 300–400 species (Pipoly et al. 1998, Stevens 2001, onwards) and 67 species
in Brazil (BFG 2015).
3.1 Clusia candelabrum Planchon & Triana (1860: 348) (Fig. 3A–B)
Type:—BRAZIL. Amazonas: Rio Uaupés prope Panuré, October 1852–January 1853, fl., Spruce 2431 (holotype P: barcode 01901134,
isotype: GOET, barcode 003940).
= Clusia grandifolia Engler (1888: 429).
Hemiepiphytes up to 5 m, roots stilt. Exudate creamy-white. Leaf blade coriaceous, galls usually present, elliptical-
oboval to oblanceolate, 16–24 × 8.1–12.5 cm, apex rounded, base acute; exudate channels conspicuous on both
surfaces, distant 2–4 mm; secondary veins conspicuous on both surfaces, 39–58 pairs, distant 2.8–4 mm; petiole
24.4–36.9 mm long. Inflorescence: staminate plants with clustered flowers; pistillate plants with solitary or clustered
flowers; pedicels 2.0–2.4 cm long. Sepals 4–5, 8.2–13.9 × 9.2–14.8 mm, white. Petals 5, 25–30 mm long, white and
red. Staminate flowers with more than 50 fertile resiniferous stamens, united at the base, with sessile anthers at the
top; pistillode conspicuous and higher than stamens; pollen mixed with oil or resin. Pistillate flowers with more than
20 resiniferous staminodes, 3.5–4.0 mm long, united at the base; stigma 5, 5–7 mm long, with conspicuous space at
the apex. Fruit oblong to cylindrical, 3.2–4.0 × 3–3.5 cm; sepals, petals, staminodes and stigma persistent. Seeds with
orange aril.
Phenology:—Flowering May to September; fruiting January, April and October to November.
Distribution and habitat:—Brazil, Colombia, Ecuador, Peru and Venezuela. In Brazil, it occurs in the north
region (states of Amazonas, Rondônia and Roraima). In VNP, it has been found in flooded areas.
Specimens examined:—BRAZIL. Roraima: Caracaraí, Viruá National Park, 1º25’11’’N, 60º51’20.6’’W, 18
January 2011, fr., Cabral et al. 345 (INPA).
Additional specimens:—BRAZIL. Amazonas: Camanau/Curiau, 28 June 1987, fl., Grenard et al. 9786 (INPA);
Presidente Figueiredo, Rio Uatumã, 1º2’S, 59º60’W, 19 March 1986, fr., Ferreira et al. 6877 (INPA); Lábrea, Bacia
do Rio Purus, 07º16’S, 64º47’W, 29 October 1968, fr., Prance et al. 8060 (INPA); Ibid., 9 July 1971, fl., Prance et al.
14021 (INPA); Rio Negro, 0º14’S, 66º48’W, 15 May 1973, fl., Silva et al. 1644 (INPA); Manaus-Itacoatiara, 02º05’S,
60º05’W, 4 April 1967, fr., Prance et al. 4774 (INPA); Ibid., 02º46’S, 59º38’W, 14 October 1966, fr., Prance et al.
3158 (INPA). Rondônia: Rio Madeira, 06º59’S, 62º42’W, 18 September 1962, fl., Duarte 7104 (INPA).
Taxonomic notes:—Clusia candelabrum is easily distinguishable from other Clusia species that occur in VNP.
Its leaf blade is larger than in the other species, with the exception of C. insignis. Furthermore, C. candelabrum is the
only species in which tertiary veins are conspicuous.

FIGURE 3. Clusia candelabrum (A. Staminate flower; B. Fruit), Clusia insignis (C. Habit with pistillate and staminate flower; D.
Staminate flower; E. Fruit; photos from specimens collected in Gran Sabana, Venezuela, Amaral & Bittrich 91-26) and Clusia lopezii (F.
Pistillate flower; G. Staminate flower; H. Fruit).
3.2 Clusia insignis Martius (1829: 164) (Fig. 3C–E)

Lectotype (designated here):—BRAZIL. Amazonas: Oppidum Barra do Rio Negro (=Manaus), October 1818, fl., Martius s.n., (lectotype:
M, barcode 0086179, isolectotypes: M, barcodes 0089137, 0086180).
Epiphytes, stranglers, or trees up to 30 m. Exudate yellow to cream. Lamina coriaceous, oboval, oboval-oblong or
oboval-cuneate, 13–25 × 6–15 cm, apex rounded, base attenuate to cuneate; exudate channels inconspicuous on
both surfaces; secondary veins conspicuous on both surfaces, 35–50 pairs, distant 1.5–3 mm; petiole 2.5–5 cm long.
Inflorescence or flowers pendant: staminate plants with clustered or solitary flowers; pistillate plants with solitary
flowers; pedicels 1–2(–3) cm long. Sepals 4, 18–20 × 17–20 mm, whitish to reddish. Petals 7–9, 40–60 × 30–35 mm,
dark red to vinaceous adaxially, white abaxially. Staminate flowers with numerous stamens, in 8–12 series, united at
the base; staminodes, united until the middle, forming a central flat disk, up to 1.5 cm wide, covered with yellow resin,
pistillode absent. Pistillate flowers with numerous resiniferous staminodes, ca. 6 mm long, in 6–8 series surrounding
the base of the ovary; stigmas 15–21, ca. 4 mm long, connivent. Fruit globose, 2.5 × 3 cm; sepals and staminodes
persistent, stigmatic area ca. 14 mm wide. Seeds red, with yellow aril.
Phenology:—Flowering August to December; fruiting November to January.
Distribution and habitat:—Brazil and Venezuela. In Brazil, in the north region (states of Amapá, Amazonas,
Pará and Roraima). In VNP, it grows on white-sand vegetation.
Specimens examined:—BRAZIL. Roraima: Caracaraí, Viruá National Park, margem direita da Estrada Perdida
em direção à primeira bueira (sul), a cerca de 800 m da bifurcação para a entrada do Parque, 01º28’22.9”S, 60º58’7.9”W,
12 January 2015, fr., Amaral & Bittrich 2015/13 (INPA!).
Additional specimens:—BRAZIL. Roraima: Plateau of Serra Tepequem, elevation 1,200 m, 18 February 1967,
fl., Prance et al. 4505 (NY, UEC!).
Taxonomic notes:—This species is easy to distinguish from all other Clusia species in VNP by its yellowish
exudate, the long petiole abruptly broadened at the base and the large leaf blades.
3.3 Clusia lopezii Maguire (1951: 61) (Fig. 3F–H)

Type:—COLOMBIA. Vaupes: Rio Negro, s.d., fl., Schultes & Lopez 9388 (holotype: NY, barcode 00380995, isotype: GH, barcode
00067427).
Shrubs, trees or small trees up to 7 m. Exudate white. Lamina subcoriaceous to coriaceous, oboval to elliptical, 6.4–
13.7 × 2.9–8 cm, apex obtuse and rounded, base attenuate and acute; exudate channels conspicuous on the abaxial
surface, barely visible on the adaxial surface, distant 4.8–8.0 mm; secondary veins conspicuous on both surfaces,
18–48 pairs, distant 1.8–3 mm; petiole 14.6–30.9 mm long. Inflorescence: staminate plants with clustered flowers;
pistillate plants with clustered or solitary flowers; pedicels 1.0–2.0 mm long. Sepals 4–6, 5.7–13.4 × 6.4–14.0 mm,
white to slightly pink. Petals 4–8, 10.7–28.4 × 9.7–23.4 mm, pinkish-white to white. Staminate flowers with stamens
more than 50, forming a synandrium with sessile anthers on top, 1.5–2 mm long, resin absent; androecium 7–8 mm
wide, forming a circular, convex disk; staminodes numerous; pollen not mixed with oil or resin. Pistillate flowers
with ca. 70 staminodes, 3–5 mm long, arranged around the base of the ovary, densely clustered, with resin at the apex;
stigmas 4–6, 2.3–3.8 mm long, with a conspicuous space between them. Fruit ovoid to ellipsoid, 1.9–3.8 × 1.3–2.4 cm;
sepals, petals, staminodes and stigmas persistent. Seeds 4.3–5.0 × 3.0–3.3 mm (immature).
Phenology:—Flowering September to October; fruiting July, December to January.
Distribution and habitat:—Brazil, Colombia and Venezuela. In Brazil, it occurs in the northern region (states of
Amazonas and Roraima). In VNP, it is found in flooded areas and white-sand vegetation.
60º59’08’’W, 4 December 2009, fr., Cabral et al. 113 (INPA); Ibid., 21 July 2010, fr., Cabral et al. 265 (INPA); Ibid.,
16 September 2010, fl., Cabral et al. 290 (INPA); Ibid., 11 October 2010, fl., Cabral et al. 293 (INPA); Ibid., 20
January 2011, fr., Cabral et al. 363 (INPA).
Taxonomic notes:—Clusia lopezii has an androecium in the form of a circular, convex disk, somewhat similar
as in C. microstemon Planchon & Triana (1860: 331) and C. nitida Bittrich & F.N. Cabral in Bittrich et al. (2013: 36),
but is easily distinguishable by having resin-secreting staminodes in the center surrounded by the stamens arranged
in a broad ring. Clusia microstemon and C. nitida, on the other hand, lack staminodes in the staminate flowers, and in
these species, the pollen is mixed with fluid resin secreted by the stamens themselves, while in C. lopezii the pollen is
separated from the viscous resin.
3.4 Clusia microstemon Planchon & Triana (1860: 331) (Fig. 4A–C)
Type:—BRAZIL. Amazonas: Rio Uaupés, October 1852, fl., Spruce 2511 (holotype: P, barcode 01901205, isotypes: BR, barcode
0000008676702, C, barcode 10009454, E, barcode 00326699, G, barcode 00355089, GH, barcode 00026868, K, barcode 000820682,
LD, barcode 1060294, NY, barcode 380975).
= Clusia gaudichaudii Planchon & Triana (1860: 359), nom. illeg.
Shrubs, small trees up to 3 m or hemiepiphytes. Exudate yellowish to yellowish-brown. Lamina subcoriaceous to

coriaceous, oboval or elliptical to oblanceolate, (3.5–)4.4–14 × 2.6–8.1 cm, apex obtuse, rounded to acute or subacute,
10 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

base attenuate and acute; exudate channels conspicuous on both surfaces, distant 1.2–3.0 mm; secondary veins
conspicuous on both surfaces, 13–38 pairs, distant 1.7–2.5 mm; petiole (6.5–) 8.2–38.4 mm long. Inflorescence:
staminate plants with clustered flowers; pistillate plants with clustered or solitary flowers; pedicels 0.8–1.0 cm long.
Sepals 4, 7.0–10.6 × 7.2–10.3 mm, white to slightly pink. Petals 5–7, 13.7–19.6 × 12.1–15.4 mm, violet-pink or
white and violet-pink adaxially, white-pinkish abaxially. Staminate flowers with stamens numerous, ca. 1600, densely
compact, 1.8–2.2 mm long, forming a flattened synandrium with sessile anthers on top, 8–9 mm wide, the lateral side
of the disk with anthers, covered with resin-pollen mix during anthesis. Pistillate flower with 2 series of staminodes
around the base of the ovary, with resin at the apex; stigma 4–6, 2–4 mm long, covered with small rounded papilla.
Fruit cylindrical to ovoid, 1.0–1.4 × 1.0–1.4 cm; sepals and stigmas persistent. Seeds with yellowish-orange aril.
Phenology:—Flowering June to September; fruiting January.
Distribution and habitat:—Brazil, Colombia and Venezuela. In Brazil, it occurs in the north region (states
of Acre, Amapá, Amazonas, Pará, Rondônia and Roraima), and in the northern central-west region (state of Mato
Grosso). In VNP, it is found in white-sand vegetation and at river margins.
61º01’18’’W, 28 January 2010, fr., Cabral et al. 190 (INPA); Ibid., 24 July 2010, fl., Cabral et al. 249 (INPA); Ibid.,
24 July 2010, fl., Cabral et al. 251A (INPA); Ibid., 16 September 2010, fl., Cabral et al. 287 (INPA); Ibid., 15 January
2011, fr., Cabral et al. 337 (INPA).
Additional specimens:—BRAZIL. Amazonas: Entre Paraná da Floresta e boca do rio Branco. Praia do Gavião,
26 June 1979, fl., Maia et al. 57 (INPA).
Taxonomic notes:—For differences between Clusia microstemon and C. nitida see notes under C. nitida.
3.5 Clusia nemorosa Meyer (1818: 203) (Fig. 4D–E)

Type:—SURINAME. Ilha Wacanama: E.C. Rodschied 299 (holotype: GOET, barcode 003942).
= Clusia bicolor Martius (1832: 164).
= Clusia lhotzkyana Schlechtendal (1833: 184).
= Clusia mammosa Casaretto (1843: 60).
Trees, shrubs or small trees up to 15 m. Exudate, white or cream. Lamina subcoriaceous to coriaceous, elliptical,
oval or oboval, (6.6–)7.4–17 × 3.5–6.9(–7.2) cm, apex acuminate to slightly rounded, base acute, cuneate or convex;
exudate channels usually conspicuous on both surfaces; galls on the lamina common, ca. 6 mm wide; secondary
veins conspicuous on both surfaces, 23–51(–56) pairs, distant 2–3.5 mm; petiole 12.5–30.3 mm long. Inflorescence:
staminate plants with clustered flowers; pistillate plants with clustered or solitary flowers; peduncle recurved; pedicels
1.3–1.8 cm long. Sepals 4, 7.1–12.9 × 8–12.9 mm, white-pinkish. Petals 4, 11.1–19.5 × 10.9–19.1 mm, red-pinkish
adaxially, white-pinkish abaxially, or, less commonly, completely white on both sides. Staminate flowers with more
than 50 fertile stamens, arranged in four series around the central staminodes, all united at the base, ca. 4 mm long;
central staminodes resiniferous, 5.7 mm long, densely clustered; pollen not mixed with oil or resin. Pistillate flowers
with numerous staminodes arranged in 3 series around the base of the ovary, densely clustered, with resin at the apex;
stigmas 4–6, 2–3 mm long, sessile or subsessile. Fruit ovoid to ellipsoid, 2.2–4.0 × 2.0–3.9 cm; sepals and stigmas
persistent. Seeds in two rows per locule, bright red, with yellow to orange aril.
Phenology:—Flowering November to July; fruiting December to July.
Distribution and habitat:—Brazil, French Guiana, Guyana, Suriname, Trinidad and Tobago and Venezuela. In
Brazil, it occurs in the north region (states of Amapá, Amazonas, Pará and Roraima), in the northeast region (states of
Alagoas, Bahia, Ceará, Pernambuco and Sergipe), in the central-west region (states of Distrito Federal, Goiás and Mato
Grosso), and in the southeast region (states of Espírito Santo, Minas Gerais and Rio de Janeiro). In VNP, it is found in
white-sand vegetation, temporarily flooded areas and on river margins.
Specimens examined:—BRAZIL. Roraima: Caracaraí, Viruá National Park, elevation, 56 m, 1º16’20’’N,
60º58’10’’W, 25 November 2009, fl., Cabral et al. 6 (INPA); Ibid., elevation 65 m, 1º28’08’’N, 60º58’18’’W, 1
December 2009, fl., Cabral et al. 67 (INPA); Ibid., elevation 79 m, 1º48’19’’N, 61º02’05’’W, 21 January 2010, fr.,
Cabral et al. 151 (INPA).
Additional specimens:—BRAZIL. Amazonas: Tarumã, 7 June 1984, fl., Miranda et al. 841 (INPA); Presidente
Figueiredo, Rebio Uatumã, 1º00’S, 59º00’W, 17 July 2006, fl., Ribeiro et al. 2698 (INPA). Bahia: Acajutiba, 11º43’S,
37º59’W, 22 August 1984, fl., Lima et al. 193 (INPA).
Taxonomic notes:—Clusia nemorosa is the most abundant species of Clusia in VNP, being easily found in
white-sand vegetation. The species can be recognized by the persistent calyx and staminodes and a deciduous corolla
Clusiaceae s.l. in the Viruá National Park Phytotaxa 329 (1) © 2017 Magnolia Press • 11
in the ripe fruits. Moreover, the fact that the stigmas close completely at the apex of the fruit with no conspicuous
space between them distinguishes C. nemorosa from most other species, which show a conspicuous space between
the stigmas. Clusia insignis also has stigmas closely connivent, but they are more numerous than in C. nemorosa and
yellow instead of green.
Clusia nemorosa may also have hermaphrodite flowers in gynodioecious populations in the northeast and southeast
of Brazil. However, we found only dioecious plants in VNP. The number of stigmas can vary from 4–12, sepals from
4–6 and petals from 4–9. Ripe fruits may be green instead of brown.
FIGURE 4. Clusia microstemon (A. Habit; B. Pistillate flower; C. Staminate flower), Clusia nemorosa (D. Habit with staminate flower;
E. Pistillate flower and fruit) and Clusia nitida (F. Staminate flower; G. Pistillate flower; H. Fruit).

3.6 Clusia nitida Bittrich & F.N. Cabral (2013: 36) (Fig. 4F–H)
Type:—BRAZIL. Roraima: Caracaraí, Viruá National Park, elevation 59 m, 1º24’50.7”N, 60º59’16.5”W, 15 October 2010, fl., Cabral &
Santos 298 (holotype: INPA, barcode 237546).
Trees, small trees or shrubs up to 6 m; sometimes with prop roots. Exudate white or cream. Lamina coriaceous, oblong-
elliptic, 4–8.5(–10.8) × (1.8–)2.1–4.9(–5.3) cm, apex rounded, base attenuate; exudate channels conspicuous on both
surfaces, distant 0.3–1.0 mm; secondary veins conspicuous on both surfaces, (9–)12–26(–29) pairs, distant 1.5–4.0
mm; petiole 4.0–16.0(–17.0) mm long. Inflorescence: staminate plants with clustered flowers; pistillate plants with
clustered flowers; pedicels ca. 2 mm long. Sepals 4–5(–6) × 4–8 mm, green-whitish. Petals 5–8, 6–13 × 6–12 mm, dark
bordeaux or blood red. Staminate flower with stamens numerous, ca. 300, densely compact, 2.3–3.0 mm long, forming
a flattened synandrium with sessile anthers on top, 3–6 mm wide, the lateral of the disk without anthers, covered with
resin-pollen mix during anthesis. Pistillate flower with 3–4 series of staminodes around the base of the ovary, 1.3–2.0
mm long, secreting resin at the apex, without sterile anthers; stigma 4–5, ca. 2.5 mm long. Fruit oval to hemispherical,
1.5–2.2 × 1.3–1.9 cm; sepals and stigma persistent. Seeds 5.0–9.0 × 3.5–5.0 mm, with orange aril.
Phenology:—Flowering September to February; fruiting November to July.
Distribution and habitat:—Endemic to Brazil, in the north region (states of Amazonas and Roraima). In VNP, it
mainly occurs in white-sand vegetation and river margins.
December 2009, fl., Cabral et al. 68 (INPA); Ibid., elevation 60 m, 1º24’58’’N, 61º12’27’’W, 22 January 2010, fl.,
Cabral et al. 168 (INPA); Ibid., elevation 58 m, 1º29’91’’N, 60º97’58’’W, 23 January 2010, fr., Cabral et al. 176
(INPA); Ibid., 31 May 2010, fr., Cabral et al. 215 (INPA); Ibid., 2 June 2010, fr., Cabral et al. 226 (INPA); Ibid., 21
July 2010, fr., Cabral et al. 264 (INPA); Ibid., 12 September 2010, fr., Cabral et al. 266 (INPA); Ibid., 15 October
2010, fl., Cabral et al. 298 (INPA); Ibid., 14 January 2011, fr., Cabral et al. 332 (INPA).
Taxonomic notes:—Clusia nitida is most similar to Clusia microstemon, which sometimes occurs sympatrically.
Clusia microstemon has a wide distribution in the Brazilian, Colombian and Venezuelan Amazon. The main difference
between the two species is in the male androecium, which in C. microstemon has anthers around the androecial disk,
whereas C. nitida anthers are on top of it. For more differences, see Bittrich et al. 2013.
3.7 Clusia renggerioides Planchon & Triana (1860: 350) (Fig. 5A–C)
Lectotype (designated here):—BRAZIL. October 1852-January 1853, fl., Spruce 2895 (lectotype: P, barcode 01901175, isolectotypes:
BM, barcodes 001191242, 001191243, C, barcode 10009458, E, barcode 00326696, G, barcodes 00355311, 00355312, GH, barcode
00026871, K, barcodes 000487423, 001229779, LD, barcode 1029125, MPU, barcode 014316, NY, barcodes 00084759, 00380931,
P, barcode 01901176).
Trees, shrubs or small trees up to 15 m. Exudate white, cream, yellow-orange, sometimes dark brown. Lamina
chartaceous to coriaceous, elliptical to oblong, (5–)5.6–12(–13) × (2–)2.1–4.4 cm, apex rounded to obtuse, base acute
and cuneate; exudate channels inconspicuous; secondary veins conspicuous on both surfaces, 23–60 pairs, distant ca.
1.5 mm; petiole 8–22 mm long. Inflorescence: staminate plants with clustered or solitary flowers; pistillate plants with
clustered flowers; pedicels 1.0–1.3 cm long. Sepals 4–8, 3.4–8 × 5.5–10 mm, green. Petals 5, (7.6–)9–13.3 × 9–12.5
mm, red-pinkish and white with red center adaxially, white abaxially. Staminate flowers with or without pistillode.
Staminate flowers with pistillode: fertile stamens 7–8, arranged in series around the pistillode, pistillode 4.6–7.3 mm
long, stigma 2.3–3.1 mm long. Staminate flowers without pistillode: fertile stamens 21–39, densely clustered, forming
a pentagonal and flat androecium, 4.5–5.4 mm wide; pollen mixed with resin. Pistillate flowers with 7–9 resiniferous
staminodes, 1.7–2.4 × 2.0–2.7 mm, with horizontal antherodes at the apex; stigma (4–)5, 2–3 mm long, surface with
acute papilla or short trichomes. Fruit ellipsoid to ovoid, 1.5–2(–3.1) × 1.4–1.9 cm; stigma persistent. Seeds in two
rows per locule, with red aril.
Phenology:—Flowering November to January; fruiting November to January and July.
Distribution and habitat:—Brazil, Colombia and Venezuela. In Brazil, it occurs in the north region (states of
Amazonas, Pará, Rondônia and Roraima), and in the central-west region (state of Mato Grosso). In VNP, in white-sand
vegetation and river margins.
December 2009, fl., Cabral et al. 74 (INPA); Ibid., elevation 69 m, 1º48’45’’N, 61º02’07”W, 21 January 2010, fl.,
Cabral et al. 150 (INPA); Ibid., 22 July 2010, fr., Cabral et al. 266 (INPA).
FIGURE 5. Clusia renggerioides (A. Habit; B. Pistillate flower; C. Staminate flower), D. Platonia insignis (D. Habit; E. Flower) and
Symphonia globulifera (F. Habit; G. Flower).
Additional specimens:—BRAZIL. Amazonas: Rio Bararati, PARES Sucunduri, 26 June 2006, fl., Zartman et al.
5771 (INPA).
Taxonomic notes:—Clusia renggerioides is distinguishable from the other species of Clusia in VNP for not
having exudate channels visible on the lamina. Moreover, its bracts are early deciduous, the calyx and the staminodes
are not persistent, the stigma has acute papilla and the endocarp of the valves becomes very hard, showing transversal
sulci, sometimes also visible on the pericarp of dried sub-mature fruits.

4 Garcinia Linnaeus (1753: 443).
Type: Garcinia mangostana Linnaeus (1753: 443).
= Rheedia Linnaeus (1753: 1193).
= Verticillaria Ruiz & Pavon (1794: 81).
= Chloromyron Persoon (1807: 73).
= Lamprophyllum Miers (1855: 338).
Trees or shrubs; exudate yellow or cream. Leaves opposite or whorled; exudate channels conspicuous; petiole
bases usually strongly excavated. Inflorescence in axillary fascicles sometimes appearing after leaves have fallen
(“cauliflorous”). Flowers bisexual and/or unisexual; sepals 2–4(–5); petals 2–4(–6), imbricate or decussate. Staminate
flowers with numerous stamens. Bisexual flowers with fewer stamens, pedicels thicker than in the staminate flowers.
Stamens free or united, connate forming a central mass. Ovary 3(–2–1)-locular, often absent from staminate flowers,
ovule 1 per locule, style short or lacking, stigma expanded. Fruit a berry, mostly 1-locular, epicarp coriaceous. Seeds
1–3, cylindrical.
Pantropical; with ca. 240 species (Kearns 1998a, Stevens 2001, onwards) and five species in Brazil (BFG 2015).
4.1 Garcinia albuquerquei (M.E.Berg) Bittrich, comb. nov.

Type:—BRAZIL. Amazonas: Rio Negro, Ilha Gavião, “near mouth of Rio Branco”, 5 July-August 1967, fr., Schultes 24535 (holotype:
INPA, barcode 20707). Fig. see Van den Berg (1979).
= Rheedia albuquerquei M.E. Berg (1975: 285).
Trees up to 5 m. Exudate cream, not abundant. Branches with scars from fallen inflorescences. Lamina strongly
coriaceous, oblong to elliptical or obovoid, (5.5–)6.5–12.5 × (3.0–)3.4–8.3 cm, apex emarginate, base emarginate or
rounded; base of lamina with a prominent adaxial margined pit, sessile; exudate channels not visible; midvein strongly
prominent on both surfaces, especially on the basal part; secondary veins prominent, conspicuous on both surfaces,
15–24 pairs, distant 2–5 mm. Inflorescences axillary. Flowers not seen. Fruit elliptical, rostrate, 2.5 × 1.7 cm, stigmas
persistent. Seeds cylindrical, 10.0 × 0.4 mm, surrounded by mesocarp pulp.
Phenology:—Flowering not known; fruiting July to November.
Distribution and habitat:—In Brazil, in the north region (states of Amazonas and Roraima). In VNP, it is found
in white-sand vegetation at the margin of a tributary of the Rio Branco. The species was considered endemic to
the Middle and Upper Rio Negro region, but our collection represents a new record of this species for the state of
Roraima.
60º59’13’’W, 30 September 2011, fr., Cabral et al. 430 (UEC); Ibid., 30 September 2011, fr., Cabral et al. 431
(UEC).
Taxonomic notes:—The Neotropical species of Garcinia were formerly included in Rheedia. Robson (1958)
suggested there was no basis for maintaining Rheedia separate from Garcinia. Stevens (2007) supported Robson’s
view and considered Rheedia as a synonym of Garcinia, and Sweeney (2008) confirmed that the recognition of
Rheedia would leave Garcinia paraphyletic. As a result, new combinations of Rheedia species were necessary. This
was followed by the work of Zappi (1993), who created the combination Garcinia gardneriana (Planchon & Triana)
Zappi for Rheedia gardneriana Planchon & Triana and Hammel (1989) who created new combinations for numerous
Central American species of Rheedia.
Garcinia albuquerquei is easily recognized by its angled branches, sessile lamina and the prominent margined
pit at the base of the leaf blade. There is no description of flowers as we did not find any fertile specimens in the field.
Flowers were also not available for the original species description by Van den Berg (1975) or the revision of Rheedia
(Van den Berg 1979).
5 Platonia Martius (1829: 168), nom. cons.

Type: Platonia insignis Maritus (1829: 169).
Trees; exudate yellow or cream. Perulae present, protecting the terminal buds, resulting later in groups of scars on
the branches. Leaves opposite, secondary veins conspicuous; exudate channels conspicuous. Flowers hermaphrodite,
terminal, solitary and showy; sepals 5, petals 5, contorted. Stamens numerous, in fascicles; anthers ± locellate. Ovary
with 5 locules, with several ovules per locule, style 5-radiate, stigma 5. Fruit a berry. Seeds 5.
Neotropical; with two species (Kearns 1998b) and one species in Brazil (BFG 2015).
5.1 Platonia insignis Martius (1829: 169) (Fig. 5D–E)
Lectotype (designated by Rijckevorsel 2002: 813):—BRAZIL. Pará: s.d., fl., Martius 2596 (M, barcode 0086228).
= Aristoclesia esculenta Stuntz (1912: 58).
= Moronobea esculenta Arruda, nom. inval.
= Platonia esculenta (Arruda) Oken (1841: 1431).
Trees up to 20 m. Exudate dark yellow. Lamina subcoriaceous, oboval to elliptical, 7–11.7 × 3.7–7.8 cm, apex slightly
acuminate, acute, obtuse or rounded, base acute; exudate channels conspicuous only on the abaxial surface, with dark
lines, between the secondary veins, distant 0.3–0.9 mm; secondary veins prominent, conspicuous on both surfaces,
50–76 pairs, distant 1–1.5 mm; petiole 12.5–26.8 mm long. Flowers pedicellate, 41.1–45.5 × 29.1–42.0 mm. Sepals
7.0–8.6 × 6.0–12.0 mm, yellow-cream, not united at the base. Petals 37.1–42.0 × 38.5–42.0 mm, pink to cream, apex
cream, contorted. Stamens in 5 fascicles, with 97–110 stamens each, ca. 40 mm long; anthers 6.0–15.3 mm long,
exuding oil. Ovary 14.2–14.4 × 8.8–10.3 mm; annular nectary around the ovary base, 3.7–7.0 mm long, stigma 1.8–2.0
mm long, with a pore at the apex. Fruit oblong to cylindrical, 7.3–11.5 × 6.2–8.4 cm.
Phenology:—Flowering August to November; fruiting December to March.
Distribution and habitat:—Brazil, Colombia, French Guiana, Guyana, Suriname and Venezuela. In Brazil, it
occurs in the north region (states of Pará and Roraima), and in the northeast region (state of Maranhão). In VNP, it is
found in white-sand vegetation and terra-firme forest.
60º59’13’’W, 4 December 2009, fr., Cabral et al. 110 (INPA); Ibid., 10 October 2010, fl., Cabral et al. 292 (INPA);
Ibid., 2 November 2010, fl., Cabral et al. 327 (INPA).
Additional specimens:—BRAZIL. Amazonas: Manaus, Colônia Santo Antônio, 14 August 1968, fl., Rodrigues
et al. 8520 (INPA); Ibid., Estrada Manaus-Itacoatiara, km 10, 18 August 1970, fl., Rodrigues et al. 8909 (INPA);
Barcelos, Parque Estadual Serra do Aracá, 6 March 1984, fr., Rodrigues et al. 10602 (INPA).
Taxonomic notes:—Platonia insignis is easily recognized when flowering. In addition to a colored and abundant
exudate, the flowers are large and showy and the stamens, grouped in fascicles, are very numerous (about 100 per
fascicle). The species could be confused with Moronobea Aublet (1775: 788), the flowers of which are very similar
and are also pollinated by parakeets. However, the number of stamens per fascicle is much lower in Moronobea
than in Platonia (3–4 stames/fascile in Moronobea, vs. ∞ stamens/fascicle in Platonia). The two genera could be
combined into one as they are morphologically very similar and belong to the same clade within the tribe Symphonieae
(Clusiaceae) (Ruhfel et al. 2011).

6 Symphonia L.f. (1781: 49).
Type: Symphonia globulifera Linnaeus filius (1781: 302).
Trees with stilt roots; exudate bright yellow. Perulae present, protecting the terminal buds, resulting in groups of scars
on the branches. Leaves opposite, secondary veins conspicuous; exudate channels conspicuous. Inflorescence terminal
or axillary or solitary flowers. Flowers hermaphrodite; sepals 5, petals 5, contorted, free but tube-like closed and
forming a chamber. Stamens arranged in fascicles of 3 stamens each, all united at the base, forming a staminal tube
and with 5 carnose lobes to which the long anthers are abaxially attached. Ovary with 5 locules, style terminating in
a head of 5 radiating branches, stigmas forming cylindrical lobes, each with an apical pore. Fruit a berry. Seeds 1–8,
with hairy-like testa.
Neotropical; with ca. 22 species (Kearns 1998c) and one species in Brazil (BFG 2015).
6.1 Symphonia globulifera Linnaeus filius (1781: 302) (Fig. 5F–G)

Type:—SURINAME. “Habitat ad Surinamum”, s.d., fl., Dalberg s.n. (holotype: LINN, barcode 853.1).
= Symphonia microphylla R.E. Schultes (1955: 20).
= Symphonia utilissima R.E. Schultes (1955: 22).
Trees up to 15 m. Exudate orange cream, abundant. Lamina subcoriaceous, oboval, 5.2–9.9 × 2.4–3.7 cm, apex
acuminate, base acute; exudate channels conspicuous on the abaxial surface, sometimes conspicuous on the adaxial
surface; secondary veins prominent, conspicuous on both surfaces, (25–)34–55 pairs, distant 1–2 mm; petiole 5.9–10.6
mm long. Inflorescence terminal or axillary, with solitary or clustered flowers. Flowers pedicellate. Sepals 3.7–4.5 ×
4.9–5.7 mm, green-pinkish, free at the base. Petals 10.8–12.6 × 10.0–11.1 mm, red-pinkish, forming a chamber that

surrounds most of the basal part of the stamens and gynoecium. Filaments fused into a staminal tube with 5 triangular
lobes at the apex, each with 3 anthers attached, ca. 5 mm long, thecae producing an oil involving the pollen; staminal
tube surrounding the gynoecium; stigma region star-shaped with 5 lobes. Ovary with 5 locules; nectary well-developed
surrounding the base of the staminal tube, 2.4–3.0mm long; ovules 4 per locule. Fruit 2–4 × 1.5–3.5 cm, with style
persistent. Seeds 1–3 × 0.5–2.5 cm.
Phenology:—Flowering May to February; fruiting May to June and September to January.
Distribution and habitat:—Widely distributed in tropical America and Africa. In Brazil, it occurs in the north
region (states of Acre, Amazonas, Pará, Roraima and Tocantins), in the northeast region (state of Alagoas, Bahia,
Maranhão and Sergipe), in the central-west region (state of Mato Grosso), and in the southeast region (state of Espírito
Santo and Rio de Janeiro). In VNP, it grows in white-sand vegetation.
Specimens examined:—BRAZIL. Roraima: Caracarai, Viruá National Park, elevation 68 m, 1º48’21’’N,
61º02’00’’W, 21 January 2010, fl., Cabral et al. 154 (INPA).
Additional specimens:—BRAZIL. Acre: Mâncio Lima, Rio Moa, 15 October 1989, fr., Ferreira et al. 10112
(INPA). Amazonas: Manaus, 28 July 1961, fl., Rodrigues et al. 2269 (INPA); Ibid., Reserva Experimental de Silvicultura
Tropical, Estrada Manaus-Itacoatiara, km 39, 26 July 1977, fl., Mello et al. 65 (INPA). Pará: Jacundá, Comunidade
Nossa Senhora de Fátima, 5 February 2000, fl., Miranda et al. 571 (INPA). Mato Grosso: Aripuanã, 22 June 1974, fl.,
Rodrigues 9406 (INPA). PANAMÁ. Province of Cocle: Vicinity of La Mesa, 14 June 1975, fr., Mori 6603 (INPA).
VENEZUELA. San Carlos de Río Negro, 1º56’N, 67º06’W, 13 December 1979, fl., Clark et al. 7331 (INPA).
Taxonomic notes:—Symphonia globulifera, the only species of Symphonia occurring in Brazil, is easy to
recognize. The filaments fuse to form a staminal tube with five carnose triangular lobes at the apex and the anthers are
attached abaxially to the lobes. The flowers have petals that firmly enclose the staminal tubes and the ovary. There are
four anthers per fascicle and the ovary has four ovules per locule. The combination of these characters is found only in
this species in the Neotropics (Abdul-Salim 2002).
7 Tovomita Aublet (1775: 956).

Type: Tovomita guianensis Aublet (1775: 957).
= Beauharnoisia Ruiz & Pavón (1808: 71).
= Marialva Vandelli (1788: 37).
Trees or shrubs, usually with stilt roots; exudate white, yellow or orangish. Branches with long first basal internode.
Leaves opposite, secondary veins conspicuous; petiole at base usually with an adaxial margined pit, sometimes very
conspicuous. Inflorescences terminal, dichasial or pleiochasial, pedicels thickened in fruit. Flowers unisexual; sepals
2–4, the outer 2 completely enclosing the bud, petals 4–8, decussed. Staminate flowers with numerous stamens,
filaments free, filiforms, dorsiventrally compressed, cylindrical or clavate. Pistillate flowers with staminodes; filaments
of stamens and staminodes secreting droplets of fragrant oil. Ovary with 4–6 locules, one ovule per locule, styles 4–6,
stigmas capitate. Fruit a carnose coriaceous capsule, dehiscence septifragal, with reflexed valves exposing a bright
white, orange or red-purple endocarp. Seeds 1 per locule, with a fleshy vascularized aril, orange or yellow.
Neotropical; with ca. 70 species (Marinho et al. 2016) and 40 species in Brazil (BFG 2015).
Tovomita is a genus relatively easy to recognize by the combination of vegetative and floral characters. However,
on specific level, identification is often difficult. For the Amazonian species, no revision has been done since the
19th century. The identification keys are not clear and, like Caraipa, the genus is in urgent need for a new and more
complete taxonomic review, phylogenetic analyses and investigation of morphological characters.
7.1 Tovomita spruceana Planchon & Triana (1860: 277) (Fig. 6A–B)
Type:—FRENCH GUIANA. s.d., fl., Spruce 2428 (syntypes: BR, barcode 0000005913534, G, barcode 00355515, GH, barcode 00067917,
K, barcode 000488533, NY, barcode 00578998, P, barcode 00093878) & 3554 (syntype: P, barcode 00093880).
Trees up to 5 m. Exudate light yellow. Lamina subcoriaceous, elliptical, 12.5–21.6 × 4.9–7.6 cm, apex slightly
acuminate, acute to obtuse, base acute; exudate channels conspicuous on the abaxial surface; secondary veins prominent,
conspicuous on both surfaces, 14–16 pairs, distant 6.3–15 mm; petiole 2.1–4.4 cm long. Inflorescence pleochasial,
flowers pedicellate, cream, fragrant. Floral buds oblong, apex rounded. Sepals 2, 8–10 × 8–10 mm, longitudinally
striated, green. Petals 4, 10–12 × 5–6 mm. Staminate flowers with stamens, ca. 5 mm long, cylindrical, shorter in the
center. Pistillate flowers with numerous staminodes, ca. 4 mm long. Ovary 8–10 × 6–8 mm, 5 locules, 1 ovule per
locule; stigma 5. Fruit 4–5 × 2–3 cm, pyriform, stigma persistent. Seed 1.7–2.2 × 0.8–0.9 cm (immature), surrounded
by a fleshy orange aril.
Phenology:—Flowering October; fruiting November to December.
Distribution and habitat:—Brazil, Colombia and Venezuela. In Brazil, it occurs in the north region (states of
Amazonas, Pará and Roraima). In VNP, it grows in white-sand vegetation and terra-firme forests.
December 2009, fr., Cabral et al. 91 (INPA).
Taxonomic notes:—It is possible to differentiate Tovomita spruceana from T. umbellata Bentham (1843: 367)
only by vegetative characters. Tovomita spruceana has shorter internodes and leaf lamina and a longer petiole than T.
umbellata. When in floral buds, Tovomita spruceana is distinguished by T. umbellata by its oblong floral buds with
rounded apex (vs. ovate floral buds with acute apex in T. umbellata).
7.2 Tovomita umbellata Bentham (1843: 367) (Fig. 6C–D)

Type:—GUYANA. s.d., fl., Schomburgk 991 (syntypes: G, barcode 00355790, K barcode 000488538, NY, barcode 00579000, S, barcode
03-2040, US, barcode 00114305).
= Tovomita cephalostigma Vesque (1892: 207).
= Tovomita nigrescens Planchon & Triana (1860: 276).
Trees up to 11 m. Exudate yellow to orangish. Lamina subcoriaceous, elliptical, 5.3–16.4 × (1.8–)2.1–7.5(–9.7)

cm, apex mucronate, acute to obtuse, base acute to obtuse; exudate channels conspicuous on the abaxial surface;
secondary veins prominent, conspicuous on both surfaces, 12–16 pairs, distant 4.2–11.4 mm; petiole 7.2–20.5 mm
long. Inflorescence pleiochasial, flowers pedicellate, cream, fragrant. Floral buds ovate, apex acute. Sepals (2–)4,
3.6–4.9 × 3.0–4.2 mm, green. Petals 4(–5), 3.8–5.1 × 3.0–4.7 mm, cream. Staminate flowers with stamens 33–51, 5–8
mm long, cylindrical, with different length. Pistillate flowers not seen. Fruit 2.8–3 × 1.3–1.5 m, pyriform to oblong,
stigma persistent. Seeds not seen.
Phenology:—Flowering May to October; fruiting April to July.
Distribution and habitat:—Brazil, Guyana, Peru and Venezuela. In Brazil, it occurs in the north region (states
of Amapá, Amazonas, Pará and Roraima), in the northeast region (state of Maranhão), and in the central-west region
(state of Mato Grosso). Our collection is a new record for the state of Roraima. In VNP, it grows in terra-firme forest.
61º00’31’’W, 28 November 2009, fl., Cabral et al. 45 (INPA); Ibid., 30 May 2010, fl., Cabral et al. 250 (INPA); Ibid.,
25 July 2010, fr., Cabral et al. 260 (INPA); Ibid., 11 September 2010, fl., Cabral et al. 262-A (INPA); Ibid., 20 October
2010, fl., Cabral et al. 311 (INPA); Ibid., 24 October 2010, fl., Cabral et al. 314 (INPA).
Additional specimens:—BRAZIL. Amazonas: Manaus, Reserva Florestal Ducke, Estrada Manaus-Itacoatiara,
km 26, 02º53’S, 59º58’W, 10 October 1995, fl., Ribeiro et al. 1731 (INPA); Ibid., 02º53’S, 59º58’W, 21 April 1997, fr.,
Souza et al. 358 (INPA).
Taxonomic notes:—See taxonomic notes under Tovomita spruceana for taxonomic differences between the two
species.
Hypericaceae Jussieu
Type: Hypericum Linnaeus (1753: 783).
Small trees to shrubs or annual to perennial herbs. Exudate yellow or orange or exudate absent. Leaves simple, opposite,
without stipules, margin entire, usually with glandular dots. Inflorescence terminal or axillary, cymose, or flowers
solitary. Flowers bisexual, actinomorphic, sepals 4–5, free, imbricate; petals 4–5, free, cochleate or contorted; stamens
15 to numerous, sometimes grouped in fascicles, staminodes occasionally with nectary function; ovary superior (2–)3–
5(6–8) carpels, placentation axillar or parietal, ovules many. Fruit capsule, drupe or berry. Seeds (5–) many, cotyledons
moderate in size.
Hypericaceae is composed of 7 genera and about 700 neo- and paleotropical species and it can be divided in
three tribes: Vismieae (pantropical), Cratoxyleae (paleotropical) and Hypericeae (pantropical, especially Northern
Hemisphere) (Stevens 2001, onwards, Ruhfel et al. 2011). In Brazil, the family is represented by 52 species (BFG
2015).

FIGURE 6. Tovomita spruceana (A. Habit; B. Mature fruit), Tovomita umbellata (C. Habit; D. Flower) and Vismia cayennensis (E. Habit;
F. Flower; G. Fruit).
8 Vismia Vandelli (1788: 51).

Type: Vismia cayennensis (Jacq.) Persoon (1807: 86).
Shrubs, small trees or trees; exudate orange or yellow. Leaves opposite, lamina rusty tomentose in most species,
especially on the abaxial surface, conspicuous dark glands; secondary veins conspicuous. Inflorescence terminal,
cymose, indument tomentose. Flowers hermaphrodite, sometimes heterostylous; sepals 5, petals 5, contorted or
cochleate. Stamens 15 to numerous grouped into five fascicles opposite the petals, usually with a staminodial nectary
opposite the sepals. Ovary with 5 locules, several ovules per locule, style 5, stigma expanded. Fruit a coriaceous berry.
Seeds numerous.
Neotropical; with ca. 65 species (Robson 1998) and 29 species in Brazil (BFG 2015).
8.1 Vismia cayennensis (Jacquin) Persoon (1807: 86) (Fig. 6E–G).
Type:—FRENCH GUIANA. Not indicated by Jacquin, but undoubtedly from Cayenne.
= Hypericum cayennense Jacquin (1760: 28).
Small trees up to 1.8 m. Exudate yellow. Lamina subcoriaceous, elliptical to oval, 5.9–9 × 3.4–5.1 cm, apex acuminate,
base obtuse and rounded to acute, abaxial surface with several conspicuous dark glands; secondary veins prominent,
conspicuous on both surfaces, 12–17 pairs, distant 3.8–5 mm; petiole 6.7–10.8 mm long. Inflorescence with 8–14
flowers. Flowers pedicellate. Sepals 6.7–9.2 × 3.4–4.4 mm, green and brown. Petals 8–9 × 3–4 mm wide, yellowish-
green, covered with cream colored trichomes adaxially, with linear orange-brown glands abaxially. Fascicles of
stamens deciduous with petals, stamens numerous, filaments with white trichomes. Fruit oblong to cylindrical, 0.9–1.2
× 0.5–1.3 cm, sepals and style persistent. Seeds ca. 2 mm long, shiny.
Phenology:—Flowering and fruiting in July and December.
Distribution and habitat:—Brazil, Colombia, French Guiana, Guyana, Suriname and Venezuela. In Brazil, it
occurs in the north region (states of Acre, Amapá, Amazonas, Pará and Roraima), and in northeast region (state of
Maranhão). In VNP, it grows in white-sand vegetation.
60º58’10’’W, 1 December 2009, fl., fr., Cabral et al. 77 (INPA).
Additional specimens:—BRAZIL. Amazonas: Manaus, Reserva Florestal Ducke, Estrada Manaus-Itacoatiara,
km 26, 02º53’S, 59º58’W, 12 July 1994, fl., Hopkins et al. 1465 (INPA); Ibid., 02º53’S, 59º58’W, 19 July 1994, fr.,
Ribeiro et al. 1341 (INPA).
Taxonomic notes:—Vismia cayennensis is an easily recognizable species as it is the only Vismia species in the
VNP with both surfaces of the leaf lamina completely glabrous.
8.2 Vismia japurensis Reichardt (1878: 209) (Fig. 7A–B)

Type:—BRAZIL. Amazonas: Prov. do Alto Amazonas in silva ad ripas fluminis Japurá, s.d., fl., C.F.P. Martius s.n. (syntypes: M, barcodes
0112247, 0112245, 0112246, 0112244, 0112243).
Shrubs up to 2 m. Exudate orange. Lamina coriaceous, oval, 12.3–15.4 × 5.7–6.3 cm, apex acute and acuminate, base
rounded, with dark glands conspicuous abaxially; abaxial surface rusty tomentose, with rusty-brown stellate trichomes,
adaxial surface glabrous; secondary veins prominent, conspicuous on both surfaces, 11–13 pairs, distant 7.0–20.8 mm;
petiole 10.3–13.3 mm long. Inflorescences with ca. 15 flowers. Sepals 7.7–10.0 × 3.9–5.4 mm, rusty tomentose. Petals
11.9–15.0 × 4.8–5.0 mm, yellowish cream, with linear orange-brown glands abaxially. Fascicles of stamens deciduous
with petals, stamens numerous, ca. 50 in each fascicle. Fruit ellipsoid to cylindrical, 1–1.7 × 0.8–0.9, sepals rigid,
adpressed to the fruit, style persistent. Seeds not seen.
Phenology:—Flowering May and September–October; fruiting July and October to February.
Distribution and habitat:—Brazil, Colombia, Ecuador, Guyana, Suriname and Venezuela. In Brazil, it occurs in
the north region (states of Pará, Amazonas and Roraima), and in central-west region (state of Mato Grosso). This is a
new record for the state of Roraima. In VNP, it occurs in white-sand vegetation.
Specimens examined:—BRAZIL. Roraima: Caracaraí, Viruá National Park, 16 September 2010, fl., Cabral et
al. 281 (INPA).
Additional specimens:—BRAZIL. Amazonas: Rio Xié, 9 May 1973, fl., Silva et al. 1420 (INPA); Manaus, 13
February 1962, fr., Rodrigues et al. 4205 (INPA). Pará: Santarém, Alter do Chão, 12 December 1978, fr., Maciel et
al. 291 (INPA). GUIANA. Elevation 50 m, 5º58’N, 5º30’W, 1 January 1987, fr., Pipoly 9593 (INPA); Ibid., 02º53’S,
59º58’W, 19 July 1994, fr., Ribeiro et al. 1341 (INPA); Ibid., 02º53’S, 59º58’W, 24 October 1994, fr., Sothers et al. 236
(INPA).
Taxonomic notes:—Vismia japurensis is characterized by the broadly rounded leaf bases, the petals deciduous in
fruit and by its large rigid sepals.

FIGURE 7. Vismia japurensis (A. Habit; B. Flower), Vismia laxiflora (C. Habit; D. Flower; E. Fruit) and Vismia macrophylla (F. Habit;
G. Flower; photos from specimens collected in Serra Tepequem, Roraima, Brazil, Amaral & Bittrich 2015/01).
8.3 Vismia laxiflora Reichardt (1878: 203) (Fig. 7C–E)

Type:—VENEZUELA. 1841, fl., Schomburgk 835 [an error for 837] (syntypes: BM barcode 000624911, F, barcode 0054568F, G,
barcode 00355559).
= Vismia angustifolia Rusby (1920: 59).
= Vismia falcata Rusby (1920: 59).
Trees or small trees up to 6 m. Exudate abundant, orange. Lamina subcoriaceous, elliptic to oval, 5.7–15.8 × 2.5–6.6
cm, apex acute to slightly acuminate, base obtuse or rounded, with dark glands conspicuous abaxially; abaxial surface
rusty tomentose, with stellate trichomes, adaxial surface glabrous or pubescent, with trichomes visible on the midvein;
secondary veins prominent, conspicuous on both surfaces, 8–15 pairs, distant 6.2–14 mm; petiole 5.4–14.5 mm long.
Inflorescence with 12–30 flowers. Sepals 6.3–8.4 × 1.8–4.5 mm, rusty tomentose. Petals 7.8–9.7 × 4.4–5.6 mm, cream
yellowish, with linear orange-brown glands. Fascicles of stamens ca. 6 mm long, stamens ca. 50 in each fascicle. Fruit
ellipsoid to cylindrical, 0.8–0.9 × 0.8–0.9 cm; sepals reflex in fruit, petals and style persistent.
Phenology:—Flowering September to December; fruiting September and January.
Distribution and habitat:—Brazil, Colombia, Guyana, Trinidad and Tobago and Venezuela. In Brazil, it occurs in
the north region (states of Acre, Amapá, Amazonas, Pará and Roraima), and in the northeast region (state of Maranhão).
In VNP, it occurs in white-sand vegetation and flooded areas.
60º58’10’’W, 25 November 2009, fl., Cabral et al. 12 (INPA); Ibid., elevation 59 m, 1º25’33’’N, 60º59’15’’W, 3
December 2009, fl., fr., Cabral et al. 96 (INPA); Ibid., elevation 62 m, 1º45’90’’N, 60º97’29’’W, 29 January 2010, fr.,
Cabral et al. 202 (INPA); Ibid., 15 September 2010, fl., fr., Cabral et al. 278 (INPA).
Taxonomic notes:—Vismia laxiflora is characterized by having persistent petals in fruit. Furthermore, the sepals
are fleshy and not rigid coriaceous as in V. japurensis.
8.4 Vismia macrophylla Kunth (1821: 184) (Fig. 7F–G, Fig. 8A)
Type:—VENEZUELA. Amazonas: banks of the Río Casiquiare, s.d., fl., Humboldt & Bonpland s.n. (holotype: P, barcode 00679837).
= Vismia angusta Miquel (1844: 27).
Trees or rarely shrubs up to 2–25 m. Exudate orange to yellowish; branches densely ferrugineous-tomentose, with
stellate to dendroid trichomes. Lamina coriaceous, oblong to lanceolate or ovate, 10–42 × 6–18 cm, apex shortly
acuminate, base cordate to rounded; secondary veins prominent, 12–31, distant 7–15 mm; petiole 10–25 mm long,
ferrugineous-tomentose. Inflorescences with 5–6(–7) flowers. Flowers heterostylous. Sepals 5–6 mm long, whitish,
sparsely ferrugineous pubescent with stellate-dendroid trichomes and densely covered with tiny whitish papillae.
Petals 10–13 × 2–2.5 mm, distally reflexed and glandular-punctate. Fascicles of stamens ca. 3 mm long, stamens ca.
25 per fascicle. Fruits ovoid-subglobose to globose, 1.2–1.4 × 1.1–1.3 cm, sepals reflexed, stamen fascicles persistent.
Seeds 2.5–3.5 mm long.
Phenology:—Flowering November to June; fruiting June and November.
Distribution and habitat:—Widespread in northern South America, Brazil, Costa Rica, Mexico and Peru. In
Brazil, it occurs in the north region (states of Acre, Amazonas, Pará, Rondônia and Roraima), and in the central-west
region (state of Mato Grosso). In VNP, it grows in white-sand vegetation.
Specimens examined:—BRAZIL. Roraima: Caracaraí, Viruá National Park, Margem esquerda da Estrada
Perdida, a cerca de 500 m da bifurcação para a sede do Parque, em direção à primeira bueira (sul), 01°28’33.4”N;
60°58’5.4”W, 12 January 2015, fl., Amaral & Bittrich 2015/17 (INPA, UEC).
Additional specimens:—BRAZIL. Amazonas: Balbina, Ilha na Represa Uatumã, 2 km da base WABA, 1º00’S,
59º00’W, 6 August 2006, fr., Zartman et al. 5912 (UEC). Bahia: Camamu, povoado de Barcelos do Sul, 18 July
2005, fl., Miranda & Silva 5166 (UEC). Mato Grosso: Aripuanã, margem da MT-420 a 30 km da cidade em direção a
Juruena, 10º15’43”S, 59º16’57”W, 4 July 1997, fr., Assis et al. 486 (UEC); Chapada dos Guimarães, Parque Nacional
da Chapada dos Guimarães, 1 March 1997, fr., Nave et al. 1294 (UEC). Rondônia: Porto Velho, UHE de Samuel, Rio
Jamari, 11 February 1989, fl., Maciel & Rosário 1587 (UEC).
Taxonomic notes:—Vismia macrophylla is very easy to distinguish from other Vismia species found in VNP, by
its very large petiole and lamina and cordate to rounded leaf base. Furthermore, the stellate to dendroid trichomes do
not cover the dark glandular dots on the abaxial side of the lamina. As a result of the distally reflexed petals, the corolla
has a star-like form. At VNP, V. macrophylla is the only species that exhibits this feature.
8.5 Vismia sp. (Fig. 8B)
Shrubs up to 1.8 m. Exudate orange. Lamina subcoriaceous, elliptical, 7.7–12.8 × 2.7–4 cm, apex acute to acuminate,
base acute to cuneate, with chocolate-brown glands abaxially; secondary veins prominent, conspicuous on both
surfaces, 8–14 pairs, distant 6.1–9.7 mm; petiole 7.8–10.5 mm long. Flowers not seen. Sepals 7.8–9.0 × 3.8–4.4 mm,
chocolate-brown. Fascicles of stamens 8 mm. Fruit 0.8–0.9 × 0.6–0.7 cm, ellipsoid, fascicles of stamens sepals and
style persistent in fruit, sepals reflexed.
Phenology:—Flowering unknown; fruiting November.

FIGURE 8. Vismia macrophylla (A. Flower, photo from specimen collected in Serra Tepequem, Roraima, Brazil, Amaral & Bittrich
2015/01) and Vismia sp. (B. Fruit).
Distribution and habitat:—Currently, this plant has only been found at VNP. It occurs in the transition area
between terra-firme and white-sand vegetation.
61º01’00’’W, 30 November 2009, fr., Cabral et al. 65 (INPA).
Taxonomic notes:—The specimen collected at VNP has obvious morphological differences when compared with
the isotype of most similar species such as V. steyermarkii Robson (1990: 411) (i.e. Maguire & Steyermark 46799,
NY). This collection is likely to represent a new species as it differs from other species with fascicles of stamens
persistent in fruit. For instance, Vismia macrophylla also has persistent fascicles, but it is clearly different in vegetative
characters, such as larger leaf size (10–42 × 6–18 cm, vs. 7.7–12.8 × 2.7–4 cm in Vismia sp.), larger petiole (10–25 mm
long, vs. 7.8–10.5 mm long in Vismia sp.), cordate to rounded leaf base (vs. acute to cuneate in Vismia sp.), and the
shape of its petals (see comments on V. macrophylla above). We could only find one voucher specimen of this plant in
fruit, which is not sufficient to produce a complete diagnosis and describe it as a new species. Thus, description of this
entity will be postponed until flowering material is available for us to study.
Discussion
Floristic inventories are the starting point to assess the biodiversity and geographical distribution of taxa. Similar to
our study, previous taxonomic studies at the Viruá National Park (Costa et al. 2016a, 2016b, Dávila 2011, Melo et
al. 2014, Mota et al. 2015, Pessoa et al. 2016) have reported new species and new species records for the state of
Roraima, showing a high diversity of taxa in the area. More taxonomic studies, focusing on other plant families, are
necessary in the VNP and surrounding areas to gather basic and reliable data for the Amazon region. These studies will
provide conservation biologists with the necessary knowledge to identify priority regions for conservation and protect
the future of endangered taxa. Such studies, however, must involve professionals with an advanced knowledge of the
native flora and the collection of voucher specimens. For instance, we found out that ICMBio (2014) lists 35 species
of Clusiaceae s.l. for the park. Seventeen of these 35 species were not found in our study. ICMBio (2014) lists two
different species of Caraipa, five different species of Clusia, two of Garcinia, six of Tovomita, and two of Vismia. It is
likely that this checklist is inaccurate as no specialist in none of the three plant families we studied here were involved
in the taxonomic inventories. Furthermore, we were unable to confirm their claims as we could not locate any voucher
specimens supporting the presence of these 17 species in the VNP.
Acknowledgements
We would like to thank the financial support from PNADB - Programa Nacional de Apoio e Desenvolvimento da
Botânica/CAPES, PROCAD Amazônia - Programa de Cooperação Acadêmica/CAPES, INPA, CAPES for a current
postdoc fellowship for F.N. Cabral, and the herbaria and curators at INPA, NY and UEC. We thank Lucas Marinho
(UEFS, Feira de Santa, Brazil) for the identification and correction of the descriptions of the Tovomita species and
Bruno Amorim for reviewing and discussing the manuscript. We also thank all the people who work at Parque Nacional
do Viruá, specially Antônio Lisboa and Beatriz Lisboa for being so helpful and kind during our field work. Special
thanks to Valmiranda Souza Santos, our field assistant, and to Nállarett Marina Cardozo Dávila, a great friend and
botanist, without them this work would not have been possible. We thank the anonymous reviewers and the editor Dr.
Carlos Lehnebach for the great contributions and suggestions to improve our manuscript.
References
Abdul-Salim, K. (2002) Systematics and Biology of Symphonia L. f. (Clusiaceae). PhD Thesis. Harvard University, Cambridge, Mass,
210 pp.
APG IV. (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV.
Botanical Journal of the Linnean Society 181:1–20.
https://doi.org/10.1111/boj.12385
Aublet, J.B.C.F. (1775) XIII. Polyandria, Monogynia. Histoire des Plantes de la Guiane Françoise 1: 532–578.
Aublet, J.B.C.F. (1775) IV. Polygamia, Tetragynia. Histoire des Plantes de la Guiane Françoise 2: 956–958.
Aublet, J.B.C.F. (1775) IV. Polyadelphia, Polyandria. Histoire des Plantes de la Guiane Françoise 2: 780–792.
Bentham, G. (1843) Flora of South America. Clusiaceae. London Journal of Botany 2: 367–370.
BFG. (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085–1113.
https://doi.org/10.1590/2175-7860201566411
Bittrich, V. & Amaral, M.C.E. (1996) Pollination biology of Symphonia globulifera (Clusiaceae). Plant Systematics and Evolution 200:
101–110.
https://doi.org/10.1007/BF00984751
Bittrich, V. (2008) Chave das espécies de Clusia L. (Guttiferae) do Brasil. Available from: http://www2.ib.unicamp.br/profs/volker/
chaves/Clusia/Chave_Clusias.htm (accessed: January 2017)
Bittrich, V., Cabral, F.N. & Hopkins, M.J.G. (2013) Clusia nitida, a new species of Clusia (Clusiaceae) from the Brazilian Amazon.
Phytotaxa 100: 36–40.
Cambessèdes, J. (1825) Calophyllum. Flora Brasiliae Meridionalis 1: 320–321.
Cabral, F.N., Bittrich, V. & Amaral, M.C.E. (2016a) Two new species and one new record of Caraipa (Calophyllaceae) from Colombia.
Systematic Botany 41: 348–353.
https://doi.org/10.1600/036364416X691911
Cabral, F.N., Bittrich, V. & Amaral, M.C.E. (2016b) Four new species of Caraipa (Calophyllaceae) from the Amazon basin and the Guiana
Shield. Phytotaxa 286: 245–255.
Carvalho, J.E.U., Nazaré, R.F.R. & Nascimento, W.M.O. (2003) Características físicas e físico-químicas de um tipo de Bacuri (Platonia
insignis Mart.) com rendimento industrial superior. Revista Brasileira de Fruticultura 25: 326–328.
https://doi.org/10.1590/S0100-29452003000200036
Casaretto, G. (1843) Chrysoxylon. In: Novarum Stirpium Brasiliensium Decades, Decas VII. pp. 59–64.
Castillo, A. & Gil, C. (1991) Ernstia, segunda etapa (2nd series), vol. 1. pp. 41–42.
Costa, S.M., Barbosa, T.D.M., Bittrich, V. & Amaral, M.C.E. (2016a) Floristic survey of herbaceous and subshrubby aquatic and palustrine
angiosperms of Viruá National Park, Roraima, Brazil. Phytokeys 58: 21–48.
https://doi.org/10.3897/phytokeys.58.5178
Costa, S.M., Bittrich, V. & Amaral, M.C.E. (2016b) Lentibulariaceae from the Viruá National Park in the northern Amazon, Roraima,
Brazil. Phytotaxa 258: 1–25.
Cronquist, A. (1981) An integrated system of classiﬁcation of ﬂowering plants. Columbia University Press, New York, New York, USA,

1262 pp.
Dávila, N. (2011) Rubiaceae das campinaranas do Parque Nacional do Viruá, Roraima, Brasil. Master Thesis, Instituto de Pesquisas da
Amazônia, Manaus, Amazonas, 105 pp.
Díaz, D.M.V. (2013) Multivariate analysis of morphological and anatomical characters of Calophyllum (Calophyllaceae) in South America.
Botanical Journal of the Linnean Society 171: 587–626.
https://doi.org/10.1111/boj.12012
Endress, P. (2010) Disentangling confusions in inflorescence morphology: Patterns and diversity of reproductive shoot ramification in
angiosperms. Journal of Systematics and Evolution 48: 225–239.
https://doi.org/10.1111/j.1759-6831.2010.00087.x
Engler, H.G.A. (1888) Guttiferae: Clusia. Flora brasiliensis 12: 400–434.
Hammel, B. (1989) New combinations and taxonomies in Clusiaceae. Annals of the Missouri Botanical Garden 76: 927–929.
https://doi.org/10.2307/2399657
Harris, J.G. & Harris, M.W. (2007) Plant identification terminology: an illustrated glossary. Second edition, Spring Lake Publishing, 216
pp.
Hewson, H. (1990) Plant Indumentum – A handbook of terminology. Australian Flora and Fauna series 9: 1–25.
Hickey, M. & King, C. (2000) The Cambridge illustrated glossary of botanical terms. Second edition. Cambridge University Press, 222
pp.
ICMBio. (2014) Plano de Manejo do Parque Nacional do Viruá. Boa Vista, ICMBio, 626 pp.
Jacquin, N.J. (1760) Polyadelphia. Enumeratio Systematica Plantarum 1760: 28.
Kearns, D.M. (1998a) Garcinia. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. Flora of the Venezuelan Guayana. Volume 4. 1ª ed. St.
Louis, Missouri, EUA, pp. 295–299.
Kearns, D.M. (1998b) Platonia. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. Flora of the Venezuelan Guayana. Volume 4. 1ª ed. St.
Louis, Missouri, EUA, pp. 310–311.
Kearns, D.M. (1998c) Symphonia. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. Flora of the Venezuelan Guayana. Volume 4. 1ª ed. St.
Louis, Missouri, pp. 311–312.
Kubitzki, K. (1978) Caraipa and Mahurea (Bonnetiaceae). In: The botany of the Guyana Highlands. Part X, Memoirs of the New York
Botanical Garden, New York, pp. 82–131.
Kubitzki, K. & Holst, B.K. (1998) Caraipa. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. Flora of the Venezuelan Guayana. Volume 4.
1ª ed. St. Louis, Missouri, pp. 252–258.
Kunth, K.S. (1821) Hypericaceae. Nova Genera et Species Plantarum 5: 181–197.
Linnaeus, C. (1753) Species Plantarum, vols. 1–2. L. Salvius, Holmia [Stockholm], 1200 pp.
https://doi.org/10.5962/bhl.title.669
Linnaeus filius, C. (1781) Supplementum plantarum. Impensis Orphanotrophei, Brunsvigae [Braunschweig], 467 pp.
Linnaeus, C. (1782) Palmae. Supplementum Plantarum 1782: 454–456.
Mabberley, D.J. (2008) Mabberley’s Plant-Book: A portable dictionary of plants, their classification and uses. Third edition, Cambridge
University Press, 1021 pp.
Maguire, B. (1951) Guttiferae. Botanical Museum Leaflets 15: 55–69.
Marinho, L.C., Fiaschi, P., Santos, F.A.R. & Amorim, A.M. (2016) Three new species of Tovomita (Clusiaceae) from the Amazon River
basin and first record of papillae for Tovomita. Plant Systematics and Evolution 302: 1121–1134.
https://doi.org/10.1007/s00606-016-1321-7
Martius, C.F.P. (1829) Nova Genera et Species Plantarum 3: 164–170.
Melo, A., Guimarães, E.F. & Alves, M. (2014) Piperaceae do Parque Nacional do Viruá, Caracaraí, Roraima, Brasil. Rodriguésia 65:
455–470.
https://doi.org/10.1590/S2175-78602014000200010
Melo, A., Guimarães, E.F. & Alves, M. (2016) Synopsis of the genus Peperomia Ruiz & Pav. (Piperaceae) in Roraima State, Brazil.
Hoehnea 43: 119–134.
https://doi.org/10.1590/2236-8906-75/2015
Meyer, G.F.W. (1818) Primitiae Florae Essequeboensis, pp. 203–204.
Miers, J. (1855) Transactions of the Linnean Society of London 21: 338.
https://doi.org/10.1111/j.1096-3642.1852.tb00461.x
Miquel, F.A.W. (1844) Vismia. Linnaea 18: 27–28.
Mota, N.F.O., Campbell, L.M., Viana, P.L. & Wanderley, M.G.L. (2015) Xyridaceae of Viruá National Park, Roraima state, Brazil.
Rodriguésia 66: 523–553.
https://doi.org/10.1590/2175-7860201566219
Oken, L. (1841) Platonia. Allgemeine Naturgeschichte 3: 1429–1431.
Persoon, C.H. (1807) Synopsis Plantarum, vol. 2. Parisiis Lutetiorum, pp. 1–656.
Pessoa, E., Barros, F. & Alves, M. (2015) Orchidaceae from Viruá National Park, Roraima, Brazilian Amazon. Phytotaxa 192: 61-96.
Pipoly, J.J., Kearns, D.M. & Berry, P.E. (1998) Clusia. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. Flora of the Venezuelan Guayana.
Volume 4. 1ª ed. St. Louis, Missouri, EUA, pp. 260–294.
Pipoly, J.J. & Gustafsson, M.H.G. (2003) Clusiaceae (mangosteen family). In: Mori, S.A., Cremers, G., Gracie, C., de Granville, J.-J,
Heald, S.V., Hoff, M. & Mitchell, J.D. Guide to the Vascular Plants of Central French Guiana: Part 2. Dicotyledons. Vol. 76 (2).
New York Botanical Garden Press, New York, pp. 212–224.
Planchon, J.E. & Triana, J.J. (1860) Mémoire sur la famille des guttifères. Annales des Sciences Naturelles; Botanique, série 4 13:
306–361.
267–285.
247–295.
Reichardt, H.W. (1878) Hypericaceae. In: Martius, C.F.P. & Eichler, A.W. (Eds.) Flora brasiliensis. 12(1), fasc. 81. Frid. Fleischer,
Lipsiae, pp. 181–212.
Robson, N. (1958) Guttiferae. In: New and little known species from the Flora Zambesiaca area. Boletim da Sociedade Broteriana 32:
170–173.
Robson, N.K.B. (1990) Two new species and a new combination in Vismia (Guttiferae-Hypericoideae). Annals of the Missouri Botanical
Garden 77: 410–411.
https://doi.org/10.2307/2399556
Robson, N.K.B. (1998) Vismia. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. Flora of the Venezuelan Guayana. Volume 4. 1ª ed. St.
Louis, Missouri, pp. 322–329.
Ruhfel, B.R., Bittrich, V., Bove, C.P., Gustafsson, M.H.G., Philbrick, C.T., Rutishauser, R., Xi, Z. & Davis, C.C. (2011) Phylogeny of the
Clusioid clade (Malpighiales): Evidence from the plastid and mitochondrial genomes. American Journal of Botany 98: 1–20.
https://doi.org/10.3732/ajb.1000354
Ruhfel, B.R., Stevens, P.F. & Davis, C.C. (2013) Combined morphological and molecular phylogeny of the Clusioid Clade (Malpighiales)
and the placement of the ancient rosid macrofossil Paleoclusia. International Journal of Plant Sciences 174: 910–936.
https://doi.org/10.1086/670668
Rusby, H.H. (1920) Descriptions of Three Hundred New Species of South American plants 59: 1–170.
Rusby, H.H. (1927) Descriptions of new genera and species of plants collected on the Mulford biological exploration of the Amazon valley.
Memoirs of the New York Botanical Garden 7: 303–304.
Ruiz, H.L. & Pavón, J.A. (1794) Florae Peruvianae. Chilensis Prodromus 1794: 80–81.
Ruiz, H.L. & Pavón, J.A. (1808) Beauharnoisia. Annales Du Muséum National d’Histoire Naturelle 11: 71–76.
Schaefer, C.E.G.R., Mendonça, B.A.F. & Fernandes Filho, E.I. (2009) Geoambientes e Paisagens do Parque Nacional do Viruá - RR:
esboço de integração da geomorfologia, climatologia, solos, hidrologia e ecologia (Zoneamento Preliminar). Relatório para
ICMBio, 59 pp.
Schlechtendal, D.F.L. (1833) Guttiferae. Linnaea 8: 180–186.
Schultes, R.E. (1955) Pitch-Yelding trees of the Colombian Amazonia. Botanical Museum Leafletls 17: 20–23.
Smith, A.C. (1933) Calophyllum. Bulletin of the Torrey Botanical Club 60: 379–380.
Stevens, P.F. (2001 onwards) Angiosperm Phylogeny Website. Version 12, July 2012 [and more or less continuously updated since].
Available from: http://www.mobot.org/MOBOT/research/APweb/ (accessed 6 June 2017)
Stevens, P.F. (2007) Clusiaceae-Guttiferae. In: Kubitzki, K. (Ed.) The families and genera of vascular plants. Flowering plants. Eudicots:
Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales,
Zygophyllales, Clusiaceae alliance, Passifloraceae alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae. Springer-Verlag,
Berlin, pp. 48–66.
https://doi.org/10.1007/978-3-540-32219-1_10
Stuntz, S.C. (1912) Seeds and plants imported. U.S. Department of Agriculture Bureau of Plant Industry Bulletin 248: 58–59.
Sweeney, P.W. (2008) Phylogeny and floral diversity in the genus Garcinia (Clusiaceae) and relatives. International Journal of Plant
Sciences 169: 1288–1303.
https://doi.org/10.1086/591990

Thiers, B. (2017) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual
Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 6 June 2017)
Vandelli, D. (1788) Florae Lusitanicae et Brasiliensis Specimen 1788: 36–51.
Van den Berg, M.E. (1974) Contribuição ao estudo do gênero Vismia Vandelli (Guttiferae). Acta Amazonica 4: 15–18.
https://doi.org/10.1590/1809-43921974042015
Van den Berg, M.E. (1975) Nota prévia sobre uma Guttiferae nova da Amazônia. Acta Amazonica 5: 285.
https://doi.org/10.1590/1809-43921975053285
Van den Berg, M.E. (1979) Revisão das espécies brasileiras do gênero Rheedia L. (Guttiferae). Acta Amazonica 9: 43–74.
https://doi.org/10.1590/1809-43921979091043
Veloso, H.P.; Rangel Filho, A.L.R. & Lima, J.C.A. (1991) Classificação da vegetação brasileira, adaptada a um sistema universal. IBGE.
Rio de Janeiro, 124 pp.
Vesque, J.J. (1892) Tovomita. Epharmosis 3: 207–208.
Wurdack, K.J. & Davis, C.C. (2009) Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in the angiosperm
tree of life. American Journal of Botany 96: 1551–1570.
https://doi.org/10.3732/ajb.0800207
Xi, Z., Ruhfel, B.R., Schaefer, H., Amorim, A.H., Sugumaran, M., Wurdack, K.J., Endress, P.K., Matthews, M.L., Stevens, P.F., Mathews,
S. & Davis, C.C. (2012) Phylogenomics and a posteriori data partitioning resolve the Cretaceous angiosperm radiation Malpighiales.
Proceedings of the National Academy of Sciences of the United States 109: 17519–17524.
https://doi.org/10.1073/pnas.1205818109
Zappi, D. (1993) A New Combination in Garcinia (Guttiferae). Kew Bulletin 48: 410.
https://doi.org/10.2307/4117953
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Clusiaceae s.l. (Calophyllaceae, Clusiaceae s.s. and Hypericaceae) in the Viruá

Article in Phytotaxa · November 2017

Fernanda Cabral Volker Bittrich

Families and genera of vascular plants View project

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Clusiaceae s.l. (Calophyllaceae, Clusiaceae s.s. and Hypericaceae) in the Viruá

Keywords: Amazon, biodiversity, Guttiferae, taxonomy

Palavras-chave: Amazônia, biodiversidade, Guttiferae, taxonomia

Accepted by Carlos Lehnebach: 19 Oct. 2017; published: 24 Nov. 2017 

Material and methods

Taxonomic and morphological descriptions

 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

Key to the species of Clusiaceae s.l. in the VNP

 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

1.1 Calophyllum brasiliense Cambessèdes (1825: 320) (Fig. 2A–C)

2 Caraipa Aublet (1775: 561).

 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

2.2 Caraipa savannarum Kubitzki (1978: 104) (Fig. 2G–H)

3 Clusia Linnaeus (1753: 509).

 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

3.2 Clusia insignis Martius (1829: 164) (Fig. 3C–E)

3.3 Clusia lopezii Maguire (1951: 61) (Fig. 3F–H)

Shrubs, small trees up to 3 m or hemiepiphytes. Exudate yellowish to yellowish-brown. Lamina subcoriaceous to

10 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

3.5 Clusia nemorosa Meyer (1818: 203) (Fig. 4D–E)

12 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

14 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

4.1 Garcinia albuquerquei (M.E.Berg) Bittrich, comb. nov.

5 Platonia Martius (1829: 168), nom. cons.

6.1 Symphonia globulifera Linnaeus filius (1781: 302) (Fig. 5F–G)

16 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

7 Tovomita Aublet (1775: 956).

7.2 Tovomita umbellata Bentham (1843: 367) (Fig. 6C–D)

Trees up to 11 m. Exudate yellow to orangish. Lamina subcoriaceous, elliptical, 5.3–16.4 × (1.8–)2.1–7.5(–9.7)

18 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

8 Vismia Vandelli (1788: 51).

8.2 Vismia japurensis Reichardt (1878: 209) (Fig. 7A–B)

20 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

8.3 Vismia laxiflora Reichardt (1878: 203) (Fig. 7C–E)

8.5 Vismia sp. (Fig. 8B)

22 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

24 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

26 • Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

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Accepted by Carlos Lehnebach: 19 Oct. 2017; published: 24 Nov. 2017

• Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

• Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

• Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.

• Phytotaxa 329 (1) © 2017 Magnolia Press CABRAL ET AL.