See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/307605973

A Taxonomic Update of Neotropical Pradosia (Sapotaceae, Chrysophylloideae)

Article  in  Systematic Botany · September 2016

DOI: 10.1600/036364416X692389

CITATIONS READS
11 2,297

3 authors:

Mario Terra-Araujo De Faria

Instituto Nacional de Pesquisas da Amazônia Universidade Estadual de Londrina
24 PUBLICATIONS   277 CITATIONS    16 PUBLICATIONS   390 CITATIONS   

SEE PROFILE SEE PROFILE

Ulf Swenson
Swedish Museum of Natural History
96 PUBLICATIONS   2,471 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Taxonomy of Sapotaceae in the Neotropics View project

Pollen morphology of Neotropical Chrysophylloideae (Sapotaceae) View project

All content following this page was uploaded by Mario Terra-Araujo on 06 March 2018.

The user has requested enhancement of the downloaded file.

 A Taxonomic Update of Neotropical Pradosia (Sapotaceae, Chrysophylloideae)
Author(s): Mário H. Terra-Araujo, Aparecida D. de Faria, and Ulf Swenson
Source: Systematic Botany, 41(3):634-650.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364416X692389

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and
environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published
by nonprofit societies, associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of
BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries
or rights and permissions requests should be directed to the individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research
libraries, and research funders in the common goal of maximizing access to critical research.
Systematic Botany (2016), 41(3): pp. 634–650
© Copyright 2016 by the American Society of Plant Taxonomists
DOI 10.1600/036364416X692389
Date of publication August 26, 2016

A Taxonomic Update of Neotropical Pradosia (Sapotaceae, Chrysophylloideae)

Mário H. Terra-Araujo,1,2,4 Aparecida D. de Faria,3 and Ulf Swenson2

1
Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Botânica (PPG-BOT),
Av. André Araújo 2.936, 69067-375, Manaus, Amazonas, Brazil.
2
Department of Botany, Swedish Museum of Natural History, Box 50007, 104 05 Stockholm, Sweden.
3
Universidade Estadual de Londrina, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas,
Box 6001, 86051-980, Londrina, Paraná, Brazil
4
Author for correspondence (araujo.mht@gmail.com)

Communicating Editor: Chrissen E. C. Gemmill

Abstract—We provide a systematic update of Pradosia (Sapotaceae, Chrysophylloideae), including overall morphology, a key to all
species, comprehensive morphological descriptions, geographic distributions, and important characteristics for each species. Phyloge-
netic analyses based on molecular data demonstrated that the genus is monophyletic and includes three main clades. Twenty-three
species of Pradosia are accepted, which are mostly distributed in lowland rainforests on either white-sand or clayish soils in tropical
South America. A rotate corolla with a short tube, lack of staminodes, a drupaceous fruit with plano-convex cotyledons, an exserted
radicle below the cotyledons, and the absence of endosperm are diagnostic for the genus. Two names are reduced into synonymy, viz.
Pradosia atroviolacea Ducke, syn. of P. grisebachii (Pierre) T. D. Penn., and Pradosia verrucosa Ducke, syn. of P. glaziovii (Pierre) T. D. Penn.
The affinity of P. argentea (Kunth) T. D. Penn., a species known only from the type collection, remains uncertain and for now excluded
from the genus.

Keywords—Amazon, Atlantic forest, extinct species, phylogeny

Pradosia Liais (Sapotaceae, Chrysophylloideae) is a genus of
23 species of trees and shrubs that are mainly distributed
throughout South America across a wide variety of habitats,
including savannahs, evergreen and deciduous forests in
the Amazon, the Brazilian Atlantic coast, and the Andes
(Pennington 1991; Alves-Araújo and Alves 2012; Terra-Araujo
et al. 2012, 2013). The Amazon region and the Brazilian
Atlantic coast are major centers of diversity for the genus with
16 species (Pennington 2006; Terra-Araujo et al. 2012, 2013).
Many of the species were previously known only from the
type specimen (see Pennington 1990, 1991), or were described
based on poor-quality collections, lacking fruits and/or
flowers. This resulted in many species assessed to be either
extinct or endangered (IUCN 2015). However, recent field-
work conducted in these areas has located some of the
presumed extinct or endangered species and revealed them
being more abundant than previously reported (Terra-Araujo
et al. 2015).
The latest taxonomic revision of Pradosia was provided by
Pennington (1990), but over the past twenty-five years it has
become outdated, with newly described species and an accu-
mulation of new material that has extended the geographical
distribution of many species (Alves-Araújo and Alves 2012;
Terra-Araujo et al. 2012, 2013). Recent phylogenetic analyses
have demonstrated that Pradosia is monophyletic, provided
that the African species Pradosia spinosa Ewango & Breteler
(Ewango and Breteler 2001) is excluded from the genus
(Anderberg and Swenson 2003; Swenson and Anderberg
2005; Swenson et al. 2008). Monophyly of Pradosia was ini-
tially inferred based on three species only, and a single mor-
phological character, the drupaceous fruit, appears to be the
only synapomorphy for the genus (Swenson et al. 2008;
Terra-Araujo et al. 2012, 2013). A more detailed study,
including 18 of the 23 species in Pennington’s revision, plus
three recently described species (Alves-Araújo and Alves
2012; Terra-Araujo et al. 2012, 2013), was recently under-
taken (Terra-Araujo et al. 2015). Results confirmed that
Pradosia is monophyletic and seems to have originated in the
Middle Eocene at 47.5 Ma somewhere in the rainforests of
South America, likely in the Amazon basin, and may have
reached the Brazilian Atlantic coast for the first time around
34.4 Ma.
Historical Review
Liais (1872) described Pradosia based on a single species
(Lucuma glycyphloea (Casar.) Mart. & Eichler) published in
Martius’s Flora Brasiliensis (Miquel 1863; pp. 82–83, Table 25).
The generic name was instated in honor of “Visconde de
Prados,” a person with political prominence who lived in Rio
de Janeiro and Minas Gerais, Brazil, in the 19th century. Diag-
nostic features for Pradosia as circumscribed at the time
included opposite leaves, absence of staminodes, fruits with a
soft pericarp, and lack of endosperm. These characters were
not attributes of either Chrysophyllum L. or Lucuma Molina, so
Liais (1872) instated his genus Pradosia.
Fifty years later, Ducke (1922) noticed in northeastern
Amazonia that some trees with opposite or subopposite
leaves and folded stamens (at least in bud), formed a distinct
group within Sapotaceae, and named these Glycoxylon
Ducke. In addition to the leaf arrangement, the inner bark of
these species also shared the presence of a sweet, afterward
astringent taste. Glycoxylon refers to the inner bark with
a sweet taste, which is called “casca doce” or “pau doce”
in northern Brazil. Ducke (1922) based the genus on
Chrysophyllum inophyllum Mart. ex Miq. from Manaus in
central Amazon and subsequently added two species discov-
ered in the State of Pará (Glycoxylon huberi Ducke and
G. pedicellatum Ducke). An additional species (Ducke 1925;
G. praealtum Ducke), with alternate or subverticillate leaves,
was also described from Pará. At that time, opposite leaves,
folded stamens, and the sweet taste of the bark were no
longer valuable generic characters. Ducke himself recognized
the close relationship between his new genus Glycoxylon and
the south Brazilian species Pradosia lactescens (Vell.) Radlk.
and P. kuhlmannii Toledo.

634
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA
Eyma (1936), after a detailed review, proposed the sup-
pression of Glycoxylon and placed Pradosia within Pouteria
Aubl. Cronquist (1946a, 1946b), on the other hand, in
his revision of several American groups of Sapotaceae,
maintained Pradosia and proposed new species and combi-
nations in the genus. In a similar manner, Ducke (1942) also
recognized Pradosia and provided a synopsis of the genus,
and later an identification key together with a new species
(Ducke 1953). In contrast, Pradosia and Glycoxylon were still
considered as distinct groups by Aubréville (1964) in his
Malacanthées, recognizing the genera based on leaf arrange-
ment: opposite in Glycoxylon and alternate in Pradosia. One
year later, Baehni (1965) united Pradosia with Chrysophyllum.
Over the years, some authors have accepted Pradosia at the
generic level, but retained some species in Chrysophyllum,
Pouteria, Pometia Vell. (an illegitimate homonyn of Pometia
J. R. Forst & G. Forst, Sapindaceae; renamed Neopometia by
Aubréville), or even Lucuma (Aubréville 1961). Species of
Pradosia have further been placed in Chrysophyllum, Ecclinusa
Mart., Diploon Cronquist, Micropholis (Griseb.) Pierre, and
Pouteria, demonstrating the difficulties there are to find the
natural limits of genera in neotropical Chrysophylloideae
(Swenson et al. 2008).
Pennington (1990), in his treatment of neotropical Sapo-
taceae, united Glycoxylon with Pradosia and recognized a total
of 23 species. He distinguished the genus from other neotropi-
cal genera by the rotate corolla with a short tube, lack of
staminodes, a stipitate attachment between the anther and its
filament, and a drupe (a drupaceous fruit). However, it is
now clear that Pradosia including Glycoxylon forms a strongly
supported group (Terra-Araujo et al. 2015).
We here present a taxonomic update of Pradosia that incor-
porates novel molecular phylogenetic data, field observa-
tions, and the examination of herbarium collections, many of
which were not available to Pennington. The taxonomic
changes herein proposed include synonymy of two species
names (i.e. Pradosia atroviolacea Ducke and Pradosia verrucosa
Ducke), a review of the species geographical distribution,
and a new diagnostic key followed by a short morphological
description for each recognized species, as well as notes
regarding species circumscription and field tips.
Materials and Methods
Pradosia was revised with full nomenclatural account in Flora
Neotropica (Pennington 1990). Since few new nomenclatural changes are
necessary here, nomenclature is limited for simplicity to the current
accepted names and basionyms. This taxonomic update includes an
identification key to species and their distributions, which are based on
fieldwork conducted throughout the Neotropics by one of us (MHTA),
bibliographic review, and examination of specimens deposited in Brazil
(ALCB, CEPEC, CVRD, HRB, IAN, INPA, IPA, MBML, MG, PEUFR, RB,
SP and UEFS), USA (CA, NY, MO, PH and US), and in Europe (K, P
and S) (abbreviations follow Holmgren et al. 1990). A complete list of the
examined specimens is provided in Supplemental Appendix S1.
Morphological data were obtained from herbarium specimens and
from fresh material collected from 2008 and onwards. Morphological
features such as habit, bark patterns, flowers, fruits, and seeds were
also studied in the field. The terminology used follows Harris and
Harris (2001).
A database with species georeferences was created from label data
from 275 specimens to generate distribution maps in QGIS program
(QGIS Development Team 2014). The Pradosia phylogeny used here is a
simplified version including 21 of the 23 recognized species reported by
Terra-Araujo et al. (2015).
Here, we adopted the general lineage-based concept (GLC) (de
Queiroz 1998) in delimiting species. Under GLC, a species is formed by
635
a separately evolving metapopulation lineage, where reproductive isola-
tion, reciprocal monophyly or ecological divergence is reached indepen-
dently at different times along the evolutionary history (de Queiroz
2007). Thus, a lineage does not have to be necessarily monophyletic,
morphologically distinct, or reproductively isolated to be recognized as
a species. In practice, any property that provides evidence of lineage
separation is relevant to infer the boundaries and number of species.
For this taxonomic treatment we considered ecology, species geographi-
cal distribution, phylogenetic relationships, and morphology important
for the delimitation of the species. Two species here recognized,
P. beardii (Monach.) T. D. Penn. and P. huberi (Ducke) Ducke, have not
been tested phylogenetically because no samples for molecular data
have been available.
Results
Habit — As in other neotropical Sapotaceae, Pradosia com-
prises woody taxa ranging in habit from shrubs such as
Pradosia schomburgkiana (A. DC.) Cronquist, understory trees
around 10–15 m (most of the species), to canopy trees up to
35 m tall like P. decipiens Ducke, P. cochlearia (Lecomte) T. D.
Penn., P. glaziovii (Pierre) T. D. Penn., and P. kuhlmannii
(Figs. 1A–B). Most tree species are buttressed, usually about
1 m high, but occasionally up to 2 m (Fig. 1C). Only
P. brevipes (Pierre) T. D. Penn. has a specialized habit with
most of the plant being below the ground (geoxylic sub-
shrub), with only a few branches emerging above the ground.
Leaves — The species have entire, alternate (Fig. 1D), sub-
verticillate, verticillate (Fig. 1E), or opposite leaves (Fig. 1F);
the latter condition is found in Pradosia beardii, P. huberi and
P. schomburgkiana. The petioles are flat above (semiterete) or
canaliculate, rarely carrying paired scales attached midway
up the petiole as in P. grisebachii (Pierre) T. D. Penn. and
P. lahoziana Terra-Araujo (Fig. 1G). The midvein may be
deeply sunken or less frequently flat or raised on the upper
leaf surface. The leaf venation of the majority of species
is eucamptodromous, but eucampto-brochidodromous vena-
tion occurs when secondaries are joined by loops in the
upper half of the blade but not in the lower half (Ellis et al.
2009). The secondary veins frequently are arcuate or some-
times straight. Intersecondaries are unusual, but if present
they extend from the midvein and disappear or integrate
into a higher venation pattern as in P. kuhlmannii, P. mutisii,
and P. schomburgkiana (Fig. 1H). The tertiaries, if visible,
are usually oblique-reticulate (Fig. 1I) or oblique (Fig. 1J).
A horizontal-reticulate pattern (Fig. 1H) is a less common
condition within the genus and only found in P. kuhlmannii,
P. mutisii, and P. schomburgkiana. The leaf blade surfaces
range from glabrous to sparsely pubescent (Fig. 1K), or
densely covered in a reddish-brown indument as in Pradosia
beardii (Fig. 1L).
Trunk and Inner Bark — In addition to leaf and flower
characters, the combination of some bark features are very
useful as field characters at the species or subclade level. For
example, a smooth bark of gray-brown or yellowish-gray
color, usually bearing discolored scars (red or brown) are typi-
cal for Pradosia cochlearia, P. decipiens, P. glaziovii, P. kuhlmannii,
P. schomburgkiana, P. subverticillata, P. surinamensis, and
P. verticillata (Figs. 1M–T). In additional, a bark with sweet
taste is found in P. cochlearia, P. kuhlmannii, P. schomburgkiana,
P. subverticillata, and P. surinamensis, a character not present in
any other genera of neotropical Sapotaceae. On the other
hand, some species are characterized by a gray or brown col-
ored bark with rather deep vertical fissures like in P. granulosa
and P. longipedicellata (Figs. 1U–V). Most exude white latex
636 SYSTEMATIC BOTANY [Volume 41

Fig. 1. Habitat, leaf venation and bark variation in Pradosia (Sapotaceae, Chrysophylloideae). A–B. Habit. A. P. granulosa, an understory tree grow-
ing in rainforests of the Amazon region. B. P. kuhlmannii, a tall canopy tree from the Brazilian Atlantic coast. C. Buttresses to 1.5 m high in P. cochlearia.
D–F. Leaf arrangement. D. Alternate in P. granulosa. E. Verticillate in P. verticillata. F. Opposite in P. schomburgkiana. G. Scales on the petiole of
P. lahoziana. H–L. Lower leaf surfaces showing the midvein, secondaries or higher leaf venation and indument. H. P. schomburgkiana. I. P. longipedicellata.
J. P. granulosa. K. P. cochlearia. L. P. beardii. M–V. Bark patterns and slash of ten species. The scale bar in H–L corresponds to 1 cm. Photos: M. H. Terra-
Araujo (A–E, G–V), A.Vicentini (F).
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA
and the color of the inner bark ranges from light orange or
yellowish in P. decipiens, P. glaziovii, P. granulosa, P. lactescens,
P. lahoziana, P. ptychandra, P. subverticillata, and P. verticillata
to pinkish or reddish in P. cochlearia, P. kuhlmannii,
P. longipedicellata, P. montana, and P. schomburgkiana.
Flowers — In Sapotaceae flowers are borne in fascicles and
in Pradosia the fascicles usually appear below the apical clus-
ters of leaves, rendering the majority of the species
ramiflorous. Flowers of some species, viz. Pradosia lactescens,
P. lahoziana, and P. ptychandra (Eyma) T. D. Penn., are borne
on the trunk, making them cauliflorous (Fig. 1Q). One spe-
cies, P. longipedicellata, has a terminal inflorescence (Fig. 2A).
Flowers are bisexual and frequently 5-merous; the corolla is
rotate with a short tube and spreading to flat corolla lobes.
The color varies, ranging from white, green, and reddish to
deep wine red (Fig. 2A–F). The stamens are exserted and
equal the number of the corolla lobes. They are further usu-
ally attached in the corolla tube orifice (seldom just below
the tube orifice). The filaments are long, geniculate in bud,
becoming straight in open flowers, and are attached to the
glabrous anthers by a narrow stipe (Fig. 2C). Staminodes are
lacking, a trait considered diagnostic for the genus (Liais
1872; Ducke 1953; Pennington 1990; Swenson et al. 2008).
The ovary is 5(–6)-locular and pubescent, in contrast to the
glabrous style that is simple, without any visible stigmatic
areas (cf. Planchonella; Swenson et al. 2007).
Fruits — The fruits usually have an obovoid or ellipsoid
form, less frequently ovoid, and normally with a smooth
surface (Figs. 2G–I). Only Pradosia glaziovii (Pierre) T. D.
Penn. and P. granulosa Pires & T. D. Penn. have fruits with
small spicules covering the surface, giving the fruit a
muricate aspect (Fig. 2J). The mesocarp is soft and most of
the species are edible, having a sweet taste. In Pradosia, fruits
have been characterized as a drupe with a thin cartilaginous
endocarp (Pennington 1990, 1991; Swenson et al. 2008).
However, it is better characterized as a drupaceous fruit,
because the endocarp is rather jelly-like, partly transparent,
never hardened (Terra-Araujo et al. 2012, 2013; Figs. 2K–N).
Fruits are further considered as one-seeded, but more than
one seed have been observed in P. cochlearia, P. granulosa,
P. longipedicellata and P. restingae Terra-Araujo. The seeds are
a little laterally compressed with a smooth, shinning testa,
and with a seed scar as long as the seed itself; the cotyledons
are plano-convex, the radicle is exserted below the cotyle-
dons and there is no endosperm.
Phylogenetic Relationships — A recent phylogenetic study
of Pradosia was recently undertaken (Terra-Araujo et al.
2015). The results of this study support Pradosia as mono-
phyletic and indicate P. longipedicellata as the sister to all
other species of the genus (Fig. 3). According to this study,
the species are grouped in three clades, which were
strongly supported and are morphologically, geographi-
cally, and ecologically coherent. The Montane clade
includes three species that have narrow distributions in the
northeast corner of Colombia, Venezuela, and Ecuador
(Fig. 4A). All three species are trees having an inner bark
lacking a sweet taste, leaves with sunken midvein, fascicles
of flowers borne along the branches (ramiflorous) of old
wood below the leaves without any adjacent leaf scars
(axillary in P. colombiana), and greenish flowers. The Sweet-
bark clade includes four species that are almost exclusively
distributed in the Amazon region and have leaves with flat
or raised midvein, fascicles of flowers below the leaves,
637
and greenish flowers with small corollas. The Red-flowered
clade includes species from the Amazon region (eight spe-
cies), the Brazilian Atlantic coast (four species), the savan-
nah of central Brazil, and the Chocó region on the Pacific
coast in Colombia. Within this clade the majority of species
have non-sweet bark, leaves with sunken midvein, reddish
or wine red flowers, and larger (> 4 mm long) corolla. This
latter clade further includes plants with flowers borne on
the trunk, scales attached at the middle of the petiole, and
fruits with muricate surface, features not found in any
other clade of Pradosia.
Distribution and Habitat — Pradosia occurs throughout
South America, although Pradosia grisebachii extends to Costa
Rica and Panama. The highest diversity of Pradosia is found
in the Amazon region (10 species), followed by the Brazilian
Atlantic coast with six species (Fig. 4). Most of the species
grow in lowland rainforest on either white-sand or clayish
soils. Nine species are found in forests on white-sand soil,
locally known as campina, campinarana, and restinga. Such
patches of forests are nutrient-poor, have low pH and low
water retention, resulting in completely different ecological
conditions (Prance and Schubart 1978; Anderson 1981;
Prance 1996). One species has been recorded in permanent
flooded-forests, or swampy land in northeast Amazon
(P. huberi). Only P. brevipes occurs in savannas ranging
from 300–1000 m altitude in central Brazil. Three species,
P. colombiana (Standl.) T. D. Penn. ex T. J. Ayres & Boufford,
P. montana T. D. Penn., and P. mutisii Cronquist are found
in dry montane forest up to 1200 m altitude in north and
northwest South America.
Conservation assessments — During the past years have
Red List assessments for conservation using IUCN (2015)
guidelines and criteria have usually accompanied each
newly described or revised species. Assessments are possibly
the most authoritative list to conservation of species. By
using the IUCN criteria, species are categorized based on
multiple sources of evidence, including population size, pop-
ulation decline, range area, and range decline. For the past
century, many species of Pradosia have been, and are still,
poorly documented. In the light of the IUCN assessments,
this resulted in three extinct species, viz. P. argentea (Kunth)
T. D. Penn., P. mutisii and P. glaziovii, where others were clas-
sified as Critically Endangered or Vulnerable (IUCN 2015).
Recent fieldwork in the Amazon and the Atlantic forest have
revealed new populations of almost all species. We here
report previously unknown populations of P. cochlearia,
P. decipiens, P. glaziovii, P. kuhlmannii, P. lahoziana, P. mutisii
and P. subverticillata, which extend their distributions and
point out that some of the presumed extinct or endangered
species are more common than previously reported. For
this reason, we refrain from red list assessments because
with the current knowledge, most species would in fact be
classified as data deficient (DD) and such assessments are
not meaningful.
Taxonomic Treatment
Pradosia Liais, Clim. Géol. Géogr. Bot. Brésil 614. 1872.—
TYPE: BRAZIL, Pradosia glycyphloea (Casar.) Liais.
(= Pradosia lactescens (Vell.) Radlk.)
Tall canopy trees, treelets, or rarely geoxylic shrubs;
buttresses short or lacking, less frequently up to 2 m tall.
638 SYSTEMATIC BOTANY [Volume 41

Fig. 2. Flowers and fruits of Pradosia (Sapotaceae, Chrysophylloideae). A–F. Flowers. A. P. longipedicellata. B. P. schomburgkiana. C. P. montana.
D. P. restingae. E. P. lactescens. F. P. granulosa. G–J. Fruit. G. P. restingae. H. P. lactescens. I. P. surinamensis. J. P. granulosa. K–N. Transection of fruits show-
ing the half-transparent cartilaginous endocarp. K. P. cochlearia. L. P. granulosa. M. P. restingae. N. P. surinamensis. Photos: A. Alves-Araújo (D, G, M),
R. I. Barbosa (I, N), J. Jardim (A), M. H. Terra-Araujo (E, F, H, J, K, L), A. Vicentini (B).
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA 639

Fig. 3. Phylogeny of Pradosia (Sapotaceae, Chrysophylloideae) obtained from Bayesian analyses using three nuclear molecular markers (ITS, ETS
and RPB2) by Terra-Araujo et al. (2015) with herein adopted nomenclature. Posterior probabilities (PP) are indicated as black (PP > 0.95) and gray
(0.85 < PP < 0.95) branches.

Bark gray-brown, or yellowish-gray colored, smooth or bark non-sweet or sweet. Leaves simple, entire, sometimes
rough, shallowly fissured or scaling in thin asymmetric revolute, usually alternate or subverticillate, less frequently
plates leaving reddish, orange, or dark marks; slash of verticillate or opposite, coriaceous or chartaceous, clus-
trunk light orange, yellowish, pinkish or reddish; inner tered at branch tips or scattered along branches; upper
640 SYSTEMATIC BOTANY [Volume 41

Fig. 4. Known geographic distribution to the species of Pradosia (Sapotaceae, Chrysophylloideae). A. Sister species and Montane clade.
B. Sweet-bark clade. C. Clade “a” from Red-flowered clade. D. Clade “b” from Red-flowered clade. * = Species not included in the phylogeny
of the genus.

surface glabrous or with residual indument at the midvein
and secondary veins, lower surface glabrous, pubescent,
puberulous or tomentulose; trichomes malpighiaceous,
Y-shaped, usually brownish, less frequently whitish, golden
or gray; midvein usually sunken, less frequently flat or
raised; secondary venation eucamptodromous, eucampto-
brochidodromous or rarely brochidodromous, parallel or
slightly divergent, flat, raised or impressed above; inter-
secondaries sometimes present, but rarely well-developed;
tertiaries oblique, horizontal, reticulate, or a combination of
these patterns; higher venation usually areolate; petiole
semiterete or canaliculate, pubescent, glabrous or with
some residual indument, sometimes with small scales, 0.5–
2.0 mm long, attached midway on the petiole. Flowers
bisexual, clustered in fascicles, usually below the leaves
(ramiflorous), sometimes on the trunk (cauliflorous), rarely
clustered at the apices of short branches (terminal) or axil-
lary; pedicel glabrous, usually tomentulose. Sepals are in a
single whorl, 5(–6), usually ovate, with a rounded apex,
glabrous, tomentose or tomentulose outside, glabrous
inside, and persistent in fruit. Corolla is rotate with 5(–6)
corolla lobes; corolla tube always shorter than the lobes,
wine red, reddish, greenish or less frequently white, gla-
brous, sericeous or tomentulose outside, glabrous inside.
Staminodes absent. Disk absent. Stamens opposite and
equal the corolla lobes, glabrous; filaments geniculate in
bud, slightly exserted in open flowers; anthers glabrous,
cream or white. Ovary with 5(–6) locules, frequently
puberulous or strigose; style simple, glabrous. Fruit drupa-
ceous, ellipsoid, obovoid or ovoid, greenish, yellow or
orange when mature, smooth, lenticellate or muricate, gla-
brous, puberulous or tomentulose, usually one-seeded;
mesocarp soft and endocarp cartilaginous. Seeds usually
laterally compressed; testa smooth, brown, shinning; seed
scar adaxial, 100% of seed length, 10–25% of seed circum-
ference; embryo with plano-convex cotyledons, exserted
radicle, and no endosperm.
Identification tips — The genus is very easy to recog-
nize by its rotate corolla with a short tube, lack of
staminodes and drupaceous fruits, unlike any other
Sapotaceae in South America. Most species are distin-
guished by some vegetative features such as the taste of
the inner bark, leaf arrangement, venation, leaf surface,
and presence or absence of appendages on the petiole. If
fertile specimens are present, useful characters include
the position of the fascicles, the size and color of the
corolla, and the fruit surface. Even the ecology is usually
species specific. A combination of these characters usu-
ally leads to the right species as exemplified in Figs. 1
and 2.
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA 641

Key to the Species of PRADOSIA

1. Geoxylic subshrub in savannahs from central Brazil to Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. brevipes

1. Tree, treelets or shrubs in evergreen and deciduous forests, not in savannahs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Upper leaf surface with a flat or raised midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Secondary venation brochidodromous; intersecondaries present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaves alternate, chartaceous; intersecondaries present, short. On clayish soils in the Brazilian Atlantic coast . . . . . . . . . . P. kuhlmannii
4. Leaves opposite, coriaceous; intersecondaries long, extending towards the margin. In campina or
campinarana forests throughout Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. schomburgkiana
3. Secondary venation eucamptodromous; intersecondaries absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Petiole semiterete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Secondary veins divergent. Lower leaf surface glabrous; ramiflorous; pedicel 1.0–4.0 mm long;
corolla greenish. In swampy lands or flooded forest along the northeastern Amazon coast (estuary areas) . . . . . . . . . . . P. huberi
6. Secondary veins parallel. Lower leaf surface covered by whitish or ferruginous trichomes;
inflorescence terminal; pedicels 18–30 mm long; corolla white. In restinga forested in southern Bahia . . . . . . . . P. longipedicellata
5. Petiole slightly canaliculate or canaliculate in the upper half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Inner bark with sweet taste; leaves chartaceous, glabrous or appressed brown-puberulous on the lower
surface; petiole 0.7–1.6 cm long; pedicel 0.5–4.0 mm long; corolla 1.7–3.0 mm long; fruits 3.5–5.0 cm long . . . . . . . . P. cochlearia
7. Inner bark without a sweet taste; leaves coriaceous, glabrous; petiole 1.7–2.6 cm long;
pedicel 5.0–13.0 mm long; corolla 4.5–5.2 mm long; fruits 3.0–3.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. decipiens
2. Upper leaf surface with a sunken midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Petiole with scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Ramiflorous; pedicel 1.0–3.0 mm long; corolla 4.0–5.0 mm long; fruits 3.5–5.0 cm long. Known from
western Amazonia to Panama and Costa Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grisebachii
9. Cauliflorous; pedicel 7.0–9.0 mm long; corolla 6.4–9.5 mm long; fruits 2.5–3.0 cm long. Known from
central Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lahoziana
8. Petiole without scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Lower leaf surface sparsely gray-puberulous, glabrous or glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Intersecondaries present and extending towards the margin. In dry montane forests of Colombia,
Ecuador, and Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mutisii
11. Intersecondaries absent or short, not extending towards the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Fascicles mostly cauliflorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Pedicel 4.0–6.0 mm long; corolla usually glabrous. In rainforests on clayish soils in the
Brazilian Atlantic coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lactescens
13. Pedicel 7.0–10.5 mm long; corolla sparsely tomentulose outside. In non-flooded forests on
clayish soils in central and eastern Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ptychandra
12. Fascicles ramiflorous or axillary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Corolla ≥ 4.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Petiole 0.6–0.8 cm long. In forested and open vegetation on white-sand soils in
northeast Atlantic coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. restingae
15. Petiole ≥ 0.9 cm long. In rainforests on sandy or clayish soils throughout Amazonia . . . . . . . . . . . . . . . . . . . . . 16
16. Inner bark without a sweet taste; lower leaf surface with indument at the
midvein and secondary veins; petiole 0.9–2.0 cm long, canaliculate in the
upper half; flowers wine red; fruit muricate. In rainforest on sandy soils in
northeastern Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. granulosa
16. Inner bark with sweet taste; lower leaf surfaces glabrous; petiole 2.0–3.2 cm long,
canaliculate; flowers greenish; fruit smooth. In forest on clayish or white-sandy
soils in central and northwestern Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subverticillata
14. Corolla ≤ 4.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17. Lower leaf surface sparsely gray-puberulous; fascicle axillary; pedicel 5.0–12 mm long.
In deciduous forests in Colombia and Venezuela . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. colombiana
17. Lower leaf surface glabrous or glabrescent; fascicle ramiflorous; pedicel ≤ 2.5 mm long.
In rainforest of Amazon region and northern South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18. Pedicel 1.5–2.5 mm long. Fruit 5.0–7.0 cm long, densely lenticellate, and glabrous.
In wet forests of northern Colombia and north and northwest Venezuela . . . . . . . . . . . . . . . . P. caracasana
18. Pedicel 0.5–2.0 mm long. Fruit 2.0–4.0 cm long, not lenticellate, glabrous or
puberulous. In rainforest on sandy or clayish soils in northeastern Amazonia,
Brazil and Guyana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. surinamensis
10. Lower leaf surface densely tomentulose, puberulous or with erect two-branched pale-brownish trichomes . . . . . . . . . . . . . . . . . . . . 19
19. Corolla ≥ 5mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20. Leaves 20–40 cm long; petiole 2.0–2.6 cm long; pedicel 5.0–10 mm long; corolla ca. 7.5 mm long,
greenish; fruit smooth. In wet lowland forest of Chocó region in Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cuatrecasasii
20. Leaves 9–24 cm long; petiole 0.8–2.2 cm long; pedicels ca. 1 mm long; corolla 5.0–6.5 mm long,
reddish; fruit muricate. In rainforests on clayish soils throughout the Brazilian Atlantic coast . . . . . . . . . . . . . . . P. glaziovii
19. Corolla < 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21. Lower leaf surface with indument of erect two-branched pale-brownish trichomes; pedicel
2.0–3.0 mm long; corolla glabrous or sparsely tomentulose outside, especially the corolla tube.
In deciduous coastal forests of Ecuador and Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. montana
21. Lower leaf surface densely reddish-brown tomentulose or brown-pubescent; pedicel 1.0–1.7 mm long;
corolla sericeous or tomentulose outside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22. Leaves opposite or subverticillate, lower surface densely reddish-brown tomentulose;
tertiaries obscure; corolla 2.0–3.4 mm long, greenish. Fruit 1.2–1.6 cm, brown-tomentulose.
In scrublands or tall forests, usually on sandy soils of western corner of Guiana Shield . . . . . . . . . . . . . . . . P. beardii
22. Leaves verticillate, lower surface brown-pubescent; tertiaries visible; corolla 4.0–4.6 mm long,
reddish. Fruit 3.5–4.5 cm long, glabrous. In non-flooded forests on clayish soils of central
and eastern Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. verticillata
642 SYSTEMATIC BOTANY
Pradosia beardii (Monach.) T. D. Penn., Fl. Neotrop.
Monogr. 52: 665. 1990. ≡ Chrysophyllum beardii Monach.,
Phytologia 3: 159. 1949.—TYPE: TRINIDAD. Long Stretch
Reserve, alongside main road near Turure junction, 23
Aug 1949 (fl), R. S. Ayliffe s.n. [Monachino 526] (holotype:
NY 00273412!; isotype: U 0006709!).
Shrub 1.5–3.0 m tall, sometimes tree up to 20 m; buttresses
short; bark smooth, reddish, scaling in thin irregular plates
and leaving deep marks; slash of trunk unknown. Leaves
loosely clustered at branch tips, opposite or subverticillate,
5–16 × 2–6 cm long, oblanceolate or obovate, coriaceous,
with a revolute margin; upper surface glabrous or with resid-
ual indument at the midvein and secondary veins, lower sur-
face densely reddish-brown tomentulose; midvein sunken on
the upper surface; secondary venation eucamptodromous,
parallel, impressed above; intersecondaries absent; tertiaries
oblique to horizontal, obscure; petiole 0.5–1.3 cm long,
slightly canaliculate, pubescent, without scales. Flowers 3–
many in each fascicle, ramiflorous; pedicel 1.0–1.5 mm long,
tomentose. Sepals are tomentose outside. Corolla is 2.0–
3.4 mm long, greenish, tomentulose outside. Fruit 1.2–1.6 cm
long, ellipsoid or obovoid, smooth, and brown-tomentulose.
Distribution and habitat — This species is found in the
western corner of Guiana Shield (Fig. 4C). It occurs in scrub-
lands or tall forests, usually on sandy soils associated with
periodic flooding, although it is also found in humid forest
up to 1000 m alt. in Venezuela.
Recognition — Pradosia beardii is easily distinguished from
all other Pradosia species by its dense reddish-brown pubes-
cence covering the pedicel and the lower leaf surface
(Fig. 1L).
Pradosia brevipes (Pierre) T. D. Penn., Fl. Neotrop.
Monogr. 52: 641. 1990. ≡ Ecclinusa brevipes Pierre, Not.
Bot. Sapot. 57. 1891.—TYPE: BRAZIL. Prov. Paraná. São
Bento (Rincao das Pedras), prope Castro, 8 Jun 1880,
C. A. W. Schwacke 2894 (holotype: GOET 010954!; iso-
type: P 00649435 [fragment]!)
Geoxylic shrub to 30–100 cm tall. Leaves alternate, 8–23 ×
2.5–8.5 cm long, oblanceolate, coriaceous, upper surface gla-
brous or with some residual indument at the midvein and
secondary veins, lower surface usually pubescent; midvein
sunken on the upper surface; secondary venation eucampto-
dromous, parallel, impressed above; intersecondaries absent;
tertiaries oblique or oblique-reticulate; petiole 0.6–1.8 cm
long, canaliculate, pubescent, without scales. Flowers 3–10 in
each fascicle, ramiflorous; pedicel 5–30 mm, tomentulose.
Sepals are tomentulose outside. Corolla is 4.2–7.3 mm long,
reddish, sparsely sericeous outside. Fruit 2.5–4.0 cm long,
ellipsoid, yellowish, smooth and puberulous.
Distribution and habitat — Widespread from central Brazil,
south into Paraná State and eastern Paraguay in scrublands
(Cerrado), usually on sandy soils, from 300–1000 m alt.
(Fig. 4D).
Recognition — Pradosia brevipes is closely related to P.
glaziovii and P. granulosa, species with similar leaf shape and
reddish flowers, but they cannot be confused since P. brevipes
is the only geoxylic species in the genus, not forming treelets
or trees, and is restricted to savannahs, not occurring in wet
evergreen forests. If one confronts a fruiting specimen where
the habit is unclear, fruits of P. brevipes are smooth, not
muricate as in other members with similar leaf shape.
[Volume 41
Pradosia caracasana (Pittier) T. D. Penn., Fl. Neotrop.
Monogr. 52: 668. 1990. ≡ Oxythece caracasana Pittier,
Arb. Arbust. Venez. 1: 11. 1921.—TYPE: VENEZUELA.
Lower Cotiza, near Caracas, 800–1200 m alt., [28] Jul
1918 (fl), H. Pittier 7955 (holotype: US 00113132!; iso-
types: BKL 00002344!, F 0072112F and F 0072113F, GH,
MICH 1210173!, MO 1991133, PH 00018154 and PH
00018155, VEN).
Tree up to 10 m, but can reach 25 m tall; buttresses short.
Bark smooth, reddish-brown, shallowly fissured, sloughing
off in thin plates and leaving deep whitish marks; slash
unknown; inner bark non-sweet tasting. Leaves subverti-
cillate or alternate, 8–23 × 4–10 cm, elliptic or obovate,
chartaceous, glabrous on both sides; midvein sunken on the
upper surface; secondary venation eucamptodromous, paral-
lel, slightly impressed above; intersecondaries absent; tertia-
ries oblique-reticulate; petiole 1.0–2.7 cm long, canaliculate,
glabrous or with residual indument, without scales. Flowers
3–20 in each fascicle, ramiflorous; pedicel 1.5–2.5 mm long,
tomentulose. Sepals are tomentulose outside. Corolla is 3.5–
4.0 mm long, greenish, sericeous outside. Fruit 5.0–7.0 cm
long, ellipsoid, greenish-brown, smooth, densely lenticellate,
and glabrous.
Distribution and habitat — Pradosia caracasana is known
from northern Venezuela where it occurs in wet lowland to
montane forests, from 80–900 m alt. (Fig. 4A).
Recognition — Pradosia caracasana, P. colombiana, and P.
montana form a strongly supported group (Fig. 3). Pradosia
caracasana and P. colombiana are sister species and are some-
times confused morphologically, especially in Zulia, north-
western Venezuela, where these two species meet (Pennington
1990). However, P. caracasana differs from P. colombiana by
glabrous leaves (vs. pubescent), ramiflorous fascicles (vs.
axillary), flowers with 1.5–2.5 mm long pedicels (vs. 5–
12 mm long), and densely lenticellate fruits (vs. smooth).
They also occur in different types of forests; P. caracasana
occurs in wet evergreen forests, while P. colombiana occurs
in deciduous forests.
Pradosia cochlearia (Lecomte) T. D. Penn., Fl. Neotrop.
Monogr. 52: 655. 1990. ≡ Chrysophyllum cochlearium
Lecomte, Notul. Syst. (Paris) 4: 63. 1923.—TYPE:
FRENCH GUIANA. Charvein, 13 Nov 1913 (fl), R.
Benoist 224 (holotype: P 00649438!; isotypes: F 0071930F
[fragment]!, P 00649439!).
Tree up to 35 m tall; buttresses up to 2 m tall (Fig. 1C);
bark grayish, smooth, scaling and leaving orange-brown
patches; slash of trunk pinkish (Fig. 1M); inner bark sweet-
tasting. Leaves alternate, less frequently opposite or verti-
cillate, oblanceolate or obovate, 5–11(–15) × 2.5–4.5 cm,
chartaceous, glabrous or appressed brown-puberulous on
the lower surface (Fig. 1K); midvein raised on the upper sur-
face; secondary venation eucamptodromous, parallel, raised
above; intersecondaries sometimes present, short; tertiaries
oblique to horizontal, sometimes obscure; petiole 0.7–1.6 cm
long, slightly canaliculate, glabrous or appressed brown-
puberulous, without scales. Flowers 4–10 in each fascicle,
ramiflorous; pedicel 0.5–4.0 mm long, tomentulose. Sepals
are tomentulose outside. Corolla is 1.7–3.0 mm long, green-
ish, sparsely sericeous outside. Fruit 3.5–5.0 cm long,
ellipsoid or obovoid, yellow-greenish, smooth, glabrous or
glabrescent (Fig. 2K).
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA
Distribution and Habitat — Widespread, extending from
eastern to central Amazon in Brazil and French Guiana
(Fig. 4B). It usually grows in tall forests, non-flooded sys-
tems on sandy soils, although some plants have been col-
lected in tall forests on clayish soils near Manaus, central
Amazon. Additional specimens have been reported for
Caquetá (Colombia), and from Loreto in Peru, in upland for-
est on white-sandy soils. However, we have not been able to
examine voucher material from these areas.
Note — Raymond Benoist (1881–1970), a French botanist,
was sent on an official botanical mission to French Guiana
in 1913–1914. His collection Benoist 224 was later used by
Lecomte (1923) to describe Chrysophyllum cochlearium, a spe-
cies transferred to Pradosia by Pennington (1990). Pennington
stated that the holotype is present in the Paris herbarium. In
Paris, two sheets of Benoist 224 are present, P 00649438 and
P 00649439, of which the first mentioned has pencil draw-
ings of a flower, a corolla, and an ovary that concur with the
diagnosis (Lecomte 1923). We therefore, in accordance with
Article 9 of the International Code of Nomenclature (McNeill
et al. 2012), believe that Benoist 224 (P 00649438) constitutes
the holotype.
Recognition — Pradosia cochlearia is similar in morphology
to P. huberi and P. schomburgkiana among the other Amazonian
species with sweet bark. It differs from P. huberi in that
the secondary veins are parallel (vs. divergent), the tiny, 1.7–
3.0 mm long corolla (vs. 4.5–5.5 mm long) (see Pennington
1990), and that it grows in non-flooded forests, usually on
sandy soils (not swampy lands or flooded forests). It is fur-
ther distinguished from P. schomburgkiana in its eucampto-
dromous venation and by the absence or much shorter
intersecondaries (vs. brochidodromous venation and long
intersecondaries extending towards the margin). Pennington
(1990) recognized two subspecies, P. cochlearia subsp.
cochlearia and P. cochlearia subsp. praealta (Ducke) T. D. Penn.,
based on the presence or absence of a sericeous indument on
the lower leaf surface plus whether the tertiaries are visible
or obscure. We have examined the morphology of samples
covering the entire geographic distribution of these subspe-
cies and could not find morphological coherence supporting
subspecies in P. cochlearia.
Pradosia colombiana (Standl.) T. D. Penn. ex T. J. Ayers &
Boufford, Brittonia 40: 428. 1988. ≡ Sideroxylon colombianum
Standl., Trop. Woods 22: 13. 1930.—TYPE: COLOMBIA.
Region of Santa Marta, 250 ft alt., Aug 1898–1901 (fl),
H. H. Smith 456 (holotype: F 0072232F!; isotypes:
A 00589962 and A 00589963, BM 000952570!, BR
0000005415045!, COL 000003939!, CM 1674, E 00259453!,
F 0072232F, G 00439604!, MICH 1104683!, MO 345912!,
MPU 019068!, NY 00296951!, PH 00024050, S 05–10509!,
U 0006711!, US 00113305!).
Tree up to 8 m, but can reach 25 m tall; buttresses short;
bark red-brown, smooth, with irregular grayish spots along
the trunk; slash of trunk unknown; inner bark non-sweet
tasting. Leaves alternate, elliptic, 8–25 × 3–10 cm, chartaceous,
upper surface glabrous or with some residual indument,
lower surface sparsely gray-puberulous; midvein sunken on
the upper surface; secondary venation eucamptodromous,
parallel, not impressed above; intersecondaries sometimes
present, short; tertiaries oblique or horizontal; petiole 1.7–
6.5 cm long, slightly canaliculate, glabrescent, without scales.
643
Flowers 5–20 in each fascicle, axillary; pedicel 5.0–12 mm
long, tomentulose. Sepals are tomentulose outside. Corolla
is 3.0–4.0 mm long, greenish, sparsely sericeous outside.
Fruit 3.0–4.0 cm long, ellipsoid, gray-greenish, not lenticellate,
and puberulous.
Distribution and habitat — Pradosia colombiana occurs in
the north corner of Colombia, along the Atlantic coast, into
northwestern Venezuela, in deciduous forests from sea level
to 400 m alt. (Fig. 4A).
Recognition — This species is easy to recognize because it
is the only congener growing in deciduous forest with axil-
lary flowers and smooth fruits.
Pradosia cuatrecasasii (Aubrév.) T. D. Penn., Fl. Neotrop.
Monogr. 52: 663. 1990. ≡ Ecclinusa cuatrecasasii Aubrév.,
Adansonia 7: 144. 1967.—TYPE: COLOMBIA. Depart-
amento del Valle, Costa del Pacífico, Río Naya, Puerto
Merizalde, bosque 5–20 m alt., 20 Feb 1943 (fl), J.
Cuatrecasas 13988 (holotype: US 00153804! and US
00153803!; isotypes: COL 000003925, F 0071965F! and
F 0071966F, P 00649441!).
Large buttressed tree; bark grayish, scaling; slash of trunk
unknown; inner bark sweet-tasting. Leaves alternate, 20–
40 × 8–14 cm, oblanceolate, coriaceous, upper surface gla-
brous or with some residual indument, lower surface
densely brown-pubescent; midvein sunken on the upper
surface; secondary venation eucamptodromous, parallel,
impressed above; intersecondaries absent; tertiaries oblique,
impressed above; petiole 2.0–2.6 cm long, canaliculate,
pubescent, without scales. Flowers 5–14 in each fascicle,
born on small burls, ramiflorous; pedicel 5.0–10 mm long,
tomentulose. Sepals are tomentulose outside. Corolla ca.
7.5 mm long, greenish, sericeous outside. Fruit 5.5–6.5 cm
long, ellipsoid, green-brownish, smooth, and glabrescent.
Distribution and Habitat — This species is only known
from wet lowland forests at low altitudes along the Pacific
coast of Colombia (Chocó) (Fig. 4C).
Note — The holotype Cuatrecasas 13988 is deposited in the
Smithsonian herbarium US (Aubréville 1967), not in Paris as
stated by Pennington (1990). Further, it is mounted in two
parts, one with leaves and one with branches carrying
flowers, fully in accordance with article 8.3 of the Interna-
tional Code of Nomenclature (McNeill et al. 2012).
Recognition — Pradosia cuatrecasasii is readily recognized
by the large, strongly coriaceous leaves, with sunken midvein,
conspicuous secondary veins, and densely brown-pubescent
lower surface. There is no other species of Pradosia in west
Colombia with which it can be confused.
Pradosia decipiens Ducke, Trop. Woods 71: 17. 1942.—
TYPE: BRAZIL. Amazonas: Manaus, Cachoeira baixa
do Tarumã, mata da t. f. baixa, 2 Dec 1932 (fl), A. Ducke
385 [RB24860] (holotype: RB 00379516!; isotypes: G
00439603, K 000640454! and K 000640455!, R 000054689,
RB 00635940! and RB 00635945!, WIS 00000902MAD).
Tree up to 20 m tall; buttresses up to 0.5 m tall; bark gray-
ish, smooth, scaling in asymmetric thin plates leaving orange
marks; slash of trunk orange (Fig. 1N); inner bark non-sweet
tasting. Leaves alternate, 8–19 × 4–8 cm, oblanceolate, coria-
ceous, glabrous; midvein raised on the upper surface;
secondary venation eucamptodromous, parallel, impressed
above; intersecondaries absent; tertiaries oblique, obscure;
petiole 1.7–2.6 cm long, canaliculate in the upper half,
644 SYSTEMATIC BOTANY
glabrous, without scales. Flowers 5–12 in each fascicle,
ramiflorous; pedicel 5.0–13.0 mm long, glabrous. Sepals are
glabrous outside. Corolla is 4.5–5.2 mm long, greenish, gla-
brous. Fruit 3.0–3.5 cm long, ellipsoid, yellowish, smooth,
and glabrous.
Distribution and habitat — Two disjunct populations of
Pradosia decipiens are known: near Manaus in central
Amazon, where it has been collected in forest on white-
sandy soil in the Reserva Florestal Adolpho Ducke and in
Solano, Caquetá, southern Colombia (Fig. 4B). This species
usually grows in lowland non-flooded forests or occasionally
flooded forests on nutrient-poor white-sandy soils (campina
and campinarana).
Recognition—Pradosia decipiens, like P. cochlearia, P. kuhlmanni
and P. schomburgkiana, belongs to the Sweet-bark clade and has
similar bark structure, leaves with flat or raised midvein on
the upper surface, and green flowers, but P. decipiens differs
from these congeners in that the inner bark is not sweet in
taste. In addition, the tertiary venation is weak and not promi-
nent and the corolla is slightly longer, 4.5–5.2 mm instead of
less than 4 mm long.
Pradosia glaziovii (Pierre) T. D. Penn., Fl. Neotrop.
Monogr. 52: 643. 1990. ≡ Ecclinusa glaziovii Pierre, Not.
Bot. Sapot. 56. 1891.—TYPE: BRAZIL. Rio de Janeiro:
Itaborahy, 12 Sep 1875 (fl, fr), A. Glaziou 8229 (lectotype
chosen by Pennington [1990: 643]: P 00649442!; iso-
lectotype: BR 0000005415342 and BR0000005415014,
F 0BN004272 [photo negative image]!, G 00439602!,
K 000518106!, P 00649443!).
Pradosia verrucosa Ducke, Bol. Tecn. Inst. Agron. N. 28: 27.
1953.—TYPE: BRAZIL. Pernambuco: Recife, Estrada da
Aldeia, Km 28, mata alta, 30 Apr 1952 (fl, fr), A. Ducke &
A. Lima 80 (holotype: IAN!; isotype: US 00037033!).
syn. nov.
Tree up to 30 m tall; short buttresses; bark gray-brown,
smooth; slash of trunk light orange; inner bark non-sweet
tasting (Fig. 1O). Leaves clustered at branch tips, alternate or
verticillate, 9–24 × 4–11 cm, oblanceolate or obovate, coria-
ceous, upper surface glabrous or midvein pubescent, lower
surface puberulous; midvein sunken on upper surface;
secondary venation eucamptodromous, parallel, slightly
impressed above; intersecondaries absent; tertiaries oblique;
petiole 0.8–2.2 cm long, canaliculate in upper half, pubescent,
without scales. Flowers 2–10 in each fascicle, ramiflorous;
pedicel ca. 1 mm long, tomentulose. Sepals are tomentulose
outside. Corolla is 5.0–6.5 mm long, reddish, sericeous out-
side. Fruit 3.5–5.0 cm long, ovoid to ellipsoid, yellowish,
muricate, and brown-tomentulose.
Distribution and Habitat — Pradosia glaziovii is found
throughout the Brazilian Atlantic coast, occurring from
Pernambuco in the north to the state of Espírito Santo in
the south, occurring in wet lowland tall forests on clayish
soils (Fig. 4D).
Recognition — Pennington (1990) suggested that Pradosia
glaziovii is closely related to P. brevipes, which has been con-
firmed by our phylogenetic analysis (Terra-Araujo et al.
2015). These two species have similar leaves, but readily rec-
ognized in that P. glaziovii is a tall canopy tree confined to
wet coastal forests whereas P. brevipes is a geoxylic shrub
from the savannah in central Brazil. However, Pennington
overlooked the similarities between P. glaziovii and P. verrucosa,
[Volume 41
two species of Pradosia with muricate fruits, restricted to the
Atlantic coast, and occurring in wet forests on clayish soils.
The name glaziovii was proposed in honor of A. Glaziou
whereas verrucosa probably refers to the muricate fruit.
Because P. glaziovii is the older name it has authority over
P. verrucosa. Pradosia glaziovii is mainly characterized by being
a tall canopy tree with coriaceous leaves and with appressed
indument on the lower surface, short pedicel, reddish flowers,
muricate fruit surface, and by its occurrence in wet forest on
clayish soils.
Pradosia granulosa Pires & T. D. Penn., Fl. Neotrop.
Monogr. 52: 645. 1990.—TYPE: BRAZIL. Pará: Serra dos
Carajás, AMZA camp 4-Alfa, ca. 25 km by road north-
west of Rio Itacaiúmas, edge of forest around aban-
doned mining camp, 5°46′S, 50°36′W, ca. 225 m alt.,
6 Jun 1982 (fl), C. R. Sperling et al. 5923 (holotype: K
001096525!; isotypes: F V0072206F!, MG!, NY 01168534!,
US 00516779!).
Treelet 4–6 m tall; unbuttressed; bark gray-brownish,
rough, lenticellate, shallowly fissured; slash of trunk light
yellow (Fig. 1U); inner bark non-sweet tasting. Leaves
subverticillate, 8–22 × 5–8 cm, oblanceolate or obovate, coria-
ceous, upper surface glabrous, lower surface glabrous or
indument if present, restrict to the midvein or secondary
veins; midvein sunken on upper surface; secondary venation
eucamptodromous, parallel, straight or slightly arcuate, slightly
impressed above; intersecondaries absent; tertiaries oblique
(Fig. 1J); petiole 0.9–2.0 cm long, canaliculate in the upper
half, tomentulose or glabrescent, without scales. Flowers 10–
many in each fascicle, ramiflorous or axillary; pedicel 1.6–
3.0 mm long, tomentulose. Sepals are tomentulose outside.
Corolla is 4.5–5.5 mm long, wine red, sericeous outside (Fig. 2F).
Fruit 3.0–4.5 cm long, ellipsoid, yellow-orange, muricate, and
brown-tomentulose (Fig. 2J, 2L).
Distribution and habitat — Pradosia granulosa is known
from northeastern Amazon, occurring in the states of Goiás,
Maranhão and Pará in Brazil (Fig. 4D). It grows in open low-
land forests on sandy soils (campina forest) or occasionally
on clay, from sea level to 250 m altitude.
Recognition — This species is easily distinguished from
all other Amazonian species by its muricate fruits.
Pradosia granulosa is sister of P. brevipes and P. glaziovii in
the Red-flowered clade (Fig. 3), and apart from the mor-
phology, also has other ecological preferences, growing in
campina forests or occasionally on clay, not in Cerrado as
P. brevipes or in wet lowland tall forests on clayish soils as
P. glaziovii.
Pradosia grisebachii (Pierre) T. D. Penn., Fl. Neotrop.
Monogr. 52: 655. 1990. ≡ Ecclinusa grisebachii Pierre, Not.
Bot. Sapot. 57. 1891.—TYPE: TRINIDAD AND TOBAGO.
Trinidad: Versimmiliter ex insula Trinitatis et ex Cruegeri
colectione, Aug 1860 (fl), H. Crüger 157 (holotype: GOET
010955!; isotypes: GOET 010956 [fragment]!, K
000640458!, NY 0273462 [fragment]!, P 00649446!).
Pradosia atroviolacea Ducke, Trop. Woods 90: 25. 1947.––
TYPE: COLOMBIA. Amazonas: Leticia, mata de t.
firme, baixa ao Nordeste, 3 Nov 1945 (fl), A. Ducke 1800
(lectotype chosen by Pennington [1990: 653]: RB
00380742!; isolectotypes: A 00460602! CAS 214950!,
F V0072205F!, GH 00075855!, IAN!, K 000640438! and
K 000640439!, MG!, NY 00273667! and NY 00273668!,
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA 645

R 000075072!, RB 00635954!, US 00113358!, US 00323586!, chartaceous, glabrous; midvein flat and slightly raised on
and US 00930768!). syn. nov. upper surface; secondary venation eucamptodromous, diver-
gent and slightly raised above; intersecondaries absent; ter-
Tree up to 15 m tall; buttressed; bark grayish, smooth,
tiaries oblique-reticulate; petiole 0.5–1.5 cm long, semiterete,
sloughing off in thin plates and leaving dark marks; slash
glabrous, without scales. Flowers 2–5 in each fascicle,
of trunk unknown; inner bark non-sweet tasting. Leaves
ramiflorous; pedicel 1.0–4.0 mm long, tomentulose. Sepals
alternate or subverticillate, 8–22 × 4–8 cm, oblanceolate or
are tomentulose outside. Corolla is 4.5–5.5 mm long, green-
obovate, chartaceous to coriaceous, upper surface glabrous,
ish, glabrous outside. Fruit 2.5–4.5 cm long, obovoid to ellip-
lower surface glabrous or puberulous; midvein sunken on
soid, greenish, smooth, glabrous.
the upper surface; secondary venation eucamptodromous,
Distribution and habitat — Pradosia huberi occurs in north-
parallel, slightly impressed above; intersecondaries absent;
eastern Amazon, in the states of Amapá and Pará in Brazil
tertiaries oblique to horizontal; petiole 0.8–2.0 cm long,
and nearby Cayenne in French Guiana (Fig. 4B). In spite of
canaliculate in the upper half, glabrous; scales present,
the low number of recent collections this species is believed
paired, 1.0–2.0 mm long, fixed at the midway of the petiole.
to be a characteristic member of swampy lands or flooded
Flowers 2–many in each fascicle, ramiflorous; pedicel 1.0–
forests along the coast (estuarine areas) and associated with
3.0 mm long, tomentulose. Sepals are tomentulose outside.
riverine systems (igarapé or furo) (Ducke 1953).
Corolla is 4.0–5.0 mm long, reddish, sparsely sericeous out-
Recognition — Pradosia huberi is similar in morphology to
side. Fruit 3.5–5.0 cm long, ellipsoid, yellowish, smooth, and
P. cochlearia, but is readily recognized on its semiterete peti-
sparsely puberulous.
ole (vs. slightly canaliculate), divergent secondary veins
Distribution and habitat — Widespread species from south-
(vs. parallel), larger corolla (vs. 1.7–3.0 mm) and that it occur
western Amazon, Acre, Brazil to northwestern Amazon in
in swampy or flooded forests (vs. non-flooded forests, usu-
Colombia, reaching the north of Venezuela and Costa Rica
ally on sandy soils).
(Fig. 4C). Pradosia grisebachii grows in lowland non-flooded
forests on clayish soils from 100–700 m alt. Pradosia kuhlmannii Toledo, Arq. Bot. Estado São Paulo
Note — Hermman Crüger collection was used by Pierre 2: 29. 1946.—TYPE: BRAZIL. Rio de Janeiro: Mata
(1891) to describe Ecclinusa grisebachii, a species later trans- do morro do Jardim Botânico, 22 May 1926 (fl, fr),
ferred to Pradosia by Pennington (1990). In the herbarium of D. Constantino & P. Occhioni [RB22231] (lectotype chosen
the Universität Göttingen (GOET), there are two specimens, by Pennington [1990: 651]: RB 00560363!; isolectotypes:
GOET 010955 and GOET 010956, of which only the first is a IAN!, NY 00375787!, P 00649448!, RB 00560364!, S 05–
good sample bearing leaves and flowers while the second 10534!, U 0006713!, US 00323588!).
one has only fragments of leaves, flowers, and a pencil
Tree up to 15 m tall; buttresses well developed; bark
drawing of a flower, which agree with the diagnosis of
smooth, grayish, scaling in thin irregular plates leaving deep
Pierre. In accordance with article 9 of the International Code
orange to brown patches; slash of trunk reddish; inner bark
of Nomenclature (McNeill et al. 2012), we believe that Crüger
sweet-tasting (Fig. 1P). Leaves alternate, 4–11 × (1.4–)2–4 cm,
157 (GOET 010955) constitutes the holotype.
elliptic or oblanceolate, chartaceous, glabrous; midvein flat
Recognition — We unite here Pradosia grisebachii and
on the upper surface; secondary venation brochidodromous,
P. atroviolacea based on morphological similarities. These
parallel, not impressed above; intersecondaries present,
species are the only Pradosia having scales on the petiole
short; tertiaries horizontal-reticulate; petiole 0.8–2.0 cm long,
combined with ramiflorous fascicles. They have been distin-
semiterete, glabrous, without scales. Flowers 3–6 in each fas-
guished by the presence or absence of trichomes on the
cicle, ramiflorous or axillary; pedicel 0.5–1.0 mm long,
lower leaf surface, petiole length, and the number of flowers
tomentulose. Sepals are tomentulose outside. Corolla is 3.0–
in their fascicles (Pennington 1990). In general, P. grisebachii
4.0 mm long, greenish, glabrous outside. Fruit 2.5–3.5 cm
is known from Trinidad and Venezuela, whereas P. atrovio-
long, ellipsoid, greenish, smooth, and glabrous.
lacea occurs in western Amazonia to Costa Rica and Panama.
Distribution and habitat — Pradosia kuhlmannii occurs
Studies of the types and many new collections covering the
along the Brazilian Atlantic coast, from Pernambuco in the
geographic range of both species cannot reveal any clear
north to Rio de Janeiro State in the south, and always in wet
morphological discontinuities between them. The above-
lowland tall forests on clayish soils (Fig. 4B).
mentioned morphological characters used by Pennington
Recognition — Sterile specimens of Pradosia kuhlmannii
(1990) to distinguish them do not clearly separate the material
are easy to recognize because of its sweet tasting bark,
in two distinct morphological groups. We consider it therefore
semiterete petioles, brochidodromous venation, flat midvein,
appropriate to unite P. atroviolacea with P. grisebachii.
and horizontal tertiaries. Other members of the Sweet-bark
clade have similar character combinations, but occur in
Pradosia huberi (Ducke) Ducke, Trop. Woods 71: 16. 1942. ≡
rainforests on sandy soils in the Amazon. On the other hand,
Glycoxylon huberi Ducke, Arch. Jard. Bot. Rio de Janeiro
P. kuhlmannii grows on clayish soils in wet rainforests and is
3: 253. 1922.–– TYPE: BRAZIL. Pará: Furo Macujubim,
sympatric with P. lactescens, a species with which it has been
Ilhas de Breves, mata alagada, 16 Nov 1923 (fl, fr),
confused (see Kuhlmann 1930; Toledo 1946). However,
A. Ducke [RB3782] (lectotype chosen by Pennington
P. lactescens differs in an astringent inner bark, eucampto-
[1990: 657]: RB 00560333!; isolectotypes: G 00439601!,
dromous leaf venation, canaliculate petiole, cauliflorous
IAN!, K 000640447!, MG!, P 00649447!, S 05–10587!,
fascicles, and larger and wine-reddish flowers.
U 0006712!).
Tree up to 30 m tall; large buttresses to 2 m tall; bark Pradosia lactescens (Vell.) Radlk., Sitzungsber. Math.-
smooth, grayish; slash of trunk unknown; inner bark sweet- Phys. Cl. Königl. Bayer. Akad. Wiss. München 18: 407.
tasting. Leaves opposite, 5–15 × 3–4 cm, elliptic or obovate, 1888. ≡ Pometia lactescens Vell., Fl. Flum. 81, 2: Table 87.
646 SYSTEMATIC BOTANY
1825.—TYPE: BRAZIL: J. M. Vellozo s.n. (lectotype cho-
sen by Pennington [1990: 650]: P 00649449!).
Tree up to 15 m tall; buttresses poorly developed; bark
reddish-brown, scaling in thin plates; slash of trunk orange;
inner bark non-sweet tasting (Fig. 1Q). Leaves alternate,
5–20 × 2–6 cm, elliptic to obovate, chartaceous, glabrous;
midvein sunken on the upper surface; secondary venation
eucamptodromous, parallel, slightly sunken above; inter-
secondaries absent; tertiaries oblique; petiole 0.6–1.8 cm
long, canaliculate, glabrous or with residual appressed
indument, without scales. Flowers usually many in each fas-
cicle, cauliflorous; pedicel 4.0–6.0 mm long, tomentulose.
Sepals are tomentulose outside. Corolla is 4.8–6.5 mm long,
wine red, usually glabrous (Fig. 2E). Fruit 3.0–4.5 cm long,
obovoid to ellipsoid, yellow-orange, smooth, and glabrous
(Fig. 2H).
Distribution and habitat — Pradosia lactescens is a very
common and widespread species along the whole range of
the Brazilian Atlantic coast rainforest, from Pernambuco in
the north to Paraná State in the south. It grows in wet low-
land tall forests on clayish soils from sea level to 980 m alti-
tude (Fig. 4D).
Recognition — Pradosia lactescens is in overall morphology
most similar to P. ptychandra. These two species are cauli-
florous, have similar leaf shapes, and wine red flowers.
However, they are not closely related and are best distin-
guished by flower features (e.g. pedicel and corolla length),
which are larger in P. ptychandra. Moreover, they are allopat-
ric: P. lactescens is restricted to the Brazilian Atlantic coast,
while P. ptychandra occurs in Guyana, Suriname, and French
Guyana. On the other hand, P. lactescens has been historically
confused with P. kuhlmannii, a species also restricted to the
Brazilian Atlantic coast region. In part, the confusion came
from a swap of samples committed by Casaretto (Toledo
1946). The diagnosis was based on two different samples:
leaves of Pradosia lactescens and bark of P. kuhlmannii, a mis-
take highlighted by Toledo (1946) and which was dealt with
subsequently with new descriptions.
In the southern coast of the state of Rio Grande do Norte
in Brazil, P. lactescens could be confounded with P. restingae.
Pradosia lactescens is best recognized in having decurrent leaf
bases, being cauliflorous with 4.0–6.0 mm pedicellate, wine
reddish flowers, and growing in forests on clayish soils. In
contrast, P. restingae has cuneate leaves, being ramiflorous
with subsessile, greenish flowers, and is only known from
white-sand forests (Terra-Araujo et al. 2013).
Pradosia lahoziana Terra-Araujo, Brittonia, 64: 142.
2012.—TYPE: BRAZIL. Amazonas: Manaus, Estrada do
Aleixo, grounds of Companhia das Plantações, forest on
terra firme [03°05′ S, 59°55′ W, 50–94 m], 30 Aug 1973
(fl), G. T. Prance 18763 (holotype: INPA!; isotypes: MG!,
MO!, NY 01171401!).
Tree up to 15 m tall; unbuttressed; bark rough, lenticellate,
grayish-brown; slash of trunk light orange; inner bark non-
sweet tasting. Leaves alternate or subverticillate, 9–20 × 3–
7 cm long, oblanceolate or elliptic, chartaceous, upper
surface glabrous, lower surface sparsely tomentulose, at least
on the venation (visible with a hand-lens); secondary vena-
tion eucamptodromous; midvein sunken on the upper sur-
face; secondary veins parallel, slightly raised or flat above,
usually tomentulose, glabrescent; intersecondaries absent;
[Volume 41
tertiaries horizontal or oblique; petiole 1.6–2.6 cm long,
canaliculate, tomentulose; scales present, attached below the
blade to the middle of the petiole, 1.0–2.0 mm long, paired
(Fig. 1G). Flowers 5–10 in each fascicle, cauliflorous; pedicel
7.0–9.0 mm long, tomentulose. Sepals are tomentulose
outside. Corolla is 6.4–9.5 mm long, wine red, glabrous
outside. Fruit 2.5–3.0 cm long, obovoid, yellowish, smooth,
and tomentulose.
Distribution and habitat — Pradosia lahoziana is known
from central Amazon, where it occurs in forests north of
Manaus, in the permanent plots of the Dinâmica Biológica
de Fragmentos Florestais Project, Dimona Reserve and from
the Estação Experimental de Sivicultura Tropical-ZF2, both
from the Instituto Nacional de Pesquisas da Amazônia
(Fig. 4C). It grows in rainforest on sandy (e.g. campina and
campinarana forests) or occasionally clayish soils from
50–125 m alt.
Recognition — This species has a unique combination of
characters in the genus: having scales on the petiole below
the blade and flowers on the stem (cauliflorous). Similar
scales are present in Pradosia grisebachii, a ramiflorous spe-
cies. Other cauliflorous species such as P. ptychandra and
P. lactescens lack these scales.
Pradosia longipedicellata Alves-Araújo & M. Alves,
Brittonia 64: 24. 2012.—TYPE: BRAZIL. Bahia: Una,
Reserva Biológica do Mico-Leão (IBAMA), BA-001 Km
46, 15°09′S, 39°05′W, 9 Mar 1993 (fl, fr), J. G. Jardim et al.
92 (holotype: CEPEC!; isotype: NY 00585395!).
Tree up to10 m, but can reach 20 m tall; unbuttressed or
occasionally with buttresses to 0.5 m tall; bark grayish,
rough, lenticellate, shallowly fissured, scaling; slash of trunk
pinkish; inner bark non-sweet tasting (Fig. 1V). Leaves clus-
tered, alternate, 6–17 × 3–7 cm, ovate to elliptic, chartaceous,
both leaf surfaces covered by whitish to ferruginous tri-
chomes, glabrescent; secondary venation eucamptodromous;
midvein flat; secondary vein parallel, flat or slightly raised;
intersecondaries absent; tertiaries oblique-reticulate (Fig. 1I);
petiole 0.4–1.6 cm, semiterete, with whitish to ferruginous
trichomes, without scales. Flowers 2–8 in each fascicle, termi-
nal; fascicles in clusters, umbelliform; pedicel 18–30 mm
long, tomentulose. Sepals are tomentulose outside. Corolla is
7.0–8.0 mm long, white, glabrous (Fig. 2A). Fruit 3.0–4.0 cm
long, ellipsoid, smooth, and brown-puberulous.
Distribution and habitat—Pradosia longipedicellata is restricted
to the coastal region of southern Bahia, Brazil, and is found in
white-sandy restinga forests, from sea level to around 60 m
altitude (Fig. 4A).
Recognition — Sterile specimens of Pradosia longipedicellata
are easily distinguished from all other Brazilian Atlantic
coast species by its leaves covered by whitish to ferruginous
trichomes and flat midvein. If fertile specimens are available,
it is readily distinguished on dense, terminal inflorescences
with long pedicels (18–30 mm) and white flowers.
Pradosia montana T. D. Penn., Fl. Neotrop. Monogr. 52:
661. 1990.—TYPE: ECUADOR. Province El Oro, ca.
60 km SE of Arenillas on road to Loja, 13 Nov 1982 (fl), T.
D. Pennington & G. Tenorio 10719 (holotype: K 000640437!;
isotypes: MO 1991078, QCA 100610!, QCNE 735!).
Tree up to 8 m, but can reach 25 m tall; unbuttressed or
with slight buttress to 0.5 m tall; bark light gray, lenticellate,
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA
rough, fissured, sloughing off in small plates; inner bark
non-sweet tasting; slash of trunk pinkish. Leaves alternate or
subverticillate, 5–15 × 2–7 cm, obovate or elliptic, coriaceous,
upper surface sparse pubescent on midvein and secondary
veins, lower leaf surface with indument of erect two-
branched pale-brownish trichomes; midvein sunken on the
upper surface; secondary venation eucamptodromous, paral-
lel, slightly impressed above; intersecondaries absent; tertia-
ries oblique to horizontal; petiole 0.5–1.3 cm long, slightly
canaliculate, tomentulose, without scales. Flowers 5–10 in
each fascicle, ramiflorous; pedicel 2.0–3.0 mm long, tomen-
tulose. Sepals are tomentulose outside. Corolla is 3.0–4.0 mm
long, greenish, glabrous or sparsely tomentulose outside,
especially the corolla tube (Fig. 2C). Fruit 1.5–2.5 cm long,
obovoid or ellipsoid, smooth, and densely brown-pubescent.
Distribution and habitat — This species is restricted to
western Ecuador and northwest corner of Peru, being con-
sidered a common component of tropical deciduous forests
from sea level to 720 m altitude (Fig. 4A).
Recognition — Pradosia montana is readily distinguished
from all other species of Pradosia by the soft pubescence on
the lower leaf surface. There are two other species associated
with deciduous forests, P. colombiana and P. mutisii, but they
have glabrous or glabrescent leaves.
Pradosia mutisii Cronquist, Bull. Torrey Bot. Club 73: 470.
1946.—TYPE: COLOMBIA. (fl), J. C. Mutis 4004 (holo-
type: US 00037017!; isotype G 00439600 [fragment]!).
Tree up to 8 m, but can reach 30 m tall; buttress up to
0.5 m tall; bark dark brown; slash of trunk unknown. Leaves
clustered, alternate, 5–14 × (2.5–)3–6 cm, obovate or elliptic,
chartaceous, upper surface glabrous, lower surface glabrous
or with sparse short whitish trichomes; midvein sunken on
the upper surface; secondary venation eucamptodromous,
parallel, not impressed above; intersecondaries present
and extending towards the margin; tertiaries horizontal-
reticulate; petiole 1.1–2.2 cm long, canaliculate, glabrous,
without scales. Flowers 2–10 in each fascicle, ramiflorous;
pedicel ca. 1 mm long, glabrescent. Sepals are glabrescent
outside. Corolla is 2.5–3.0 mm long, greenish, glabrescent
outside. Fruit ca. 2 cm long, ellipsoid, greenish, smooth,
and glabrous.
Distribution and habitat — Pradosia mutisii occurs from
northwestern Colombia to northwest Peru. It is a rare spe-
cies found in tropical deciduous forest from 180–1200 m
alt. (Fig. 4A).
Recognition — This species can be mistaken for P. montana,
and their ranges overlap in the northwest corner of Peru.
However, P. mutisii is easily distinguished from P. montana
by its glabrous leaves (vs. pubescent), presence of long
intersecondaries veins (vs. absent), and glabrous fruits
(vs. pubescent).
Pradosia ptychandra (Eyma) T. D. Penn., Fl. Neotrop.
Monogr. 52: 648. 1990. ≡ Pouteria ptychandra Eyma,
Recueil Trav. Bot. Néerl. 33: 189. 1936.—TYPE: SURINAM.
Wilhelminagebergte, Eindkamp Lucie rivier, 10 Apr
1926 (fl), G. Stahel [B. W. 6943] (holotype: U 0006715!;
isotypes: G 00439599 [fragment], K 000640453 [frag-
ment]!, U 0006716!).
Tree up to 13 m, but can reach 25 m tall; unbuttressed or
with poorly developed buttress to ca. 30 cm tall; bark
647
smooth, grayish-brown, scaling in thin plates and leaving
pock-like depressions; slash of trunk cream with orange
streaks; inner bark non-sweet tasting. Leaves clustered, alter-
nate, 7–22 × 2–8 cm long, elliptic, oblanceolate or obovate,
glabrous; midvein sunken on the upper surface; secondary
venation eucamptodromous, parallel, slightly sunken above;
intersecondaries absent; tertiaries oblique; petiole 0.6–2.2 cm
long, canaliculate, glabrous or with appressed indument,
without scales. Flowers usually many in each fascicle,
cauliflorous, less frequently ramiflorous; pedicel 7.0–10.5 mm
long, tomentulose. Sepals are tomentulose outside. Corolla is
5.0–5.5 mm long, wine red, sparsely tomentulose outside.
Fruit 2.5–4.0 cm long, ovoid to ellipsoid, yellow or orange,
smooth, and glabrous.
Distribution and habitat — This species has a disjunct dis-
tribution, in central Amazon in the surroundings of Manaus,
and also known from several collections in French Guiana,
Guyana and Surinam (Fig. 4C). Pradosia ptychandra grows in
non-flooded forests on clayish soils, from 75–770 m alt.
Recognition — Pradosia ptychandra is most similar to
P. lasctescens, a species of the Atlantic coast, and are best sep-
arated on flower features (see under P. lactescens for further
discussion). It could also be confused with P. lahoziana, a
closely related species, also having wine-red flowers borne
along the trunk, but P. ptychandra lacks scales on the petiole
(present in P. lahoziana).
Pradosia restingae Terra-Araujo, Nord. J. Bot. 31: 437.
2013.—TYPE: BRAZIL. Rio Grande do Norte: Pipa,
estrada para o Santuário Ecológico, 06°13′43.7″S, 35°03′
31.7″W, 55 m alt., 26 Sep 2011 (fl, fr), A. Alves-Araújo
et al. 1373 (holotype: UFP!, isotypes: INPA!, RB, S!,
UFRN!).
Tree about 2–6 m, sometimes up to 9 m tall; slash of trunk
unknown. Leaves alternate or subverticillate, (7–)8–13 × 3–
4 cm, obovate or elliptic, chartaceous, glabrous; midvein
sunken on the upper surface; secondary venation eucampto-
dromous, parallel, slightly sunken above; intersecondaries
absent; tertiaries, visible with lens are oblique or horizontal;
petiole 0.6–0.8 cm long, canaliculate, tomentulose, without
scales. Flowers 2–10 in each fascicle, ramiflorous, usually not
on the trunk; pedicel 0.8–1.6 mm long, tomentulose. Sepals
are tomentulose outside. Corolla is 4.8–5.5 mm long, green-
ish with a flesh colored ring, tomentulose outside, especially
the corolla tube and central part of the lobes (Fig. 2D). Fruit
2.5–3.7 cm long, ovoid, yellowish, smooth, and glabrous
(Figs. 2G, 2M).
Distribution and habitat — Pradosia restingae is only
known from the southern coast of the state of Rio Grande
do Norte in Brazil, where it is found in both forested
(restinga alta) and open vegetation on white-sand soils
(dunas), from sea level to 55 m altitude (Fig. 4D).
Recognition — This species can be confused with P. lactescens;
see under P. lactescens for discussion.
Pradosia schomburgkiana (A. DC.) Cronquist, Bull. Torrey
Bot. Club 73: 311. 1946. ≡ Chrysophyllum schomburgkianum
A. DC. in Candolle A. L. P. P. de, Prodr. 8: 157. 1844.—
TYPE: GUYANA, 1838 (fl), M. R. Schomburgk 505 (lecto-
type chosen by Pennington [1990: 659]: G 00139569!;
isolectotypes: BM 000952571!, BR 0000005415137!,
E 00259450!, F V0071946F!, G 00439595 and G 00439596,
648 SYSTEMATIC BOTANY [Volume 41

K 000640445! and K 000640446!, NY 902225!, OXF, P ondary venation eucamptodromous, parallel, impressed
00649452!, US 00113129!). above; intersecondaries absent; tertiaries oblique, obscure;
petiole 2.0–3.2 cm long, canaliculate, tomentulose, without
Shrub or small tree up to 4 m, but can reach 20 m tall;
scales. Flowers usually many in each fascicle, usually rami-
unbuttressed or occasionally with large buttresses to ca. 1 m
florous but sometimes axillary; pedicel 4.0–7.0 mm long,
tall; bark grayish, smooth, scaling and leaving orange-brown
tomentulose. Sepals are tomentulose outside. Corolla is 5.5–
patches; slash of trunk reddish; inner bark sweet-tasting
6.5 mm long, greenish, sparsely sericeous outside. Fruit 3.5–
(Fig. 1R). Leaves opposite (Fig. 1F), 4–18 × 2–7 cm, oval
4.0 cm long, ellipsoid, smooth, and glabrescent.
(broadly elliptic) to obovate, coriaceous, usually glabrous on
Distribution and habitat — Pradosia subverticillata occurs
both sides, sometimes sparsely ferruginous tomentulose
from the lower Rio Negro basin near Manaus to upper Rio
below; midvein flat; secondary venation brochidodromous,
Negro around Barcelos and Issana rivers in the Upper Rio
parallel, flat or slightly raised on upper leaf surface; inter-
Negro region (Fig. 4C). Its distribution further seems to
secondaries long, extending towards the margin; tertiaries
extend to southern Colombia, in La Uribe, Meta (M. Gaitán
horizontal-reticulate; petiole 0.4–1.5 cm long, semiterete, gla-
155, COAH). However, we have not been able to examine
brous or with appressed indument, without scales. Flowers
voucher material from this area. In Brazil, along Rio Negro,
5–many in each fascicle, ramiflorous; pedicel 2–10 mm long,
it grows in campina and campinarana forests or occasionally
tomentulose to glabrous. Sepals are tomentulose to glabrous
on lowland non-flooded forests on clayish soils near Manaus,
outside. Corolla is 2.0–4.0 mm long, greenish, sparsely
in central Amazon.
tomentulose outside, especially the corolla tube (Fig. 2B).
Recognition — If Pradosia subverticillata is compared with
Fruit 1.0–2.0 cm long, obovoid to ellipsoid, greenish, smooth,
other species in the Amazon region, its morphology is most
glabrous or brown-puberulous.
similar to P. surinamensis. These two species have a sweet-
Distribution and habitat—Pradosia schomburgkiana is widely
tasting inner bark, glabrous leaves, sunken midvein above,
distributed in the Amazon region, being found in campina,
and greenish flowers, and they grow in forests on sandy
campinarana or restinga forests, from sea level in Pará State
soils. However, P. subverticillata differ from P. surinamensis by
up to 1200 m altitude in Guiana Shield (Fig. 4B).
the secondaries veins deeply impressed above and the larger
Recognition — A sweet-tasting bark combined with an
flowers, 5.5–6.5 mm long, in contrast to 2.0–3.0 mm in
opposite leaf arrangement and a brochidodromous leaf
P. surinamensis. The grayish, scaling, and sweet-tasting bark,
venation readily distinguish Pradosia schomburgkiana from
however, increase the risk to confound P. subverticillata with
all other congeners. It furthers have long intersecondaries
P. cochlearia and P. schomburgkiana, two species of the Sweet-
that extend towards the margin, a rather unusual characters
bark clade that also grow on sandy soils. The midvein is the
for Pradosia.
best vegetative character to separate these species, being
Pennington (1990) recognized two subspecies of Pradosia
sunken in P. subverticillata and raised in P. cochlearia and
schomburgkiana (subsp. schomburgkiana and subsp. sericea),
P. schomburgkiana.
which are sympatric in the Serra do Aracá region. The sub-
species are distinguished by the presence and absence,
Pradosia surinamensis (Eyma) T. D. Penn., Fl. Neotrop.
respectively, of a dense indument on the lower leaf surface.
Monogr. 52: 652. 1990. ≡ Pouteria surinamensis Eyma,
Both subspecies were included in a recent phylogenetic anal-
Recueil Trav. Bot. Néerl. 33:189. 1936.—TYPE: SURINAM.
ysis using molecular data, but there was no phylogenetic
Boven-Suriname Rivier bij Goddo, 23 Jan 1926 (fl),
signal (Terra-Araujo 2013) supporting subspecies, or correla-
G. Stahel 39 (holotype: U 0006719!; isotypes: G 00439591,
tion between morphological variation, habitat or geography.
K 000640450!, MO 345916!, RB 00544069!, U 0006720! and
Furthermore, both subspecies have the same ecological pref-
U 0006721!).
erences and grow in campina and campinarana forests.
Species of Sapotaceae in the Neotropics are frequently
Tree up to 15 m, but can reach 30 m tall; buttressed 0.30–
poorly collected and their full morphological variation and
1.0 m tall; bark grayish, smooth, scaling and leaving orange
distribution range is therefore unclear. We propose that the
patches; slash of trunk light orange; inner bark sweet-tasting.
taxon limits within P. schomburgkiana requires a closer exami-
Leaves spaced, loosely clustered, alternate, 4–16 × 2–6 cm,
nation to better understand whether it contains more than
elliptic or obovate, coriaceous, upper surface glabrous, lower
one species.
surface glabrous or glabrescent; midvein sunken on the
upper surface; secondary venation eucamptodromous, slightly
Pradosia subverticillata Ducke, Trop. Woods 71: 13.
impressed above; intersecondaries absent; tertiaries oblique-
1942.—TYPE: BRAZIL. Amazonas: Manaus, Rio Tarumã,
reticulate, obscure; petiole 0.5–1.9 cm long, canaliculate,
cachoeira baixa, mata de t. f. em solo arenoso, 27 Jun
tomentulose, without scales. Flowers 1–4 in each fascicle,
1941 (fl), A. Ducke 812 (lectotype chosen by Pennington
ramiflorous; pedicel 0.5–2.0 mm long, tomentulose. Sepals are
[1990: 646]: RB 00544076!; isolectotypes: B 10 0413399!,
sparsely tomentulose outside. Corolla is 2.0–3.0 mm long,
F 0072208F!, IAN!, K 000640456!, MG!, MO 1991009!,
greenish, with sparsely tomentulose tube. Fruit 2.0–4.0 cm
NY 00273673! and NY 01168598!, R 000075081!, RB
long, obovoid to ellipsoid, yellowish, smooth, not lenticellate,
00642343!, US 00113361! and US 00323587!).
glabrous or puberulous (Fig. 2I, 2N).
Tree up to 4 m, but can reach 20 m tall; bark grayish, Distribution and habitat — This species is known from
smooth, scaling and leaving irregular brown deep patches; northeastern Amazonia, in Roraima and Pará State, Brazil,
slash of trunk yellowish; inner bark sweet-tasting (Fig. 1S). Guyana and Surinam where it is found in lowland
Leaves clustered at branch tips, alternate or subverticillate, non-flooded forest from 60–600 m. alt. (Fig. 4D). In
10–21 × 3–8 cm, oblanceolate or obovate, coriaceous, both Brazil, it has been collected usually on sandy soils
surfaces glabrous; midvein sunken on the upper surface; sec- (campinarana forest).
2016] TERRA-ARAUJO ET AL.: UPDATE OF PRADOSIA
Recognition — Pradosia surinamensis belongs to the Red-
flowered clade and is morphologically similar to P. subver-
ticillata. Its relationships with P. subverticillata are discussed
under the latter.
Pradosia verticillata Ducke. Trop. Woods 71: 12. 1942.—
TYPE: BRAZIL. Amazonas: Manaus, Estação do Aleixo
além do V. Municipal, restos de mata de terra firme,
argilosa, 14 Oct 1941 (fl, fr), A. Ducke 811 (lectotype cho-
sen by Pennington [1990: 664]: RB 00379868!; iso-
lectotypes: F 0072209F!, IAN!, K 000640440!, MG!, MO
1991017, NY 01365138!, R 000075080!, RB 00635941!
and RB 00635944!, US 00113362!, US 00930762! and
US 00930763!).
Tree up to 15 m, but can reach 30 m tall; buttresses up to
0.5 m tall; bark greenish, smooth, scaling and leaving brown
deep patches; slash of trunk orange; inner bark non-sweet
tasting (Fig. 1T). Leaves clustered at branch tips, verticillate
(Fig. 1E), 13–31 × 4–11 cm, oblanceolate, coriaceous, upper
surface glabrous or with some residual indument, lower sur-
face brown-pubescent, denser on the venation; midvein
sunken on the upper surface; secondary venation eucampto-
dromous, parallel, impressed above; intersecondaries absent;
tertiaries oblique; petiole 1.0–4.1 cm long, canaliculate,
pubescent, without scales. Flowers 5–10 in each fascicle,
ramiflorous; pedicel 1.0–1.7 mm long, tomentulose. Sepals
are tomentulose outside. Corolla is 4.0–4.6 mm long, red-
dish, sericeous outside. Fruit 3.5–4.5 cm long, obovoid,
smooth, and glabrous.
Distribution and habitat — Pradosia verticillata is pres-
ently known as disjunct, with one population in the lower
Rio Negro basin near of Manaus in Brazil, and one in La
Fumée Mountain, French Guiana (Fig. 4D). It grows in
lowland non-flooded forest on clayish soils, from 59–
400 m alt.
Recognition — A very distinct species from all other con-
geners because of the verticillate clusters of leaves near the
branch apices, an impressed venation, and a dark brown
indument on the lower leaf surface.
Doubtful Taxon—
Pradosia argentea (Kunth) T. D. Penn., Fl. Neotrop. Monogr.
52: 660. 1990. ≡ Nycterisition argenteum Kunth in Humbolt,
Bonpland & Kunth. Nov. Gen. Sp. 3: 238. 1819.—TYPE:
PERU. [Crescit prope Jaen de Bracamoros (Regno Novo-
Granatensi), 320 m alt.] (fl), A. J. A. Bonpland & F. W. H. A.
von Humboldt s.n. (holotype: P 00670922!; isotype: G
00439608 [fragment]!).
Pradosia argenta is only known from the type collection,
dating back to 1819 in Peru, having flowers but no fruits.
Since it has never been relocated, P. argentea was treated as
extinct by the Red List in Peru (IUCN 2015). Pennington
(1990) in his monograph placed it in Pradosia because of
5-merous, rotate flowers with long-exserted stamens, and
lack of staminodes. It was further considered closely related
to P. schomburgkiana due to a brochidodromous venation
with long intersecondaries, but readily distinct from the lat-
ter in having alternate leaves with a fine silvery indument.
The overall flower morphology is coherent with Pradosia, but
since fruits and molecular data are unknown, it remains
unclear if P. argentea is a member of the genus.
649
Acknowledgments. This study was part of the Ph.D. thesis of M. H.
Terra-Araujo. The directors at the herbaria ALCB, CA, CEPEC, CVRD,
HRB, IAN, INPA, IPA, MBML, MG, NY, PEUFR, PH, RB, S, SP, UEFS and
US are acknowledged for permission to examine the herbarium material.
We thank Alberto Vicentini, Ana Andrade, Anderson Alves-Araújo,
Eduardo Prata, Fernanda A. Carvalho, Flávio Costa, Flávio Obermüller,
Jefferson Carvalho-Sobrinho, José Ribamar, Jomar Jardim, Nallarett
Dávila, Nory Daniel, Rafaela Forzza, Ricardo Perdiz, Saci and Xavier
Cornejo for their help in the field. This research was financed through a
scholarship to the first author by Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq, 143693/ 2008-5) and Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES, BEX 6161/ 11-1).
Literature Cited
Alves-Araújo, A. and M. Alves. 2012. Two new species and a new com-
bination of Neotropical Sapotaceae. Brittonia 64: 23–29.
Anderberg, A. A. and U. Swenson. 2003. Evolutionary lineages in
Sapotaceae (Ericales): A cladistic analysis based on ndhF sequence
data. International Journal of Plant Sciences 164: 763–773.
Anderson, A. B. 1981. White-sand vegetation of Brazilian Amazonia.
Biotropica 13: 199–210.
Aubréville, A. 1961. Notes sur les Sapotacées Africanes et Sud-
Américaines. Adansonia 1: 24–26.
Aubréville, A. 1964. Les Sapotace es taxonomie et phytogeographie.
Adansonia 1: 1–157.
Aubréville, A. 1967. Sapotacées nouvelles de la cote Colombienne du
Pacifique. Adansonia 7: 141–148.
Baehni, C. 1965. Mémoires sur les Sapotacées. 3. Inventairedes genres.
Boissiera 11: 1–262.
Cronquist, A. 1946a. Studies in the Sapotaceae V. The South American
species of Chrysophyllum. The Journal of the Torrey Botanical Society
73: 286–311.
Cronquist, A. 1946b. Studies in Sapotaceae VI. The Journal of the Torrey
Botanical Society 73: 465–471.
de Queiroz, K. 1998. The general lineage concept of species, species
criteria, and the process of speciation: A conceptual unification and
terminological recommendations. Pp. 57–75 in Endless forms: Species
and speciation, eds. D. J. Howard and S. H. Berlocher. Oxford, UK:
Oxford University Press.
de Queiroz, K. 2007. Species concepts and species delimitation. System-
atic Biology 56: 879–886.
Ducke, A. 1922. Plantes nouvelles ou peu connues de la région
amazoniènne, Sapotaceae. Archivos do Jardim Botânico do Rio de
Janeiro 3: 230–238.
Ducke, A. 1925. Plantes nouvelles ou peu connues de la région
amazonienne III. Archivos do Jardim Botânico do Rio de Janeiro 4:
158–166.
Ducke, A. 1942. New and noteworthy Sapotaceae of Brazilian Amazonia.
Tropical Woods 71: 7-25.
Ducke, A. 1953. As espécie brasileiras do gênero Pradosia Liais
(Fam. Sapotaceae). Boletim Técnico do Instituto Agronômico do Norte
28: 21-34.
Ellis, B., D. C. Daly, L. J. Hickey, K. R. Johnson, J. D. Mitchell, P. Wilf, and
S. L. Wing. 2009. Manual of leaf architecture. Ithaca, New York:
The New York Botanical Garden Press and Comstock Publishing
Associates.
Eyma, P. J. 1936. Notes on Guiana Sapotaceae. Recueil des Travaux
Botaniques Néerlandais 33: 156–210.
Ewango, C. E. N. and F. J. Breteler. 2001. Présence du genre Pradosia
(Sapotaceae) en Afrique: description d’une nouvelle espèce,
P. spinosa. Adansonia 23: 147–150.
Harris, J. G. and M. W. Harris. 2001. Plant identification terminology: An
illustrated glossary. Spring Lake, Utah: Spring Lake Publishing.
Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index
Herbariorum. Part I: The herbaria of the world. Eighth edition. New
York: New York Botanical Garden Press.
IUCN. 2015. IUCN red list of threatened species. Version 2015–3. <www
.iucnredlist.org>. Downloaded on 9 September 2015.
Kuhlmann, J. G. 1930. Contribuição para o conhecimento de algumas
novas espécies da região amazônica e uma do Rio de Janeiro, bem
como algumas notas sobre espécies já conhecidas. Arquivos do Jardim
Botânico do Rio de Janeiro 5: 201–211.
Lecomte, H. 1923. Quelques espèces nouvelles de la famille des
Sapotacées. Notulae Systematicae 4: 62–64. (Paris).
650 SYSTEMATIC BOTANY
Liais, E. 1872. Climats, géologie, faune et géographie botanique du Brésil.
Paris: Garnier Frères.
McNeill, J., F. R. Barrie, W. R. Buck, V. Demoulin, W. Greuter, D. L.
Hawksworth, P. S. Herendeen, S. Knapp, K. Marhold, J. Prado, W. F.
Prud’Homme van Reine, G. F. Smith, J. H. Wiersema, and N. J.
Turland (eds.). 2012. International Code of Nomenclature for algae,
fungi, and plants (Melbourne Code): Adopted by the Eighteenth Inter-
national Botanical Congress Melbourne, Australia, July 2011. Regnum
Vegetabile 154. Königstein: Koeltz Scientific Books.
Miquel, F. A. W. 1863. Sapotae. Pp. 38–117 in Flora brasiliensis vol. 7, eds
C. F. P. Martius and A. W. Eichler. Leipzig & München: Frid. Fleischer.
Pennington, T. D. 1990. Flora Neotropica Monograph 52: Sapotaceae.
New York: New York Botanical Garden.
Pennington, T. D. 1991. The genera of Sapotaceae. Kew: Royal Botanic Garden.
Pennington, T. D. 2006. Flora da Reserva Ducke, Amazonas, Brasil:
Sapotaceae. Rodriguésia 57: 251–366.
Prance, G. T. 1996. Islands in Amazonia. Philosophical Transactions of the
Royal Society of London. Series B, Biological Sciences 351: 823–833.
Pierre, L. 1891. Notes botaniques, Sapotacées .II. Paris: Paul Klincksieck.
Prance, G. T. and H. O. R. Schubart. 1978. Notes on the vegetation of
Amazonia I. A preliminary note on the origin of the open white
sand campinas of the Lower Rio Negro. Brittonia 30: 60–63.
QGIS Development Team. 2014. QGIS geographic information system.
Open Source Geospatial Foundation Project. < http://qgis.osgeo.org >.
Accessed October 2014.
View publication stats
[Volume 41
Swenson, U. and A. A. Anderberg. 2005. Phylogeny, character evolu-
tion and classification of Sapotaceae (Ericales). Cladistics 21:
101–130.
Swenson, U., J. Munzinger, and I. V. Bartish. 2007. Molecular phylogeny
of Planchonella (Sapotaceae) and eight new species from New
Caledonia. Taxon 56: 329–354.
Swenson, U., J. E. Richardson, and I. V. Bartish. 2008. Multi-gene phylog-
eny of the pantropical subfamily Chrysophylloideae (Sapotaceae):
Evidence of generic polyphyly and extensive morphological homo-
plasy. Cladistics 24: 1006–1031.
Terra-Araujo, M. H., A. D. Faria, and A. Vicentini. 2012. A new species
of Pradosia (Sapotaceae) from Central Amazonia. Brittonia 64:
139–142.
Terra-Araujo, M. H. 2013. Phylogeny, biogeography and taxonomy of
Pradosia (Sapotaceae, Chrysophylloideae). Ph.D. dissertation. Manaus,
Amazonas: Instituto Nacional de Pesquisas da Amazônia.
Terra-Araujo, M. H., A. D. Faria, A. Alves-Araújo, and M. Alves. 2013.
A new species of Pradosia from the Atlantic forest, Brazil. Nordic
Journal of Botany 31: 437–441.
Terra-Araujo, M. H., A. D. Faria, A. Vicentini, S. Nylinder, and U.
Swenson. 2015. Species tree phylogeny and biogeography of
the Neotropical genus Pradosia (Sapotaceae, Chrysophylloideae).
Molecular Phylogenetics and Evolution 87: 1–13.
Toledo, J. F. 1946. Observações criticas sôbre nomes de algumas plantas
Brasileiras. Arquivos Botânicos do Estado de São Paulo 2: 29–31.