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Two new species and a new combination of Neotropical Sapotaceae


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This paper provides descriptions and illustrations of two new species of Sapotaceae (Pradosia longipedicellata and Chromolucuma apiculata) from Bahia, Brazil, a new combination (Chromolucuma congestifolia), and a new identification key for Chromolucuma. Pradosia longipedicellata is distinguished from other congeners by its clustered fascicles of flowers on pedicels 2–3 cm long at the apices of the shoots, and a corolla that is 7–8 mm long. Chromolucuma apiculata is distinguished by glabrous shoots and leaves, short petioles (10–15 mm long), and stipules 5–8 mm long with apiculate apices, while its close relative C. congestifolia has a lower leaf surface with trichomes usually restricted to the midrib and stipules 3–6 mm long with acute apices. Due to intense deforestation in the Amazonian and the Atlantic forests, preliminary IUCN Red List assessments are provided. Pradosia longipedicellata and Chromolucuma congestifolia are proposed the IUCN status Vulnerable, while the data for C. apiculata are still insufficient (Data Deficient).
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Two new species and a new combination of Neotropical Sapotaceae
Programa de Pós-Graduação em Biologia Vegetal PPGBV/UFPE, CEP 50670901, Recife-PE,
Brazil; e-mail:
Laboratório de Morfo-Taxonomia Vegetal, Departamento de Botânica, Universidade Federal de
Pernambuco UFPE, CEP 50670901, Recife-PE, Brazil
Abstract. This paper provides descriptions and illustrations of two new species of
Sapotaceae (Pradosia longipedicellata and Chromolucuma apiculata) from Bahia,
Brazil, a new combination (Chromolucuma congestifolia), and a new identication
key for Chromolucuma.Pradosia longipedicellata is distinguished from other cong-
eners by its clustered fascicles of owers on pedicels 23 cm long at the apices of the
shoots, and a corolla that is 78 mm long. Chromolucuma apiculata is distinguished
by glabrous shoots and leaves, short petioles (1015 mm long), and stipules 58mm
long with apiculate apices, while its close relative C. congestifolia has a lower leaf
surface with trichomes usually restricted to the midrib and stipules 36 mm long with
acute apices. Due to intense deforestation in the Amazonian and the Atlantic forests,
preliminary IUCN Red List assessments are provided. Pradosia longipedicellata and
Chromolucuma congestifolia are proposed the IUCN status Vulnerable, while the data
for C. apiculata are still insufcient (Data Decient).
Key Words: Brazil, Chrysophylloideae, conservation, diversity, Neotropics.
Resumo. São descritas e ilustradas duas novas espécies de Sapotaceae (Pradosia
longipedicellata e Chromolucuma apiculata), ocorrentes no estado da Bahia, Brasil,
além de uma nova combinação (Chromolucuma congestifolia) e uma nova chave de
identicação para Chromolucuma. Pradosia longipedicellata é reconhecida entre as
espécies do gênero pela presença de fascículos agrupados no ápice dos ramos, ores
longo-pediceladas (23 cm compr.) e corola 7-8 mm compr. Chromolucuma apiculata é
distinta pelos seus ramos e folhas glabros, pecíolos mais curtos (1015 mm long) e
estípulas (58 mm long) com ápices apiculados, enquanto que C. congestifolia tem face
abaxial foliar geralmente com tricomas restritos à nervura central e estípulas mais curtas
(36 mm long) com ápices agudos. Dada à intensa devastação das orestas Amazônica e
Atlântica, uma prévia classicação segundo as normas da IUCN Red List é proposta para
as espécies. Pradosia longipedicellata e Chromolucuma congestifolia são consideradas
como pertencentes ao status da IUCN como Vulneráveis, enquanto que dados sobre C.
apiculata ainda são insucientes para tal proposta (Dados Decientes).
Tropical America is one of the regions with
the highest species diversity of angiosperms in
the world (Mittermeier et al., 1999;Myerset
al., 2000), which also is where the sapodilla
family, Sapotaceae, has its greatest diversity
(Pennington, 2004). One of the most important
centers of diversication for the family in the
Neotropics is the Amazonian forest (Pennington
1991,2006; Anderberg & Swenson, 2003;
Swenson & Anderberg, 2005). However, other
biomes, such as the Atlantic forest, also show a
high species richness. The Atlantic forest, most
of which is in Brazil, is considered one of the
richest biogeographic regions in the world in
terms of plant species diversity and endemism,
but it has been heavily impacted by human
Brittonia, 64(1), 2012, pp. 2329 ISSUED: 1 March 2012
© 2011, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
development (Mittermeier et al., 1999;Myerset
al., 2000). According to Myers et al. (2000), the
Atlantic forest is a hotspot of diversity, and for
this reason it is considered one of the highest
priority areas for conservation. In most of
northeastern Brazil, the Atlantic forest is
restricted to a strip of vegetation along the
coast with an average elevation of up to 100 m
(Velloso et al., 1991).
Sapotaceae consists of 58 genera and about
1250 species in subtropical and tropical areas
of the world (Pennington, 1991; Govaerts et
al., 2001; Swenson et al., 2007a,b). For the
New World, Pennington (1990) monographed
the family and later provided a generic
classication for the family (Pennington,
1991). Recent phylogenetic studies have
demonstrated that Penningtons concepts of
genera do not always correspond with natural
groups (Anderberg & Swenson, 2003; Bartish
et al., 2005; Swenson et al., 2007a,2008).
Subsequently, Swenson and Anderberg
(2005) provided a new subfamilial classica-
tion, recognizing Chrysophylloideae, Sapo-
toideae, and Sarcospermatoideae, and also
indicated that genera such as Chrysophyllum
and Pouteria sensu Pennington (1990, 1991)
are polyphyletic (Bartish et al., 2005;Swenson
et al., 2007a,2008).
While carrying out a taxonomic survey of
Pouteria in the Atlantic forest, new species
of Pouteria (Alves-Araújo & Alves, in
press), Pradosia,andChromolucuma have
been discovered. This paper aims to
describe two novelties, one in each of the
latter two genera, both of which are
exclusively Neotropical and belong in the
Chrysophylloideae (Swenson & Anderberg,
2005; Swenson et al., 2008).
Pradosia is a genus of 23 species of trees
and geoxylic shrubs, all conned to tropical
America, except for the African species P.
spinosa (Ewango & Breteler, 2001), which
probably belongs to Pouteria (Swenson et al.,
2008). Pradosia is characterized by spirally
arranged leaves, usually eucamptodromous
leaf venation, and the absence of stipules.
The owers are bisexual with exserted sta-
mens and no staminodes. The fruit is different
from most other genera in the family, because
it develops into a one-seeded drupe with a
thin cartilaginous endocarp (Pennington,
1991). The presence of a drupe is the most
important character to distinguish Pradosia
from Elaeoluma, which has a berry.
Chromolucuma is a small genus of two tree
species in South America (Pennington, 1990).
Two important diagnostic characters are yellow
latex and persistent stipules. The inorescence
is axillary and bears unisexual, pedicellate
owers. The stamens are included in the
corolla, and staminodes are always present
(Pennington, 1991). Presence of stipules is an
important feature for both Chromolucuma and
Ecclinusa, but the latter genus has white latex
(not yellow), sessile owers, and lacks stamin-
odes. Recent phylogenetic analyses, based on
molecular and morphological data, indicate that
Chromolucuma is related to some species of
Pouteria (Swenson et al., 2008); however, only
one species of Chromolucuma was included in
this study. Although owers of the species
described below are still unknown, the presence
of yellow latex, persistent stipules, and a
distinct pedicel give us a reliable indication
that it is a member of Chromolucuma.
Pilz (1981) described Pouteria congestifolia
and called attention to the presence of stipules
and yellow latex, which are uncommon for the
genus. Despite this fact, Pennington (1990,
1991) kept the species in Pouteria when he
recognized Chromolucuma. Hence, we here
transfer P. congestifolia to Chromolucuma and
provide a new identication key for the genus.
New Species
Pradosia longipedicellata Alves-Araújo &
M. Alves, sp. nov. Type: Brazil. Bahia:
Una, Reserva Biológica do Mico-Leão-
Dourado, BA-001 Km 46, 15º09S,
39º05W, 9 Mar 1993 (,fr),J. G. Jardim,
A. M. Amorim, S. C. de SantAna, E. B. dos
Santos & J. L. Lage 92 (holotype: CEPEC;
isotype: NY). (Figs. 1AD,2AC,3)
Pradosiae brevipedi similis, sed habitu arboreo, 10
20 m alto, ramis juvenibus sericeis, pilis candidis vel
auratis, lenticellis absentibus, foliis 713 cm longis,
chartaceis, ellipticis vel ovatis, petiolis 11.5 cm longis,
inorescentiis oribus 24, pedicellis 1.83cmlongis,
calyce et corolla 56 mm longis, staminibus 56mmlongis,
ovario loculis 57, stylis 56 mm longis imprimis differt.
Trees 1020 m tall, shoot without lenticels,
tomentulose with whitish to golden tri-
chomes. Leaves spirally arranged, ovate to
.1. AD. Pradosia longipedicellata.A. Habit with details of lower leaf indument. b Outer (above) and inner
(below) sepals. C. Corolla and stamens. D. Gynoecium. EH.Chromolucuma apiculata.E. Habit. F. Stipules from the
outside (left) and inside (right). G. Fruit. H. Seed, view of testa (left) and seed scar (left). (AD, from Jardim et al. 92,
CEPEC; EH from Voeks 88, CEPEC.)
elliptic, 712 × 4.86 cm, chartaceous, both
surfaces tomentulose with whitish to golden
trichomes, upper surface sometimes shiny,
base cuneate, apex acute, margin at; venat-
ion eucamptodromous, midrib slightly sunken
on the upper surface; petiole 914 mm long,
channeled, tomentulose with whitish to
golden trichomes. Inorescences at apices of
.2. AC. Pradosia longipedicellata.A. Holotype (Jardim et al. 92, CEPEC). B.Inorescence and owers of
living material (photo by J. G. Jardim). C.Inorescence of the voucher (Jardim et al. 92, CEPEC). DE.
Chromolucuma apiculata.D. Holotype (Voeks 88, CEPEC). E. Stipule (arrow).
short branches; fascicles in clusters, umbelli-
form, 24owers (5-merous) per fascicle;
pedicels 23 cm long, tomentulose with whitish
to golden trichomes. Sepals 56 mm long, the
outer slightly shorter than the inner, ovate,
tomentulose on the outer surface with whitish
to golden trichomes, glabrous on the inner
surface, apex acute to ± rounded, margin entire.
Corolla cup-shaped, white, 78 mm long, tube
ca. 2 mm long; lobes 56 mm long, ovate,
margin entire, apex truncate to rounded, gla-
brous. Stamens ca. 6 mm long, xed in the
middle of the tube; laments glabrous, ca. 4 mm
long, upper half deexed, whitish; anthers 1.8
2 mm long, glabrous. Ovary 57-locular, 5
6 mm long, ferruginous pilose; style 56mm
long, glabrous, green; stigma simple. Fruit 1-
seeded, 34 cm long, ellipsoid, style persistent
in fruit, villous to tomentulose of ferruginous
trichomes; seeds laterally compressed, 20
22 mm long; testa smooth, shiny, brownish;
seed scar 1820 mm long, 23mmwide.
Distribution and conservation status.Pradosia
longipedicellata is currently known only from
the coastal Atlantic forest of southern Bahia
in northeastern Brazil (Fig. 3). It has a
narrow distribution with seven known sub-
populations, often near cocoa plantations. It
is a naturally rare species with few individuals
in each location. Following the IUCN Red List
criteria (IUCN, 2001,2008), P. longipedicellata
is assigned a preliminary threat status of
Vulnerable (VU A1cd,B1).
Phenology.Pradosia longipedicellata has
been recorded with owers and fruits almost
throughout the year (March to November).
refers to the long pedicel of the ower.
Additional specimens examined. BRAZIL. Bahia:
Una, Reserva Biológica do Mico-Leão-Dourado,
15º09'S, 39º05'W, 9 Nov 1993 (, fr), A. Amorim et al.
1424 (CEPEC, NY); Marambaia, 15 Jul 1995 (fr), A. M.
Amorim et al. 6067 (CEPEC); Estrada Ubaitaba-Maraú, 5
Sep 1999 (fr), A. M. Carvalho et al. 6730 (CEPEC, MO,
NY); Una, Estação Experimental do CEPLAC, 24 Oct
1980 (fr), A. Rylands 481980 (CEPEC); Ilhéus, Fazenda
Guanabara, 16 Oct 1980 (fr), L. A. Mattos-Silva et al.
1166 (CEPEC); Estrada Itacaré-Serra Grande, Maraú, 12
Jan 1967 (fr), R. P. Belém 3079 & R. S. Pinheiro
(CEPEC); Itacaré, 9 May 1968 (fr), R. P. Belém 3507
(CEPEC); Estrada Ubaitaba-Maraú, 24 May 1990 (), T.
S. Santos et al. 4554 (CEPEC); Una, Reserva Biológica
do Mico-Leão-Dourado, 15º09'S, 39º05'W, 20 Sep 1998
(fr), S. C. Sant'Ana et al. 668 (CEPEC, MO, NY).
Pradosia longipedicellata is recognized by
its clustered fascicles at the apices of short
shoots and pedicels 23 cm long. However,
there are other species of Pradosia that have
owers with long pedicels, such as P. brevipes
(Pierre) T. D. Penn., P. colombiana (Standl.)
T. D. Penn., and P. ptychandra (Eyma) T. D.
Penn. (Pennington, 1990). Pradosia long-
ipedicellata is easily distinguished from P.
ptychandra, which has shorter sepals (~2 mm
long) and a shorter corolla (~5 mm long), and
from P. colombiana, which has longer
petioles (2045 mm) and a shorter corolla
(~3 mm). The owers of P. longipedicellata
resemble that of P. brevipes, but the rst is a
tree that lacks lenticels on the shoots, and it is
only known from the Atlantic forest. In
contrast, P. brevipes is a geoxylic subshrub
that usually has shoots with lenticels, and it is
known from the Brazilian Cerrado. In other
words, P. longipedicellata and P. brevipes
differ in habit and are allopatric.
Chromolucuma apiculata Alves-Araújo &
M. Alves, sp. nov. Type: Brazil. Bahia:
Ilhéus, Fazenda Barra do Manguinho, Rodo-
via BA-001 Km 11, 14º47S, 39º02'W, 31
Jan 1985 (fr), R. Voeks 88 (holotype:
CEPEC). (Figs. 1EH,2DE,3)
.3. Location of the Atlantic forest, together with
known distribution of Pradosia longipedicellata
(circles), Chromolucuma apiculata (square), and C.
congestifolia (triangles).
Chromolucumae rubriorae afnis, a qua lenticellis
praesentibus, stipulis lanceolatis, ubi juvenibus tomento-
sis, ubi adultis glabris, apice apiculato, foliis coriaceis vel
crassis, glabris, petiolis 1015 mm longis, fructibus 15
18 mm longis, seminibus 1516 mm longis, testis
laevibus et nitidis imprimis differt.
Trees 1012 m tall with whitish bark and
yellow latex; shoots glabrous with lenti-
cels; stipules 58 mm long, lanceolate, rst
tomentose with ferruginous trichomes,
glabrescent; apex apiculate. Leaves spirally
arranged, oblanceolate, 1621 x 5.5
7.5 cm, coriaceous, glabrous, lower surface
glaucous, base cuneate, apex acute, margin
slightly revolute; petiole 1015 mm long,
not channeled, glabrous. Fascicles at apices
of branches, axillary, 12-owered; pedicel
1018 mm, glabrous. Sepals ca. 4 mm
long, ovate, glabrous to glabrescent on the
outer surface, glabrous on the inner sur-
face, apex obtuse, margin entire, sometimes
ciliate. Flowers unknown. Fruit 1-seeded,
2530 mm long, spherical to ellipsoid,
glabrous, purple; seeds 1518 mm long,
slightly laterally compressed; testa smooth,
shiny, brownish; seed scar 1416 mm long,
ca. 4 mm wide.
Distribution and conservation status.
Chromolucuma apiculata is known only from
the coastal Atlantic forest of Bahia, Brazil
(Fig. 3). This is the rst record of the genus
occurring outside the Amazonian region and is
an example of a disjunct distribution between
the Amazonian and Atlantic forests. The spe-
cies is only known from the type collection and
we refrain from giving it a preliminary con-
servation assessment, and instead classify it as
Data Decient (DD).
Phenology.Fruiting in January; owering
period unknown.
Etymology.This species is named for the
characteristic apiculate stipules.
Among Chromolucuma,C. apiculata is the
only species that has young shoots and leaves
that are completely glabrous, along with short
petioles (1015 mm long), and stipules that
are 58 mm long and apiculate.
New Combination
Chromolucuma congestifolia (Pilz) Alves-
Araújo & M. Alves, comb. nov. Pouteria
congestifolia Pilz, Ann. Missouri Bot.
Gard. 68: 191. 1981. Type: Panamá. Coclé:
El Valle de Antón, [16 Mar 1946] (), P.
H. Allen 3426 (holotype: MO; isotypes:
BM, F, G). (Fig. 3)
Distribution and conservation.Chromolu-
cuma congestifolia is known from the moun-
tain region between Costa Rica and Panama,
and the Amazonian forest in Brazil and
French Guiana (Fig. 3). Because of human
impact, increasingly more forest is frag-
mented, especially in the Brazilian Amazon.
Ten populations of C. congestifolia are
known, three of which are in protected areas.
Considering this information, C. congestifolia
is assigned a preliminary threat status of
Vulnerable (VU A1c,B1,B2c).
Phenology.Chromolucuma congestifolia
has been recorded with owers and fruits
from February to September.
Additional specimens examined. COSTA RICA.
Alajuela: Reserva Biológica Monteverde, Laguna de
Minor Vargas, 10°18'36"N, 84°42'36"W, 14 Sep 1989
(), E. Bello 1269 (MO); Laguna de Poco Sol, 10°21'N,
84°39'36"W, 30 Sep 1989 (), E. Bello 1300 (MO);
Reserva Biológica de San Ramon, 10º18N, 84º34W, 30
May1 Jun 1986 (), B. Hammel et al. 15262 (MO, CR);
without exact locality, 21 Feb 1984 (fr), T. D Pennington
& L. J. Poveda 11546 (K). Cartago: Paraiso, 09°
44'05"N, 83°46'42"W, 14 Mar 2000 (), L. Acosta & V.
H. Ramírez 627 (MO). Heredia: North of Quebrada
Tigre from Finca El Plástico, 10º18N, 84º02W, 14 Feb
1986 (), M. H. Grayum & P. Sleeper 6494 (MO).
PANAMA. Chiriquí: Along Quebrada de Arena, 12
Mar 1982 (fr), S. Knapp et al. 4068 (MO). Veraguas:
Near Cerro Tute-Arizona, above Santa Fe and Alto de
Piedra, 08°30'N, 81°10'W, 5 Feb 1988 (), G. McPherson
12050 (MO).
FRENCH GUIANA. Saul, Eaux Claires, 9 Feb 1993,
(fr), T. D. Pennington et al. 13840 (K).
BRAZIL. Amazonas: Manaus-Itacoatiara, Reserva
Florestal Ducke, km 26, 02º53S, 59º58W, 1976, J. A.
Souza s.n. (INPA); Manaus-Itacoatiara, Reserva Florestal
Ducke, km 26, 02º53S, 59º58W, 22 Aug 1997, P. A. C.
L. Assunção 627 (INPA, K).
Among Chromolucuma species, C. con-
gestifolia is distinguished by its shorter
stipules (36 mm long) and a lower leaf
surface that usually has long trichomes
restricted to the midrib. In contrast, C.
baehniana has a sericeous lower leaf sur-
face, and C. rubriora and C. apiculata
have glabrous leaves. For more detailed
information about Chromolucuma species,
see the following key.
1. Lower leaf surface sericeous; shoots without lenticels (Guiana, Venezuela). ........C. baehniana
1. Lower leaf surface glabrous (except for midrib); shoots with lenticels.
2. Leaves chartaceous; stipules 24 cm long; pedicels 48 cm long (Venezuela, northern
Brazil)...................................... C. rubriora
2. Leaves coriaceous; stipules <1 cm long; pedicels <2 cm long.
3. Stipules apiculate; petioles 1015 mm long; fascicles 12-owered; fruit 1-seeded (Brazilian
Atlanticforest).........................................C. apiculata
3. Stipules acute; petioles 2060 mm long; fascicles 520-owered; fruit 2-seeded (Costa Rica,
Panama,FrenchGuiana,northernBrazil) ............ ..........C. congestifolia
The rst author thanks PPGBV/Capes for
nancial support. The authors thank André
Amorim (Curator of CEPEC), James C.
Solomon (Curator of MO), and the staff of
CEPEC and MO for the support. The
authors would also like to thank Jorge
Fontella for helping with the Latin diag-
nosis, Regina Carvalho for the illustration,
Nathan Smith for English revision, and Ulf
Swenson for improving the paper for high
quality suggestions.
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Key to the species of Chromolucuma
... Latex in Sapotaceae is usually white, but one clade of Neotropical species has yellow or yellowish latex (Fig. 3E), diagnostic of Chromolucuma (Pennington, 1991;Alves-Araújo & Alves, 2012a). A few species may also have cream latex, one being P. eugeniifolia, occurring in clade Q. Pennington (1991) stressed that American Pouteria spp. ...
... Our phylogenetic analyses, based on ribosomal and nuclear sequence data, group Sarcaulus-clade Q with maximum Bayesian (PP 1.0) and weak jackknife (JK 64%) support in one large clade. It includes all species of Pouteria not discussed above, plus three widely accepted genera, Chromolucuma, Pradosia and Sarcaulus (Aubréville, 1964a;Pennington, 1990Pennington, , 1991Pennington, , 2006Pennington, , 2007Alves-Araújo & Alves, 2012a;Terra-Araujo et al., 2013, 2015. Different character combinations readily distinguish these three genera, but in our analyses, they are embedded among the clades of Pouteria. ...
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Generic limits of Chrysophyllum and Pouteria (Chrysophylloideae, Sapotaceae) have been found to be untenable. We here search for natural lineages in Neotropical Chrysophylloideae by sampling 101 terminals for molecular sequences of nuclear ribosomal DNA (external and internal transcribed spacer), the nuclear gene RPB2 and 17 morphological characters. Data were analysed with Bayesian inference and parsimony jackknifing. Morphological traits were finally optimized onto the tree to identify the most coherent characters. The resulting phylogenetic tree suggests that the limits of the well-known genera Chrysophyllum and Pouteria must be amended. Diploon, Ecclinusa and Elaeoluma can be maintained and Chrysophyllum sections Ragala section Prieurella and the satellite gen- era Achrouteria, Cornuella, Martiusella and Nemaluma merit generic resurrection. Lucuma may be restored if the type species belongs to the clade. The accepted genera Chromolucuma, Pradosia and Sarcaulus gain strong clade support, but are embedded in a core clade of Pouteria and may be relegated to the subgeneric level if morphologi- cal studies cannot provide evidence concurring with narrow generic concepts. Circumscriptions of Micropholis and Chrysophyllum sections Chrysophyllum and Villocuspis remain unclear and must be explored by using an extended taxon sampling. We predict that yet-to-be-analysed species of Pouteria sections Franchetella, Gayella, Oxythece and Pouteria and members of the currently accepted genera Chromolucuma, Pradosia and Sarcaulus will fall inside the core clade of Pouteria when analysed.
... Since discussions about the correct positioning of Pouteria species are still ongoing, taxonomic surveys regarding AUtALVES-ARAÚJOHOR 56 • Phytotaxa 461 (1) © 2020 Magnolia Press to Sapotaceae species from neotropics have revealed several novelties (Alves-Araújo & Alves 2011, 2012a, 2012b, terra-Araújo et al. 2012, 2013, Popovkin et al. 2017, Alves-Araújo & Mônico 2017, Sossai et al. 2017, Alves-Araújo 2018a, 2018b, Vasconcelos et al. 2020. Considering it, the more we know about Sapotaceae diversity, the better will be the understanding of new perspectives on phylogeny paths. ...
Pouteria alvesii (Chrysophylloideae, Sapotaceae) a new species from Atlantic Forest is here described, and it is morphologically related to Pouteria pachyphylla, which occurs in Amazon Rainforest. Herein, description, illustration, comments on geographic distribution and phenology information are provided. Based on its distribution and habitat, the new species is assigned as Vulnerable, according to the IUCN criteria.
... In a narrow sense, Pouteria is restricted to South America (Swenson et al. 2008, Faria et al. 2017) and distributed across a wide variety of habitats, including lowland rainforests on both white-sand and clayish soils (Pennington 1990(Pennington , 2006. Even if Pouteria s.l. has been found untenable to upheld, new species have been described from the Brazilian Atlantic forests (Alves-Araujo & Alves 2011, 2012a, 2012b, Popovkin et al. 2016, Alves-Araujo 2018, Alves-Araujo & Mônico 2018. In Central Amazonia, density and species richness for Pouteria were recently established for approximately 1000 km 2 study plots of terra firme forest from the Área de Relevante Interesse Ecológico Projeto Dinâmica Biológica de Fragmentos Florestais (ARIE-PDBFF, in Portuguese). ...
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Pouteria kossmanniae (Sapotaceae, Chrysophylloideae), a new species from Central Amazonia, is here described. It is known from the Manaus region, Amazonas, Brazil, and is found in non-flooded upland forests, known as terra firme. Illustrations are provided in addition to geographic distribution, with a comparison of the morphological and spectral (near-infrared) differences between P. kossmanniae and its very similar species Pouteria macrophylla, Pouteria manaosensis, and Pouteria rodriguesiana. The species is naturally common in some urban forest fragments, but since only a low number of subpopulations is known, we assign P. kossmanniae the preliminary conservation status of "Endangered".
... Over the past few years, new species and new occurrence records of Pouteria have been reported from the Brazilian Atlantic forest (Alves-Araújo and Alves 2011, 2012a, 2012bPopovkin et al. 2016;Mônico et al. 2017;Alves-Araújo 2018;Alves-Araújo and Mônico 2018). In the highly diverse Amazonia forest (ter Steege et al. 2016), Pouteria is amongst the 10 top most species-rich tree genera (ter Steege et al. 2016;Cardoso et al. 2017), but still remains poorly studied in this biome. ...
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This is the first record of Pouteria franciscana Baehni (Chrysophylloideae, Sapotaceae) in Amapá state, Brazil. We provide a brief description as well as a distribution map, illustrations, and a table with useful features to distinguish P. franciscana from its morphologically related Amazonian species. Using geographic data and applying IUCN criteria, we assign the status as Least Concern for P. franciscana .
... Over the past few years, new species and new occurrence records of Pouteria have been reported from the Brazilian Atlantic forest (Alves-Araújo and Alves 2011, 2012a, 2012bPopovkin et al. 2016;Mônico et al. 2017;Alves-Araújo 2018;Alves-Araújo and Mônico 2018). In the highly diverse Amazonia forest (ter Steege et al. 2016), Pouteria is amongst the 10 top most species-rich tree genera (ter Steege et al. 2016;Cardoso et al. 2017), but still remains poorly studied in this biome. ...
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This is the first record of Pouteria franciscana Baehni (Chrysophylloideae, Sapotaceae) in Amapá state, Brazil. We provide a brief description as well as a distribution map, illustrations, and a table with useful features to distinguish P. franciscana from its morphologically related Amazonian species. Using geographic data and applying IUCN criteria, we assign the status as Least Concern for P. franciscana.
... Nos Neotrópicos, a diversidade da família é estimada em cerca de 450 espécies , além da inclusão contínua de novas espécies (ex. Pennington 2006; Alves-Araújo e Alves 2011Alves , 2012Terra-Araujo et al. 2012, 2015. Em florestas inundáveis e de terra-firme da Amazônia está sempre entre as dez famílias botânicas mais representativas (Ferreira e Prance 1998;Milliken 1998;Oliveira e Daly 1999;ter Steege et al. 2000;Hopkins 2007). ...
Sapotaceae is a pantropical family divided in the three subfamilies Chrysophylloideae, Sapotoideae and Sarcospermatoideae, together comprising approximately 1,250 species and 58 genera. Traditional generic limits are notoriously problematic within family, due to extensive homoplasy, which have been resulting in different proposals of classification. There is strong phylogenetic support for the subfamily Chrysophylloideae, being considered the largest grouping in Sapotaceae and the best represented in Neotropics. However, their internal generic limits are still not entirely clear, mainly for Pouteria and Chrysophyllum, which are paraphyletic or polyphyletic genera. Considering that new approaches are necessary in the search for diagnostic characters to support the circumscription of genera, this study focus on taxonomic and systematic issues in the Chrysophylloideae, especially to genera from Amazon. The purpose of this study is to increase the morphological knowledge about fruits, seeds and, especially seedlings of species Neotropical Chrysophylloideae (Sapotaceae), with characters evaluation useful under a phylogenetic perspective. Thus, we provide comprehensive morphological descriptions, taxonomic notes, identification keys and illustrations of fruits, seeds and seedlings for 36 species in eight genera of Chrysophylloideae: Chromolucuma, Chrysophyllum, Ecclinusa, Elaeoluma, Micropholis, Pouteria, Pradosia and Sarcaulus. We collected ripe fruits mainly in the Amazonas, Bahia and Roraima states, Brazil and soon after we described 220 quantitative and qualitative morphological characters (30 of fruits, 33 of seeds and 157 of seedlings), based on previous studies for Sapotaceae. We traced 78 characters in the molecular phylogeny recently proposed for Neotropical Chrysophylloideae, of which eight have been identified as useful for diagnostic purposes: a) presence/absence of “marzipan” odor, b) presence/absence of pedicel on fruit, c) type of cotyledons, d) presence/absence of endosperm, e) cotyledon consistency, f) hypocotyl elongation, g) type of seedling (innovative character), and h) presence/absence of stipule on seedlings. Our results supported recent proposals of generic delimitation to subfamily, because such as Ecclinusa, Micropholis, Prieurella, Ragala and the easily distinguished Chromolucuma, Pradosia and Sarcaulus genera had morphological support for the clades.
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A new species from central Amazonia, Chromolucuma brevipedicellata (Sapotaceae, Chrysophylloideae) is described and illustrated. It occurs in terra firme (non-flooded) forests near Manaus, Brazil. We compare C. brevipedicellata to similar species in terms of diagnostic morphology, as well as near-infrared spectral differences of the leaves, and provide, an identification key for its distinction. Although the new species is common in the permanent plots of the Área de Relevante Interesse Ecológico Projeto Dinâmica Biológica de Fragmentos Florestais, it has a narrow distribution and is assessed as being endangered following criteria of the IUCN Red List.
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Sapotaceae is historically known as having a tricky and challenging taxonomy due to tangled morphologic heterogeneity. Consequently, this resulted in a large number of described genera and binomials. After Pennington’s Flora Neotropica work, several of those nomenclature issues were resolved. Nevertheless, many binomials remain unsolved and up for typification. Thus, following the International Code of Nomenclature for Algae, Fungi and Plants, we propose 74 new lectotype designations, four of these are second-step typifications.
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Sapotaceae is a large family of angiosperms which has predominantly tree species, and worldwide distribution in subtropical and tropical regions. The Neotropics are one of the greatest centers of diversity of Sapotaceae, and in Brazil, the family comprises 12 genera and 234 species, of which 103 species are endemic. We present here a taxonomic treatment for the native species of Sapotaceae from the State of Paraná, with an identification key, descriptions, geographic distribution, phenology, illustrations, and current conservation status for all taxa. Twenty-two species were documented in Paraná, belonging to seven genera. The richest genus was Pouteria with 11 species, followed by Chrysophyllum with five species and Pradosia with two species. The genera Diploon, Ecclinusa, Manilkara, and Sideroxylon have only one species each. Pouteria guianensis and Pouteria ramiflora are new records for the state. The Atlantic Rain Forest is the richest vegetation type with six genera and 13 species, followed by the Seasonal Forest with three genera and nine species. The Araucaria Forest and Cerrado have two genera each, the former with three species and the latter with two species. Out of 22 species, one has been mentioned as endangered (EN), one is vulnerable (VU), one is near threatened (NT), nine are least concern (LC), and ten have not been evaluated (NE).
A new species assigned to the genus Pouteria is described and illustrated. Pouteria citriodora is morphologically related to Pouteria procera but differs from itmainly by its axillary and isolated flowers (3-15-flowered in P. procera), calyx aestivation quincuncial (convolute/contorted in P. procera), longer tube corolla (5.0-7.3mmlong vs. 1.75-3.5mmlong in P. procera), and fruits with whitish dots on the surface and a very distinctive Citrus-like smell. Herein environmental, geographic distribution and conservation data for the new species are provided. In addition, a key to species of Pouteria with 5-merous flowers from Bahia state-Brazil is presented.
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The phylogeny of the Sapotaceae was investigated by DNA sequence analysis of the chloroplast gene ndhF. Three or possibly four main evolutionary lineages were identified. Sarcosperma is the sister group of all other Sapotaceae, which together form two strongly supported monophyletic groups. One large clade with strong support is comprised of the tribes Isonandreae, Mimusopeae, and Sideroxyleae, and also includes the genus Capurodendron of tribe Chrysophylleae. The second large clade with strong support is formed by the tribes Chrysophylleae and Omphalocarpeae, as well as Diploon of tribe Sideroxyleae. Weak support was found for a position of the genus Xantolis as sister to the Chrysophylleae-Omphalocarpeae, but this genus could equally well constitute a separate evolutionary lineage. The relationships between many of the genera within each of the larger clades are still unclear, but based on the results, a discussion of diagnostic characters in the Sapotaceae from a cladistic perspective is presented.
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We present a hypothesis of phylogenetic relationships in Planchonella (Sapotaceae, Chrysophylloideae) using nrDNA (ITS) data analysed with parsimony jackknifing and Bayesian inference. Results from these two approaches yield similar majority-rule consensus trees. Planchonella (formerly included in Pouteria) is a monophyletic genus of trees and shrubs with 60 species currently recognised. They are distributed in Australia (13 spp.), New Caledonia (32 spp.), the Pacific islands (10 spp.), and New Guinea (5 spp.), but more molecular research is needed in order to identify all members in New Guinea and Malesia. Three strongly supported clades and several subclades were recovered by our analysis. Two clades are restricted to New Caledonia. Within these are eight recently discovered species that are described here (P. crenata, P. glauca, P. latihila, P. luteocostata, P. mandjeliana, P. povilana, P. roseoloba, P. rufocostata), each with a conservation assessment. Our molecular phylogeny shows that some of these taxa, initially believed to be conspecific or closely related, represent separate lineages that deserve species rank. Two of these species are restricted to areas with near-future mining activities, which is why urgent conservation assessments are needed. Several subspecies or varieties of P. myrsinifolia and P. cotinifolia are recognised in Australia. Neither of these two species is monophyletic in its present circumscription, and for this reason P. cotinifolia var. pubescens is given species rank here as P. pubescens. We also amend the generic description of Planchonella, a genus best distinguished on a character combination of stamens positioned just below (rarely in) the tube orifice, a multi-seeded fruit, and foliaceous cotyledons embedded in endosperm.
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We present the first cladistic study of the largely tropical family Sapotaceae based on both morphological and molecular data. The data were analyzed with standard parsimony and parsimony jackknife algorithms using equally and successive weighted characters. Sapotaceae are confirmed to constitute two main evolutionary lineages corresponding to the tribes Isonandreae-Mimusopeae-Sideroxyleae and Chrysophylleae-Omphalocarpeae. The Sideroxyleae are monophyletic, Isonandreae are polyphyletic as presently circumscribed, and as suggested by the analyses, the subtribe Mimusopeae-Mimusopinae has evolved within the Mimusopeae-Manilkarinae, which hence is also paraphyletic. Generic limits must be altered within Sideroxyleae with the current members Argania, Nesoluma and Sideroxylon. Argania cannot be maintained at a generic level unless a narrower generic concept is adopted for Sideroxylon. Nesoluma cannot be upheld in a narrow or broad generic concept of Sideroxylon. The large tribe Chrysophylleae circumscribes genera such as Chrysophyllum, Pouteria, Synsepalum, and Xantolis, but the tribe is monophyletic only if the taxa from Omphalocarpeae are also included. Neither Chrysophyllum nor Pouteria are monophyletic in their current definitions. The results indicate that the African taxa of Pouteria are monophyletic and distinguishable from the South American taxa. Resurrection of Planchonella, corresponding to Pouteria section Oligotheca, is proposed. The African genera Synsepalum and Englerophytum form a monophyletic group, but their generic limits are uncertain. Classification of the Asian genus Xantolis is particularly interesting. Morphology alone is indecisive regarding Xantolis relationships, the combined unweighted data of molecules and morphology indicates a sister position to Isonandreae-Mimusopeae-Sideroxyleae, whereas molecular data alone, as well as successive weighted combined data point to a sister position to Chrysophylleae-Omphalocarpeae. An amended subfamily classification is proposed corresponding to the monophyletic groups: Sarcospermatoideae (Sarcosperma), Sapotoideae (Isonandreae-Mimusopeae-Sideroxyleae) and Chrysophylloideae (Chrysophylleae-Omphalocarpeae), where Sapotoideae circumscribes the tribes Sapoteae and Sideroxyleae as well as two or three as yet unnamed lineages. Morphological characters are often highly homoplasious and unambiguous synapomorphies cannot be identified for subfamilies or tribes, which we believe are the reason for the variations seen between different classifications of Sapotaceae. © The Willi Hennig Society 2005.
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We present a molecular phylogeny of 26 out of the 28 currently accepted genera in the subfamily Chrysophylloideae (Sapotaceae) using parsimony, parsimony jackknifing, and Bayesian inference. A data matrix of 8984 characters was obtained from DNA sequences of seven chloroplast loci, two nuclear loci, indels coded as binary characters, and morphology. Our phylogenetic reconstruction suggests that Chrysophyllum, Pouteria, and Pradosia, as well as some sections within Chrysophyllum and Pouteria, are all polyphyletic. These taxa were previously described largely on the basis of unique combinations of states for a set of morphological characters. Mapping some of these characters onto one of the most parsimonious trees indicates that the symplesiomorphic flower in the subfamily was probably 5-merous, had stamens inserted in the tube orifice, staminodes, seeds with foliaceous cotyledons, exserted radicle, and endosperm. These characters have subsequently been lost multiple times and cannot be used as synapomorphies to support broad generic concepts. Despite the high degree of homoplasy some well-defined clades can be described on the basis of alternative character state combinations. Also, many of these well-supported clades appear to be restricted to particular geographical areas (e.g. all taxa in Australasia form a monophyletic group). Hence, we suggest that the segregate genera Aningeria, Malacantha, and Martiusella may ultimately be resurrected, and probably also Donella and Gambeya, but their circumscriptions are still unclear. One species, Chrysophyllum cuneifolium, may have originated from a hybridization event between continents where the maternal genome (cpDNA) comes from South America and the nuclear genome comes from Africa. © The Willi Hennig Society 2008.
Trees or shrubs, rarely geoxylic suffrutices or lianas, sometimes spiny; branching usually sympodial; latex nearly always present in trunk, branches and fruits, usually white, rarely yellow or blue; indumentum nearly always of malpighiaceous hairs (simple in Delpydora). Leaves alternate, spirally arranged or distichous, less frequently opposite or verticillate, simple, entire or very rarely spinous-toothed; petiole rarely bearing a pair of minute stipels; stipules + or 0. Inflorescence fasciculate or flowers occasionally solitary, axillary, ramiflorous or cauliflorous; fascicles occasionally arranged along short leafless axillary, panicle-like shoots; fascicle base sometimes developing into short, densely scaly brachyblasts. Flowers bisexual or unisexual (plants monoecious or dioecious), actinomorphic; calyx a single whorl of 4–6 free or partly fused, imbricate, sometimes quincuncial sepals, or 6–11 sepals in a closely imbricate spiral, or with 2 whorls of 2–4 sepals and then the outer whorl valvate or only slightly imbricate; corolla rotate, cyathiform or tubular, sympetalous; tube shorter than, equalling or exceeding the petals; petals 4–18, entire, lobed or partly divided or divided to the base into 3 segments and then median segment entire, 2 lateral or dorsal segments entire, laciniate or shallowly or deeply divided; stamens 4–35(−43), fixed in lower or upper half of corolla tube or at the base of the lobes, rarely free, in a single whorl opposite the corolla lobes, or, when more numerous than the corolla lobes, some opposite and some alternate with the corolla lobes, or sometimes several stamens clustered opposite each lobe, or arranged in 2–3 alternating whorls within the corolla tube, exserted or included; filaments often geniculate in bud, free, rarely fused into a staminal tube, or partially fused to the staminodes; anthers often extrorse; staminodes 0–8(−12) in a single whorl alternating with the stamens or fixed in the corolla lobe sinuses, simple or variously lobed, toothed or divided, sometimes petaloid; disk annular or patelliform, surrounding the ovary base and sometimes fused with it, or absent; ovary superior, l–15(−30)locular, loculi usually uniovulate, rarely 2−5-ovulate, placentation axile, basi-ventral or basal; style simple, included or exserted; style-head simple or minutely lobed. Fruit a berry or rarely a drupe, or tardily dehiscent by a single lateral valve; pericarp fleshy or less frequently leathery or woody; seeds 1-many, globose, ellipsoid, oblong, often strongly laterally compressed, testa usually smooth and shining, free from the pericarp, less frequently roughened, wrinkled or pitted and then often adherent to the pericarp; hilum adaxial, basi-ventral or basal, narrow or broad, sometimes extending to cover most or all of the seed; embryo vertical, oblique or horizontal, with thin foliaceous or thick flat or plano-convex, usually free, cotyledons; radicle included or exserted; endosperm + or 0. x = 10, 11, 12, 13, 14.
The Flora of Panama Sapotaceae treatment by Blackwell is updated. Pouteria congestifolia Pilz and P. leptopedicellata Pilz are newly described and the combination Pouteria buenaventurensis (Aubr.) Pilz is made. As now revised, the Panamanian flora includes 44 species in 11 genera, an increase from the 24 species in 6 genera known to Blackwell. This revision is based on collections made in the past decade.
Current generic limits in Chrysophylloideae (Sapotaceae) from Australia, New Caledonia and the Pacific islands have been shown not to correspond to monophyletic groups. In particular, revisions of generic boundaries are necessary for Pouteria and Niemeyera. We present the first cladistic study of a large representative sample from these areas based on (i) nuclear ribosomal DNA (nrDNA) sequence data, and (ii) combined data of nrDNA and morphology. The data were analyzed with parsimony jackknifing using equal weights and gaps coded as binary characters. Our results from the two data sets are highly congruent and morphological data often increase support as well as tree resolution. A basal polytomy prevents hypotheses of intergeneric relationships, but several groups receive strong support, and hence, four segregates of Pouteria (Beccariella, Planchonella, Sersalisia and Van-royena) are resurrected. Four others, Albertisiella, Bureavella, Iteiluma and Pyriluma are rejected. Niemeyera is redefined as a small genus confined to Australia. Generic limits within the sister group to Niemeyera are still unclear, a group that includes Leptostylis and Pycnandra. Furthermore, Van-royena may have originated from an intergeneric hybridization event. Traditionally used and newly identified morphological characters are scrutinized for their diagnostic value. For instance, the position of stamen insertion within the corolla tube is a strong indication of generic relationship. Unique synapomorphies are rare and genera must be distinguished on character state combinations. Following the results, several taxonomic combinations are necessary (Beccariella brownlessiana, B. macrocarpa, B. singuliflora, B. vieillardii, Pichonia daenikeri, Planchonella asterocarpon, P. dothioense, P. myrsinifolia, P. myrsinodendron and P. xylocarpa). © The Willi Hennig Society 2007.