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A Taxonomic Survey of Pouteria (Sapotaceae) from the Northern Portion of the Atlantic Rainforest of Brazil

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Abstract— Pouteria s. s. (Chrysophylloideae) includes ca. 200 species, representing the largest genus of Sapotaceae. In Brazil, 120 species are recorded in Amazonia and in the Atlantic Rainforest where the genus is centered. This paper presents a taxonomic survey of the 34 species of Pouteria s. s. from the northern portion of the Atlantic Rainforest of Brazil, and includes an identification key, nomenclatural updates, new synonyms, descriptions, and detailed information about geographic distribution and phenology for all taxa, and illustrations. Species of Pouteria are mainly separated based on leaf venation, trichome type, stamen and staminode morphology, and fruit traits. Out of the 34 species recognized, 18 are endemic to the Atlantic Rainforest of Brazil, 10 are disjunctly distributed with Amazonia, and six are widespread in South America. According to the IUCN red list criteria, 20 of the 34 species treated here are threatened; of these, 10 are vulnerable, seven are endangered, and three are critically endangered.
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Systematic Botany (2014), 39(3): pp. 915–938
©Copyright 2014 by the American Society of Plant Taxonomists
DOI 10.1600/036364414X681428
Date of publication 05/27/2014
A Taxonomic Survey of Pouteria (Sapotaceae) from the Northern Portion
of the Atlantic Rainforest of Brazil
Anderson Alves-Arau
´jo,
1,4
Ulf Swenson,
2
and Marccus Alves
3
1
Laborato
´rio de Sistema
´tica e Gene
´tica Vegetal, Departamento de Cie
ˆncias Agra
´rias e Biolo
´gicas, CEUNES/Universidade
Federal do Espı
´rito Santo – UFES, CEP 29932-540, Sa
˜o Mateus- Espı
´rito Santo, Brazil.
2
Swedish Museum of Natural History - NRM, Phanerogamic Department, Box 50007, 104 05 Stockholm, Sweden.
3
Laborato
´rio de Morfo-Taxonomia Vegetal, Departamento de Bota
ˆnica, Universidade Federal de
Pernambuco – UFPE, CEP 50670-901, Recife- Pernambuco, Brazil.
4
Author for correspondence (sapotae@gmail.com)
Communicating Editor: Lucia G. Lohmann
Abstract—Pouteria. s. s. (Chrysophylloideae) includes ca. 200 species, representing the largest genus of Sapotaceae. In Brazil, 120 species are
recorded in Amazonia and in the Atlantic Rainforest where the genus is centered. This paper presents a taxonomic survey of the 34 species of
Pouteria s. s. from the northern portion of the Atlantic Rainforest of Brazil, and includes an identification key, nomenclatural updates, new
synonyms, descriptions, and detailed information about geographic distribution and phenology for all taxa, and illustrations. Species of
Pouteria are mainly separated based on leaf venation, trichome type, stamen and staminode morphology, and fruit traits. Out of the 34 species
recognized, 18 are endemic to the Atlantic Rainforest of Brazil, 10 are disjunctly distributed with Amazonia, and six are widespread in South
America. According to the IUCN red list criteria, 20 of the 34 species treated here are threatened; of these, 10 are vulnerable, seven are
endangered, and three are critically endangered.
Keywords—Biodiversity, Chrysophylloideae, conservation, IUCN, Neotropics.
The Atlantic Rainforest of Brazil is a global hotspot of
diversity and endemism, representing one of the most impor-
tant centers of biodiversity in the Neotropics, and constitut-
ing a top priority for conservation (Mittermeier et al. 1999;
Myers et al. 2000). This biome is distributed across 17 Brazil-
ian states and constitutes a mosaic of vegetation types,
including the “Brejos de Altitude” (i.e. inland forests above
600 m) in the northern portions to highlands in the southern
regions, in addition to the coastal forests from the eastern
shores of Brazil (Velloso et al. 1991). Unfortunately, logging,
clearings, and plantations have led to intensive fragmention
and destruction of ca. 88–93% of its original forest cover
(Metzger 2003; Myers et al. 2000; Ribeiro et al. 2009;
Stehmann et al. 2009).
Sapotaceae is one of the most important plant families in
the Atlantic Rainforest of Brazil (Pennington 1990, 2006;
Swenson and Anderberg 2005). This family consists of
58 genera and about 1,250 species in subtropical and tropical
regions of the world (Pennington 1991; Govaerts et al. 2001;
Swenson et al. 2007a, b).
The center of diversity of Sapotaceae is in the Neotropics,
where the majority of species occur in moist lowland forests
(Pennington 2006). In Brazil, Carneiro et al. (2013) docu-
mented 10 genera and 231 species of Sapotaceae, of which 87
species are registered in northeastern Brazil. However, many
species of Sapotaceae are exploited for timber, which has led
to a decrease in the size of the populations of several species,
especially those that are naturally uncommon (Pennington
2006). The exact degree of destruction of the various popula-
tions is, however, yet to be documented. In addition, most
species still lack the IUCN red list assessments necessary for
appropriate conservation measurements.
Pennington (1990) produced a monograph of Neotropical
members of the family, which was followed by a revised
generic classification (Pennington 1991). More recently, three
different subfamilies were recognized by Swenson and
Anderberg (2005): Sarcospermatoideae, Sapotoideae, and
Chysophylloideae. Pouteria Aubl. represents one of the most
diverse genera within subfamily Chrysophylloideae. The
genus was originally divided into nine sections, six of which
were confined to the New World and the remaining three
were restricted to Africa and Australasia (Pennington 1990,
1991). However, recent molecular phylogenetic studies have
demonstrated that Pennington’s sections were not monophy-
letic as originally circumscribed, suggesting that the genus
should be circumscribed more narrowly and should only
include the Neotropical taxa (Anderberg and Swenson 2003;
Bartish et al. 2005; Swenson and Anderberg 2005; Swenson
et al. 2007a, 2008). Despite the reduction in the circumscrip-
tion of the genus, Pouteria still includes ca. 200 species, and
remains as the largest genus in the family (Pennington 1990,
2006). The natural boundaries of Pouteria have not been fully
resolved yet, and further phylogenetic and taxonomic studies
are needed to clarify the relationships within Pouteria, and
between Pouteria and its closest relatives. Indeed, taxonomic
novelties are common in the genus (Alves-Arau
´jo and Alves
2011, 2012).
In Brazil, 121 species of Pouteria were documented (Alves-
Arau
´jo 2014), of which 41 names were registered for north-
eastern Brazil, and 30 were cited for the Atlantic Rainforest.
Nevertheless, the real diversity and endemism of Pouteria in
the Atlantic Rainforest remains to be confirmed. The objec-
tive of this study is to provide a taxonomic survey of all
species of Pouteria from the northern portions of the Atlantic
Rainforest of Brazil, to better characterize its taxa, contribute
to its taxonomy, and evaluate the status of threat of the indi-
vidual species.
Materials and Methods
The northern portion of the Atlantic rainforest of Brazil extends from
the State of Bahia to Ceara
´, including the “Brejos de Altitude” and sea-
sonal forests from Chapada Diamantina in Bahia, and the coastal forests
in the eastern shores (Velloso et al. 1991; Mittermeier et al. 1999; Myers
et al. 2000; Ribeiro et al. 2009; Stehmann et al. 2009) (Fig. 1).
Fieldwork was performed from March 2007 to December 2009 in 38
different vegetation fragments along the study area. Vouchers were
915
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deposited at UFP and duplicates sent to CEPEC, NY, RB, and S. In
addition, most of the main herbaria with holdings of Sapotaceae and
the Atlantic Rainforest were visited and/or consulted on-line (*):
ALCB, ASE, BC*, BHCB, BM*, BR*, C, CEPEC, EAC, EAN, F*, FI*, G,
GB, HRB, HST, HUEFS, IPA, JPB, K, M, MA*, MAC, MO, NY, P,
PEUFR, RB, S, TEPB, UESC, UFP, UFRN, US*, W*, WAG*, and Z
(acronyms following Thiers 2011). Type specimens, herbarium
vouchers, and literature were also consulted in order to confirm spe-
cies identifications (Pennington 1990, 1991, 2006). The conservation
status of individual species was determined according to the IUCN
red list criteria (IUCN 2012).
Morphological terms follow Hickey and King (2000) and Harris and
Harris (2001). In addition special terminology was adopted to the fol-
lowing characters: (i) “eucampto-brochidodromous venation” was used
to designate a composite character that includes the eucamptodromous
and brochidodromous venation types of Rickey’s (1979) within a single
leaf; (ii) “4-merous” and “5-merous” flowers was used to designate
flowers with 4 sepals or 5 sepals, respectively, as the number of corolla
lobes, stamens and staminodes is often different from the number of
sepals; and (iii) “punctate scars on the leaf surface” refers to the tri-
chome scars that are left after the senescence of trichomes. Illustrations
are provided for some species for which no diagnostic illustrations were
Fig. 1. Map of the study area (grey): northern portion of the Atlantic Rainforest of Brazil.
916 SYSTEMATIC BOTANY [Volume 39
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available to date; for the remaining taxa, see illustrations presented in
Pennington (1990), Pennington and Edwards (2005), and Alves-Arau
´jo
and Alves (2011).
Results
Thirty-four species of Pouteria were documented from the
northern portion of the Atlantic Rainforest of Brazil after an
exhaustive and detailed analysis of type collections and over
4,800 specimens. Of the 34 taxa recognized, 18 are endemic
from the Atlantic rainforest, while 10 are disjunctly distributed
in Amazonia (P. bangii,P. cuspidata,P. durlandii subsp.
durlandii,P. egregia,P. franciscana,P. gallifructa,P. glauca,
P. guianensis,P. oblanceolata,andP. venosa subsp. amazonica),
and six are widespread in South America (P. caimito,
P. gardneri,P. macrophylla,P. procera,P. ramiflora,and
P. reticulata). In addition, seven new synonyms are proposed
and 20 species are considered threatned according to the
IUCN red list criteria (IUCN 2012). In particular, 10 are classi-
fied as VU-vulnerable (P. andarahiensis,P. ciliata,P. coelomatica,
P. franciscana,P. gallifructa,P. glauca,P. macahensis,
P. microstrigosa,P. oblanceolata,andP. subsessilifolia), seven as
EN-endangered (P. butyrocarpa,P. confusa,P. oxypetala,
P. pachycalyx, P. aff. stenophylla, P. trifida, and P. velutinicarpa),
three as CR-critically endangered (P. atlantica,P. bapeba,and
P. grandifolia), and 14 as LC-least concern (P. bangii,
P. beaurepairei,P. caimito,P. cuspidata,P. durlandii subsp.
durlandii,P. egregia,P. gardneri,P. guianensis,P. macrophylla,
P. nordestinensis, P. procera,P. ramiflora,P. reticulata,P. venosa
subsp. amazonica).
Taxonomic Treatment
POUTERIA Aubl., Hist. Pl. Guiane 1: 8586. 1775.
Treelets to trees; lenticels present or absent. Leaves spirally
arranged to distichous, lanceolate, oblanceolate, oblong,
elliptic to obovate; base cuneate, acute, attenuate, truncate,
obtuse to rounded; apex acute, attenuate, acuminate, cuspi-
date, apiculate, rounded, obtuse, retuse to emarginate; mar-
gin plane to strongly revolute; venation brochidodromous,
eucamptodromous, or eucampto-brochidodromous. Flowers
4– 5-merous; ramiflorous, axillary or on leafless shoots, gyno-
monoecious, uni- or bisexual. Calyx 4–5 sepals, persistent on
fruits. Corolla cyathiform to tubular, lobes 48. Stamens 48,
always included in the corolla. Staminodes 48, inserted
between corolla lobes. Ovary 116-locular. Fruits 16-
seeded; seeds shiny, brownish, vinaceous to blackish; seed
scar narrow to wide, or even covering up to three fourths of
the surface.
The genus is widespread in Brazil where 120 species occur,
often associated with wet riparian forests within Cerrado,
Amazonia, and the Atlantic Rainforest; some species extend
to drier environments such as the Caatinga and Cerrado.
Key to the Species of POUTERIA From the Northern Portion of the Atlantic Rainforest of Brazil
1. Leaf venation eucamptodromous ................................................................................................ 2
2. Flowers 5-merous . ....................................................................................................... ... 3
3. Lower leaf surface glaucous . . . ........................................................................................... 4
4. Leaves coriaceous, leaf margin revolute, corolla lobes with margins lacerate . ................................... 1.P. andarahiensis
4. Leaves membranaceous to chartaceous, leaf margin plane, corolla lobes
with margins entire or ciliate . . ...................................................................................... 5
5. Sepals 2– 3 mm long, corolla lobes with margins entire . . ................................................... 16.P. gardneri
5. Sepals 4– 5 mm long, corolla lobes with margins ciliate . . ................................................ 21.P. macrophylla
3. Lower leaf surface glabrous, glabrate, pubescent, velutinous to tomentose ...................................................... 6
6. Abaxial side of leaves with punctate scars on the leaf surface ..................................................... 27.P. procera
6. Abaxial side of leaves without punctate scars ............................................................................ 7
7. Flowers sessile, if pedicellate, then pedicels £0.2cmlong .............................................................. 8
8. Lenticels present, corolla cyathiform, staminodes with apex trifid . . . ....................................... 32.P. trifida
8. Lenticels absent, corolla tubular, staminodes without apex trifid ..................................... ... 6.P. butyrocarpa
7. Flowers pedicellate, with pedicels > 0.2 cm long ...................................................................... 9
9. Lower leaf surface velutinous to tomentose, pedicels 12 cm long, anthers pilose ............................ 2.P. atlantica
9. Lower leaf surface glabrate, puberulent or pubescent, pedicels 0.3 0.8 cm long, anthers glabrous ....................... 10
10. Corolla lobes with margins ciliate, ovary 1-locular, stigma punctiform, not lobed ..................... 9.P. coelomatica
10. Corolla lobes with margins entire, ovary 2 5-locular, stigma 2– 5-lobed, not punctiform . . ........................ 11
11. Lower leaf surface puberulent, corolla ca. 2 mm long, stamens inserted
at the lower half of the corolla tube . ................................................... 22.P. microstrigosa
11. Lower leaf surface glabrate, corolla 3.5– 7.0 mm long, stamens inserted
at the upper half of the corolla tube . ......................................... 12.P. durlandii subsp. durlandii
2. Flowers 4-merous . ....................................................................................................... .. 12
12. Lower leaf surface tomentose . .......................................................................................... 13
13. Leaves chartaceous, fruits glabrous . . . ................................................................... 14.P. franciscana
13. Leaves coriaceous, fruits velutinous to lanuginous ....................................................... 33.P. velutinicarpa
12. Lower leaf surface glabrous, glabrate to pubescent . . ....................................................................... 14
14. Corolla 6– 8-lobed ................................................................................................. 15
15. Lower leaf surface glabrous, glaucous, ovary stipitate . . . ........................................... 30.P. aff. stenophylla
15. Lower leaf surface glabrate, pubescent, velutinous to tomentose
(if glabrous, never glaucous), ovary sessile . ................................................................... 16
16. Style constricted at base . . ................................................................................. 17
17. Lenticels present, leaves 7.0 16.0 cm long, corolla lobes with obtuse apices,
ovary with ferruginous trichomes . . ....................................................... 18.P. grandiflora
17. Lenticels absent, leaves 21.0 32.0 cm long, corolla lobes with acute apices,
ovary with golden trichomes . ..... ......................................................... 25.P. oxypetala
2014] ALVES-ARAU
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16. Style straight, and not constricted at the base . . . ............................................................ 18
18. Pedicels 2.8– 3.6 cm long, sepals coriaceous ................................................ 26.P. pachycalyx
18. Pedicels <2 cm long, sepals chartaceous . . . ..................................... 34.P. venosa subsp. amazonica
14. Corolla 4-lobed . . ............................................................................................... 19
19. Anthers pilose ............................................................................................. 20
20. Petioles 1.5–1.9 cm long, pedicels 0.5 1.0 cm long, style papillate . . . .................................... 8.P. ciliata
20. Petioles 2–5 cm long, pedicels 0.1–0.2 cm long, style not papillate . . ............................. 23.P. nordestinensis
19. Anthers glabrous . ......................................................................................... 21
21. Corolla lobes with margin entire .......................................................................... 22
22. Pedicels ca. 0.1 cm long, margin of sepals lacerate . . . ........................................... 10.P. confusa
22. Pedicels > 0.2 cm long, margin of sepals entire . . ........................................................ 23
23. Stamens inserted at the upper half of the corolla tube, 4–10 mm long,
staminodes subulate ................................................................. 28.P. ramiflora
23. Stamens inserted at the lower half of the corolla tube, 2.0–2.5 mm long,
staminodes deltoid to lanceolate .......................................................... 3.P. bangii
21. Corolla lobes with margin ciliate .......................................................................... 24
24. Pedicels 0.5–1.0 cm long . . ........................................................................... 25
25. Sepals of equal length, margins ciliate, staminodes subulate
and glabrous . ..................................................................... 19.P. guianensis
25. Sepals with unequal length, the outer ones shorter than inner ones,
margins entire, staminodes lanceolate, papillate to ciliate ............................... 14.P. franciscana
24. Pedicels £0.2cmlong ............................................................................... 26
26. Margins of inner sepals lacerate, staminodes ciliate ....................................... 5.P. beaurepairei
26. Margins of inner sepals entire, staminodes glabrous . . . ............................................... 27
27. Fruits covered by pilose bristles, style 1–1.5 cm long . . . ................................ 15.P. gallifructa
27. Fruits glabrous to puberulent, style 0.4–0.8 cm long . . . ................................... 7.P. caimito
1. Leaf venation brochidodromous or eucampto-brochidodromous ..................................................................... 28
28. Flowers4-merous ............................................................... .......................................... 29
29. Corolla 6– 8-lobed .................................................................................................... 30
30. Style with a basal constriction ......................................................................... 18.P. grandiflora
30. Style without a basal constriction . . . ........................................................ 34.P. venosa subsp. amazonica
29. Corolla 4-lobed . . .................................................................................................... 31
31. Lower leaf surface glaucous . . ..................................................................................... 32
32. Leaf margin plane, corolla tubular, stamens inserted at the upper half of the corolla tube ...................... 17.P. glauca
32. Leaf margin slightly revolute, corolla cyathiform, stamens inserted at the lower
half of the corolla tube . . . .......................................... ................................ 13.P. egregia
31. Lower leaf surface glabrous, glabrescent to pubescent ................................................................. 33
33. Lenticels present, sepals ca. 1 mm long, ovary 2-locular . . .................................................... 13.P. egregia
33. Lenticels absent, sepals 3–10 mm long, ovary 4-locular ................................................ 19.P. guianensis
28. Flowers5-merous ............................................................... .......................................... 34
34. Flowers on leafless shoots . . ............................................................................... 20.P. macahensis
34. Flowers axillary, never on leafless shoots . . . ............................................................................. 35
35. Leaves coriaceous . . .............................................................................................. 36
36. Lower leaf surface tomentose, petioles 0.1–0.3 cm long, corolla whitish,
staminodes subulate to digitiform . . ........................................................... 31.P. subsessilifolia
36. Lower leaf surface strigillose and glaucous, petioles 1 2 cm long,
corolla greenish, staminodes bifid to falcate (rarely lanceolate) ........................................ 11.P. cuspidata
35. Leaves chartaceous .............................................................................................. 37
37. Sepals velutinous inside . . . ..................................................................... 24.P. oblanceolata
37. Sepals glabrous inside ....................................................................................... 38
38. Lower leaf surface with punctate scars .......................................................... 29.P. reticulata
38. Lower leaf surface without punctate scars . ................................................................ 39
39. Sepals velutinous to tomentose outside, staminodes digitiform to narrowly
lanceolate, fruits brownish, smooth . . ......................................................... 2.P. bapeba
39. Sepals pubescent outside, staminodes lanceolate, fruits yellowish, rugulose . . ....... 12.P. durlandii subsp. durlandii
1. POUTERIA ANDARAHIENSIS T. D. Penn., Fl. Neotrop. 52: 269
270. 1990.—TYPE: BRAZIL. Bahia, Ibiguara, road to
Andarahy, [12] Oct 1942, R. Fro
´es 1020. (holotype: NY!;
isotype: A).
Treelets to trees 414 m tall, shoots tomentose, trichomes
ferruginous, lenticels present. Leaves 3.48.5
+
1.8–3.5 cm,
spirally arranged, obovate to elliptic, coriaceous, glabrous
to glabrate, upper surface sometimes shiny, lower surface
glaucous; margin revolute; venation eucamptodromous;
petiole 0.40.8 cm long, channeled, pubescent. Flowers
5-merous, 2–3 per fascicle, axillary, unisexual; pedicel 0.3
0.5 cm long, puberulent. Sepals 3– 6 mm long, ovate to
lanceolate, chartaceous, glabrous inside, pubescent outside
(mainly at the apex), apex acute; margin ciliate. Corolla
cyathiform, 2.5–3.0 mm long; tube 0.5 1.0 mm long; lobes
1.5–2.0 mm long, ovate, glabrous, greenish, apex rounded,
margin lacerate. Stamens 1.31.5 mm long, inserted at the
upper half of the corolla tube; filaments and anthers gla-
brous. Staminodes 510 mm long, lanceolate, glabrous.
Ovary 2-locular, 0.51.0 mm long, pilose, trichomes golden;
stipe absent; style ca. 1 mm long, glabrous, basal restriction
absent; stigma punctiform to truncate. Fruit 1-seeded, 1.2
1.4 cm long, globoid, smooth, glabrous, black; seeds 1.0–
1.2 mm long, smooth; seed scar 0.8 1.0 cm long, narrow.
Figures 2A– B.
Representative Specimens—BRAZIL. Bahia: A
´gua Quente, Pico das
Almas, 1 Dec 1988, R. Harley et al. 26550 (CEPEC, MO, NY); Andaraı
´,
estrada p/ Mucuge
ˆ, 12 Jan 1983, L. Mattos-Silva et al. 1609 (CEPEC);
Ibicoara, 17 Jan 1996, L. Fe
´lix & O. Moura 4693 (HST); Jacobina, Tombador,
6 Sep 1999, E. Melo et al. 2991 (HUEFS, MO); Morro do Chape
´u, 29 Sep
2002, J. Jardim et al. 4015 (CEPEC); Rio de Contas, Cachoeiras do Fraga, 22
Oct 1997, M. Alves et al. 1386 (IPA, UFP).
918 SYSTEMATIC BOTANY [Volume 39
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Fig. 2. A –O. Species of Pouteria from the northern portion of the Atlantic Rainforest of Brazil. A– B. P. andarahiensis. A. Flower. B. Corolla, stamens,
and staminodes. C– D. P. bapeba. C. Flower. D. Corolla, stamens, and staminodes. E– F. P. beaurepairei. E. Flower. F. Corolla, stamens, and stami-
nodes. G– H. P. butyrocarpa. G. Fruit. H. Seed, frontal (above) and lateral views (below). I– J. P. caimito. I. Flower. J. Corolla, stamens, and staminodes.
K. P. coelomatica. Corolla, stamens, and staminodes. L–M. P. cuspidata. L. Flower. M. Corolla, stamens, and staminodes. N O. P. franciscana. N. Flower.
O. Corolla, stamens, and staminodes. Drawings based on Harley et al. 26550 (A B), T. Santos 1249 (C D), J. Jardim et al. 4475 (E–F), A. Alves-Arau
´jo
et al. 1185 (G– H), A. Alves-Arau
´jo 1124 (I– J), M. Monteiro 23500 (K), F. Santos 622 (L M), and T. Santos 1358 (N O).
2014] ALVES-ARAU
´JO ET AL.: POUTERIA FROM THE ATLANTIC RAINFOREST 919
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Distribution—It is endemic to Chapada Diamantina
(Bahia) in seasonal forests associated with “Campos
rupestres” and “Carrasco” vegetation.
Phenology—Flowering specimens have been observed
in September to December, fruiting specimens in October
to April.
Taxonomic Comments—It can be distinguished from
P. subsessilifolia by its glaucous lower leaf surface, longer
petioles, and lacerate margin of the corolla lobes.
Conservation Status—Pouteria andarahiensis has a
restricted geographical distribution and according to IUCN
(2012) criteria, it is assessed as vulnerable [VU A2(d), B1].
2. POUTERIA ATLANTICA Alves-Arau
´jo & M. Alves, Syst. Bot.
36(4): 1004–1007. 2011.—TYPE: BRAZIL. Bahia, Arataca,
RPPN Palmeira/IESB, 17 Dec 2005, J. Jardim et al. 4909
(holotype: CEPEC!; isotype: NY!).
Trees 812 m tall, shoots velutinous to tomentose, tri-
chomes whitish to golden, lenticels absent. Leaves 1016
+
5.5–7.6 cm, spirally arranged, obovate to elliptic,
chartaceous, upper surface tomentulose to glabrescent
(mainly on the midrib), lower surface velutinous to tomen-
tose; margin plane; venation eucamptodromous; petiole 1.5
2.6 cm long, channeled, velutinous to tomentose. Flowers
5-merous, 1–3 per fascicle, axillary, bisexual; pedicel 12 cm
long, velutinous to tomentose. Sepals 4–5 mm long, ovate,
chartaceous, densely pilose on both surfaces, apex obtuse;
margin entire. Corolla cyathiform, 34 mm long; tube 1.0
1.2 mm long; lobes 2–3 mm long, ovate, densely pilose out-
side, glabrous inside, greenish, apex obtuse; margin slightly
lacerate. Stamens 2.83.2 mm long, inserted at the upper
half of corolla tube; filaments glabrous; anthers pilose.
Staminodes 2.02.2 mm long, lanceolate, pilose. Ovary
2-locular, 1.0–1.2 mm long, pilose, trichomes whitish to
golden; stipe absent; style 1.7– 2.0 mm long, glabrous, basal
restriction absent; stigma slightly 2-lobate. Fruit 1-seeded,
1.0– 2.5 cm long, obovoid, smooth, pilose (immature) to gla-
brescent (mature), brownish; seeds 0.8– 1.8 cm long, smooth;
seed scar 0.6–1.4 mm long, narrow.
Representative Specimens—BRAZIL. Bahia: Una, ReBio Mico-Lea
˜o-
Dourado [15090S, 39050W], 25 Jul 1996, S. Santana et al. 624 (CEPEC, G,
HRB, MO, NY); 01 Jun 2000, S. Santana et al. 888 (CEPEC, NY).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (southern Bahia).
Phenology—Flowering specimens have been observed in
December and June, fruiting specimens in June to July.
Taxonomic Comments—It differs from P. gardneri by its
densely pilose shoots, leaves, and sepals, velutinous to
tomentose pedicel, and pilose anthers.
Conservation Status—Pouteria atlantica shows a restricted
geographical distribution and, despite the fact that there are
different populations within legally protected areas, the spec-
imens were collected near forest edges, which are at high risk
of further disturbance, especially by logging. Therefore, fol-
lowing IUCN (2012) criteria, it is assessed as critically endan-
gered [CR A1(cde), B1, D2)].
3. POUTERIA BANGII (Rusby) T. D. Penn., Fl. Neotrop. 52: 288.
1990. Sideroxylon bangii Rusby, Bull. New York Bot.
Gard. 4: 407. 1907.—TYPE: BOLIVIA. without precise
locality, without date, M. Bang 1953 (holotype: NY!;
isotypes: F!, GH, K!, M!, MO!, NY!, US!, W).
Trees 8–15 m tall, shoots pubescent, trichomes ferruginous
to golden, lenticels absent. Leaves 6.823.0
+
2.7–7.8 cm,
distichous to spirally arranged, elliptic to lanceolate,
chartaceous to coriaceous, upper surface glabrous to
glabrate, lower surface glabrate to pubescent; margin plane;
venation eucamptodromous; petiole 1.5–2.5 cm long, terete,
glabrate to pubescent. Flowers 4-merous, 614 per fascicle,
axillary, bisexual; pedicel 0.51.0 cm long, puberulent. Sepals
1.5–2.0 mm long, ovate, chartaceous, glabrous inside, puber-
ulent outside, apex obtuse; margin entire. Corolla
cyathiform, 2.53.5 mm long; tube ca. 2 mm long; lobes
0.75– 1.5 mm long, ovate, glabrous, greenish to pinkish, apex
slightly acute to obtuse; margin entire. Stamens 1.5–2.5 mm
long, inserted at the lower half of corolla tube; filaments and
anthers glabrous. Staminodes 0.51.0 mm long, deltoid to
lanceolate, glabrous. Ovary 2-locular, ca. 1 mm long, pilose,
trichomes ferruginous; stipe absent; style 1.01.5 mm long,
glabrous, basal restriction absent; stigma punctiform to
2-lobate. Fruit 1-seeded, 35 cm long, ellipsoid to ovoid,
smooth, glabrous, yellowish; seeds 24 cm long, rugulose;
seed scar 1.83.6 cm long, narrow.
Representative Specimens—BRAZIL. Alagoas: Murici, Est. Ecol.
Murici, 11 Feb 2009, A. Alves-Arau
´jo et al. 1152 (UFP); Bahia: Itabuna/
Ilhe
´us, A
´rea do CEPEC, 19 Jun 2008, A. Alves-Arau
´jo 1014 (CEPEC);
Uruc¸ uca, Parque Est. Serra do Conduru
´, 27 Mar 2009, A. Alves-Arau
´jo &
L. Lanev 1205 (UFP); Ceara
´: Guaramiranga, 2 Jun 2004, V. Gomes et al. 2063
(EAC); Pernambuco: Igarassu, Usina Sa
˜o Jose
´, Mata da Piedade, 26 Dec
2008, A. Alves-Arau
´jo & B. S. Amorim 1087 (IPA, UFP).
Distribution—It is disjunct between Atlantic (from Ceara
´
to Rio de Janeiro) and Amazonian forests (Bolivia, Brazil,
Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname,
and Venezuela).
Phenology—Flowering specimens have been collected and
observed in October to February, fruiting specimens in
December to June.
Taxonomic Comments—It can be distinguished from
P. microstrigosa by its greenish to pinkish and 4-merous
flowers; and from P. coelomatica by the entire margin of
corolla lobes and stamens being inserted at the lower half of
the corolla tube.
Conservation Status—Pouteria bangii is widely distributed
in the Amazon and Atlantic Rainforest in the South America
and according to IUCN (2012) criteria, it is assessed as of least
concern (LC).
4. POUTERIA BAPEBA T. D. Penn., Fl. Neotrop. 52: 328 329.
1990.—TYPE: BRAZIL. Bahia, Itamaraju-Sa
˜oPaulinho,
[17 Sep 1968], J. Almeida & T. Santos 58 (holotype: FHO;
isotype: CEPEC!).
Trees 10–15 m tall, shoots glabrous to velutinous to tomen-
tose, trichomes brownish, lenticels absent. Leaves 13– 22
+
7–
10 cm, spirally arranged, obovate to oblanceolate, chartaceous,
upper surface glabrous, lower surface tomentulose; margin
slightly revolute to plane; venation brochidodromous; petiole
1.6– 2.2 cm long, channeled, tomentose. Flowers 5-merous, 3–8
per fascicle, axillary, gyno-monoecious or bisexual; pedicel
0.3– 0.5 cm long, tomentose. Sepals 3.5– 4.0 mm long, ovate,
chartaceous, glabrous inside, velutinous to tomentose outside,
apex rounded to obtuse; margin entire. Corolla cyathiform, 6–
8 mm long; tube 3–4 mm long; lobes 3– 4 mm long, rotund,
glabrous, greenish, apex rounded to truncate; margin slightly
lacerate. Stamens 2.0 2.5 mm long, inserted at the upper half
of corolla tube; filaments and anthers glabrous. Staminodes
2.0– 2.5 mm long, digitiform to narrowly lanceolate, glabrous.
Ovary 5-locular, 1.8– 2.0 mm long, pilose, trichomes brownish;
stipe absent; style 2.0–2.5 mm long, glabrous, basal restriction
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absent; stigma 5-lobed. Fruit 1–2-seeded, 3– 4 cm long,
obovoid to ellipsoid, smooth, velutinous, brownish; seeds
2.0– 2.5 cm long, smooth; seed scar 1.0– 2.2 mm long, wide.
Figures 2C–D.
Representative Specimens—BRAZIL. Bahia: Belmonte, Est. Exp. de
Belmonte-CEPLAC, 18 Sep 1970, T. Santos 1108 (CEPEC); Ipiau
´, estrada
p/ Jequie
´, 31 Oct 1970, T. Santos 1249 (CEPEC, FHO); Jussari, Rod.
Jussari/Palmira-RPPN Serra do Teimoso, 14 Jan 2000, J. Jardim et al. 2362
(CEPEC); 9 Feb 1998, W. Thomas et al. 11785 (CEPEC, NY); Santa Cruz de
Cabra
´lia, Est. Ecol. Pau-Brasil, 10 Dec 1987, F. Santos 775 (CEPEC).
Espı
´rito Santo: Reserva Florestal da CRVD, Est. Flamengo, 30 Nov 1981,
I. Silva 273 (K).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (southern Bahia and
northern Espı
´rito Santo).
Phenology—Flowering specimens have been observed
in September to October, fruiting specimens in October
to November.
Taxonomic Comments—It can be distinguished from
P. butyrocarpa by its longer pedicels (0.3–0.5 cm long),
digitiform to narrowly lanceolate staminodes, smooth and
velutinous fruits.
Conservation Status—Pouteria bapeba has a restricted geo-
graphical distribution and the area of occurrence has a high
risk of disturbance, especially by logging. Following IUCN
(2012) criteria, it is assessed as critically endangered [CR (A1
(cde), B1, D2].
5. POUTERIA BEAUREPAIREI (Glaz. & Raunk.) Baehni, Candollea 9:
241. 1942.—TYPE: BRAZIL. Rio de Janeiro, [29 Jan 1887],
M. Glaziou 16241 [holotype: P! (P00640530); isotypes:
BR!, F! (F0BN004230, F0072025F), G!, IAN, K!, MO!,
NY!, P! (P00640532, P00640531)].
Treelets to trees 810 m tall, shoots glabrate to pubescent,
trichomes whitish to golden, lenticels absent. Leaves 1015
+
3.5–5.0 cm, spirally arranged (rarely distichous), oblanceo-
late to lanceolate, chartaceous (rarely membranaceous),
glabrous; margin plane; venation eucamptodromous; petiole
1–2 cm long, terete, glabrous. Flowers 4-merous, 13 per
fascicle, axillary, bisexual; pedicel ca. 0.1 cm long, pubescent.
Sepals 3.5–4.0 mm long, ovate, membranaceous, glabrous
inside, pubescent outside, apex rounded to obtuse; margin
lacerate (inner ones). Corolla tubular, 3.54.5 mm long; tube
2.5–3.0 mm long; lobes 1.0 1.5 mm long, rotund, ciliate,
greenish, apex truncate; margin entire. Stamens 2.02.5 mm
long, inserted at the middle of corolla tube; filaments and
anthers glabrous. Staminodes ca. 1 mm long, lanceolate, cili-
ate. Ovary 4-locular, 1.82.0 mm long, pilose, trichomes
golden; stipe absent; style 2.02.5 mm long, glabrous, basal
restriction absent; stigma 4-lobed. Fruit 12-seeded, 34 cm
long, obovoid to globoid, smooth, glabrous to puberulent,
yellowish to brownish; seeds 1.5–2.0 cm long, smooth; seed
scar 1.21.7 cm long, wide. Figures 2EF.
Representative Specimens—BRAZIL. Bahia: Camaca
˜, RPPN Serra
Bonita, entrada para a RRPN e torre de transmissa
˜o, 13 Feb 2005, J. Jardim
et al. 4475 (CEPEC, HUEFS); Ilhe
´us, CEPEC, 16 Mar 1983, T. Santos 3854
(CEPEC, K); Porto Seguro, Est. Vera Cruz, 13 Oct 2006, A. Amorim et al.
6427 (CEPEC, HUEFS); Una, Rod. Una/Canavieiras, 20 Jul 1994, J. Jardim
505 et al. (CEPEC, K).
Distribution—The species is widely distributed through
the Atlantic Rainforest (from Bahia to Santa Catarina).
Phenology—Flowering specimens have been observed in
February to July, fruiting specimens in March to October.
Taxonomic Comments—It can be distinguished from
Pouteria subcaerulea Pierre ex Dubard, a cerrado species, by
its habit (geoxylic in P. subcaerulea and arboreal in P.
beaurepairei), glabrous leaves, and shorter pedicels. In the
Atlantic Rainforest, P. beaurepairei can be recognized by the
4-merous flowers associated with a ciliate corolla, glabrous
stamens, and ciliate staminodes.
Conservation Status—It is widely distributed in the Atlantic
Rainforest, so even considering the human impact on it,
according to IUCN (2012) criteria Pouteria beaurepairei is
assessed as least concern (LC).
6. POUTERIA BUTYROCARPA (Kuhlmann) T. D. Penn., Fl. Neotrop.
52: 501. 1990. Lucuma butyrocarpa Kuhlmann, Arq. Inst.
Biol. Veg. 3: 46. 1936.—TYPE: BRAZIL. Espı
´rito Santo,
Rio Doce, Colonia e Goytacazes, [7 Apr] [14] May 1934,
J. Kuhlmann 341 (lectotype, designated by Pennington
1990: 501, RB!; isolectotype: G!).
Trees 20–35 m tall, shoots pubescent, trichomes ferrugi-
nous, lenticels absent. Leaves 1328
+
3.4–11.0 cm, spirally
arranged, oblanceolate, chartaceous to coriaceous, glabrous;
margin plane; venation eucamptodromous; petiole 1.3
3.2 cm long, terete, pubescent. Flowers 5-merous, 35 per
fascicle, axillary, bisexual, sessile. Sepals 3.55.0 mm long,
ovate to lanceolate, chartaceous, glabrous inside, pubescent
outside, apex acute or obtuse; margin entire. Corolla tubular,
6– 8 mm long; tube 4–5 mm long; lobes 2– 3 mm long, oblong,
glabrous, greenish, apex rounded; margin entire. Stamens
2.5–3.0 mm long, inserted at the upper half of corolla tube;
filaments and anthers glabrous. Staminodes 2.0– 2.5 mm
long, lanceolate, glabrous. Ovary 5-locular, 23 mm long,
pilose, trichomes ferruginous; stipe absent; style 3.54.5 mm
long, glabrous, basal restriction absent; stigma 5-lobed. Fruit
1– 4-seeded, 3 5 cm long, ovoid to globoid, verruculose, gla-
brous, yellowish; seeds 2.74.0 cm long, smooth; seed scar
2.5–3.5 cm long, wide. Figures 2G H.
Representative Specimens—BRAZIL. Bahia: Arataca, 18 Jan 2006,
W. Thomas et al. 14516 (CEPEC, NY); Camaca
˜, RPPN Serra Bonita, 2 Apr
2009, A. Alves-Arau
´jo et al. 1185 (CEPEC, UFP); Entre Bom Gosto e
Olivenc¸ a, 15 Mar 1943, R. Fro
´es 20025 (NY); Fazenda Canaa
˜, Rio Mucuri,
19 Oct 1946, R. Bondar s. n. (G, NY-375364); Jussari, Rod. Jussari/Palmira-
Serra/Faz.Teimoso, 29 Mar 2002, J. Paixa
˜o 165 & Dove Backe (CEPEC, NY);
Maraca
´s, 5 May 1979, S. Mori et al. 11799 (CEPEC, NY). Espı
´rito Santo:
Colatina, Rio Doce, 19 Sep 1930, J. Kuhlmann 359 (CEPEC, MO, RB).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (Bahia and Espı
´rito Santo).
Phenology—Flowering specimens have been observed
in September to December, fruiting specimens in October
to May.
Taxonomic Comments—It can be distinguished from
P. bapeba by its 5-merous and sessile flowers, and verruculose
and glabrous fruits.
Conservation Status—Pouteria butyrocarpa has a restricted
geographical distribution and appears to be rare based on the
paucity of known specimens. Following IUCN (2012) criteria,
it is assessed as endangered [EN A1(cd), B2(a)].
7. POUTERIA CAIMITO (Ruiz & Pavo
´n) Radlk., Sitzungsber.
Math.-Phys. Cl. Ko
¨nigl. Bayer. Akad. Wiss. Mu
¨nchen
12 (3): 333. 1882. Achras caimito Ruiz & Pavo
´n, Fl. Peruv.
3: 18, t. 240. 1802.—TYPE: PERU. without precise local-
ity, without date, J. Pavo
´n[holotype: MA!; isotypes: BC!,
BR!, F! (F0042252F, F0042253F), FI!, G!].
Treelets to trees 3–8 m tall, shoots glabrous to glabrate,
lenticels absent. Leaves 517
+
2– 4 cm, spirally arranged,
elliptic to lanceolate, membranaceous to chartaceous, glabrous;
2014] ALVES-ARAU
´JO ET AL.: POUTERIA FROM THE ATLANTIC RAINFOREST 921
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margin plane; venation eucamptodromous; petiole 0.5–2.0 cm
long, terete, glabrous. Flowers 4-merous, 13 per fascicle, axil-
lary or ramiflorous, bisexual; pedicel 0.10.2 cm long, gla-
brous. Sepals 35 mm long, ovate to elliptic, chartaceous,
glabrous inside, glabrate outside, apex obtuse; margin entire.
Corolla tubular, 4–7 mm long; tube 3–5 mm long; lobes 1–
2 mm long, ovate to rotund, glabrous, greenish, apex obtuse to
truncate; margin ciliate. Stamens 2.5–4.5 mm long, inserted at
the middle of corolla tube; filaments and anthers glabrous.
Staminodes 1.0–1.5 mm long, lanceolate, glabrous. Ovary 4-
locular, 2.0–2.5 mm long, pilose, trichomes whitish to golden;
stipe absent; style 0.4–0.8 cm long, glabrous, basal restriction
absent; stigma 4-lobate. Fruit 1–4-seeded, 3–8 cm long, ellip-
soid to globoid, smooth, glabrous to puberulent, yellowish;
seeds 2–3 cm long, smooth; seed scar 1.8–2.7 mm long, nar-
row. Figures 2I– J.
Representative Specimens—BRAZIL. Bahia: Alcobac¸a, 16 Sep 1978, T.
Santos 3327 (CEPEC, NY); Ilhe
´us, CEPEC, 20 Jan 1971, R. Pinheiro 1011
(CEPEC, RB); Uruc¸ uca, estrada Uruc¸uca-Parque Est. Serra do Conduru
´,
27 Mar 2009, A. Alves-Arau
´jo 1203 (UFP). Paraı
´ba: Areia, Fazenda Sabura
´,
4 Oct 1993, L. Fe
´lix 6167 (EAN); Pernambuco: Igarassu, Usina Sa
˜o Jose
´,
Mata dos Macacos, 12 Dec 2002, I. Sa
´e Silva et al. 148 (PEUFR, UFP);
Paudalho, Mata do Condomı
´nio Santa Maria, 12 Feb 2009, A. Alves-Arau
´jo
1337 (UFP); Recife, Mata de Dois Irma
˜os, 13 Feb 1990, M. Guedes 2254
(ASE). Minas Gerais: Belo Horizonte, Campus da UFMG, 5 Dec 2008, A.
Alves-Arau
´jo 1124 (UFP).
Distribution—It is widespread in the Neotropics and usu-
ally cultivated because of its edible fruits. In the northern
Atlantic Rainforest, it is known from wet forests (from
Paraı
´ba to Bahia).
Phenology—Flowering specimens have been collected and
observed in August to November, fruiting specimens in Sep-
tember to December.
Taxonomic Comments—Pouteria caimito is easily confused
with P. guianensis and P. gallifructa, from which it can be
distinguished by its large (3– 8 cm), glabrous to puberulent,
and yellowish fruits.
Conservation Status—Pouteria caimito has a wide distribu-
tion in the New World tropics, and according to IUCN (2012)
criteria, is assessed as of least concern (LC).
8. POUTERIA CILIATA Alves-Arau
´jo & M. Alves, Nord. J. Botany
30: 399406. 2012.—TYPE: BRAZIL. Pernambuco, Sa
˜o
Lourenc¸o da Mata, Estac¸a
˜o Ecolo
´gica de Tapacura
´,3
Aug 2001, T. da Silva & K. Almeida 67 (holotype: PEUFR!;
isotype: NY!).
Trees 8–12 m tall, shoots glabrous, lenticels present. Leaves
12–17
+
5.0–6.5 cm, spirally arranged, lanceolate, chartaceous,
glabrous; margin plane; venation eucamptodromous; petiole
1.5– 2.0 cm long, channeled, glabrous. Flowers 4-merous, 10-
many per fascicle, axillary, gyno-monoecious or bisexual; ped-
icel 0.5–1.0 cm long, glabrous. Sepals 2.53.0 mm long, ovate,
membranaceous, glabrous inside, sericeous outside, apex
rounded to acute; margin ciliate (inner ones). Corolla tubular,
3.5– 5.5 mm long; tube 2– 4 mm long; lobes 1.5– 2.0 mm long,
rotund, glabrous, whitish, apex truncate; margin ciliate. Sta-
mens 2.3–2.7 mm long, inserted at the middle of corolla tube,
glabrous; filaments glabrous; anthers pilose. Staminodes ca.
1 mm long, lanceolate, ciliate. Ovary 4-locular, 1.0 1.5 mm
long, pilose, trichomes whitish to golden; stipe absent; style 2
3 mm long, papillate, basal restriction absent; stigma 4-lobed.
Fruit 1– 2-seeded, 3– 4 cm long, ovoid to ellipsoid, glabrous,
yellowish; seeds 1.52.0 cm long, smooth, shiny, brownish;
seed scar 1.21.7 mm long, narrow.
Representative Specimens—BRAZIL. Bahia: Muritiba, May 1950, G.
Pinto 50/33 (ALCB); Ceara
´: Guaramiranga, Pico Alto, 20 Mar 2009, E.
Silveira & Otı
´lia s. n. (EAC-45012).
Distribution—It is endemic to the northern Atlantic
Rainforest (Ceara
´, Pernambuco and Bahia).
Phenology—Flowering specimens have been observed in
March to May, fruiting specimens in April to August.
Taxonomic Comments—Pouteria ciliata is similar to the
Amazonian species Pouteria fimbriata and P. penicillata from
which it differs by the shorter petioles (1.5 2.0 cm) and lon-
ger pedicels (0.5–1.0 cm). In the Atlantic Rainforest, it can be
recognized by whitish and 4-merous flowers, with a ciliate
corolla and pilose anthers.
Conservation Status—Pouteria ciliata is poorly represented
in the collections. The extent of occurrence (EOO) of the spe-
cies is rather large, but since most of the forests between
these points are devastated, the area of occupancy (AOO) is
reduced to less than 20 km
2
. Therefore, according to IUCN
(2012) criteria it is assessed as vulnerable [VU A1(c), D2].
9. POUTERIA COELOMATICA Rizzini, Rodrigue
´sia 28: 174. 1976.—
TYPE: BRAZIL. Bahia, Itamaraju, Fazenda Boa Vista,
24 Jul 1971, M. Monteiro 23500 (holotype: RB!; isotype:
PEUFR!).
Trees 10–20 m tall, shoots puberulent, trichomes ferrugi-
nous, lenticels absent. Leaves 8– 17
+
3.56.3 cm, spirally
arranged, oblanceolate to lanceolate, chartaceous to coria-
ceous, upper surface glabrous to puberulent on midrib, lower
surface pubescent; margin plane; venation eucamptodromous;
petiole 1.6–3.5 cm long, terete, puberulent. Flowers 5-merous,
3– 8 per fascicle, axillary, bisexual; pedicel 0.4– 0.8 cm, pubes-
cent. Sepals 1.8–2.6 mm long, ovate to orbicular, chartaceous,
glabrous inside, puberulent outside, apex obtuse to rounded;
margin ciliate. Corolla cyathiform, 3–4 mm long; tube 0.5–
1.0 mm long; lobes 2.5– 3.0 mm long, ovate, glabrous, greenish,
apex rounded; margin ciliate. Stamens 1– 2 mm long, inserted
at the upper half of corolla tube; filaments and anthers gla-
brous. Staminodes 0.8–1.0 mm long, lanceolate, glabrous.
Ovary 1-locular, 2– 3 mm long, pilose, trichomes ferruginous;
stipe absent; style 1 mm long, glabrous, basal restriction
absent; stigma punctiform. Fruit 1-seeded, 2.0–2.5 cm long,
obovoid, smooth, glabrate to pubescent, yellowish; seeds 1.5–
2.0 cm long, smooth; seed scar 1.0– 1.5 cm long, narrow.
Additional Specimens—BRAZIL. Espı
´rito Santo: Conceic¸a
˜o da Barra,
25 Oct 1993, O. Pereira et al. 5144 (RB, VIES); Guarapari, Setiba, 24 Sep
1992, L. Fabris 811 (RB); Guarapari, Parque Est. Paulo Ce
´sar Vinha, 23 Aug
1998, A. Assis 555 (RB); Presidente Kennedy, 18 May 1994, C. Farney 3368
(RB); Vila Velha, Interlagos, 11 Oct 1996, O. Zambom 304 (RB); Rio de
Janeiro: Jacarepagua
´, Represa de Camorim, 19 Jan 1987, A. Peixoto et al.
4003 (MO); Nova Iguac¸u
´, ReBio do Tingua
´, 6 Dec 1995, P. Fara
´g 189 (RB).
Distribution—It is endemic to the Atlantic Rainforest
where it is known from wet forests (from Bahia to Rio
de Janeiro).
Phenology—Flowering specimens have been observed in
July to September, fruiting specimens in August to January.
Taxonomic Comments—It can be confused with P. bangii
but differs by the ciliate margin of the corolla lobes, stamens
inserted near to TO, and 1-locular ovary.
Conservation Status—Pouteria coelmoatica has a restricted
geographical distribution and according to IUCN (2012)
criteria, it is assessed as vulnerable [VU A1(c), D2].
10. POUTERIA CONFUSA Alves-Arau
´jo & M. Alves, Nord. J. Bot-
any 30: 399–406. 2012.—TYPE: BRAZIL. Bahia, Uruc¸uca,
Escola Me
´dia de Agropecua
´ria da regia
˜o Cacaueira,
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Reserva Grego
´rio Bondar, 20 May 1994, W. Thomas et al.
10424 (holotype: CEPEC!; isotypes: G!, MO!, NY!).
Treelets to trees 4–10 m tall, shoots glabrous to tomentose,
trichomes golden to ferruginous, lenticels present. Leaves
12–16
+
5.5–7.5 cm, distichous, obovate to elliptic,
membranaceous to chartaceous, upper surface glabrous, usu-
ally shiny, lower surface glabrous to strigillose; margin
plane; venation eucamptodromous; petiole 1.02.3 cm long,
channeled, glabrous to tomentose. Flowers 4-merous, 25 per
fascicle, axillary, gyno-monoecious or bisexual; pedicel ca.
0.1 cm long, tomentose. Sepals 2.53.0 mm long, ovate,
chartaceous, tomentose both surfaces (outer ones glabrous
inside); apex rounded; margin lacerate. Corolla cyathiform,
3–4 mm long; tube 2 3 mm long; lobes ca. 1 mm long,
rotund, glabrous, greenish, apex rounded to truncate; margin
entire. Stamens ca. 2 mm long, inserted at the middle of
corolla tube; filaments and anthers glabrous. Staminodes 1
2 mm long, lanceolate, glabrous. Ovary 4-locular, 1.01.2 mm
long, pilose, trichomes whitish to golden; stipe absent; style
1.0–1.5 mm long, glabrous, basal restriction absent; stigma 4-
lobed. Fruit 14-seeded, 34 cm long, ovoid, smooth, gla-
brous, yellowish; seeds 2.02.8 cm long, smooth; seed scar
0.4–0.6 cm long, narrow.
Representative Specimens—BRAZIL. Bahia: Itacare
´, Fazenda das
Almas, 27 Jan 1982, T. Santos et al. 3724 (CEPEC); Itapebi, Fazenda
Lombardia/Fazenda Ventania BR-101, 19 Aug 1971, T. Santos 1863
(CEPEC); Santo Anto
ˆnio de Jesus, Rod. S. Miguel das Matas/Amargosa,
30 Jan 1993, J. Pirani & J. Kallunki 2715 (CEPEC).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (Bahia).
Phenology—Flowering specimens have been observed in
January to March, fruiting specimens in July.
Taxonomic Comments—It can be easily distinguished from
P. oblanceolata by the distichous leaves, outer sepals that are
glabrous inside, and stamens inserted at the middle portion
of the corolla tube. Pouteria confusa can be confused with
Chrysophyllum gonocarpum (Mart. & Eich. ex Miq.) Engl., from
which it differs by the wider (5.57.5 cm) and shiny adaxial
leaf surface, 4-merous flowers, and presence of staminodes.
Conservation Status—Pouteria confusa is recorded from
four different populations which are all restricted to wet
vegetation. The individuals are rare and sparsely distributed.
Most of the forests between those occurrences are now dev-
astated and P. confusa shows a restricted extent of occurrence
(EOO <5,000 km
2
) and area of occupancy (AOO > 500 km
2
).
According to IUCN (2012), it is assessed as endangered [EN
A1(cde), B1].
11. POUTERIA CUSPIDATA (A. DC.) Baehni, Candollea 9: 231
232. 1942. Sideroxylon cuspidatum A. DC., Prodr. 8: 183.
1844.—TYPE: GUYANA. without precise locality, 1838,
R. H. Schomburgk [318] 518 (holotype: G!; isotypes: BM,
BR, F!, G!, K!, U, W).
Sideroxylon cuspidatum A. DC. var. ellipticum Miq. in Martius, Fl
bras. 7: 56. 1863.—LECTOTYPE, designated here: GUYANA.
1841, R. H. Schomburgk [172] 147 [G!; isolectotypes:
BM! (BM000952538, BM000952541), BR, F! (F0072239F,
F0072240F), P! (P00640559, P00640558, P00640557), US!].
Pouteria cuspidata subsp. dura (Eyma) T. D. Penn., Fl. Neotrop.
52: 352. 1990. Sideroxylon cuspidatum var. crassifolium
Miq. in Mart., Fl. bras. 7: 56. 1863 —LECTOTYPE, desig-
nated here: GUYANA. Banks of Pomeroon R., [1843],
R. H. Schomburkg 1470 (K; isolectotype: P). syn. nov.
Neoxythece schulzii Aubre
´v., Adansonia 4: 231. 1964.—TYPE:
SURINAME. Middle Wayombo, Sep 1955, Schulz 7327
(holotype: P!; isotypes: K!, U).
Neoxythece wurdackii Aubre
´v., Adansonia 5: 201. 1965.—
TYPE: PERU. Amazonas, Prov. Bagua, right bank of R.
Santiago, Oct 1962, Wurdack 2409 (holotype: US!;
isotypes: F!, K!, NY!).
Pouteria cuspidata subsp. robusta (Mart. & Eich.) T. D. Penn.,
Fl. Neotrop. 52: 351. 1990. Sideroxylon robustum Mart. &
Eich. in Mart., Fl. bras. 7: 56. 1863.—LECTOTYPE, desig-
nated here: [VENEZUELA]. Cassiquiari: Vasiva et
Pacimoni, [Jan 1854], R. Spruce 3331 [K! (K000641132);
isolectotypes: BM!, BR!, E!, F!, G!, GOET, K
(K000641133), NY!, OXF, P! (P00640564, P00640563,
P00640562, P00640565), RB!]. syn. nov.
Treelets to trees 420 m tall, shoots strigillose, trichomes
golden, lenticels usually present. Leaves 8.513
+
3.8
6.0 cm, spirally arranged, obovate to oblanceolate, coriaceous,
upper surface glabrous to glabrate, lower surface strigillose,
glaucous; margin revolute; venation brochidodromous; peti-
ole 1–2 cm long, slightly channeled, glabrate. Flowers
5-merous, 6–20 per fascicle, axillary, bisexual; pedicel 0.5
0.9 cm long, strigillose. Sepals 34 mm, deltoid, chartaceous,
glabrous inside, strigillose outside, apex acute; margin entire.
Corolla cyathiform, 23 mm long; tube 1 mm long; lobes ca.
2 mm long, ovate, glabrous, greenish, apex rounded to cuspi-
date; margin slightly lacerate. Stamens 0.91.1 mm long,
inserted at the upper half or the middle of corolla tube; fila-
ments and anthers glabrous. Staminodes 1.01.2 mm long,
bifid to falcate (rarely lanceolate), glabrous. Ovary 2-locular,
ca. 1 mm long, pilose, trichomes golden; stipe absent; style ca.
1 mm long, glabrous, basal restriction absent; stigma 2-lobed.
Fruit 1-seeded, 2.8–3.5 cm long, ovoid, smooth, strigillose,
orange; seeds 1.5– 1.8 cm long, smooth; seed scar 1.4 1.7 cm
long, narrow. Figures 2LM.
Representative Specimens—BRAZIL. Bahia: Canavieiras, estrada
Canavieiras/Una, 18 Feb 2003, P. Fiaschi et al. 1332 (CEPEC); Ilhe
´us,
CEPEC, 12 Apr 2004, R. Sambuichi 558 (CEPEC); Itacare
´, 20 Nov 1991,
A. Amorim et al. 374 (CEPEC); Porto Seguro, Estac¸a
˜o Vera Cruz, 01 Aug
1999, L. Paraguassu
´et al. 124 (ALCB); Salvador, Jardim Bota
ˆnico, 12 Mar
2003, E. Queiroz & P. Carvalho 417 (HRB); Uruc¸uca, Faz. Lagoa do
Conjunto/Faz. Santa Cruz, 1– 12 Jul 1991, W. Thomas et al. 8800 (CEPEC,
NY); Santa Cruz de Cabra
´lia, Est. Ecol. Pau-Brasil, 15 Jul 1987, F. Santos
622 (CEPEC, NY).
Nomenclature—The type of Pouteria cuspidata subsp. dura
is the specimen collected by R. H. Schomburkg 1470 from
Guyana which was initially thought to be deposited at P
(Eyma 1936), and later thought to be deposited at K
(Pennington 1990). However, no specimens were located at
either herbarium and based on Pennington’s (1990) informa-
tion, it’s probable that the holotype deposited at B was
destroyed. Although we cannot check the R. H. Schomburkg
1470 voucher, a new synonym is proposed based on Schulz
7327 and Wurdack 2409 which were considered by Pennington
(1990) as synonyms of Pouteria cuspidata subsp. dura. Subspe-
cies of Pouteria cuspidata have been published based mainly on
the leaf apex, presence or absence of glaucous lower leaf sur-
face and, finally, position of staminodes. However, there are
huge variations concerning overlapping of all these characters
and it is hardly possible to identify the Pouteria cuspidata sam-
ples with certainty. For this reason, and after detecting several
vouchers which show intermediary features, the names of
subspecies are being synonymized.
2014] ALVES-ARAU
´JO ET AL.: POUTERIA FROM THE ATLANTIC RAINFOREST 923
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Distribution—It’s known distribution is in Atlantic (Bahia,
Espı
´rito Santo, and Rio de Janeiro) and Amazonian forests
(Brazil, Colombia, Guyana, Peru, and Venezuela).
Phenology—Flowering specimens have been observed
in September to February, fruiting specimens in October
to August.
Taxonomic Comments—It can be easily recognized by its
coriaceous leaves with a glaucous lower surface, and bifid to
falcate (rarely lanceolate) staminodes. There is huge variation
concerning leaf size and shape, and position of stamens on
the corolla tube. Pouteria cuspidata can be confused with P.
egregia and Manilkara species as well, but its 5-merous
flowers provide distinctive characteristic to distinguish it
from them.
Status—Pouteria cuspidata is widely distributed in the
Amazon and Atlantic Rainforest in South America and
according to IUCN (2012) criteria, it is assessed as of least
concern (LC).
12. POUTERIA DURLANDII (Standl.) Baehni subsp. DURLANDII,
Candollea 9: 422– 423. 1942. Lucuma durlandii Standl.,
Trop. Woods 4: 5. 1925.—TYPE: GUATEMALA. Pete
´n,
El Paso, 1925, W. Durland s. n. (holotype: US-1208271!).
Trees 10– 30 m tall, shoots pubescent, trichomes golden, lenti-
celsabsent.Leaves8–21
+
3.5– 7.5 cm, spirally arranged, oblan-
ceolate to elliptic, chartaceous, upper surface glabrous, lower
surface glabrate; margin plane; venation eucamptodromous
or eucampto-brochidodromous; petiole 1–2 cm long, terete,
glabrous. Flowers 5-merous, 2– 4 per fascicle, axillary or
ramiflorous, unisexual; pedicel 0.4–0.8 cm, pubescent. Sepals
2– 4 mm long, ovate, chartaceous, glabrous inside, pubescent
outside, apex acute; margin entire. Corolla cyathiform, 3.5–
7.0 mm long; tube 1.5–2.0 mm long; lobes 2– 5 mm long, ovate,
glabrous, greenish, apex rounded; margin entire. Stamens 2–
5 mm long, inserted at the upper half of corolla tube; filaments
and anthers glabrous. Staminodes 1.0–3.2 mm long, lanceo-
late, glabrous. Ovary 2 5-locular, 2–3 mm long, puberulent,
trichomes golden; stipe absent; style 1.5–4.0 mm long,
glabrous, basal restriction absent; stigma 2–5-lobed. Fruit 1–
5-seeded, 4–6 cm long, globoid, rugulose, glabrous to glabrate,
yellowish; seeds 3.5–4.5 cm long, slightly rugulose; seed scar
2.2– 3.2 cm long, wide. Figures 3A–B.
Representative Specimens—BRAZIL. Alagoas: Mata Grande, 22 Oct
1981, L. Gonc¸alves 248 (RB); Bahia: Camaca
˜, 6 Apr 1979, S. Mori & T. Santos
11713 (CEPEC, NY); Jussari, RPPN Serra do Teimoso, 14 Jan 2000,
J. Jardim et al. 2414 (CEPEC, NY); Santo Anto
ˆnio de Jesus, Rod. Sa
˜o Miguel
das Matas/Amargosa, 30 Jan 1993, J. Pirani & J. Kallunki 2726 (CEPEC,
K, MO, NY); Una, ReBio Mico-Lea
˜o, 26 Apr 1994, A. Carvalho et al. 4545
(CEPEC, NY); Pernambuco: Igarassu, Usina Sa
˜o Jose
´, 9 Mar 2010,
A. Alves-Arau
´jo et al. 1331 (UFP); Recife, Jardim Bota
ˆnico, 19 May 2010,
A. Alves-Arau
´jo et al. 1335 (UFP).
Distribution—It is widespread in the Neotropics in wet
forests (in northern Atlantic Rainforest, from Pernambuco to
Bahia), different from Pouteria durlandii (Standl.) Baehni subsp.
pubicarpa T. D. Penn. which is found only in Venezuela
and Peru.
Phenology—Flowering specimens have been collected and
observed in September to December, fruiting specimens in
November to April.
Taxonomic Comments—It can be distinguished from the
other subspecies by the rugulose and glabrous surface of the
fruits. The latter feature is associated with 5-merous flowers,
glabrous anthers, and 25-locular ovary can distinguish
P. durlandii subsp. durlandii from P. nordestinensis.
Conservation Status—Pouteria durlandii subsp. durlandii is
widely distributed in the New World tropics and following
IUCN (2012) criteria, it is assessed as of least concern (LC).
13. POUTERIA EGREGIA Sandwith, Bull. Misc. Inform 1931: 479.
1931.—TYPE: GUYANA. Essequibo river, Moraballi
Creek, [08] Nov 1929, Sandwith 573 (holotype: K (not
found); isotypes: F!, G!, NY!, P!, RB!, U!, US!, W).
Trees 10–25 m tall, shoots glabrous, lenticels present.
Leaves 5–10
+
2.0–3.6 cm, spirally arranged, oblanceolate to
elliptic, chartaceous, upper surface glabrous, lower surface
glabrescent, glaucous or not; margin slightly revolute; vena-
tion brochidodromous; petiole 0.41.2 cm long, channeled,
glabrous. Flowers 4-merous, 48 per fascicle, axillary, bisex-
ual; pedicel 0.2– 0.8 cm long, pubescent. Sepals ca. 1 mm
long, ovate, chartaceous, glabrous inside, glabrescent out-
side, apex acute to rounded; margin ciliate. Corolla
cyathiform, 1.62.0 mm long; tube 0.81.0 mm long; lobes
0.8–1.0 mm long, glabrous, whitish, apex rounded; margin
entire. Stamens 12 mm long, inserted at the lower half of
corolla tube; filaments and anthers glabrous. Staminodes
0.5–1.0 mm long, lanceolate, glabrous. Ovary 2-locular, 0.5
1.2 mm long, pilose, trichomes golden; stipe absent; style
ca. 1 mm long, glabrous, basal restriction absent; stigma
punctiform. Fruit 1-seeded, 2–3 cm long, ellipsoid, smooth,
glabrous, yellowish; seeds 1.5– 2.6 cm long, smooth; seed scar
2– 3 cm long, narrow.
Representative Specimens—BRAZIL. Bahia: Una, ReBio Mico-Lea
˜o, 20
Jan 1999, J. G. Jardim et al. 1965 (ALCB, CEPEC, HRB, K, MO, NY); Una,
ReBio Mico-Lea
˜o, 22– 28 Oct 1999, J. G. Jardim et al. 2266 (CEPEC, NY);
Pernambuco: Igarassu, Usina Sa
˜o Jose
´, Mata da Piedade, 27 Sep 2001,
S. G. Freire & H. C. Silva 19 (PEUFR, UFP). Amazonas: Sa
˜o Gabriel, 14 Jan
1942, R. L. Fro
´es 518 (G); Para
´: Rodovia Bele
´m-Brası
´lia, Km 93, 11 Sep
1959, M. Kuhlmann & S. Jimbo 221 (HUEFS, RB, UFP).
Distribution—Pouteria egregia has a disjunct distribution
in the Atlantic (Bahia and Pernambuco) and Amazonian
(Brazil, Colombia, French Guiana, Guyana, Suriname, and
Venezuela) forests.
Phenology—Flowering specimens have been observed in
October to December, fruiting specimens in December
to February.
Taxonomic Comments—Vouchers from the Amazonian
region always bear a glaucous lower leaf surface which is
not true for the ones from the Atlantic forest. It can be distin-
guished from Pouteria cuspidata by its 4-merous flowers.
Conservation Status—Pouteria egregia is widely distrib-
uted in the Amazon and Atlantic Rainforest in the South
America and according to IUCN (2012) criteria, it is assessed
as of least concern (LC).
14. POUTERIA FRANCISCANA Baehni, Candollea 9: 262. 1942.—
TYPE: BRAZIL. Acre, Rio Acre, Seringal Sa
˜o Francisco,
Oct 1911, Ule 9692 (holotype: L; isotypes: G!, MG,
K!, NY!).
Trees 10–30 m tall, shoots glabrous to tomentose, tri-
chomes golden to brownish, lenticels absent. Leaves 15
37
+
7.0–11.6 cm, spirally arranged, lanceolate, chartaceous,
both surfaces glabrous to tomentose; margin plane; venation
eucamptodromous; petiole 1.23.0 cm long, channeled,
tomentose to glabrous. Flowers 4-merous, 612 per fascicle,
axillary, bisexual; pedicel 0.71.0 cm long, tomentose. Sepals
2.7–3.0 (inner ones) to 45 mm long (outer ones), ovate,
chartaceous, glabrous inside, velutinous to tomentose out-
side, apex acute (inner) to rounded (outer); margin entire.
924 SYSTEMATIC BOTANY [Volume 39
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Fig. 3. A –P. Species of Pouteria from the northern portion of the Atlantic Rainforest of Brazil. A–B. P. durlandii subsp. durlandii. A. Branch. B. Seed.
C. P. gallifructa. Fruit. D– E. P. glauca. D. Flower. E. Corolla, stamens, and staminodes. F– H. P. grandiflora. F. Corolla, stamens, and staminodes. G. Ovary.
H. Ovary, cross section. I– J. P. macahensis. I. Flowers on leafless shoot. J. Fruits on leafless shoot. K M. P. macrophylla. K. Flower. L. Ovary. M. Corolla,
stamens, and staminodes. N– P. P. microstrigosa. N. Floral branch. O. Ovary. P. Ovary, cross section. Drawings based on A. Alves-Arau
´jo et al. 1335 (A– B),
A. Alves-Arau
´jo & B. Amorim 1303 (C), T. Santos 2993 (D–E), M. Salzmann 316 (F H), A. Alves-Arau
´jo 1094 (I), A. Alves-Arau
´jo 1009 (J), T. Santos 1229 (K–
M), and S. Mori et al. 13974 (NP).
2014] ALVES-ARAU
´JO ET AL.: POUTERIA FROM THE ATLANTIC RAINFOREST 925
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Corolla tubular, 6–8 mm long; tube 4.0 5.5 mm long; lobes
2.0–2.5 mm long, slightly unguiculate, glabrous, whitish,
apex rounded to irregular; margin ciliate. Stamens 2.7
3.0 mm long, inserted at the middle of corolla tube; filaments
and anthers glabrous. Staminodes 1.52.0 mm long, lanceo-
late, papillate to ciliate. Ovary 4-locular, 2.53.0 mm long,
pilose, trichomes golden; stipe absent; style 56 mm long,
upper third glabrous, basal restriction absent; stigma slightly
4-lobed. Fruit 1-seeded, 56 cm long, ovoid to ellipsoid,
smooth, glabrous, yellowish; seeds 23 cm long, smooth;
seed scar 1.72.5 cm long, wide. Figures 2NO.
Representative Specimens—BRAZIL. Bahia: Camaca
˜, Estrada de Pau
Brasil, 19 Jan 1971, T. Santos 1358 (CEPEC). BRAZIL. Acre: Boca do Acre,
Rio Purus, 16 Sep 1976, G. Prance et al. 2342 (NY, P); Amazonas: Humaita
´,
14 Sep to 9– 11 Oct 1934, B. Krukoff 6211 (NY, RB); Para
´: Rio Tajapuru, Ilha
Sa
˜o Sebastia
˜o e Nazare
´, 21 July 1948, G. Black 48–3009 (G, NY); Rondo
ˆnia:
Estrada Nova Vida para Vilhena, 6 Sep 1963, B. Maguire et al. 56595 (NY).
Distribution—Pouteria franciscana has a disjunct distribu-
tion in the Atlantic (Bahia) and Amazonian (Acre, Amazonas,
Para
´, and Rondo
ˆnia) forests in Brazil.
Phenology—Atlantic Rainforest flowering specimens
have been observed in January, other additional flowering
specimens in August to September, fruiting specimens
in October.
Taxonomic Comments—Vouchers from the Amazonian
region sometimes have entire corolla lobe margins. It can be
distinguished from Pouteria pisquiensis Baehni, an Amazo-
nian species, by its longer petioles, velutinous to tomentose
sepals (outside) and shorter pedicels. Among Atlantic
Rainforest species, P. franciscana stands out by its long-
pedicellate flowers (0.7– 1.0 cm long) and outer sepals shorter
than inner ones.
Conservation Status—Despite its disjunct distribution,
Pouteria franciscana is hardly found in the collections and
appears to be a rare species in the field. Therefore it is
assessed as vulnerable [VU A2(bcd)].
15. POUTERIA GALLIFRUCTA Cronq., Lloydia 9(4): 285286. 1946.
Pouteria torta (Mart.) Radlk. subsp. gallifructa (Cronq.)
T. D. Penn., Fl. Neotrop. 52: 487. 1990.—TYPE: BELIZE.
Temash River, Mar 1935, W. Schipp 1323. (holotype: NY!;
isotypes: A, BM, F!, GH, M!, MICH, MO!).
Trees 10–30 m tall, shoots glabrous to pubescent, tri-
chomes ferruginous, lenticels absent. Leaves 619
+
5.0
12.5 cm, spirally arranged, oblanceolate, chartaceous, upper
surface glabrous, lower surface pubescent (rarely glabrous),
margin plane; venation eucamptodromous; petiole 13 cm
long, slightly channeled, pubescent (rarely glabrous).
Flowers 4-merous, 13 per fascicle, ramiflorous, bisexual;
pedicel ca. 0.1 cm long, puberulent. Sepals 515 mm long,
ovate to elliptic, chartaceous, glabrous inside, puberulent
outside; apex obtuse; margin entire. Corolla tubular, 8
15 mm long; tube 510 mm long; lobes 35 mm long, ovate,
glabrous, greenish to yellowish, apex obtuse to truncate; mar-
gin ciliate. Stamens 12 mm long, inserted at the middle of
corolla tube; filaments and anthers glabrous. Staminodes ca.
1 mm long, lanceolate, glabrous. Ovary 4-locular, 2.02.5 mm
long, pilose, trichomes ferruginous; stipe absent; style 1.0
1.5 cm long, glabrous, basal restriction absent; stigma 4-lobed.
Fruit 1–4-seeded, 3–5 cm long, conic, covered by pilose bris-
tles, reddish to brownish; seeds 2–3 cm long, smooth; seed
scar 1.3–2.5 cm long, narrow. Figure 3C.
Representative Specimens—BRAZIL. Alagoas: Ibateguara, Coimbra,
2 May 2002, M. Oliveira & A. Grilo 1575 (UFP); Bahia: Itabuna, 25 Jul
1967, R. Pinheiro 136 (CEPEC, MO); Santa Cruz de Cabra
´lia, 17 Oct 1978,
S. Mori et al. 10771 (CEPEC, NY, RB); Una, ReBio Mico-Lea
˜o, 15 Jul 1993,
J. Jardim et al. 223 (CEPEC, MO, NY, RB); Pernambuco: Igarassu, Usina
Sa
˜oJose
´, Mata da Piedade, 2 Mar 2009, A. Alves-Arau
´jo & A. Melo 1172
(UFP); Recife, Mata de Dois Irma
˜os, 22 May 2008, A. Alves-Arau
´jo &
A. Melo 997 (UFP); Rio Formoso, ReBio Saltinho, 28 May 2009, A. Alves-
Arau
´jo & B. Amorim 1303 (UFP); Sergipe: Areia Branca, PARNA Serra de
Itabaiana, 5 Jan 2009, A. Alves-Arau
´jo et al. 1097 (UFP).
Distribution—It has a disjunct distribution between Atlantic
Rainforest (from Pernambuco to Bahia) and Central America.
Phenology—Flowering specimens have been collected and
observed in November to March, fruiting specimens in Janu-
ary to July.
Taxonomic Comments—The vegetative parts are similar
to Pouteria guianensis and P. caimito but only the fruits of P.
gallifructa bear pilose bristles. Pouteria torta subsp. torta is a
typical treelet from Brazilian Cerrado areas and P. torta
subsp. tuberculata occurs in the Amazonian region. The
firstoneshowsavillouslowerleafsurfacewhichisdiffer-
ent on the latter (pilose). Both of them, however, bear
malpighiaceous trichomes on leaf surfaces and pilose fruits.
The pilose bristles on the fruits and pubescent lower leaf
surface of P. gallifructa are the most distinguishible and stable
features to recognize it as a distinct species.
Conservation Status—Pouteria gallifructa shows two
regions of distribution (Central America and Northeastern
Brazil). However, the areas where it can be found are
impacted, mainly by logging. Therefore it is assessed as vul-
nerable [VU A2(bcd)].
16. POUTERIA GARDNERI (Mart. & Miq.) Baehni, Candollea 9:
233. 1942. Chrysophyllum gardneri Mart. & Miq. in Martius,
Eichler & Urban, Fl. bras. 7: 102. 1863.— LECTOTYPE,
designated here: BRAZIL. Piauı
´, 1841, M. Gardner 2659
(M!; isolectotypes: F!, G!, K!, NY!, P!, US!).
Pouteria peduncularis (Mart. & Eich.) Baehni Candollea 9: 357.
1942. Lucuma ?peduncularis Mart. & Eich. in Martius,
Eichler & Urban, Fl. bras. 7: 73. 1863.—LECTOTYPE,
designated here: [BRAZIL]. [Bahia], [Jacobina], [without
date], Blanchet 3598 (G!; isolectotypes: C!, F!, M!, P!, RB!,
U).syn.nov.
Trees 12–20 m tall, shoots pubescent, trichomes whitish to
golden, lenticels present. Leaves 4.813.0
+
1.8– 4.0 cm, spi-
rally arranged, elliptic to oblanceolate, membranaceous to
chartaceous, glabrous to pubescent, lower surface glaucous;
margin plane; venation eucamptodromous; petiole 0.5–1.0 cm
long, terete, pubescent. Flowers 5-merous, 13 per fascicle,
axillary, gyno-monoecious or bisexual; pedicel 0.6–2.5 cm
long, puberulent. Sepals 2–3 mm long, ovate, chartaceous,
glabrous inside, puberulent outside, apex obtuse; margin
entire. Corolla cyathiform, 3– 4 mm long; tube 1.0–1.5 mm
long; lobes 2.0– 2.5 mm long, ovate, glabrous, greenish, apex
obtuse; margin entire. Stamens 2.0–2.5 mm long, inserted at
the upper half of corolla tube; filaments and anthers glabrous.
Staminodes 1.5– 2.0 mm long, lanceolate, glabrous. Ovary 2–3-
locular, 1.5–2.0 mm long, pubescent, trichomes whitish to
golden; stipe absent; style 1.0–1.5 mm long, glabrous, basal
restriction absent; stigma slightly 2-lobate. Fruit 1-seeded, 1–
2 cm long, ellipsoid to globoid, smooth, glabrous to pubescent,
yellowish; seeds 1.0–1.5 cm long, smooth; seed scar 0.8– 1.2 cm
long, narrow.
Representative Specimens—BRAZIL. Alagoas: Quebrangulo, ReBio da
Pedra Talhada, 12 May 2009, A. Alves-Arau
´jo et al. 1257 (UFP); Murici,
Estac¸a
˜o Ecolo
´gica de Murici, 10 Feb 2009, A. Alves-Arau
´jo et al. 1142
926 SYSTEMATIC BOTANY [Volume 39
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(UFP); Bahia: Cravola
ˆndia, Estrada Cravola
ˆndia/Ubaı
´ra, 5 May 2007, J.
Jardim et al. 5098 (CEPEC, NY); Ceara
´: Ubajara, Serra do Ibiapaba, 28 Jan
1968, Andrade-Lima 68– 5197 (IPA); Paraı
´ba: Joa
˜o Pessoa, UFPB, 22 Apr
2007, F. Queiroz 06 (JPB); Pernambuco: Igarassu, Usina Sa
˜o Jose
´, Mata da
Piedade, 21 Nov 2008, A. Alves-Arau
´jo et al. 1073 (IPA, UFP); Sergipe:
Areia Branca, PARNA da Serra de Itabaiana, 7 Jan 2009, A. Alves-Arau
´jo
& S. Martins 1116 (UFP).
Distribution—It is widely distributed in South America
from wet forests (northern Atlantic Rainforest from Ceara
´to
Bahia) to “Matas ciliares” in the Brazilian Cerrado.
Phenology—Flowering specimens have been collected and
observed in September to December, fruiting specimens in
October to May.
Taxonomic Comments—It differs from P. atlantica by the
presence of lenticels on the shoots, glaucous lower leaf sur-
face, and glabrous corolla and stamens. The name Pouteria
peduncularis has been published based mainly on the length
of the pedicels found on a voucher from Bahia state, Brazil.
However, after analyzing several vouchers from that location
and surrounding areas, it is easily confirmed that it is not a
different taxon but variation found within the range of
Pouteria gardneri.
Conservation Status—Pouteria gardneri is widely distrib-
uted in the South America and according to IUCN (2012)
criteria, it is assessed as of least concern (LC).
17. POUTERIA GLAUCA T. D. Penn., Fl. Neotrop. 52: 263265.
1990.—TYPE: PERU. Loreto, Rı
´o Nanay, Mishana, Jul
1977, Solomon 3511 (holotype: MO!).
Trees 8–30 m tall, shoots glabrous to glabrate, lenticels
present. Leaves 14.0–25.0
+
3.0– 6.8 cm, spirally arranged,
lanceolate to oblanceolate, chartaceous, glabrous, lower sur-
face glaucous; margin plane; venation brochidodromous
or eucampto-brochidodromous; petiole 0.81.6 cm long,
channeled (rarely winged), glabrous to glabrate. Flowers 4-
merous, 23 per fascicle, axillary, bisexual; pedicel 0.5
0.7 cm long, glabrous to glabrate. Sepals 1.8–2.0 mm long,
ovate, chartaceous, glabrous inside, glabrous to glabrate out-
side, apex rounded to obtuse; margin ciliate (inner sepals).
Corolla tubular, 4–5 mm long; tube ca. 2 mm long; lobes 2
3 mm long, ovate, glabrous, greenish, apex obtuse to trun-
cate; margin ciliate (rarely entire). Stamens 1.82.0 mm long,
inserted at the upper half of corolla tube; filaments and
anthers glabrous. Staminodes ca. 2 mm long, lanceolate, gla-
brous. Ovary 24-locular, 0.8–1.0 mm long, puberulent, tri-
chomes golden; stipe absent; style ca. 1 mm long, glabrous,
basal restriction absent; stigma 24-lobate. Fruit 1-seeded, 3
4 cm long, ellipsoid, smooth, pubescent, yellowish; seeds
2.0–2.5 cm long, smooth; seed scar 1.8 2.4 cm long, narrow.
Figures 3D– E.
Representative Specimens—BRAZIL. Bahia: Belmonte, 18 Apr 1975,
T. Santos 2993 (CEPEC, MO). Roraima: Serrinha, Rio Mucajaı
´, 31 Jan
1967, G. Prance et al. 4216 (MO). ECUADOR. Pastaza: Pozo petrolero
“Garza” de TENNECO, noreste de Montalvo, 2–12 Jul 1989, Vlastimil
Zak & S. Espinoza 4406 (MO, NY); PERU. Amazonas: Bagua, 20 Oct 1995,
R. Vasquez et al. 20349 (MO); Pasco: Oxapampa, Dist. Huancabamba,
21 May 2004, J. Perea et al. 1321 (MO); Loreto: Maynas, Iquitos, Rio Nanay,
13 Dec 1977, M. Rimachi 3289 (MO).
Distribution—It has a disjunct distribution between Atlantic
(Bahia) and Amazonian forests (Bolivia, Brazil, French Guiana,
Peru, and Suriname).
Phenology—Atlantic Rainforest flowering specimens have
been observed in April, other additional flowering specimens
in January to March, fruiting specimens in April to July.
Taxonomic Comments—Pouteria glauca differs from
P. macrophylla by its 4-merous flowers and glabrous style.
Conservation Status—Despite its disjunct distribution,
Pouteria glauca is not easily found in the collections and
appears to be a rare species in the field. Therefore it is
assessed as vulnerable [VU A2(bcd)].
18. POUTERIA GRANDIFLORA (A. DC.) Baehni, Candollea 9: 391.
1942. Lucuma grandiflora A. de Candolle in A. P. de
Candolle, Prodr. 8: 169. 1844.—TYPE: BRAZIL. Bahia,
without exact locality, [1830], M. Salzmann 316 (lectotype,
designated by Baehni 1942: 39, G!; isolectotype: P!).
Trees 5–6 m tall, shoots pubescent, trichomes ferruginous,
lenticels present. Leaves 7–16
+
3–6 cm, spirally arranged,
oblanceolate to elliptic, coriaceous, glabrous to glabrate;
margin revolute; venation eucamptodromous (rarely
brochidodromous); petiole 0.6–3.0 cm long, terete, glabrous.
Flowers 4-merous, 2–10 per fascicle, axillary, bisexual; pedi-
cel 0.6–1.4 cm long, puberulent. Sepals 5–12 mm long, ovate
to orbicular, chartaceous, glabrous inside, puberulent outside,
apex obtuse; margin entire (outer sepals) to basally lacerate
(inner ones). Corolla tubular, 9–16 mm long; tube 611 mm
long; lobes 6–8, 3– 5 mm long, ovate to rotund, papillate with
trichomes outside, greenish, apex obtuse; margin entire. Sta-
mens 6–8, inserted at the upper half of corolla tube, 3–4 mm
long; filaments papillate; anthers glabrous. Staminodes 6 8,
3– 4 mm long, lanceolate to subulate, papillate. Ovary 6–10-
locular, 4–5 mm long, pilose, trichomes ferruginous; stipe
absent; style 6–10 mm long, pubescent basally, basal constric-
tion present; stigma 6–10-lobed. Fruit 1– 6-seeded, 3–5 cm
long, globoid to obovoid, smooth, puberulent (immature) to
glabrous (mature), yellowish; seeds 1.5–3.0 cm long, rugu-
lose; seed scar covering up to three fourths of the surface.
Figures 3F–H.
Representative Specimens—BRAZIL. Bahia: Cairu, Garapua
´, 13 Aug
1993, M. Guedes s. n. (ALCB-26017); Camac¸ari, Jaua
´, 15 Aug 1992,
T. Barbosa 2 (ALCB); Conde, Fazenda do Bu, 11 Sep 1996, T. Jost &
M. Ferreira 398 (ALCB, IPA); Conde, Sı
´tio do Conde, 17 Apr 2004, T. Jost
et al. 626 (ALCB, HRB, HUEFS); Salvador, Dunas do Abaete
´, 18 Aug 1996,
M. Ferrucci et al. 1093 (CEPEC, G, HRB, K); Salvador, Dunas do Abaete
´,
2 Apr 2008, J. Cairo & F. Queiroz s. n. (ALCB-83281); Salvador, Lagoa do
Abaete
´, 24 Oct 1997, M. Guedes 5534 (ALCB, CEPEC); Salvador, Dunas de
Itapoa
˜, 30 Sep 1985, M. Guedes & P. Souza 962 (ALCB, CEPEC); Salvador,
Itapoa
˜, 12 Dec 1985, L. Norblick & I. Brito 4471 (CEPEC, K).
Distribution—It is endemic to coastal areas usually with
white-sandy soils (restingas) in eastern Brazil (Bahia).
Phenology—Flowering specimens have been observed
in September to February, fruiting specimens in October
to April.
Taxonomic Comments—It is usually confused with
Pouteria venosa subsp. amazonica but P. grandiflora has smaller
and never bullate leaves and the style bears a basal constric-
tion; P. velutinicarpa differs mainly by its glabrous fruits
when mature.
Conservation Status—Pouteria grandiflora has a narrowly
restricted geographical distribution and the area of occur-
rence has a high risk of disturbance, especially by logging.
Following IUCN (2012) criteria, it is assessed as critically
endangered [CR A1(bcd), B2].
19. POUTERIA GUIANENSIS Aubl., Hist. Pl. Guiane 1: 86. 1775.—
TYPE: FRENCH GUIANA. Without exact locality, with-
out date, Aublet s. n. (holotype: P!; isotype: BM).
Pouteria hispida Eyma, Recueil Trav. Bot. Ne
´erl. 33: 177. 1936.—
TYPE: SURINAME. Nr. Wonotombo, R. Corantyn, [13]
Oct 1916, BW 2863 (holotype: U; isotypes: G!, K, MO!,
NY!). syn. nov.
2014] ALVES-ARAU
´JO ET AL.: POUTERIA FROM THE ATLANTIC RAINFOREST 927
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Pouteria torta (Mart.) Radlk. subsp. glabra T. D. Penn., Fl.
Neotrop. 52: 484. 1990.—TYPE: ECUADOR. Zamora-
Chinchipe, Yantzaza, [15] Nov 1982, T. Pennington &
Tenorio 10745 (holotype: K!; isotypes: NY!, QCA).
syn. nov.
Trees up to 40 m tall, shoots glabrate to pilose, trichomes
ferruginous, lenticels absent. Leaves 6–35
+
2.5–12.0 cm,
spirally arranged, oblanceolate to elliptic, chartaceous to cori-
aceous, upper surface glabrous to pilose on midrib, lower
surface glabrous to pubescent; margin plane; venation
eucamptodromous (rarely eucampto-brochidodromous);
petiole 1–7 cm long, channeled, glabrous to pilose. Flowers
4-merous, 3–10 per fascicle, axillary or ramiflorous, bisexual;
pedicel 0.5–0.6 cm, pubescent. Sepals 3 10 mm long, ovate,
chartaceous, glabrous inside, pubescent outside, apex obtuse
to rounded; margin ciliate. Corolla tubular, 5 15 mm long;
tube 4–10 mm long; lobes 1 5 mm long, ovate to oblong,
glabrous, greenish, apex rounded to truncate; margin ciliate.
Stamens 3.0–6.5 mm long, inserted at the middle of corolla
tube; filaments and anthers glabrous. Staminodes 0.5
2.0 mm long, subulate, glabrous. Ovary 4-locular, 35 mm
long, pilose, trichomes ferruginous; stipe absent; style 3
12 mm long, glabrous, basal restriction absent; stigma 4-
lobed. Fruit 1–4-seeded, 3–9 cm long, globoid, glabrous to
puberulent, yellowish to orange; seeds 25 cm long, smooth;
seed scar 1.03.1 cm long, narrow to wide.
Representative Specimens—BRAZIL. Bahia: Ilhe
´us, CEPEC, 8 Apr
1981, J. Hage & H. Brito 586 (CEPEC); Lenc¸o
´is, Caminho p/ Mata de
Remanso, 30 Jan 1997, S. Atkins et al. 4721 (ALCB, CEPEC, K); Una, ReBio
Mico-Lea
˜o, 6 Jun 1997, J. Jardim et al. 1085 (CEPEC, NY); Valenc¸a, RPPN
A
´gua Branca, 30 Oct 2004, P. Fiaschi et al. 2620 (CEPEC, K). Amazonas:
Manaus, Reserva Ducke, 7 May 1958, L. Coelho s. n. (INPA-6491, P); Minas
Gerais: Araguari, Bosque John Kennedy, 5 Sep 1992, G. Arau
´jo 956 (MO,
NY); Parana
´: Morretes, 4 Oct 1968, G. Hatscbach 19899 (MBM, MO, NY, Z);
Rio de Janeiro: Rio de Janeiro, Tijuca, 10 Jun 1971, M. Glaziou s. n. (P).
Distribution—It is widely distributed in wet forests in the
Neotropics [Amazonian and Atlantic (Bahia) forests, and gal-
lery forest in Cerrado].
Phenology—Flowering specimens have been observed in
January to April, fruiting specimens in March to October.
Taxonomic Comments—It can be confused with P. caimito
and P. gallifructa, but it differs from them mainly by its longer
pedicels and subulate staminodes. Both names, Pouteria
hispida and P. torta subsp. glabra were published mainly
based on the trichome length on the upper leaf surface. The
first one bears longer trichomes instead of shorter trichomes
or rarely absent on upper leaf surface on the latter. After
analyzing their types and several samples from different
regions, and comparing to the type of Pouteria guianensis,it
is clearly a case of vegetative variation that can be confirmed
when the fruits are compared. The fruits bear a continuum
from completely glabrous to pubescent surfaces and it is a
common feature in P. guianensis.
Conservation Status—Pouteria guianensis is widely distrib-
uted in South America and according to IUCN (2012) criteria,
it is assessed as of least concern (LC).
20. POUTERIA MACAHENSIS T. D. Penn., Fl. Neotrop. 52: 321322.
1990. Lucuma macahensis Glaziou, Bull. Soc. Bot. France
57 (Mem. 3): 439. 1910. nom. nud.—TYPE: BRAZIL. Rio
de Janeiro, [Alto Macahe
´], [10 Jan 1891], Glaziou 18353
(holotype: P!; isotypes: BR, G!, K!, M!).
Trees 10–20 m tall, shoots glabrous to puberulent, tri-
chomes pale, lenticels present. Leaves 7– 22
+
3.8–10.0 cm,
spirally arranged, elliptic to oblanceolate, chartaceous to cori-
aceous, upper surface glabrous, lower surface puberulous
(mainly on the midrib); margin slightly revolute; venation
brochidodromous; petiole 0.5– 3.2 cm long, slightly channeled,
glabrous. Flowers 5-merous, 1– 2 per fascicle, on leafless
shoots, unisexual; pedicel 0.1 0.6 cm long, pubescent. Sepals
ca. 2 mm, lanceolate, chartaceous, glabrous inside, puberu-
lent outside, apex acute to attenuate; margin entire. Corolla
cyathiform, ca. 2 mm long; tube ca. 1 mm long; lobes ca.
1 mm long, ovate to oblanceolate, glabrous, greenish, apex
rounded; margin entire. Stamens 58 mm long, inserted at
the upper half of corolla tube; filaments and anthers gla-
brous. Staminodes 45 mm long, lanceolate, glabrous. Ovary
2-locular, 12 mm long, pubescent, trichomes golden; stipe
absent; style ca. 1 mm long, glabrous, basal restriction absent;
stigma slightly 2-lobed. Fruit 1-seeded, 2.56.0 cm long,
obovoid to pyriform, scurfy surface, brownish; seeds 1.0
2.4 cm long, smooth; seed scar 0.8– 2.2 cm long, narrow.
Figures 3I–J.
Representative Specimens—BRAZIL. Alagoas: Piac¸abuc¸ u, Mata da
Marreca, 12 Aug 1987, G. Esteves et al. 1841 (MAC); Bahia: Itacare
´, trilha
apo
´s travessia da balsa, 15 Jun 2008, A. Alves-Arau
´jo et al. 1009 (UFP);
Santa Cruz de Cabra
´lia, Est. Ecol. Pau-Brasil, 2 Jul 1978, S. Mori 12218
(CEPEC); Pernambuco: Tamandare
´, restinga de Ariquinda
´, 9 Apr 2003,
S. Lira et al. 548 (IPA); Sergipe: Areia Branca, 5 Jan 2009, A. Alves-Arau
´jo
et al. 1094 (UFP). Espı
´rito Santo: Linhares, Rio Doce, 1 Oct 1930,
J. Kuhlmann 441 (RB); Rio de Janeiro: Rio das Ostras, ReBio Unia
˜o,
27 Mar 1997, P. Oliveira 775 (BHCB).
Distribution—It is endemic to coastal areas usually with
white-sandy soils (restingas) in eastern Brazil (from Pernambuco
to Rio de Janeiro).
Phenology—Flowering specimens have been collected
and observed in January to April, fruiting specimens in
March to October.
Taxonomic Comments—It differs from P. ramiflora by the
presence of lenticels on shoots, brochidodromous leaf vena-
tion, and 5-merous flowers.
Conservation Status—Pouteria macahensis has a restricted
geographical distribution and the area of occurrence has a
high risk of disturbance. Following IUCN (2012) criteria, it is
assessed as vulnerable [VU A1(bcd), B2].
21. POUTERIA MACROPHYLLA (Lam.) Eyma, Recueil Trav. Bot.
Ne
´erl. 33: 164. 1936. Chrysophyllum macrophyllum Lam.,
Tab. Encycl. 2: 44. 1794.—TYPE: FRENCH GUIANA.
[Cayenne, without date] Stoupe s. n. (holotype: P!).
Trees 10–30 m tall, shoots puberulent, trichomes golden,
lenticels present. Leaves 1319
+
4.3–8.0 cm, spirally
arranged, elliptic to oblanceolate, chartaceous, upper surface
glabrate, lower surface pubescent, glaucous; margin plane;
venation eucamptodromous; petiole 1.52.4 cm long, slightly
channeled, pubescent. Flowers 5-merous, 28 per fascicle,
axillary, bisexual; pedicel 1.01.5 cm long, puberulent. Sepals
4–5 mm long, ovate, chartaceous, glabrous inside, puberu-
lent outside, apex rounded (inner sepals) to acute (outer
sepals); margin entire (outer sepals) to lacerate or ciliate
(inner sepals). Corolla tubular, 57 mm long; tube 3– 4 mm
long; lobes 23 mm long, lanceolate, glabrous, greenish, apex
rounded; margin ciliate. Stamens 1.82.8 mm long, inserted
at the upper half of corolla tube; filaments and anthers gla-
brous. Staminodes 2.7–3.0 mm long, subulate, papillate.
Ovary 5-locular, 1.82.0 mm long, puberulent, trichomes
golden to ferruginous; stipe absent; style 4.55.0 mm long,
papillate, basal restriction absent; stigma 5-lobed. Fruit 12-
seeded, 2–3 cm long, globoid, smooth, glabrous, yellowish;
928 SYSTEMATIC BOTANY [Volume 39
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
seeds 1.7–2.4 cm long, smooth; seed scar 1.4 2.1 cm long,
wide, covering one third of the surface. Figures 3K M.
Representative Specimens—BRAZIL. Bahia: Itaibo
´, 29 Oct 1970, T. Santos
1229 (CEPEC, MO); Una, ReBio Mico-Lea
˜o, 11 Aug 1999, L. Mattos-Silva
et al. 4007 (CEPEC, NY); Ceara
´: Granja, Serra de Ubatuba, 10 Sep 2005,
A. Castro 1624 (EAC); Ubajara, Ibiapaba, 11 Dec 1981, J. Jangoux et al. 1819
(MO, NY); Ubajara, Ibiapaba, 24 Sep 1998, A. Fernandes & E. Nunes s. n.
(EAC-27712). Maranha
˜o: Estrada de Carolina para Estreita, 9 Aug 1964,
G. Prance & N. Silva 58616 (G, NY, RB); Loreto, 20 Nov 2005, A. Miranda
et al. 5273 (HST, IPA); Maracassume
´, 23 Aug 1933, R. Fro
´es 1863 (P, S); Santa
Quite
´ria, Faz. Marflora, 7 Sep 1993, B. Pereira 2525 (RB).
Distribution—It is widely distributed in the Neotropics
[Amazonian and Atlantic (Bahia, Ceara
´) forests, and gallery
forests in Cerrado].
Phenology—Flowering specimens have been observed
in August to November, fruiting specimens in September
to February.
Taxonomic Comments—It differs from P. glauca by its 5-
merous flowers, subulate staminodes, and papillate style.
Conservation Status—Pouteria macrophylla is widely dis-
tributed in South America and according to IUCN (2012)
criteria, it is assessed as of least concern (LC).
22. POUTERIA MICROSTRIGOSA T. D. Penn., Fl. Neotrop. 52: 278.
1990.—TYPE: BRAZIL. Minas Gerais, Marlieria, Parque
Estadual do Rio Doce, [18] Sep 1975, E. P. Heringer &
G. Eiten 15090 (holotype: MO!; isotype: UB).
Trees 15– 40 m tall, shoots glabrous to puberulent, tri-
chomes ferruginous, lenticels usually present. Leaves 8.5
16.0
+
3.0–5.5 cm, spirally arranged, elliptic to oblanceolate,
chartaceous to coriaceous, upper surface glabrous to
glabrate, lower surface puberulent; margin plane; venation
eucamptodromous; petiole 1.01.5 cm long, channeled, gla-
brous. Flowers 5-merous, 2–15 per fascicle, axillary, bisexual;
pedicel 0.3– 0.7 cm long, glabrate. Sepals 1–2 mm long, ovate,
chartaceous, glabrous on both surfaces or glabrate only out-
side, apex rounded; margin entire. Corolla cyathiform, ca.
2 mm long; tube ca. 1 mm long; lobes ca. 1 mm long, ovate,
glabrous, greenish, apex acute to obtuse; margin entire. Sta-
mens ca. 1 mm long, inserted at the lower half of corolla tube;
filaments and anthers glabrous. Staminodes ca. 1 mm long,
lanceolate, glabrous. Ovary 2-locular, ca. 1 mm long, pilose,
trichomes ferruginous; stipe absent; style ca. 1 mm long, gla-
brous, basal restriction absent; stigma 2-lobed. Fruit 1-
seeded, 2.5–3.0 cm long, ellipsoid to ovoid, smooth, glabrous,
brownish; seeds 1.5–2.5 cm long, smooth to rugulose; seed
scar 1.32.2 cm long, narrow. Figures 3NP.
Representative Specimens—BRAZIL. Bahia: Una, Fazenda Maruim,
12 May 1981, S. Mori et al. 13974 (CEPEC, NY). Santa Cruz de Cabra
´lia, 3 Apr
1997, M. Guedes et al. 6692 (ALCB, CEPEC). Espı
´rito Santo: Linhares,
Reserva Florestal da CVRD, 14 Nov 1974, J. Spada 396 (MO).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (Bahia, Espı
´rito Santo,
and Minas Gerais).
Phenology—Flowering specimens have been observed in
November, fruiting specimens in April to May.
Taxonomic Comments—It differs from P. bangii mainly by
its 5-merous flowers and shorter stamens.
Conservation Status—Pouteria microstrigosa has a
restricted geographical distribution and the area of occur-
rence has a high risk of disturbance. Following IUCN (2012)
criteria, it is assessed as vulnerable [VU A2(cd), B1].
23. POUTERIA NORDESTINENSIS Alves-Arau
´jo & M. Alves, Nord.
J. Botany 30: 399 –406. 2012.—TYPE: BRAZIL. Pernambuco,
Paudalho, Aldeia, Km 17, 23 Jul 2010, A. Alves Arau
´jo
et al. 1338.(holotype:UFP!;isotypes:CEPEC!,G!,K!,
NY!, RB!, S!).
Trees 10–20 m tall, shoots pubescent, trichomes ferrugi-
nous to golden, lenticels absent. Leaves 716
+
3–8 cm,
spirally arranged, lanceolate to oblanceolate, chartaceous,
glabrous to pubescent both surfaces; margin plane; venation
eucamptodromous; petiole 2–5 cm long, terete, pubescent.
Flowers 4-merous, 28 per fascicle, axillary or ramiflorous,
bisexual; pedicel 0.10.2 cm long, puberulent. Sepals 2.5
3.5 mm long, ovate, chartaceous, glabrous inside, puberulent
outside, apex obtuse; margin ciliate. Corolla tubular, 2.5
4.0 mm long; tube 1.52.5 mm long; lobes 1.0–1.5 mm long,
rotund, glabrous, greenish, apex obtuse to truncate; margin
ciliate. Stamens 2.02.5 mm long, inserted at the middle of
corolla tube; filaments glabrous; anthers pilose. Staminodes
0.9–1.0 mm long, rotund to deltoid, pilose. Ovary 4-locular,
0.5–1.0 mm long, pilose, trichomes golden; stipe absent; style
1.0–2.5 mm long, glabrous, basal restriction absent; stigma
slightly 4-lobed. Fruit 14-seeded, 3– 6 cm long, globoid,
smooth, glabrous (mature), yellowish; seeds 24 cm long,
smooth; seed scar 1.43.2 cm long, narrow.
Representative Specimens—BRAZIL. Alagoas: Coruripe, Usina
Coruripe, 17 Dez 1999, R. Lyra-Lemos & I. Bayma 4389 (MAC);
Quebrangulo, ReBio da Pedra Talhada, 12 May 2009, A. Alves-Arau
´jo
et al. 1282 (UFP); Murici, Est. Ecol. de Murici, 10 Feb 2009, A. Alves-Arau
´jo
et al. 1134 (UFP); Bahia: Esplanada, Fazenda do Bu, 18 Jun 1996,
H. Bautista 1805 et al. (CEPEC, IPA); Pernambuco: Buı
´que, PARNA do
Catimbau, 11 Jun 2009, A. Alves-Arau
´jo et al. 1318 (UFP); Igarassu, Usina
Sa
˜oJose
´, Mata da Zambana, 28 Jul 2007, A. Alves-Arau
´jo et al. 472 (IPA,
UFP); Sa
˜o Lourenc¸ o da Mata, Tapacura
´, 17 Mar 2001, T. Silva & K. Almeida
45 (PEUFR); Sa
˜o Vicente Fe
´rrer, Mata do Estado, 9 May 2000, E. Ferraz
et al. 916 (PEUFR).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (from Pernambuco
to Bahia).
Phenology—Flowering specimens have been collected and
observed in November to February, fruiting specimens in
May to September.
Taxonomic Comments—Pouteria nordestinensis differs from
P. durlandii subsp. durlandii by its longer petioles (25 cm),
shorter pedicels (0.10.2 cm long), pilose anthers, and
4-locular ovary.
Conservation Status—Although it is such an adaptable
species, Pouteria nordestinensis is endemic to the Atlantic
rainforest and it is so far only known from northeastern Brazil.
This region is one of the most devastated areas in the Atlantic
forest because of logging, grazing, and plantations of sugar-
cane and cocoa. The area of occurrence is considered as
decreasing but according to the IUCN (2012) criteria, Pouteria
nordestinensis is assigned as of least concern (LC).
24. POUTERIA OBLANCEOLATA Pires, Bol. Tecn. Inst. Agron. N. 38:
38, t. 12, 22, 23. 1960.—TYPE: BRAZIL. Para
´: Fazenda
Uriboca: Companhia Pirelli S. A., [Aug 1958], J. Pires
7105 (holotype: IAN; isotypes: G!, NY!, US!).
Trees 10–20 m tall, shoots glabrous to tomentose, tri-
chomes yellowish to ferruginous, lenticels present. Leaves
10– 15
+
5.5–6.5 cm, spirally arranged, oblanceolate to ellip-
tic, chartaceous, glabrous; margin plane; venation eucampto-
brochidodromous; petiole 2.02.5 cm long, terete, glabrous.
Flowers 5-merous, 12 per fascicle, axillary, bisexual; pedicel
0.3–0.5 cm long, velutinous. Sepals 4 5 mm long, ovate,
chartaceous, velutinous inside, glabrous to glabrate outside;
apex rounded; margin entire. Corolla cyathiform, 67 mm
long; tube 2–3 mm long; lobes 34 mm long, lanceolate,
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glabrous, greenish, apex acute; margin entire. Stamens 2
3 mm long, inserted at the lower half of corolla tube; fila-
ments and anthers glabrous. Staminodes 0.51.0 mm long,
lanceolate, glabrous. Ovary 5-locular, 1.01.2 mm long,
pilose, trichomes whitish to golden; stipe absent; style 2
3 mm long, glabrous, basal restriction absent; stigma 5-lobed.
Fruit 1-seeded, 3– 5 cm long, obovoid, smooth, glabrous, yel-
lowish; seeds 1.5–2.5 cm long, slightly rugulose; seed scar
1.4–2.3 cm long, narrow. Figures 4A E.
Representative Specimens—BRAZIL. Bahia: Una, Fazenda Sa
˜o Rafael,
10 Dec 1968, T. Santos et al. 292 (CEPEC, MO); Una, ReBio Mico-Lea
˜o,
9 Mar 1993, J. Jardim et al. 103 (CEPEC, MO, NY). Amazonas: Distrito
Agropecua
´rio, Reserva 1501, 12 Jun 1991, M. Freitas et al. 780 (NY);
Distrito Agropecua
´rio, Reserva 1501, 16 Apr 1991, M. Freitas et al.
728 (NY).
Distribution—It has a disjunct distribution between the
Atlantic (southern Bahia) and Amazonian forests (Brazil,
French Guiana, and Peru).
Phenology—Flowering specimens have been observed in
December to January, fruiting specimens in March.
Taxonomic Comments—It can be distinguished from P.
confusa mainly by its 5-merous flowers, velutinous sepals
(inside), and stamens inserted at the lower half of the
corolla tube.
Conservation Status—Despite its disjunct distribution,
Pouteria oblanceolata appears to be rare based on the collec-
tions. Therefore it is assessed as vulnerable [VU A2(cd),
B1].
25. POUTERIA OXYPETALA T. D. Penn., Fl. Neotrop. 52: 411412.
1990.—TYPE: BRAZIL. Sa
˜o Paulo, km 245, Estrada
Ubatuba para Caraguatatuba (Serra), 26 Sep 1970, H.
Leita
˜o-Filho 1048 (holotype: NY!).
Treelets to trees 410 m tall, shoots puberulent, trichomes
grayish, lenticels absent. Leaves 2132
+
6.8–11.0 cm, spi-
rally arranged, lanceolate, chartaceous to coriaceous, upper
surface glabrous, lower surface glabrous or sericeous; margin
revolute; venation eucamptodromous; petiole 3.07.5 cm
long, terete, pubescent to glabrous. Flowers 4-merous, 38
per fascicle, axillary, bisexual; pedicel 1.01.5 cm long,
puberulent. Sepals 812 mm long, broadly lanceolate,
chartaceous, glabrous inside, puberulent outside, apex acute;
margin entire (outer ones) to basally lacerate (inner ones).
Corolla tubular, 14–16 mm long; tube 9 10 mm long; lobes
6, 5–6 mm long, lanceolate, sparsely pilose outside, greenish,
apex acute; margin papillate. Stamens 6, 2.53.0 mm long,
inserted at the upper half of corolla tube; filaments papillate;
anthers glabrous. Staminodes 6, 23 mm long, lanceolate,
papillate. Ovary 1012-locular, 3–4 mm long, pilose, tri-
chomes golden; stipe absent; style 910 mm long, pubescent
basally, basal constriction present; stigma 68-lobed. Fruit
not seen. Figures 4F–H.
Representative Specimens—BRAZIL. Bahia: Ilhe
´us, Rod. Olivenc¸a/
Maruim, 10 May 1981, S. Mori & B. Boom 13922 (CEPEC, NY); Ilhe
´us, Km
28 da Rod. Una/Ilhe
´us. Vila Brasil, Rio Jacuipe, Mar 1987, M. Sobral &
L. Mattos-Silva 5533 (CEPEC, NY); Ilhe
´us, 5,4 Km a W Olivenc¸a, Rod. Vila
Brasil/Maruim, 5 Feb 1994, W. Thomas et al. 10282 (CEPEC, NY); Itamarı
´-
Indaia
´, Mesquita, 25 Aug 2011, B. Amorim et al. 1021 (UFP, NY); Valenc¸a,
estrada Valenc¸a/Guaibim, 4 Sep 2004, P. Fiaschi et al. 2506 (CEPEC, NY).
Minas Gerais: Caratinga, Faz. Montes Claros, Est. Ecol. de Caratinga,
4 Sep 1998, J. Lombardi et al. 2321 (BHCB).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (from Bahia to Sa
˜o Paulo).
Phenology—Flowering specimens have been observed
in August to December, fruiting specimens in December
to May.
Taxonomic Comments—It differs from Pouteria pachycalyx
by its inner sepals with basally lacerate margins and lanceo-
late corolla lobes with an acute apex.
Conservation Status—Pouteria oxypetala has a restricted
geographical distribution and the area of occurrence has a
high risk of disturbance. Following IUCN (2012) criteria, it is
assessed as endangered [EN A1(cd), B2 (biii)].
26. POUTERIA PACHYCALYX T. D. Penn., Fl. Neotrop. 52: 409411.
1990.—TYPE: BRAZIL. Espı
´rito Santo, Linhares, Reserva
Florestal CVRD, [19 Jan 1978], J. Spada 26 (holotype:
MO!; isotype: CVRD!).
Trees 6–25 m tall, shoots pubescent, trichomes ferrugi-
nous, lenticels absent. Leaves 2538
+
12– 16 cm, spirally
arranged, oblanceolate to lanceolate, chartaceous to coria-
ceous, glabrous to pubescent (only on midrib); margin
slightly revolute; venation eucamptodromous; petiole 4.5
6.0 cm long, terete, pubescent to glabrous. Flowers 4-merous,
2–4 per fascicle, axillary, bisexual; pedicel 2.8– 3.6 cm long,
pubescent. Sepals 1214 mm long, oblanceolate to orbic-
ular, coriaceous, glabrous inside, puberulent outside, apex
rounded; margin entire (outer ones) to lacerate (inner ones).
Corolla tubular, 1721 mm long; tube 1214 mm long; lobes
6–8, 5 7 mm long, rotund, glabrous, greenish, apex truncate
to obtuse; margin papillate. Stamens 68, 2.53.0 mm long,
inserted at the upper half of corolla tube; filaments papillate;
anthers glabrous. Staminodes 68, 2–3 mm long, lanceolate,
papillate. Ovary 1012-locular, 56 mm long, pubescent,
trichomes ferruginous; stipe absent; style 1214 mm long,
glabrous, basal constriction absent; stigma 6– 8-lobed. Fruit
1–6-seeded, 35 cm long, globoid to obovoid, smooth,
puberulent (immature) to glabrate (mature), yellowish;
seeds 1.5–2.2 cm long, rugulose; seed scar covering up to
three fourths of the surface. Figures 4IK.
Representative Specimens—BRAZIL. Bahia: Una, ReBio Mico-Lea
˜o,
BA-001, KM-46, Ilhe
´us/Una, 30 Mar 1994, A. Amorim 1632 (CEPEC, NY);
Una, ReBio Mico-Lea
˜o, BA-001, KM-46, Ilhe
´us/Una, 18 Jun 1997, A.
Amorim 2043 (CEPEC, NY); Uruc¸ uca, 26 Jun 1979, S. Mori 12050 (CEPEC,
NY). Espı
´rito Santo: Linhares/Sooretama, Rebio Sooretama, 03 Jul 2012,
A. Alves-Arau
´jo et al. 1382 (VIES); Minas Gerais: Caratinga, Est. Ecol. de
Caratinga, 24 Mar 1995, J. Gomes 229 (BHCB); Caratinga, Est. Ecol. de
Caratinga, 20 Aug 1994, J. Lombardi 606 (BHCB); Caratinga, Est.
Ecol. Caratinga, 26 Feb 1992, L. Costa et al. s. n. (BHCB-22279); Caratinga,
Est. Ecol. Caratinga, 15 Apr 1992, L. Costa et al. s. n. (BHCB-22424).
Distribution—It is endemic to the Atlantic Rainforest
where it is restricted to wet forests (Bahia, Espı
´rito Santo,
Minas Gerais, and probably also occurs in Rio de Janeiro).
Phenology—Flowering specimens have been observed in
March to April, fruiting specimens in March to August.
Taxonomic Comments—It can be distinguished from P.
oxypetala by longer pedicels, coriaceous outer sepals, and
lacerate margin of inner sepals.
Conservation Status—Like Pouteria oxypetala, P. pachycalyx
has a restricted geographical distribution and the area of
occurrence has a high risk of disturbance. Following IUCN
(2012) criteria, it is assessed as endangered [EN A1(cd), B2
(biii)].
27. POUTERIA PROCERA (Mart.) K. Hammer, Verz. Landwirtsch.
Ga
¨rt.Kulturpfl.2:1046.1986.Lucuma procera Mart., Flora
22(Beibl. 1): 57. 1839.—TYPE: BRAZIL. Ilhe
´us, Serra do
Mar, [183?], Martius 398 [holotype: M!; isotypes: BM, BR,
F! (F0072055F, F0077358F, F0077359F, F0077360F), G!
(2 sheets), GH, HBG, K! (K000641429, K000641430), MO!,
NY!, OXF, P! (P00648023, P00648024, P00648025), TUB, W].
930 SYSTEMATIC BOTANY [Volume 39
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Fig. 4. A –O. Species of Pouteria from the northern portion of the Atlantic Rainforest of Brazil. A –E. P. oblanceolata. A. Branch. B. Sepal, outer surface.
C. Sepal, inner surface. D. Corolla, stamens, and staminodes. E. Ovary. F –H. P. oxypetala. F. Flower. G. Sepal, outer surface. H. Ovary, cross section. I –K.
P. pachycalyx. I. Flower. J. Sepal, outer surface. K. Sepal, inner surface. L–O. P. aff. stenophylla. L. Floral branch. M. Corolla, stamens, and staminodes.
N. Corolla lobe, stamen, and staminode. O. Ovary. Drawings based on J. Jardim et al. 103 (A E), M. Sobral & L. Mattos-Silva 5533 (F H), A. Amorim 2043
(I– K), and M. Fonseca 934 (L–O).
2014] ALVES-ARAU
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Trees 10–45 m tall, shoots glabrous, lenticels present.
Leaves 6.0– 9.6
+
2.9–3.5 cm, spirally arranged, elliptic
to lanceolate, chartaceous, glabrous to glabrate with
punctate scars on lower surface; margin plane; venation
eucamptodromous; petiole 1.21.6 cm long, channeled, gla-
brous. Flowers 5-merous, 311 per fascicle, axillary, bisexual;
subsessile. Sepals 1.52.0 mm (outer ones) to 2.53.0 mm
(inner ones) long, ovate, chartaceous, sparsely pilose inside,
pubescent outside, apex rounded; margin ciliate. Corolla
tubular, 3.85.0 mm long; tube 2.83.0 mm long; lobes 1
2 mm long, ovate, glabrous, greenish, apex rounded to trun-
cate; margin entire. Stamens 1.0– 1.2 mm long, inserted at the
upper half of corolla tube; filaments and anthers glabrous.
Staminodes ca. 1 mm long, subulate, glabrous. Ovary 5-locular,
2– 3 mm long, pilose, trichomes ferruginous; stipe absent;
style 2– 3 mm long, glabrous, basal restriction absent; stigma
slightly 5-lobed. Fruit 12-seeded, 34 cm long, obovoid,
puberulent, yellowish; seeds 23 cm long, smooth; seed scar
1.0–1.2 cm long, narrow.
Representative Specimens—BRAZIL. Bahia: Governador Mangabeira,
18 Oct 2007, E. Queiroz 2971 (HRB); Ilhe
´us, CEPLAC, 19 Oct 2007, M. Lopes
et al. 1532 (CEPEC); Jussarı
´, RPPN Serra do Teimoso, 4 Apr 2002, J. Paixa
˜o
et al. 198 (CEPEC, MO, NY); Mucuri, Rio Mucuri, 10 Oct 1998, G.
Hatschbach et al. 68393 (BHCB, CEPEC, G, MBM, Z); Palmeiras, Pai Ina
´cio,
25 Sep 1994, M. Guedes et al. 762 (ALCB); Una, Maruim, 13 May 1981, S.
Mori et al. 13992 (CEPEC, NY). Espı
´rito Santo: Colatina, Rio Doce, 24 Sep
1931, J. Kuhlmann 388 (RB); Piauı
´:Sa
˜o Raimundo Nonato, PARNA Serra
da Capivara, 11 Jul 2001, A. Moura 251 (JPB); Rio de Janeiro: Rio de
Janeiro, Canta Galo, Nov 1889, Peckolt 199 (P).
Distribution—It is widely distributed in South America
(Bolivia, Brazil, Colombia, Peru and Venezuela) and in the
northern Atlantic Rainforest (Bahia).
Phenology—Flowering specimens have been observed in
September to January, fruiting specimens in October to April.
Taxonomic Comments—It can be distinguished from
Pouteria reticulata by its subsessile flowers, outer sepals
shorter than inner ones, and 5-locular ovary.
Conservation Status—Pouteria procera is widely distrib-
uted in the Amazon but is known from only one area in the
Atlantic Rainforest. Despite this, according to IUCN (2012)
criteria, it is assessed as of least concern (LC).
28. POUTERIA RAMIFLORA (Mart.) Radlk., Sitzungsber. Math.
Phys. Cl. Ko
¨nigl. Bayer. Akad. Wiss. Mu
¨nchen 12 (3):
333. 1882. Labatia ramiflora Mart., Flora 21(2), Beibl. 2(4):
93. 1838.—TYPE: BRAZIL. Minas Gerais, Aug 1818,
Martius s. n. (holotype: M!).
Trees 10–25 m tall, shoots tomentose, trichomes golden,
lenticels absent. Leaves 3.5–16.0
+
2.0–6.5 cm, spirally
arranged, lanceolate to oblanceolate, chartaceous, glabrous
to pubescent; margin plane; venation eucamptodromous;
petiole 0.52.0 cm long, terete, glabrous to pubescent.
Flowers 4-merous, 3– 10 per fascicle, axillary or on leafless
shoots, bisexual; pedicel 0.20.4 cm long, tomentose to
puberulent. Sepals 23 mm long, ovate, chartaceous, gla-
brous inside, pubescent outside, apex rounded; margin
entire. Corolla cyathiform, 24 mm long; tube 1.02.5 mm
long; lobes 1.0–1.5 mm long, ovate, glabrous to pilose, green-
ish, apex rounded; margin entire. Stamens 410 mm long,
inserted at the upper half of corolla tube; filaments and
anthers glabrous. Staminodes 24 mm long, subulate, gla-
brous. Ovary 2(3)-locular, 12 mm long, pilose, trichomes
golden; stipe absent; style 12 mm long, glabrous, basal
restriction absent; stigma 23-lobed. Fruit 1-seeded, 2–4 cm
long, obovoid to globoid, smooth, glabrous to pubescent,
greenish to yellowish; seeds 1.52.6 cm long, smooth; seed
scar 1.0–1.6 cm long, wide.
Representative Specimens—BRAZIL. Ceara
´: Caucaia, Mata da
Cruzinha Cauı
´pe, 26 Mar 1998, A. Moura 160 (JPB); Fortaleza, UFC-Pici,
18. Nov 2009, A. Alves-Arau
´jo & A. Castro 1330 (UFP). Bahia: Barreiras, 21
Mar 1981, G. Pinto 189/81 (CEPEC, HRB, RB); Jacobina, Serra da Jacobina,
18 Nov 1986, L. Queiroz 1177 (IPA); Rio de Contas, Serra da Marsalina, 18
Nov 1996, R. Harley et al. 4438 (ALCB, CEPEC, HRB); Maranha
˜o:
Maracassume
´, 31 Aug 1932, R. Fro
´es 1841 (G, P, MO); Pernambuco:
Ibimirim, 1 Nov 1986, G. Webster 25731 (IPA, MO); Piauı
´:Sa
˜o Raimundo
Nonato, PARNA Serra da Capivara, 20 Sep 2001, A. Moura 257B (JPB).
Distribution—It is widely distributed in the Cerrado vege-
tation but also reaches the Amazonian and, currently, Atlantic
forests (Ceara
´).
Phenology—Atlantic Rainforest flowering specimens have
been collected and observed in November to March, other
additional flowering specimens in March to November,
fruiting specimens in April to November.
Taxonomic Comments—This species has great plasticity in
leaf size, shape, and indumentum. However, in the Atlantic
rainforest population, P. ramiflora differs from P. macahensis
by the absence of lenticels, eucamptodromous leaf venation,
and 4-merous flowers.
Conservation Status—Pouteria ramiflora is widely distrib-
uted in South America and according to IUCN (2012) criteria,
it is assessed as of least concern (LC).
29. POUTERIA RETICULATA (Engl.) Eyma, Recueil Trav. Bot.
Ne
´erl. 33: 183. 1936. Chrysophyllum reticulatum Engl.,
Bot. Jahrb. Syst. 12: 522. 1890.—LECTOTYPE, designated
here: BRAZIL. Rio de Janeiro, [without date], M. Glaziou
12070. [holotype: B [destroyed]; P! (P00648043);
isolectotypes: C!, F!, K!, P! (P00648044)].
Pouteria reticulata (Engl.) Eyma subsp. surinamensis T. D.
Penn., Fl. Neotrop. 52: 299. 1990.—TYPE: GUYANA.
Kanuku Mountains, Wabuwak, [15] Oct 1948, Forest
Department British Guyana 5779 (holotype: K!; isotype:
NY!). syn. nov.
Trees 7–10 m tall, shoots pubescent, trichomes ferrugi-
nous, lenticels present. Leaves 6.011.5
+
3.8– 5.2 cm, disti-
chous, oblanceolate to lanceolate, chartaceous, glabrate with
punctate scars on the lower surface; margin plane; venation
brochidodromous; petiole 0.82.4 cm long, channeled, gla-
brous. Flowers 5-merous, 48 per fascicle, axillary, bisexual;
pedicel 0.3–0.4 cm long, glabrous to glabrate. Sepals 1 2 mm
long, ovate to elliptic, chartaceous, glabrous both surfaces,
apex obtuse; margin entire. Corolla cyathiform, 1.01.5 mm
long; tube 0.5–0.75 mm long; lobes 0.5 0.75 mm long, ovate,
hirsute, greenish, apex obtuse; margin ciliate. Stamens 0.5
1.0 mm long, inserted at the upper half of corolla tube; fila-
ments and anthers glabrous. Staminodes 0.5– 1.0 mm long,
lanceolate, glabrous. Ovary 1-locular, 0.8–1.0 mm long,
puberulent, trichomes golden; stipe absent; style 0.81.0 mm
long, glabrous, basal restriction absent; stigma punctiform.
Fruit 1-seeded, 1.52.8 (4) cm long, obovoid, smooth to
slightly rugulose, glabrous to puberulent, yellowish to
brownish; seeds 1.0–1.5 cm long, smooth; seed scar 0.8
1.3 cm long, narrow.
Representative Specimens—BRAZIL. Alagoas: Coruripe, Usina
Coruripe, 6 Dec 2001, M. Machado 143 (MAC); Bahia: Una, ReBio Mico-
Lea
˜o, 27 Jul 1993, A. Carvalho et al. 4313 (ALCB, CEPEC, HRB, G, MO,
NY); Ceara
´: Serra do Baturite
´,Sı
´tio B. Ina
´cio de Azevedo, 15 Nov 1939,
J. Euge
ˆnio 947 (RB); Piauı
´:Sa
˜o Raimundo Nonato, PARNA Serra da
Capivara, Baixa Grande, 9 Jul 2001, A. Moura 252 (JPB, MO, NY); Rio
Grande do Norte: Martins, Serra dos Martins, 20 Oct 1978, A. Fernandes &
Matos s. n. (EAC-5,061).
932 SYSTEMATIC BOTANY [Volume 39
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Distribution—It is widespread in the Neotropics. In the
northern Atlantic Rainforest, Pouteria reticulata is a common
species (from Ceara
´to Bahia).
Phenology—Flowering specimens have been observed in
October to June, fruiting specimens in October to July.
Taxonomic Comments—It was previously divided into two
subspecies [P. reticulata (Engl.) Eyma subsp. reticulata and
P. reticulata subsp. surinamensis T. D. Penn.] by Pennington
(1990) which were weakly distinguished by few and polymor-
phic characters (e.g. indumentum, length of pedicels). Such
features are actually variable and may occur in the same indi-
vidual depending on the population as was detected on the
type of P. reticulata subsp. surinamensis and other vouchers.
Thus, they are here considered the same taxon. Among Atlantic
forest species, it can be confused with P. procera but differs
from it by its longer pedicels, sepals of the same size, and 1-
locular ovary.
Conservation Status—Pouteria reticulata is widely distrib-
uted in the New World tropics and according to IUCN (2012)
criteria, it is assessed as of least concern (LC).
30. POUTERIA aff. STENOPHYLLA Baehni, Candollea 9: 390. 1942.—
LECTOTYPE, designated here: BRAZIL. Rio de Janeiro,
Serra da Estrela, [1924], S. Riedel 06. US! (US-702351!);
isolectotypes: F [fragment]!, G!, US-1484168!.
Treelets to trees 6– 10 m tall, shoots pubescent, trichomes
ferruginous, lenticels usually present. Leaves 7–13
+
1–4 cm,
spirally arranged, lanceolate to elliptic, coriaceous, glabrous,
lower surface glaucous; margin slightly revolute; venation
eucamptodromous; petiole 1– 2 cm long, terete, glabrous to
pubescent. Flowers 4-merous, 1– 3 per fascicle, axillary, bisex-
ual; pedicel 0.3– 0.5 cm long, terete, pubescent. Sepals 6–7 mm
long, orbicular, chartaceous, pubescent on distal half inside,
pubescent outside (outer ones), apex obtuse; margin entire
(outer sepals) to lacerate (inner ones). Corolla tubular, 11–
13 mm long; tube 4–5 mm long; lobes 67, 78 mm long,
ovate to rotund, papillate, yellowish to greenish, apex trun-
cate; margin entire. Stamens 67, 23 mm long, inserted
at the upper half of corolla tube; filaments papillate; anthers
glabrous. Staminodes 67, 2.8– 3.0 mm long, lanceolate,
papillate. Ovary 6–8-locular, 34 mm long, pilose, tri-
chomes golden; stipitate, stipe <1 mm long; style 57 mm
long, pilose basally, papillate apically, basal constriction
present; stigma 68-lobed. Fruit immature, 1.5–2.0 cm long,
globoid, tomentose, reddish to brownish; seeds not seen.
Figures 4L– O.
Representative Specimens—BRAZIL. Bahia: Prado, Mata da Ange
´lica,
margem Rodovia Prado-Cumuruxatiba, 19 Oct 1997, M. Fonseca 934
(ALCB, CEPEC); Prado, Faz. Riacho das Ostras, 30 Apr 2007, S. Rezende
& A. Brita 1833 (BHCB). Espı
´rito Santo: Sa
˜o Mateus, Liberdade, 05 Jun
2012, A. Alves-Arau
´jo et al. 1381 (VIES).
Distribution—It is endemic to coastal areas usually with
white-sandy soils (restingas) in eastern Brazil (from Bahia to
Rio de Janeiro).
Phenology—Flowering specimens have been observed in
April to October, fruiting specimens not seen.
Taxonomic Comments—Pouteria aff. stenophylla differs
from typical species due to leaf variation which consequently
affects its shape (lanceolate to elliptic leaves instead of nar-
rowly oblanceolate ones). Additionally, molecular data
(Alves-Arau
´jo et al. unpublished data) have demonstrated
that there are significant differences between samples from
typical Pouteria stenophylla and this related taxon. Natural
populations are currently under analyses. Here, it can be
distinguished from P. grandiflora by its glaucous lower leaf
surface and ovary with a short stipe.
Conservation Status—Pouteria aff. stenophylla has a
restricted geographical distribution and the area of occur-
rence has a high risk of disturbance. Following IUCN (2012)
criteria, it is assessed as endangered [EN A1(cd), B2].
31. POUTERIA SUBSESSILIFOLIA Cronq., Bull. Torrey Bot. Club 73:
468. 1946.—TYPE: BRAZIL. Bahia, Andarahy, basin of
Paraguassu
´river, Serra do Corne
´lio, 13 Oct 1942, R. Fro
´es
1023. (holotype: A; isotype: NY!).
Subshrubs, treelets to trees 2–6 m tall, shoots tomentose,
trichomes ferruginous, lenticels absent. Leaves 3.5–8.5
+
2.0
4.5 cm, spirally arranged, elliptic to elliptic, coriaceous, upper
surface glabrous to glabrate, lower surface tomentose; mar-
gin revolute; venation brochidodromous (rarely eucampto-
brochidodromous); petiole 0.1– 0.3 cm long, terete, tomentose.
Flowers 5-merous, 3 8 per fascicle, axillary, unisexual; pedicel
0.3– 0.8 cm long, puberulent. Sepals 23 mm long, ovate,
chartaceous, glabrous inside, pubescent outside, apex acute;
margin ciliate. Corolla cyathiform, 2.0–3.5 mm long; tube
1.0– 1.5 mm long; lobes 1–2 mm long, ovate, glabrous, whitish,
apex rounded; margin entire. Stamens 1.5–2.0 mm long,
inserted at the upper half of corolla tube; filaments and
anthers glabrous. Staminodes 5–10 mm long, subulate to
digitiform, glabrous. Ovary 2-locular, 0.5–1.0 mm long, pilose,
trichomes golden; stipe absent; style 1.0–1.5 mm long, gla-
brous, basal restriction absent; stigma punctiform. Fruit 1-
seeded, 2.0–2.5 cm long, ovoid to globoid, smooth, glabrous;
seeds 1.7–2.0 cm long, smooth, purple to black; seed scar 1.2–
1.6 cm long, wide. Figures 5A–B.
Representative Specimens—BRAZIL. Bahia: Andaraı
´, Serra de
Andaraı
´-Capa bode, 30 Oct 1978, G. Martinelli et al. 5400 (CEPEC, RB);
Andaraı
´, Faz. Volta da Pedra, 9 Nov 1997, M. Guedes et al. 5504 (CEPEC);
Entre Breja
˜o e Ibiquara, Nov 1942, R. Fro
´es 20144 (NY); Mucuge
ˆ, Chapada
Diamantina, Parque Municipal de Mucuge
ˆ, 25 Oct 2008, N. Roque et al.
1888 (ALCB); Mucuge
ˆ, estrada p/ Jussiape
ˆ, 22 Dec 1979, S. Mori &
F. Benton 13164 (CEPEC, NY); Mucuge
ˆ, Projeto Sempre-Viva, 25 Mar
2000, A. Giulietti et al. 1966 (HUEFS); Mucuge
ˆ, Serra do Pai Ina
´cio,
10 Mar 1974, Andrade-Lima 74– 7749 (IPA).
Distribution—It is endemic to Chapada Diamantina
(Bahia), occurring in wet forests as well as in the “Campos
rupestres” vegetation, mainly in swampy places.
Phenology—Flowering specimens have been observed
in October to January, fruiting specimens in November
to March.
Taxonomic Comments—It differs from P. andarahiensis
mainly by its shorter petioles (0.10.3 cm long), corolla with
margin entire, and 2-locular ovary.
Conservation Status—Pouteria subsessilifolia has a
restricted geographical distribution and according to IUCN
(2012) criteria, it is assessed as vulnerable [VU A2(cd), B2].
32. POUTERIA TRIFIDA Alves-Arau
´jo & M. Alves, Syst. Bot. 36(4):
1004–1007. 2011.—TYPE: BRAZIL. Bahia, Ilhe
´us,
CEPLAC-CEPEC, A
´rea de cultivo de cacau - cabruca, 15
Dec 1997, J. Jardim et al. 1165 (holotype: CEPEC!;
isotypes: MO!, NY!).
Trees up to 10 m tall, shoots pilose, trichomes ferruginous,
lenticels present. Leaves 15–29
+
8– 10 cm, distichous to
spirally arranged, elliptic to lanceolate, chartaceous, upper
surface glabrous, lower surface glabrous to glabrescent; mar-
gin plane; venation eucamptodromous; petiole 2.75.0 cm
long, channeled, glabrous to glabrescent. Flowers 5-merous,
2– 5 per fascicle, axillary, unisexual; pedicel 0.10.2 cm long,
2014] ALVES-ARAU
´JO ET AL.: POUTERIA FROM THE ATLANTIC RAINFOREST 933
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Fig. 5. A– P. Pouteria species from the northern part of the Atlantic rainforest, Brazil. A– B. P. subsessilifolia. A. Floral branch. B. Corolla, stamens, and
staminodes. C– P. P. venosa subsp. amazonica. C. Floral branch. D. Flower. E. Corolla, stamens, and staminodes. F. Ovary. G. Ovary, cross section. H P.
Leaf variation of P. venosa subsp. amazonica. H–K. Plane leaves. L –P. Bullate leaves with margin revolute (L– N) and strongly revolute (O–P). Drawings
based on R. Fro
´es 1023 (A– B), A. Alves-Arau
´jo et al. 1144 (H), Paixa
˜o et al. 1236 (I–K), R. Pinheiro 2245 (L–N), and J. Jardim et al. 1154 (OP).
934 SYSTEMATIC BOTANY [Volume 39
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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
velutinous to tomentose. Sepals 23 mm long, ovate,
chartaceous, partially pilose inside, pilose outside, apex
acute; margin entire. Corolla cyathiform, 1.52.5 mm long;
tube 0.2–0.3 mm long; lobes 1.3 2.2 mm long, ovate, green-
ish, apex rounded to acuminate; margin entire. Stamens
unknown (male flowers not seen). Staminodes 0.2– 0.25 mm
long, apex trifid, glabrous. Ovary 5-locular, 0.51.0 mm long,
pilose, trichomes golden; stipe absent; style 0.51.0 mm long,
glabrous, basal restriction absent; stigma punctiform. Fruit
immature, globoid, smooth, pilose; seeds not seen.
Distribution—It is known from the type collection in wet
forest (southern Bahia).
Phenology—Flowering and fruiting specimens have been
observed in December.
Taxonomic Comments—It can be distinguished from
Pouteria durlandii subsp. durlandii by its trifid staminodes
and exclusively 5-locular ovary.
Conservation Status—Pouteria trifida has a restricted geo-
graphical distribution and the area of occurrence shows a
high risk of disturbance. Following IUCN (2012) criteria, it is
assessed as endangered [EN A1(cd), B2].
33. POUTERIA VELUTINICARPA Alves-Arau
´jo & M. Alves, Nord. J.
Botany 30: 399–406. 2012.—TYPE: BRAZIL. Bahia,
Encruzilhada, 8 Aug 1984, J. Lima & M. Santos 156 (holo-
type: HRB!; isotypes: CEPEC!, NY!, RB!).
Tree 8–15 m tall, shoots tomentose, trichomes ferruginous,
lenticels absent, small furrows present. Leaves spirally
arranged, lanceolate, 9–12
+
4– 5 cm, coriaceous, sometimes
bullate, upper surface glabrate to pubescent, lower surface
tomentose (mainly on midrib), trichomes ferruginous, base
attenuate to acute, apex acute to obtuse (sometimes emargin-
ate), margin revolute; venation eucamptodromous, second-
ary veins 7– 15 pairs, slightly arcuate, tertiary veins reticulate;
petiole 1.52.0 cm long, terete, tomentose. Flowers unknown;
pedicel tomentose (in fruit). Sepals 5–6 mm long, ovate to
orbicular, puberulent outside, apex obtuse, inner ones auricu-
late, margin basally lacerate, puberulent outside except on
the margins, outer ones with margin entire. Fruit 1–6-seeded
(8-locular), 3– 5 cm long, globoid, velutinous-tomentose, brown-
ish, pulp well-developed, yellowish, apex emarginated; seeds
not laterally compressed, 1.52.5 cm long, rugulose, shiny,
brownish; scar covering up to three fourths of the seed surface.
Distribution—It is known from the type collection in wet
forest (southern Bahia).
Phenology—Fruiting specimens have been observed
in August.
Taxonomic Comments—It can confused with P. grandiflora
but can be identified by its tomentose shoots and fruits with a
persistent velutinous to laguninous indumentum.
Conservation Status—Pouteria velutinicarpa has a restricted
geographical distribution and the area of occurrence shows a
high risk of disturbance. Following IUCN (2012) criteria, it is
assessed as endangered [EN A1(cd), B2].
34. POUTERIA VENOSA (Mart.) Baehni subsp. AMAZONICA T. D.
Penn., Fl. Neotrop. 52: 399. 1990.—TYPE: VENEZUELA.
Bolı
´var, 20 km S of Upata [El Dorado], [23] Aug 1966,
J. W. Meijeraan 22 (holotype: WAG!).
Trees 5–20 m tall, shoots pubescent, trichomes ferrugi-
nous, lenticels present. Leaves 718
+
3– 8 cm, spirally
arranged, oblanceolate to oblong, chartaceous to (usually)
coriaceous, sometimes bullate, glabrous to glabrate; margin
revolute to strongly revolute; venation eucamptodromous
(rarely eucampto-brochidodromous); petiole 13 cm long,
terete, glabrous. Flowers 4-merous, 26 per fascicle, axillary,
bisexual; pedicel 0.61.6 cm long, puberulent. Sepals 6
13 mm long, ovate to orbicular, chartaceous, glabrous inside,
puberulent outside, apex obtuse; margin entire (outer sepals)
to basally lacerate (inner ones). Corolla tubular, 1018 mm
long; tube 7– 13 mm long; lobes 6 8, 3 5 mm long, ovate to
rotund, papillate with trichomes outside, greenish, apex
obtuse; margin entire. Stamens 68, inserted at the upper
half of corolla tube, 34 mm long; filaments papillate; anthers
glabrous. Staminodes 6–8, 34 mm long, lanceolate, papil-
late. Ovary 6–16-locular, 45 mm long, pilose, trichomes
ferruginous; stipe absent; style 610 mm long, papillate,
basal restriction absent; stigma 68-lobed. Fruit 16-seeded,
3– 5 cm long, globoid to obovoid, smooth to slightly rugu-
lose, puberulent (immature) to glabrate (mature), yellowish;
seeds 1.5–2.2 cm long, rugulose; seed scar covering up to
three fourths of the surface. Figures 4CP.
Representative Specimens—BRAZIL. Alagoas: Murici, ESEC Murici, 10
Feb 2009, A. Alves-Arau
´jo et al. 1144 (UFP); Bahia: Ilhe
´us, Faz. Barra do
Manguinho, 5 Feb 1982, L. Mattos-Silva et al. 1395 (CEPEC); Jaguaquara,
Rod. Rio/Bahia, 3 Oct 1972, R. Pinheiro 1955 (CEPEC); Paraı
´ba: Joa
˜o
Pessoa, Cabo Branco, 4 Jun 2009, A. Alves-Arau
´jo et al. 911 (UFP);
Mataraca, Mineradora Millenium, 27 May 2008, A. Alves-Arau
´jo et al.
1006 (UFP); Pernambuco: Igarassu, Usina Sa
˜o Jose
´, Mata da Piedade, 21
Nov 2008, A. Alves-Arau
´jo et al. 1074 (IPA, UFP); Paudalho, Aldeia, Mata
do Cond. Haras de Aldeia, 15 Oct 2010, A. Alves-Arau
´jo 1338 (UFP);
Recife, Jardim Bota
ˆnico, 19 Oct 2010, A. Alves-Arau
´jo et al. 1336 (UFP);
Rio Grande do Norte: Tibau do Sul, Praia de Pipa, 3 Apr 2010, A. Alves-
Arau
´jo 1333 (UFP).
Distribution—It has a disjunct distribution between the
Atlantic (eastern Brazil from Rio Grande do Norte to Sa
˜o
Paulo) and the Amazonian forests (Brazil, Colombia, French
Guiana, Guyana, Suriname, and Venezuela).
Phenology—Flowering specimens have been collected and
observed in October to March, fruiting specimens in October
to June.
Taxonomic Comments—Pennington (1990) cited the same
distribution for P. venosa subsp. venosa and P. venosa subsp.
amazonica. The latter is distinguished by a set of features such
as type of indumentum, larger fruits, and seed scar. The
phototype of P. venosa subsp. venosa was examined but it
was not enough to indicate if both names are the same taxon
with considerable morphological variation. Pouteria venosa
subsp. amazonica is commonly confused with Pouteria grandi-
flora, but differs from it by its larger leaves and the style does
not bear a basal constriction.
Conservation Status—Pouteria venosa subsp. amazonica is
widely distributed in the Amazon and Atlantic Rainforest in
the South America and according to IUCN (2012) criteria, it is
assessed as of least concern (LC).
Acknowledgments. The authors would like to thank Capes-PPGBV
for financial support; all directors, curators, and staff from the herbaria
visited or consulted; Scott Heald for reviewing the English of this manu-
script; Regina Carvalho for high quality illustrations; and Morfo-
Taxonomia Vegetal Laboratory members and collaborators.
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Appendix 1.
Species Numeric List—01 - Pouteria andarahiensis T. D. Penn.;
02 - P. atlantica Alves-Arau
´jo & M. Alves; 03 - P. bangii (Rusby) T. D. Penn.;
04 - P. bapeba T. D. Penn.; 05 - P. beaurepairei (Glaz. & Raunk.) Baehni; 06 - P.
butyrocarpa (Kuhlm.) T. D. Penn.; 07 - P. caimito (Ruiz & Pav.) Radlk.; 08 -
P. ciliata Alves-Arau
´jo & M. Alves; 09 - P. coelomatica Rizzini; 10 - P.
confusa Alves-Arau
´jo & M. Alves; 11 - P. cuspidata (A. DC.) Baehni; 12 -
P. durlandii (Standl.) Baehni subsp. durlandii;13 - P. egregia Sandwith; 14 -
P. franciscana Baehni; 15 - P. gallifructa Cronquist; 16 - P. gardneri (Mart. &
Miq.) Baehni; 17-P. glauca T. D. Penn.; 18-P. grandiflora (A. DC.) Baehni;
19 - P. guianensis Aubl.; 20 - P. macahensis T. D. Penn.; 21 - P. macrophylla
Eyma; 22 - P. microstrigosa T. D. Penn.; 23 - P. nordestinensis Alves-Arau
´jo &
M. Alves; 24 - P. oblanceolata Pires; 25 - P. oxypetala T. D. Penn.; 26 - P.
pachycalyx T. D. Penn.; 27 - P. procera (Mart.) K. Hammer; 28 - P. ramiflora
(Mart.) Radlk.; 29 - P. reticulata (Engl.) Eyma; 30 - P. aff. stenophylla Baehni;
31 - P. subsessilifolia Cronquist; 32 - P. trifida Alves-Arau
´jo & M. Alves; 33 -
P. velutinicarpa Alves-Arau
´jo & M. Alves; 34 - P. venosa subsp. amazonica
T. D. Penn.
Index of Collectors—Numbers between parentheses corresponds to
the species’ number in the taxonomic treatment and species numeric list.
Vouchers without collector’s numbers are provided between square
brackets followed by collection number, when available.
Agra, M. F. s. n. [JPB-19.577] (34), [JPB-19.578] (34); Allema
˜o, F. 935
(28); Almeida, J. & T. S. Santos 58 (4); Almeida, K. 164 (23); Almeida, K. &
E. S. Silva 107 (23); Almeida, K. & E. Santos 13 (23); Almeida, L. s. n.
[EAC-2.867] (28), [EAC-3.047] (28), [EAC-3.048] (29); Alves, L. J. et al. 188
(28), 400 (7); Alves, M. et al. 1082 (29), 1386 (1); Alves-Arau
´jo, A. 337 (7),
1080 (23), 1124 (7), 1332 (12), 1333 (34), 1338 (34); Alves-Arau
´jo, A. & A.
Melo 997 (15), 1165 (34), 1170 (16), 1172 (15); Alves-Arau
´jo, A. & A. S.
Castro 1330 (28); Alves-Arau
´jo, A. & B. S. Amorim 1084 (16), 1085 (23),
1087 (3), 1089 (3), 1090 (34), 1284 (34), 1287 (16), 1292 (16), 1293 (34), 1300
(23), 1303 (15), 1305 (34), 1310 (3); Alves-Arau
´jo, A. & L. Lanev 1203 (7),
1204 (15), 1205 (3), 1207 (6); Alves-Arau
´jo, A. & M. Alves 1314 (34);
Alves-Arau
´jo, A. & T. Pontes 1218 (15), 1220 (16), 1223 (34), 1226 (34),
1228 (3), 1229 (16); Alves-Arau
´jo, A. et al. 220 (3), 234 (34), 299 (16), 300
(34), 472 (23), 525 (34), 539 (34), 607 (3), 831 (34), 911 (34), 970 (20), 1005
(15), 1006 (34), 1009 (20), 1014 (3), 1014 (3), 1055 (34), 1071 (3), 1073 (16),
1074 (34), 1075 (34), 1081 (3), 1094 (20), 1097 (15), 1103 (20), 1116 (16), 1125
(15), 1126 (15), 1132 (23), 1134 (23), 1136 (15), 1142 (16), 1144 (34), 1152 (3),
1153 (34), 1154 (16), 1174 (34), 1175 (34), 1180 (16), 1181 (3), 1185 (6), 1186
(15), 1187 (23), 1189 (6), 1216 (23), 1244 (3), 1245 (15), 1257 (16), 1258 (23),
1259 (23), 1282 (23), 1283 (15), 1318 (23), 1321 (23), 1331 (12), 1335 (12), 1336
(34), 1339 (23), 1341 (23), 1342 (34), 1343 (16), 1370 (34), 1371 (34), 1372 (34),
1381 (30); Amaral, C. & E. Santos 46 (34); Amorim, A. M. 1632 (26), 1901
(34), 2043 (26), 2230 (29), 2246 (29), 2700 (29); Amorim, A. M. et al. 374 (11),
422 (34), 553 (28), 1413 (15), 1516 (34), 1632 (34), 2157 (28), 2224 (6), 2275
(6), 2663 (6), 3883 (1), 3924 (20), 6427 (5), 6596 (7), 6774 (18), 6969 (7), 7686
(7); Amorim, B. S. et al. 1021 (25); Anderson, W. R. et al. 36685 (28), 37078
(28); Andrade, F. S. & M. M. Santos 06 (28); Andrade-Lima &F.Ferreira
04 (3); Andrade-Lima 46-269 (34), 49-265 (23), 49-288 (15), 49-289 (15),
49-347 (23), 50-492 (34), 55-2061 (16), 59-3340 (3), 61-3983 (23), 63-4153
(20), 68-5197 (16), 70-5733 (20), 74-7749 (31); Andrade-Neto, M. s. n.
[EAC-15.831] (21), [EAC-15.832] (29); Arau
´jo, F. J. F. s. n. [EAC-32.308]
(28); Arau
´jo, G. B. & A. Menezes 609 (34); Arau
´jo, G. B. & J. W. A. Silva
300 (34); Arbo, M. M. et al. 7569 (31); Argo
ˆlo, A. et al. 08 (34), 10 (34);
Assis, J. S. 341 (28); Atkins, S. et al. 4706 (28), 4708 (28), 4721 (19), 5637 (1);
Aublet s. n. [P] (19); B. W. 6548 (3); Bale
´e, W. L. & B. G. Ribeiro 14 (34),
955 (3); Bang, A. M. 1953 (3); Barbosa, E. et al. 264 (5); Barbosa, M. R. 1184
(34), 1661 (34); Barbosa, M. R. & A. C. Moura 1457 (34); Barros, F. 1372 (5);
Barroso, G. M. & Elsie 227 (28); Baumgratz, J. F. et al. 190 (34); Bautista,
H. P. 1466 (28), 1478 (28), 1530 (28); Bautista, H. P. & E. P. Queiroz 4041
(34); Bautista, H. P. & G. C. P. Pinto 1043 (34); Bautista, H. P. et al. 1805
(23), 3420 (1), 4034 (1); Bayma, I. A. 1026 (34), 1028 (34), s. n. [MAC-24.656]
(34); Bayma, I. A. & A. C. Santos 879 (34); Bele
´m, R. P. 3679 (16); Bele
´m, R.
P. & M. Magalha
˜es 772 (34), 1067 (20), 1069 (34); Bele
´m, R. P. & R. S.
Pinheiro 2375 (34), 2886 (27), 3000 (20), 3047 (34); Berto, J. s. n. [MAC-
20.236] (34); Bezerra, I. F. F. s. n. [UFP-4.359] (34); Bezerra-Silva, M. T.
01 (34); Blanchet, J. S. 264 (28), 3598 (16); Bondar, G. s. n. [K] (18),
[CEPEC-38.075] (18), [G] (6), [NY-375364] (6); Borgo, M. et al. 367 (5);
Brownsberg (BW1726) (3), (BW3315) (3), (BW6623) (3); Cairo, J. P. F. &F.
Queiroz s. n. [ALCB-83.281] (18); Calou, C. s. n. [EAC-16.100] (29);
Cantarelli, J. et al. 594 (20); Cardoso, D. 240 (1), 832 (28); Cardoso, D. &
A. M. Bastos 2183 (28); Cardoso, D. & J. M. O. Santos 924 (34); Carmo, L.
P. et al. 135 (16), 179 (16); Carneiro, C. E. et al. 82 (34); Carneiro, E. 455
(34); Carneiro, J. 1198 (18); Carneiro-Torres, D. S. et al. 145 (28);
Carvalhal, I. 01 (18); Carvalho, A. M. 735 (26), 4673 (29), 4832 (29), 4843
936 SYSTEMATIC BOTANY [Volume 39
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(29), 5251 (29), 5490 (29); Carvalho, A. M. & S. Faria 2548 (34); Carvalho, A.
M. & T. Plowman 1389 (20), 1604 (34); Carvalho, A. M. et al. 188 (34), 486
(34), 1367 (7), 3295 (34), 3794 (16), 4154 (28), 4313 (29), 4545 (12), 4641 (3),
4689 (11); Carvalho, P. A. et al. 06 (22); Carvalho-Sobrinho, J. G. 117 (16);
Castro, A. S. F. 620 (28), 765 (28), 935 (28), 1055 (21), 1624 (21), 1664 (29),
1868 (29), 1882 (28), 2109 (28), 2160 (29; Castro, A. S. F. s. n. [EAC-23.440]
(28), [EAC-26.048] (28); Cavalcante, F. & E. Nunes s. n. [EAC-16.241] (28);
Cavalcante, F. et al. 38 (34); Cavalcante, F. et al. 62 (34); Cavalcanti,A.D.C.
110 (16), 136 (16); Cervi, A. et al. 7101 (23); Cestaro, L. A. 99-007 (34), 99-
168 (34), 99-193 (34); Chagas, F. & Silva 1681 (5 ); Chagas, F. & Silva s. n.
[FUEL-10.955] (5), [FUEL-11.786] (5); Chagas-Mota 1425 (34), 1426 (34),
2018 (34); Chagas-Mota & L. M. Lea
˜o 1867 (34 ), 1868 (16); Chaves, E. M.
F. &E.M.Se
´rvio Jr. 574 (28); Chiappeta, A. 033 (34), 673 (34); Cid, C. A.
et al. 5886 (3), 8337 (3); Coelho, P. s. n. [IPA-47.899] (34); Conceic¸a
˜o, A. A.
et al. 1491 (28); Costa, A. 201 (23), 1269 (34); Costa, A. s. n. [MAC-26.207]
(34); Costa, A. L. 550 (18); Costa, A. L. s. n. [ALCB-3.239] (7); Costa, J. &
A. Pereira 1894 (1), 1897 (28); Costa, J. & C. B. N. Costa 824 (31); Costa,
J. et al. 233 (31), 346 (34); Costa-e-Silva, M. B. 257 (23); Cotias,A.L.et al.
s. n. [ALCB-81.017] (18); Cruz, A. & E. Santos 100 (34); Cruz, A. et al. 62
(34); Curran, H. M. 44 (27); D. A. F. 185/80 (3); D. A. F. 66/79 (3); Daly,
D. C. et al. D449 (15), 9305 (3); Dantas, A. et al. 130 (34); dePaula, J. E.
1422 (34); doCarmo, L. P. et al. 173 (15), 174 (3); dosAnjos,B.A.et al. 101
(28); DuBocage, A. s. n. [IPA-56.479] (34); DuBocage, A. et al. 320 (34);
Ducke, A. s. n. [EAC-1.015] (28), [Herb.Amaz.-15.041] (3), [RB-22.213] (15),
[RB-22.216](3);Ducke,A.&A.Lima97(1