Reestablishment, New Records, and A Key For The Species Of: Aspidosperma (Apocynaceae) From The Brazilian Amazon
Reestablishment, New Records, and A Key For The Species Of: Aspidosperma (Apocynaceae) From The Brazilian Amazon
doi: 10.1590/0102-33062018abb0168
ABSTRACT
As a result of systematic study of the Neotropical genus Aspidosperma (Apocynaceae), we reestablish the species
A. centrale and A. duckei, and report A. steinbachii and A. tambopatense as new records for Brazil. We provide taxonomic
descriptions of these species along with plates, distribution maps, and information on their conservation status,
habitat and phenology. We also provide an identification key for all the species of Aspidosperma from the Brazilian
Amazon.
1 Pós-graduação em Biologia Vegetal, Departamento de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas,
13083-862, Campinas, SP, Brazil
2 Pós-graduação em Ciências Biológicas (Botânica), Instituto de Biociências de Botucatu, Universidade Estadual Paulista “Júlio de Mesquita Filho”,
18618-970, Botucatu, SP, Brazil
3 Departamento de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas, 13083-862, Campinas, SP, Brazil
* Corresponding author: [email protected]
Andreza Stephanie de Souza Pereira, Ana Carolina Devides Castello, André Olmos Simões and Ingrid Koch
During ongoing studies with Aspidosperma, we found stereomicroscope. The terminology used follows Woodson
that A. centrale Markgr. and A. duckei Huber ex Ducke (1951), Radford et al. (1974) and Marcondes-Ferreira (1988).
should be reestablished, and confirmed the first records Data on distribution, habitat and phenology were obtained
of A. steinbachii Markgr. and A. tambopatense A.H.Gentry from herbarium vouchers and field notes. Information on
in Brazil. Thus, we aim to present updated taxonomic leaf and fruit coloring is based on dried material. Distribution
descriptions, plates and distribution maps for these species, maps were created using ArcGis 10.1 (ESRI 2012) using only
and provide information on their conservation, habitat and confirmed occurrences. We assessed the conservation status
phenology. In addition, since these species are restricted of the species in Brazil using the GeoCAT tool (Bachman
to the Amazon phytogeographic domain, we provide an et al. 2011), while the area of occupancy (AOO) was based
identification key for the species of Aspidosperma from the on the default cell width (2 km), as recommended by the
Brazilian Amazon. IUCN Red List guidelines (IUCN 2017).
Taxonomic treatment
Materials and methods
Reestablishment of Aspidosperma centrale and A. duckei
The study was based on material from the herbaria
collections of CEN, ESA, IAN, INPA, MG, R, RB, SPF and Aspidosperma centrale and A. duckei were considered
UEC [acronyms according to Thiers (2018)], and images synonyms of other taxa by the last two revisions of the genus
of material from the databases Atrium Biodiversity (Woodson 1951; Marcondes-Ferreira 1988). As a result of
Information System (herbarium CUZ) (AABP Atrium 2018), our ongoing research with Aspidosperma, we have come to
FMNH Botany Collections (herbarium F) (FMNH 2012), and accept these species, and thus provide updated descriptions
INCT-Virtual Herbarium of Flora and Fungi (herbaria NY with taxonomic and nomenclatural notes.
and RON) (speciesLink 2018). Species identifications were
confirmed through the analysis of protologues and types, 1. Aspidosperma centrale Markgr., Notizbl. Bot. Gart.
or images of types available at JSTOR Global Plants website Berlin-Dahlem 12(115): 560. 1935. Type: BRAZIL.
(herbaria E, G, K, L, P, PH, S, TDC and U) (Ithaka 2018), Amazonas: Parintins, Lago José-Assú, mata de terra firme,
and other online collections (herbaria A, GH and MO). 16/IX/1932, A. Ducke s.n. (lectotype: RB! [No. 24571,
Material was measured using a digital caliper and barcode RB00535019], designated here; isolectotype: RB!
the software ImageJ (Abràmoff et al. 2004). Vegetative [fragment] [No. 24571, barcode RB00535147]).
and reproductive structures were observed under a Figs. 1, 2.
Figure 1. Map indicating (A) South America and (B) the distribution of Aspidosperma centrale Markgr. and A. duckei Huber ex Ducke.
Trees 12-39.5 m high; trunk straight. Branches margin revolute, discolorous, venation craspedodromous,
cylindrical, slightly suberous, sparsely lenticellate, pubescent adaxial surface dull or lustrous, dark brown, pubescent to
to glabrescent, brown or gray, without cataphylls; latex glabrescent, primary vein immersed or flattened, secondary
red. Leaves alternate, distributed along the branches; veins immersed, tertiary veins inconspicuous, abaxial
petioles 1.2-2 cm long, flattened on the adaxial surface, not surface dull, white, velutinous, primary vein prominent,
winged, pubescent to glabrous; blades 7.11-17.5 × 2-6.9 cm, secondary veins prominulous, 18-25 pairs, tertiary veins
coriaceous, flattened, oblong or obovate, apex acute, obtuse inconspicuous. Inflorescences 9.21-11.7 cm long, terminal
or retuse, base attenuate or oblique, margin entire, base and or axillary, cyme corymbiform, rigid, tomentose. Flower
Figure 2. Aspidosperma centrale Markgr. A. reproductive branch; B. flower; C. inner part of corolla tube; D. detail of gynoecium; E.
follicle [Photos. A: J. R. Nascimento et al. 553 (NY-photo); B-D: E. Soares 221 (INPA); E: C. V. Castilho et al. 83 (INPA)].
buds with corolla lobes twisted. Flowers 6.5-7.1 mm long; Nomenclatural notes: Among the four materials reported
pedicel 0.7-0.9 × 1 mm long, tomentose. Calyx 1.7-2 × 1.5- by Markgraf (1935) in the protologue of Aspidosperma
1.7 mm, campanulate, green, without colleters, tomentose centrale, Woodson (1951) selected “Ducke 21593” from
externally, tomentose at the apex internally; lobes 5, 1 × herbaria U and US as the lectotype. However, this material
0.7-1 mm, equal, deltoid, apex acute. Corolla 5-6 × 1-1.5 should not have been chosen as a lectotype since Markgraf
mm, salverform, yellow; tube 2.5-3.2 mm long, wall not indicated “Ducke 24571” as the type using the phrase
thickened at the mouth, glabrous externally, canescent “Original der Art”. Additionally, the number “24571”
below the insertion of the anthers internally; lobes 5, is not the collector number of A. Ducke, but rather the
2.5-3 × 0.4 mm, erect, filiform, apex acute, glabrescent. catalog number of two specimens in herbarium RB (No.
Stamens 5, 2.2 mm long, included; filaments adnated to the 24571, barcodes RB00535019 and RB00535147). Therefore,
corolla tube, 1.7 mm long, canescent; anthers 0.5 mm long, the collections from herbarium RB with the catalog
free from each other and from the style-head, positioned number “24571” must be considered the type material of
above the style-head, ovate, apex acute or apiculate, base Aspidosperma centrale. To avoid confusion, we elected one of
cordate. Carpels 2, 1.4-1.6 mm long; ovary 0.5-0.7 × 0.7 these types as a lectotype, and choose the material “Ducke
mm, superior, hemisyncarpous, globoid, glabrous; style s.n.”, RB No. “24571” with barcode “RB00535019”, because
0.5 mm long, cylindrical; style-head 0.4 mm long, main it was the best preserved.
body globoid, with 2 apical appendages, ca. 0.2 mm long, Taxonomic notes: Aspidosperma centrale was synonymized
glabrous. Follicles 2 or 1 by abortion, 10.5-13.1 × 5.5-8.2 with A. album (Vahl) Benoist ex Pichon by Woodson (1951),
cm, flattened, dolabriform or suborbicular, sulcate, stipitate, but later Marcondes-Ferreira (1988) considered it a synonym
mucronate, midrib conspicuous, lenticels inconspicuous, of A. spruceanum Benth. ex Müll.Arg. Here, due to observed
brown, pubescent to glabrescent. Seeds 6-6.6 cm diam., differences, we are accepting Aspidosperma centrale. This
winged, orbicular, yellow, glabrous; seminal nucleus central, species resembles two Amazonian species, Aspidosperma
without radial lines, nucleus 2.6 cm diam. sandwithianum Markgr. and A. spruceanum, mainly in leaf
Material examined: BRAZIL. Amazonas: Manaus, characteristics, but differs from A. sandwithianum by having
Distrito Agropecuário, 90 km NNE de Manaus, Reserva smaller (ca. 1.2-2 vs. ca. 2.2-4 cm long), and pubescent
1501 (km 41), 6/XII/1991, A. A. Oliveira et al. 261 (INPA, to glabrous (vs. tomentose) petioles, and by flowers with
SPF); Manaus, Reserva Florestal Ducke, próximo ao Igarapé corolla tube canescent below the insertion of the anthers
Sempre Viva, 26/IX/1957, E. Ferreira 109-57 (INPA); internally (vs. tomentose). Furthermore, Aspidosperma
Manaus, Reserva Florestal Ducke, Manaus-Itacoatiara, centrale differs from A. spruceanum by having leaves with
km 26, 26/VII/1994, J. R. Nascimento et al. 553 (INPA, secondary veins immersed on the adaxial surface (vs.
NY-photo, SPF); Manaus, Reserva Florestal Adolfo prominulous or flattened), flowers with smaller pedicels
Ducke, Rodovia Manaus-Itacoatiara, km 26, trilha (0.7-0.9 vs. 3.7-4 mm long), and brown follicles (vs. yellow).
L-O7, km 3,5, C. V. Castilho et al. 83 (INPA); Maués,
along Rio Apoquitaua, just above mouth of Rio Pacoval, 2. Aspidosperma duckei Huber ex Ducke, Arch. Jard. Bot.
27/VII/1983, J. L. Zarucchi 3210 (INPA); Presidente Rio de Janeiro 3: 244. 1922. Type: BRAZIL. Pará: Óbidos,
Figueiredo, Rio Urubu, Cachoeira de Iracema, 22/IX/1949, Serra da Escama, 22/IX/1910, A. Ducke s.n. (lectotype: MG!
R. L. Fróes 25361 (INPA, NY-photo). Pará: Oriximiná, Porto [barcode MG011040], designated by Woodson (1951: 139);
Trombetas, próximo à área industrial, 9/X/1986, E. Soares isolectotypes: G [barcode G00169271] [digital image!],
221 (INPA); Porto de Moz, Rio Xingu, margem esquerda R! [barcode R000002228], RB! [No. 15815, barcode
do rio, região onde foi feito um levantamento estatístico RB00535035]).
florestal pelo IAN, SPVEA e FAO, 18/IX/1955, R. L. Fróes Figs. 1, 3.
32377 (IAN); Santarém, Reserva Florestal de Curuá-Una, Trees 10-40 m high; trunk straight. Branches cylindrical,
Planalto Alto II a 1,5 km do Flanco, 7/X/1963, Tressel 22 not suberous, sparsely lenticellate, glabrescent, brown,
(INPA). without cataphylls; latex white. Leaves alternate, distributed
Distribution and habitat: Aspidosperma centrale occurs along the branches; petioles 1.4-2.9 cm long, flattened on the
in Brazil and Colombia. In Brazil, it is found in the North adaxial surface, not winged, pubescent to glabrous; blades
Region (states of Amazonas and Pará), occurring in forests 6.7-17.3 × 3.7-8.3 cm, chartaceous, flattened, elliptic or
on hilly terrain (terra firme forest) in the Amazon rainforest. obovate, apex acute or acuminate, base cuneate or oblique,
Phenology: Flowers from April to September and fruits margin entire, base and margin flattened, discolorous,
from July to December. venation eucamptodromous, adaxial surface dull, dark
Conservation status: Aspidosperma centrale has an brown, glabrescent, primary vein immersed, secondary veins
AOO of 68 km2 and is considered endangered according immersed, tertiary veins inconspicuous, abaxial surface
to IUCN guidelines (IUCN 2017). It has 16 confirmed dull, light brown, glabrescent, primary vein prominent,
records for Brazil (.kml file available at https://figshare. secondary veins prominent, 12-17 pairs, tertiary veins
com/s/81cefc2ee83949c54a21). inconspicuous. Inflorescences ca. 4 cm long, subterminal,
cyme corymbiform, rigid, tomentose. Flower buds with glabrescent internally; lobes 5, 7.9-11 × 2-2.8 mm, patent,
corolla lobes not twisted. Flowers 19-21 mm long; pedicel oblong, apex rounded, tomentose. Stamens 5, 4.6-6.7
2.5-3.1 × 0.6-1.2 mm long, tomentose. Calyx 2.2-3 × 2-2.7 mm long, included; filaments adnated to the corolla tube,
mm, campanulate, green, without colleters, tomentose 3.4-5.2 mm long, glabrescent; anthers 1.2-1.5 mm long,
externally, glabrous internally; lobes 5, 1.8-2.2 × 0.8-1 free from each other and from the style-head, positioned
mm, equal, deltoid or triangular, apex acute. Corolla 14.8- above the style-head, ovate, apex apiculate, base cordate.
18.4 × 1.9-2.5 mm, salverform, white; tube 6.9-7.4 mm Carpels 2, 2.9-3.2 mm long; ovary 0.76-1.1 × 0.6-1.4 mm,
long, wall thickened at the mouth, tomentose externally, superior, hemisyncarpous, globoid, tomentose; style 1.4-
Figure 3. Aspidosperma duckei Huber ex Ducke. A. branch; B. flower; C. inner part of corolla tube; D. detail of calyx and gynoecium;
E. follicle [Photos. A-D: A. Ducke s.n. (RB No. 11402); E: A. A. Santos 3641 (RB)].
1.9 mm long, cylindrical; style-head 0.6-0.7 mm long, main its tomentose ovary (vs. glabrous). Here, we agree with
body globoid, with 2 apical appendages, ca. 0.3 mm long, Duarte and accept Aspidosperma duckei.
glabrous. Follicles 2 or 1 by abortion, 20.1-25.2 × 11.3-13.8
cm, flattened, dolabriform, smooth, stipitate, mucronate, New records for Brazil
midrib inconspicuous, lenticels inconspicuous, brown,
glabrescent. Seeds 8.3-9.7 cm diam., winged, orbicular, Our research revealed that the records for Aspidosperma
yellow, glabrous; seminal nucleus central, without radial steinbachii and A. tambopatense are new for Brazil.
lines, nucleus 2.8 cm diam. Aspidosperma steinbachii was accepted by Woodson (1951),
Material examined: BRAZIL. Acre: Cruzeiro do Sul, but Marcondes-Ferreira (1988) synonymized this species
margem esquerda, do Rio Juruá, Igarapé Viseu, 21/III/1992, and A. tambopatense with A. spruceanum and A. parvifolium
C. A. Cid Ferreira et al. 10882 (INPA). Amazonas: Parintins, A.DC., respectively. However, since Marcondes-Ferreira’s
30/VIII/1932, A. Ducke s.n. (RB No. 24574). Mato Grosso: revision was not actually published, both species remain
Aripuanã, Gleba Aripuanã a 25 km da cidade em direção accepted, which is supported by our observations of
sudoeste, 5/VII/1997, G. F. Árbocz et al. 4180 (ESA, UEC). morphological characteristics.
Pará: Óbidos, 20/X/1919, A. Ducke s.n. (RB No. 11402);
Oriximiná, Mineração Rio do Norte, próximo à Vila 1. Aspidosperma steinbachii Markgr., Notizbl. Bot. Gart.
Madezati, 17/VIII/1989, E. Soares 522 (INPA); Santarém, Berlin-Dahlem 9(90): 1158. 1927. Type: BOLIVIA. Santa
estrada de Belterra, 6/X/1962, A. P. Duarte 7016 (RB). Cruz: Sara, bosques de Buena Vista, 450 m, 2/X/1925,
Rondônia: Ariquemes, 21 km SE of Ariquemes on hwy. J. Steinbach 7261 (lectotype: K [barcode K000587712]
BR-364, then 1 km E on “Linea 45”, 17/III/1987, M. Nee [digital image!], designated here; isolectotypes: A [barcode
34422 (INPA); Porto Velho, BR-364 sentido Jaci Paraná- 00057226] [digital image!], E [barcode E00259701] [digital
Abunã, 20/VI/2012, A. A. Santos 3641 (RB). image!], F [barcode V0092485F] [digital image!], G [barcode
Additional material examined: PERU. Madre de Dios: G00169334] [digital image!], GH [barcode 00057227] [digital
Manu, 10/IX/1989, R. B. Foster & S. H. Beltran 13158 (UEC). image!], MO, NY [barcode 00297995] [digital image!], PH
Distribution and habitat: Aspidosperma duckei occurs in [No. 653008, barcode PH00004864] [digital image!], RB!
Bolivia, Brazil and Peru. In Brazil, it is found in the North [fragment] [No. 452001, barcode RB00535084], S [No.
(states of Acre, Amazonas and Rondônia) and in the Central S04-1797] [digital image!], U [barcode U0000491] [digital
West (state of Mato Grosso) Regions, occurring in forests on image!]; photo: F [No. negative 4431] [digital image!]).
hilly terrain (terra firme forest) in the Amazon rainforest, and Figs. 4, 5.
in the transition zone between the Amazon and the Cerrado. Trees 6-15 m high; trunk straight. Branches cylindrical,
Phenology: Flowers from September to October and not suberous, sparsely lenticellate, pubescent to glabrescent,
fruits from September to May. brown or black, without cataphylls; latex red. Leaves
Conservation status: Aspidosperma duckei has an AOO alternate, distributed along the branches; petioles 1.5-
of 108 km2 and is considered endangered according to 3.1 cm long, flattened on the adaxial surface, not winged,
IUCN guidelines (IUCN 2017). It has 19 confirmed records tomentose to pubescent; blades 10-16 × 3-4.7 cm,
for Brazil (.kml file available at https://figshare.com/ chartaceous, flattened, oblong or elliptic, apex acute or
s/0b298a757a82f5315dba). acuminate, base cuneate, attenuate or oblique, margin
Nomenclatural notes: Woodson (1951) designated the entire, base and margin frequently revolute, discolorous,
material “Ducke 11040” deposited in herbaria G, P and US as venation craspedodromous, adaxial surface dull or
the lectotype, which is one of the five gatherings mentioned lustrous, dark brown, glabrous, primary vein prominulous,
by Ducke (1922) in the protologue of Aspidosperma duckei. secondary veins prominent or prominulous, tertiary veins
However, most of the numbers of Ducke’s collections are inconspicuous, abaxial surface dull, light brown, pubescent
known to actually be catalog numbers of herbaria MG or RB. to glabrous, primary vein prominulous, secondary veins
In the present case, Ducke mentioned in the introduction prominent or prominulous, 25-30 pairs, tertiary veins
of the publication that specimen numbers indicate the inconspicuous. Inflorescences 10.3-12 cm long, terminal or
catalog numbers of the herbarium MG. Thus, the lectotype axillary, cyme corymbiform, not rigid, tomentose. Flower
designated by Woodson is actually “Ducke s.n.”, and refers buds with corolla lobes twisted. Flowers 7-9.5 mm long;
to the specimen deposited in MG (barcode MG011040). pedicel 1-2.5 × 1 mm long, tomentose. Calyx 3.5-3.8 × 2
Taxonomic notes: Aspidosperma duckei was considered mm, campanulate, green, without colleters, tomentose on
a synonym of A. macrocarpon Mart. & Zucc. by Woodson both surfaces; lobes 5, 2.5-3.2 × 1-1.3 mm, equal, deltoid
(1951) and Marcondes-Ferreira (1988). However, Duarte or ovoid, apex acute or obtuse. Corolla 4.5-5.5 × 1-1.2 mm,
(1970), in a preliminary version of what would be a new salverform, yellow; tube 3 mm long, wall not thickened
revision for the genus, gave indications that Aspidosperma at the mouth, glabrous externally, pubescent below the
duckei could be a “good species”, distinguishing it from A. insertion of the anthers internally; lobes 5, 1.5-2.5 × 0.5
macrocarpon by its larger petioles and fruit stipes, and by mm, erect, filiform, apex acute, glabrescent. Stamens 5,
Figure 4. Map indicating (A) South America and (B) the distribution of Aspidosperma steinbachii Markgr. and A. tambopatense A.H.Gentry.
2-2.4 mm long, included; filaments adnated to the corolla Bolivia (Markgraf 1927), occurring in moist forests in the
tube, 1.6-2 mm long, pubescent; anthers 0.4 mm long, free Amazon rainforest, but later, Woodson (1951) reported
from each other and from the style-head, positioned above it also for Peru. This is the first record of Aspidosperma
the style-head, ovate, apex acute or obtuse, base cordate. steinbachii for Brazil, where it is found between southeastern
Carpels 2, 1.7-1.95 mm long; ovary 0.5 × 0.7 mm, superior, Rondônia and northwestern Mato Grosso states, occurring
hemisyncarpous, ovoid, glabrous; style 0.5-0.75 mm long, in forests on hilly terrain (terra firme forest) in the Amazon,
cylindrical; style-head 0.7 mm long, main body globoid, and in the transition zone between the Amazon and the
with 2 apical appendages, ca. 0.2 mm long, pubescent to Cerrado.
glabrescent. Follicles 2 or 1 by abortion, 7-7.11 × 3.55-3.78 Phenology: Flowers from July to October and fruits
cm, flattened, dolabriform or suborbicular, sulcate, stipitate, in November.
mucronate, midrib conspicuous, lenticels inconspicuous, Conservation status: Aspidosperma steinbachii has an
blackish-green, velutinous. Seeds 5.8 cm diam., winged, AOO of 16 km2 and is considered endangered according to
orbicular, yellow, glabrous; seminal nucleus central, without IUCN guidelines (IUCN 2017). It has only four confirmed
radial lines, nucleus 1.5-2 cm diam. records for Brazil (.kml file available at https://figshare.
Material examined: BRAZIL. Mato Grosso: Juruena, Rio com/s/7922a4239a40c3b300d0).
Juruena, arredores da cidade, 13/VII/1977, M. G. Silva & Nomenclatural notes: According to Melchior (1926),
J. Maria 3325 (IAN, MG, RB, SPF). Rondônia: Corumbiara, who edited the publication “Plantae Steinbachianae”, the
21/XII/1996, H. S. Pereira & C. R. Souza 2112-084-1996 material collected by J. Steinbach in Bolivia were managed by
(RON-photo); Porto Velho, estrada para Alvorada, linha herbarium B, among which was the holotype of Aspidosperma
5, beira de estrada, 29/IX/2013, N. C. Bigio et al. 1101 (RB, steinbachii. However, since this specimen was destroyed in
RON-photo); Vilhena, no meio do pasto, ponto 1220, 6/ 1943, we elected its best-preserved “isotype” as a lectotype,
XII/2013, N. C. Bigio et al. 1231 (RON-photo). the material “Steinbach 7261” from herbarium K.
Additional material examined: BOLIVIA. La Paz: Taxonomic notes: Aspidosperma steinbachii shares some
Province of S. Yungas, basin of Rio Bopi, San Bartolome (near morphological features with both A. cruentum Woodson
Calisaya), 1-22/VII/1939, B. A. Krukoff 10290 (NY-photo). and A. melanocalyx Müll.Arg., but differs from the former
Santa Cruz: Sara, Buenavista, 27/XI/1925, J. Steinbach 7356a mainly by having smaller (7-7.11 vs. 19-22 cm long) and
(F-negative). PERU. Madre de Dios: Manu province, Río Los blackish-green (vs. yellow) follicles, and from the latter by
Amigos II, 14/VI/1995, P. Núñez et al. 16862 (CUZ-photo). its pubescent to glabrous leaves on the abaxial surface (vs.
Distribution and habitat: Aspidosperma steinbachii usually velutinous), and by its non-rigid inflorescences
was initially described with a distribution restricted to (vs. rigid).
2. Aspidosperma tambopatense A.H.Gentry, Ann. Missouri = Aspidosperma occidentale Markgr. (1940: 133),
Bot. Gard. 71(4): 1075. 1984. Type: PERU. Madre de Dios: non Malme. Type: BRAZIL. Acre: Rio Acre, IV/1911,
Tambopata Reserve, 26 km S of Puerto Maldonado on E E. H. G. Ule 9700 (syntypes: K [barcode K000587675]
side of Rio Tambopata, 12/XI/1979, G. S. Hartshorn 2421 [digital image!], MG! [barcode 014536]), nom. illeg., pro
(holotype: MO [barcode MO-2958528] [digital image!]; syn.
isotypes: CR, F [barcode V0421499F] [digital image!], MO Trees 8-40 m high; trunk straight. Branches cylindrical,
[barcode MO-1069092] [digital image!], USM). not suberous, densely lenticellate, glabrescent to glabrous,
Figs. 4, 6. brown, with cataphylls; latex white. Leaves alternate,
Figure 5. Aspidosperma steinbachii Markgr. A. reproductive branch; B. flower; C. inner part of corolla tube; D. detail of calyx and
gynoecium; E. detail of gynoecium [Photos. A: N. C. Bigio 1101 (RON-photo); B-E: M. G. Silva & J. Maria 3325 (IAN)].
concentrated at the apex of branches; petioles 1.2-2.9 cm primary vein immersed, secondary veins flattened, tertiary
long, subcaniculate or flattened on the adaxial surface, veins conspicuous, abaxial surface dull, light green, glabrous,
not winged, glabrescent to glabrous; blades 6.4-13 × 2.4- primary vein prominent, secondary veins flattened, 16-25
5.6 cm, chartaceous, flattened, elliptic or obovate, apex pairs, tertiary veins inconspicuous. Inflorescences 4-5.2 cm
acute or acuminate, base attenuate or oblique, margin long, subterminal, cyme corymbiform, not rigid, glabrescent.
entire, base and margin flattened, discolorous, venation Flower buds with corolla lobes not twisted. Flowers 4.4-7.3
eucamptodromous, adaxial surface dull, dark green, glabrous, mm long; pedicel 0.4-1.9 × 0.5-0.6 mm, tomentose. Calyx
Figure 6. Aspidosperma tambopatense A.H.Gentry. A. reproductive branch; B. flower bud; C. corolla lobes; D. detail of gynoecium;
E. inner part of corolla tube; F. follicle; G. seed [Photos. A: C. A. Cid 3079 (NY-photo); B-E: J. Bosco 112 (UEC); F-G: B. A. Krukoff
5470 (NY-photo)].
1.7-2.6 × 1.2-1.4 mm, campanulate, green, without colleters, at low water, Seringal União, within proposed Reserva
tomentose externally, glabrous internally; lobes 5, 0.8-1.2 Indígena Praia do Carapanã (Kaxinawá Indians), Seringal
× 0.4-0.7 mm, equal, deltoid, apex acute. Corolla 3.7-4.7 Mucuripe, Colocação Remanso, 19/IX/1994, D. C. Daly et al.
× 1.2-1.6 mm, tubular, white; tube 3.5 mm long, wall not 8244 (INPA, NY-photo); Xapuri, Reserva Extrativista “Chico
thickened at the mouth, tomentose externally, pubescent Mendes”, Seringal Cachoeira, BR-317, ramal cachoeira 16
below the insertion of the anthers internally; lobes 5, 0.8- km, 14/XI/2009, H. Medeiros 235 (RB); 45 km from Rio
1 × 0.6-0.8 mm, patent, deltoid, apex acute, tomentose. Branco on Rio Branco–Porto Velho road, 9/X/1980, S. R.
Stamens 5, 2.1-2.6 mm long, included; filaments adnated Lowrie et al. 447 (INPA, RB).
to the corolla tube, 1.9-2.2 mm long, pubescent; anthers Additional material examined: BOLIVIA. Pando:
0.6-0.8 mm long, free from each other and from the style- Manuripi, Empresa, 7/X/1977, W. Terceros 1394 (INPA).
head, positioned above the style-head, ovate, apex acute, Distribution and habitat: Aspidosperma tambopatense
base cordate. Carpels 2, 1.3-2.4 mm long; ovary 0.8 × 0.8 was originally reported only for Peru, occurring in moist
mm, superior, hemisyncarpous, ovoid, tomentose; style forests along the base of the Andes (Gentry 1984), but later
0.3-1 mm long, cylindrical; style-head 0.1-0.3 mm long, its distribution was found to extend to Bolivia (Parker III
main body ellipsoid, with 2 apical appendages, ca. 0.3 mm & Bailey 1991; Killeen et al. 1993). This is the first record
long, pubescent. Follicles 2 or 1 by abortion, 5.7-7 × 3.7-4.5 of Aspidosperma tambopatense for Brazil, being distributed
cm, flattened, dolabriform, smooth, sessile or substipitate, throughout the territory of the state of Acre, occurring
mucronate, midrib inconspicuous, lenticels conspicuous,
mainly in forests on hilly terrain (terra firme forest) in the
brown, glabrescent. Seeds 6.1 cm diam., winged, orbicular,
Amazon rainforest, but also in forests on level terrain (várzea
yellow, glabrous; seminal nucleus central, without radial
forest).
lines, nucleus 2.1 cm diam.
Phenology: Flowers from September to December and
Material examined: BRAZIL. Acre: Brasiléia, Bom Futuro,
fruits from October to December.
Reserva Extrativista Chico Mendes, km 52 of Brasiléia-Assis
Conservation status: Aspidosperma tambopatense has an
Brasil road, 18 km on ramal (side road) “Tocandeira”, 23/
AOO of 48 km2 and is considered endangered according to
IX/2003, D. C. Daly et al. 11974 (RB); Brasiléia, estrada
para Assis Brasil km 8, 1/XI/1980, C. A. Cid 3079 (INPA); IUCN guidelines (IUCN 2017). It has 12 confirmed records
Capixaba, Projeto de Assentamento Extrativista (PAE) for Brazil (.kml file available at https://figshare.com/s/
São Luis do Remanso, Colocação Estrangeiro, 30 km W of cd094331833eedd5d39b).
Capixaba, tem 15-21 km N on new unpaved access road, Nomenclatural notes: During our studies, we realized
30/IX/2003, D. C. Daly et al. 12039 (RB); Cruzeiro do Sul, that Aspidosperma occidentale of Markgraf (1940) and A.
Rio Juruá, margem esquerda ao lado do Igarapé Viseu, a 5 tambopatense of Gentry (1984) represent the same taxon.
km da margem, 4/XI/1991, C. A. Cid Ferreira et al. 10568 However, the name Aspidosperma occidentale is a later
(NY-photo); Rio Branco, Campus Universitário, 25/X/1983, homonym of the validly published name A. occidentale of
A. Rosas & M. B. Guimarães 32 (INPA); Rio Branco, Campus Malme (1927), thus making it an illegitimate name. As a
da UFAC, 22/IX/1989, J. Bosco 112 (UEC); Sena Madureira, result, we consider Aspidosperma occidentale of Markgraf a
basin of Rio Iaco (tributary of Rio Purus), Fazenda São pro synonym of the validly published name A. tambopatense.
Jorge I, property of Acre Brasil Verde, timber concession Taxonomic notes: Aspidosperma tambopatense is similar
of Laminados Triunfo Ltda., 107 km NW of Rio Branco to A. williamii Duarte, mainly in vegetative characteristics,
on BR-264, then ca. 22 km on Toco Preto access road, 5/ but it can be distinguished by having leaves with flattened
VII/2008, D. C. Daly et al. 13179 (RB); [Sena Madureira], secondary veins on both surfaces (vs. prominulous), deltoid
near mouth of Rio Macauhan (tributary of the Rio Yaco), corolla lobes (vs. orbicular), and sessile or substipitate
13/VIII/1933, B. A. Krukoff 5470 (NY-photo); Tarauacá, follicles (vs. stipitate), with an inconspicuous midrib (vs.
Reserva Indígena Praia do Carapanã, Rio Tarauacá, river conspicuous).
4. Leaves with prominulous secondary veins on both surfaces; flowers with orbicular corolla lobes; follicles stipitate, with
a conspicuous midrib��������������������������������������������������������������������������������������������������������������� A. williamii (Figs. 10A-B, 12B)
4’. Leaves with flattened secondary veins on both surfaces; flowers with deltoid corolla lobes; follicles sessile or substipitate,
with an inconspicuous midrib������������������������������������������������������������������������������������������������� A. tambopatense (Figs. 6, 12C)
5. Leaves 1.4-1.8× longer than wide, white on the abaxial surface; flowers more than 4 mm long, glabrescent externally;
follicles glabrescent, brown��������������������������������������������������������������������������������������������������A. subincanum (Figs. 7B-C, 12D)
5’. Leaves 3× longer than wide, yellow on the abaxial surface; flowers up to 3 mm long, tomentose externally; follicles
pubescent, yellow����������������������������������������������������������������������������������������������������������������������������� A. ulei (Figs. 10C-E, 12E)
6. Leaves with reticulodromous venation; inflorescences leaf-opposed; seed nuclei basal and apical���������������������������������
������������������������������������������������������������������������������������������������������������������������������������������������������������� A. cuspa (Figs. 7D, 12F)
6’. Leaves with craspedodromous, brochidodromous or eucamptodromous venation; inflorescences axillary, subterminal
or terminal; seed nuclei central or lateral������������������������������������������������������������������������������������������������������������������������������ 7
7. Leaves with craspedodromous venation���������������������������������������������������������������������������������������������������������������������������� 8
7’. Leaves with brochidodromous or eucamptodromous venation������������������������������������������������������������������������������������ 21
8. Leaves with immersed secondary veins on the abaxial surface���������������������������������������������������������������������������������������� 9
8’. Leaves with proeminent, prominulous or flattened secondary veins on the abaxial surface�������������������������������������� 12
9. Branches with white latex; flowers more than 13 mm long, corolla lobes more than 8 mm long, ovary tomentose; seed
nuclei with radial lines�������������������������������������������������������������������������������������������������������������� A. schultesii (Figs. 7F-G, 12G)
9’. Branches with red latex; flowers up to 11 mm long, corolla lobes up to 5 mm long, ovary glabrous; seed nuclei without
radial lines����������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 10
10. Leaves with crenulate margin; corolla ca. 3 mm wide�������������������������������������������� A. leucocymosum (Figs. 10F-G, 12H)
10’. Leaves with entire margin; corolla ca. 2 mm wide������������������������������������������������������������������������������������������������������� 11
11. Leaves canescent on the abaxial surface; follicles yellow������������������������������������������A. obscurinervium (Figs. 7H-J, 12I)
11’. Leaves glabrous on the abaxial surface; follicles brown��������������������������������������������A. desmanthum (Figs. 10H-J, 12J)
12. Leaves with 39-41 pairs of secondary veins������������������������������������������������������������������������������������������������������������������ 13
12’. Leaves with 18-34 pairs of secondary veins����������������������������������������������������������������������������������������������������������������� 14
13. Leaves 2× longer than wide, glabrous on the abaxial surface; inflorescences in corymbiform cymes���������������������������
������������������������������������������������������������������������������������������������������������������������������������������������ A. araracanga (Figs. 8A-B, 12K)
13’. Leaves 2.6-3.2× longer than wide, velutinous on the abaxial surface; inflorescences in panicles���������������������������������
������������������������������������������������������������������������������������������������������������������������������������������A. verruculosum (Figs. 10K-M, 12L)
14. Leaves with prominent, prominulous or flattened secondary veins on the adaxial surface�������������������������������������� 15
14’. Leaves with immersed secondary veins on the adaxial surface����������������������������������������������������������������������������������� 18
15. Flowers up to 3.5 mm long; follicles pyriform; seeds chartaceous�������������������������A. pachypterum (Figs. 10N-Q, 12M)
15’. Flowers more than 5.2 mm long; follicles dolabriform or suborbicular; seeds membranaceous������������������������������ 16
16. Leaves brown on the abaxial surface, tertiary veins inconspicuous on the adaxial surface; follicles blackish-green���
�������������������������������������������������������������������������������������������������������������������������������������������������������A. steinbachii (Figs. 5, 12N)
16’. Leaves white on the abaxial surface, tertiary veins conspicuous on the adaxial surface; follicles brown or yellow ����
����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 17
17. Leaves dark green on the adaxial surface; flowers with corolla lobes ca. 2.5 mm long; follicles brown�������������������������
������������������������������������������������������������������������������������������������������������������������������������������������������ A. album (Figs. 10R-T, 12O)
17’. Leaves olive green or brown on the adaxial surface; flowers with corolla lobes ca. 4 mm long; follicles yellow�����������
���������������������������������������������������������������������������������������������������������������������������������������������A. spruceanum (Figs. 11A-C, 12P)
18. Flowers with tubular corolla, lobes up to 1.5 mm long���������������������������������������������������� A. eteanum (Figs. 8C, E, 12Q)
18’. Flowers with salverform corolla, lobes more than 2.5 mm long��������������������������������������������������������������������������������� 19
19. Leaves 2-2.2× longer than wide; flowers with corolla lobes smaller or equal to the tube (0.8-1×)��������������������������������
���������������������������������������������������������������������������������������������������������������������������������������������������A. neblinae (Figs. 11D-F, 12R)
19’. Leaves 2.4-3.8× longer than wide; flowers with corolla lobes larger than the tube (1.1-1.7×)���������������������������������� 20
20. Leaf petioles ca. 2.2-4 cm long, tomentose; flowers with corolla tube tomentose below the insertion of the anthers
internally����������������������������������������������������������������������������������������������������������������������������� A. sandwithianum (Figs. 8D, 12S)
20’. Leaf petioles ca. 1.2-2 cm long, pubescent to glabrous; flowers with corolla tube canescent below the insertion of
the anthers internally����������������������������������������������������������������������������������������������������������������������� A. centrale (Figs. 2, 12T)
21. Leaf margins with deep revolute base��������������������������������������������������������������������������������������������������������������������������� 22
21’. Leaf margins with flattened base���������������������������������������������������������������������������������������������������������������������������������� 24
22. Leaves with eucamptodromous venation�����������������������������������������������������������������������A. auriculatum (Figs. 11G, 12U)
22’. Leaves with brochidodromous venation����������������������������������������������������������������������������������������������������������������������� 23
23. Branches angular; leaves subopposite or opposite; flowers with glabrous ovary�������� A. salgadense (Figs. 11H-I, 12V)
23’. Branches cylindrical; leaves usually alternate; flowers with tomentose ovary������������������A. oblongum (Figs. 8F, 12W)
24. Leaves with brochidodromous venation������������������������������������������������������������������������������������������������������������������������ 25
24’. Leaves with eucamptodromous venation��������������������������������������������������������������������������������������������������������������������� 28
25. Branches densely lenticellate; follicles verrucose, with inconspicuous lenticels���������� A. brasiliense (Figs. 8G-H, 12X)
25’. Branches sparsely lenticellate; follicles smooth, with conspicuous lenticels�������������������������������������������������������������� 26
26. Leaf petioles more than 2 cm long; follicles botuliform, inflated������������������������������� A. cylindrocarpon (Figs. 9A, 12Y)
26’ Leaf petioles up to 1.5 cm long; follicles dolabriform or falciform, flattened������������������������������������������������������������� 27
27. Leaves with 30-38 pairs of secondary veins; flowers up to 9 mm long, calyx with 6-7 lobes�����������������������������������������
�����������������������������������������������������������������������������������������������������������������������������������������������A. darienense (Figs. 11J-K, 12Z)
27’. Leaves with 23-25 pairs of secondary veins; flowers more than 17 mm long, calyx with 5 lobes���������������������������������
��������������������������������������������������������������������������������������������������������������������������������������������� A. inundatum (Figs. 11L-M, 13A)
28. Leaves more than 20 cm long; flowers with corolla lobes ca. 6 mm long; follicles velutinous���������������������������������������
���������������������������������������������������������������������������������������������������������������������������������������������� A. myristicifolium (Figs. 9B, 13B)
28’. Leaves up to 17 cm long; flowers with corolla lobes up to 5 mm long; follicles pubescent to glabrescent��������������� 29
29. Leaves with prominent secondary veins on the abaxial surface; follicles more than 11 cm long ������������������������������ 30
29’. Leaves with flattened or immersed secondary veins on the abaxial surface; follicles up to 9 cm long��������������������� 31
30. Leaves glabrescent on the abaxial surface; flowers with tomentose ovary��������������������������������� A. duckei (Figs. 3, 13C)
30’. Leaves pubescent on the abaxial surface; flowers with glabrous ovary�������������������� A. macrocarpon (Figs. 9C-D, 13D)
31. Leaves usually revolute, velutinous on the abaxial surface; flowers with tomentose ovary�������������������������������������������
�����������������������������������������������������������������������������������������������������������������������������������������������A. carapanauba (Figs. 9E-F, 13E)
31’. Leaves flattened, pubescent on the abaxial surface; flowers with glabrous ovary����������������������������������������������������� 32
32. Flowers with salverform corolla, glabrous externally, lobes equal to or larger than the tube (1-1.6×); follicles suborbicular,
smooth�����������������������������������������������������������������������������������������������������������������������������������������A. rigidum (Figs. 9G-H, 13F)
32’. Flowers with tubular corolla, tomentose externally, lobes smaller than the tube (0.2-0.5×); follicles dolabriform,
verrucose or spinescent�������������������������������������������������������������������������������������������������������������������������������������������������������� 33
33. Inflorescences axillary; flowers with corolla ca. 6 mm long, lobes up to 0.5 mm long��� A. nitidum (Figs. 11N-P, 13G)
33’. Inflorescences terminal; flowers with corolla ca. 7.5 mm long, lobes more than 1 mm long������������������������������������ 34
34. Leaves coriaceous, 1.7-2× longer than wide; flowers with calyx lobes equal; follicles spinescent����������������������������������
������������������������������������������������������������������������������������������������������������������������������������������������� A. excelsum (Figs. 11Q-S, 13H)
34’. Leaves chartaceous, 2.8-3× longer than wide; flowers with calyx lobes unequal; follicles verrucose����������������������������
�������������������������������������������������������������������������������������������������������������������������������������������� A. marcgravianum (Figs. 9I-J, 13I)
Figure 7. Branches, leaves, fruits and seeds of Aspidosperma species from the Brazilian Amazon. A. A. multiflorum A.DC.; B-C. A.
subincanum Mart.; D. A. cuspa (Kunth) S.F.Blake; E-G. A. schultesii Woodson; H-J. A. obscurinervium Azambuja [Photos. A-C: A. C. D.
Castello; D: A. L. Scudeler; E-I: A. S. de S. Pereira; J: D. C. Daly & N. C. Bigio (Herbarium RON archive)].
Figure 8. Branches, leaves, fruits and seeds of Aspidosperma species from the Brazilian Amazon. A-B. A. araracanga Marc.-Ferr.; C,
E. A. eteanum Markgr.; D. A. sandwithianum Markgr.; F. A. oblongum A.DC.; G-H. A. brasiliense A.S.S.Pereira & A.C.D.Castello [Photos.
A, C-D, F: A. S. de S. Pereira; B: U. Mehlig; E: J. L. L. de Abreu; G-H: A. C. D. Castello].
Figure 9. Branches, leaves, fruits and seeds of Aspidosperma species from the Brazilian Amazon. A. A. cylindrocarpon Müll.Arg.; B.
A. myristicifolium (Markgr.) Woodson; C-D. A. macrocarpon Mart. & Zucc.; E-F. A. carapanauba Pichon; G-H. A. rigidum Rusby; I-J. A.
marcgravianum Woodson [Photos. A: E. Y. Kataoka; B, G-H: R. Aguilar; C-D: A. S. de S. Pereira; E: W. Milliken; F: D. Sasaki; I: A. S. de
S. Pereira; J: R. C. dos Santos].
Figure 10. Branches, leaves, fruits and seeds of Aspidosperma species from the Brazilian Amazon. A-B. A. williamii Duarte; C-E. A. ulei
Markgr.; F-G. A. leucocymosum Kuhlm.; H-J. A. desmanthum Benth. ex Müll.Arg.; K-M. A. verruculosum Müll.Arg.; N-Q. A. pachypterum
Müll.Arg.; R-T. A. album (Vahl) Benoist ex Pichon [Photos. A: W. Rodrigues 8978 (RB); B: M. A. Freitas et al. 806 (NY-photo); C-D: E.
H. G. Ule 8451 (L-photo, U-photo); E: L. Coradin & M. dos R. Cordeiro 914 (CEN); F-G: A. Ducke s.n. (P barcode P00645102-photo);
H: G. T. Prance et al. 6741 (NY-photo); I: R. Spruce s.n. (E barcode E00259707-photo); J: M. G. Silva & C. Rosario 4031 (NY-photo);
K-L: R. Spruce 3328 (TDC-photo); M: A. Ducke s.n. (NY barcode 01172496-photo); N-Q: R. Spruce 3345 (NY-photo, P-photo); R-S:
Forest Department of British Guiana 2109 (K-photo); T: L. C. Richard s.n. (P barcode P00645147-photo)].
Figure 11. Leaves and fruits of Aspidosperma species from the Brazilian Amazon. A-C. A. spruceanum Benth. ex Müll.Arg.; D-F. A.
neblinae Monach.; G. A. auriculatum Markgr.; H-I. A. salgadense Markgr.; J-K. A. darienense Woodson ex Dwyer; L-M. A. inundatum
Ducke; N-P. A. nitidum Benth. ex Müll.Arg.; Q-S. A. excelsum Benth. [Photos. A-B: R. Spruce 2265 (F-photo); C: M. F. Silva et al. 1704
(INPA); D-E: B. Maguire et al. 37084 (NY-photo); F: B. Maguire et al. 37284 (S-photo); G: A. Ducke s.n. (RB No. 22450); H-I: A. Ducke
s.n. (RB No. 22456); J: N. T. Silva 1325 (NY-photo); K: M. J. Pires & N. T. Silva 1390 (MG); L: A. Ducke s.n. (R barcode R000007307); M:
A. Ducke s.n. (RB No. 15814); N-O: R. Spruce 1657 (E-photo); P: A. Ducke s.n. (K barcode K000587691-photo); Q-R: R. H. Schomburgk
468 (K-photo); S: B. Maguire 1398 (RB)].
Figure 12. Flowers of Aspidosperma species from the Brazilian Amazon. A. A. multiflorum A.DC.; B. A. williamii Duarte; C. A. tambopatense
A.H.Gentry; D. A. subincanum Mart.; E. A. ulei Markgr.; F. A. cuspa (Kunth) S.F.Blake; G. A. schultesii Woodson; H. A. leucocymosum
Kuhlm.; I. A. obscurinervium Azambuja; J. A. desmanthum Benth. ex Müll.Arg.; K. A. araracanga Marc.-Ferr.; L. A. verruculosum Müll.
Arg.; M. A. pachypterum Müll.Arg.; N. A. steinbachii Markgr.; O. A. album (Vahl) Benoist ex Pichon; P. A. spruceanum Benth. ex Müll.
Arg.; Q. A. eteanum Markgr.; R. A. neblinae Monach.; S. A. sandwithianum Markgr.; T. A. centrale Markgr.; U. A. auriculatum Markgr.;
V. A. salgadense Markgr.; W. A. oblongum A.DC.; X. A. brasiliense A.S.S.Pereira & A.C.D.Castello; Y. A. cylindrocarpon Müll.Arg.; Z. A.
darienense Woodson & Dwyer [Photos. A: A. M. Miranda et al. 5310 (RB); B: A. A. Oliveira et al. 111 (ESA); C: J. Bosco 112 (UEC); D:
A. T. G. Dias 398 (MG); E: J. M. Pires et al. 16800 (UEC); F: B. A. S. Pereira & A. Alvarenga 3638 (UEC); G: N. T. Silva 5375 (MG); H: A.
Ducke s.n. (G barcode G00190818-photo); I: P. A. C. L. Assunção & E. da C. Pereira 190 (UEC); J: G. T. Prance et al. 5325 (MG); K: J. M.
Pires 11911 (RB); L: K. Kubitzki et al. P21710 (INPA); M: S. A. Mori & C. Gracie 21790 (INPA); N: M. G. Silva & J. Maria 3325 (IAN);
O: N. T. Silva 5157 (MG); P: L. F. Coêlho 538 (MG); Q: A. Ducke s.n. (RB No. 22445); R: B. Maguire et al. 42125 (IAN); S: J. M. Pires &
N. T. Silva 11907 (IAN); T: E. Soares 221 (INPA); U: J. M. Pires et al. 5085 (IAN); V: R. L. Fróes 30286 (IAN); X: A. S. de S. Pereira et
al. 96 (MG); W: A. L. Scudeler et al. 199 (UEC); Y: A. A. Santos et al. 1479 (UEC); Z: N. T. Silva 1325 (IAN)].
Figure 13. Flowers of Aspidosperma species from the Brazilian Amazon. A. A. inundatum Ducke; B. A. myristicifolium (Markgr.)
Woodson; C. A. duckei Huber ex Ducke; D. A. macrocarpon Mart. & Zucc.; E. A. carapanauba Pichon; F. A. rigidum Rusby; G. A. nitidum
Benth. ex Müll.Arg.; H. A. excelsum Benth.; I. A. marcgravianum Woodson [Photos. A: G. A. Black 48-2946 (IAN); B: C. Figueiredo et al.
426 (UEC); C: A. Ducke s.n. (RB No. 11402); D: R. C. Mendonça et al. 1616 (UEC); E: G. F. Árbocz 4033 (ESA); F: A. Ducke 2132 (INPA);
G: P. Acevedo-Rodríguez et al. 8366 (UEC); H: W. W. Thomas et al. 5098 (INPA); I: J. M. Pires 7141 (IAN)].