Flora of Espírito Santo: Humiriaceae

Abstract

Belonging to Malpighiales, Humiriaceae has about 65 species in eight genera, distributed almost exclusively in the Neotropics, occurring from southern Brazil to Central America. In Brazil, 35 species have been recorded (ca. 54% of the family’s diversity), wherein the greatest diversity is found in the Amazon (27 spp.), Atlantic Forest (9 spp.) and Cerrado (6 spp.) domains. This work is part of the “Flora do Espírito Santo” project, which aims to contribute to the knowledge of the state’s floristic diversity. Field expeditions were carried out from January/2019 to March/2020. Humiriaceae, in the state of Espírito Santo, is represented by six species distributed in four genera: Humiria, Humiriastrum, Sacoglottis and Vantanea. Among the registered genera, Vantanea is the most diversified, with three species. In the dense rain forest, all Humiriaceae species recorded here were found, followed by pioneer formations of restingas, with four species and seasonal semideciduous forest, with one species. The main morphological characters that assist in the delimitation of Humiriaceae species in Espírito Santo, are androecium characteristics, such as length (homodynamous or heterodynamous), number of stamens and the number of their series, as well as anther morphology.

Full text

Rodriguésia 73: e00932021. 2022
http://rodriguesia.jbrj.gov.br
DOI: http://dx.doi.org/10.1590/2175-7860202273019
Abstract
Belonging to Malpighiales, Humiriaceae has about 65 species in eight genera, distributed almost exclusively
in the Neotropics, occurring from southern Brazil to Central America. In Brazil, 35 species have been
recorded (ca. 54% of the family’s diversity), wherein the greatest diversity is found in the Amazon (27 spp.),
Atlantic Forest (9 spp.) and Cerrado (6 spp.) domains. This work is part of the “Flora do Espírito Santo”
project, which aims to contribute to the knowledge of the state’s floristic diversity. Field expeditions were
carried out from January/2019 to March/2020. Humiriaceae, in the state of Espírito Santo, is represented
by six species distributed in four genera: Humiria, Humiriastrum, Sacoglottis and Vantanea. Among
the registered genera, Vantanea is the most diversified, with three species. In the dense rain forest, all
Humiriaceae species recorded here were found, followed by pioneer formations of restingas, with four
species and seasonal semideciduous forest, with one species. The main morphological characters that
assist in the delimitation of Humiriaceae species in Espírito Santo, are androecium characteristics, such
as length (homodynamous or heterodynamous), number of stamens and the number of their series, as
well as anther morphology.
Key words: Atlantic Forest, Brazilian Southeast, Malpighiales, taxonomy.
Resumo
Pertencente a Malpighiales, Humiriaceae possui cerca de 65 espécies em oito gêneros, distribuídas quase
exclusivamente na região Neotropical, ocorrendo do sul do Brasil à América Central. No Brasil, foram
registradas 35 espécies (ca. 54% da diversidade da família), sendo a maior diversidade encontrada nos
domínios da Amazônia (27 spp.), Mata Atlântica (9 spp.) e Cerrado (6 spp.). Este trabalho faz parte do
projeto “Flora do Espírito Santo”, que visa contribuir para o conhecimento da diversidade florística do
estado. As expedições de campo foram realizadas de janeiro/2019 a março/2020. Humiriaceae, no estado do
Espírito Santo, está representada por seis espécies distribuídas em quatro gêneros: Humiria, Humiriastrum,
Sacoglottis e Vantanea. Dentre os gêneros registrados, Vantanea é o mais diversificado, com três espécies.
Na Floresta Ombrófila Densa foram encontradas todas as espécies de Humiriaceae aqui registradas, seguidas
de formações pioneiras de restingas, com quatro espécies, e floresta estacional semidecidual, com uma
espécie. Os principais caracteres morfológicos que auxiliam na delimitação das espécies de Humiriaceae
no Espírito Santo, são características do androceu, como comprimento (homodínamos e heterodínamos),
número de estames e o número de suas séries, além da morfologia das anteras.
Palavras-chave: Mata Atlântica, Sudeste brasileiro, Malpighiales, taxonomia.
Flora of Espírito Santo, Brazil
Flora of Espírito Santo: Humiriaceae
Álvaro Nepomuceno1,2,5, Renara Nichio-Amaral 1,3 & Anderson Alves-Araújo1,4
1 Universidade Estadual de Feira de Santana - UEFS, Prog. Pós-graduação em Botânica - PPGBot, Feira de Santana, BA, Brasil.
2 ORCID: <https://orcid.org/0000-0002-4643-8177>.
3 ORCID: <https://orcid.org/0000-0002-6407-8478>.
4 Universidade Federal da Bahia, Campus de Ondina – Instituto de Biologia, Herbário ALCB, Salvador, Bahia, Brasil ORCID: <https://orcid.org/0000-0001-
5810-5145>.
5 Author for correspondence: alvaronepomuceno567@gmail.com

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Introduction
Humiriaceae belongs to Malpighiales (APG
IV 2016) and has 65 species in eight genera, of
which Endopleura and Hylocarpa are represented
by only one species (Wurdack & Zartman 2019).
The family is almost exclusively Neotropical,
occurring mainly in the tropical forests from
southern Brazil to Central America (Cuatrecasas
1961), except for Sacoglottis gabonenses (Baill.)
Urban (449: 1877) which occurs along the African
west coast (Cuatrecasas 1961).
Humiriaceae is composed mostly of shrubs
or treelets, with alternate leaves having serrate
(e.g., Humiriastrum and Sacoglottis) or entire
(e.g., Humiria and Vantanea) leaf margins. The
most important reproductive structure for genera
and/or species delimitation is the morphological
variation of the androecium (Wurdack & Zartman
2019).
In Brazil, 35 species are recorded (ca. 54%
of the family’s diversity) and the greatest diversity
is found in the Amazon domain (27 spp.), followed
by the Atlantic Forest (nine spp.) and Cerrado (six
spp.) domains (BFG 2018).
Available taxonomic information about
Humiriaceae in southeastern Brazil can be
found in Pereira (1966), with three genera and
three species for the state of Guanabara (Rio
de Janeiro); Giordano (1996), with four genera
and nine species for the state of Rio de Janeiro;
Silvestre (1981) for the Fontes do Ipiranga State
Park, São Paulo; Mendonça (2004), with three
genera and four species for the Flora de Grão-
Mogol - Minas Gerais; and Fantinati et al. (2004),
with three genera and three species for Flora da
Serra do Cipó - Minas Gerais.
For Espírito Santo state, Dutra et al. (2015)
listed five species distributed in four genera.
However, until now, there is no direct taxonomic
tool for recognizing these species. As part of the
“Flora do Espírito Santo” project, this work aims
to contribute to the knowledge of the state floristic
diversity. To do so, taxonomic descriptions,
an identification key, photographs, and the
geographical distribution of species throughout
the state are provided.
Material and Methods
Study area
Espírito Santo state has an area of
approximately 46,078 km2, equivalent to 0.53% of
the area in Brazil. Located in the eastern portion of
the Southeast Region, the state is bordered by the
state of Bahia to the north, the Atlantic Ocean to
the east, the state of Rio de Janeiro to the south and
the state of Minas Gerais to the west. The state is
located in the tropical region, presenting a hot and
rainy climate, with average annual temperatures
of 20°–25 °C and a volume of precipitation above
1,400 mm per year, especially concentrated in the
summer (Governo do estado do Espírito Santo
2010; Garbin et al. 2017).
Although the phytogeographic knowledge
of Espírito Santo is scarce and there are
inconsistencies in some classifications of the
vegetation, Garbin et al. (2017) divide the state
into five main types of vegetation: pioneer
formations of Restingas, Semideciduous Seasonal
Forest, Dense Ombrophilous Forest, Open
Ombrophilous Forest and Ecological Refuges.
This is the most current classification of the
vegetation of Espírito Santo.
Field and laboratory studies
Field expeditions were carried out from
January/2019 to March/2020 and samples were
dried according to the usual fieldwork and
herborization procedures (Bridson & Forman
1998). The vouchers were incorporated, primarily,
into the VIES collection (Thiers, continuously
updated). Specimens from the herbaria VIES,
SAMES, MBML, and CVRD were consulted
and analyzed, in addition, online samples of the
herbaria cited in the examined materials of the
species were analyzed.
The identification of taxa was performed
using the specialized literature (e.g., Cuatrecasas
1961; Cuatrecasas & Huber 1999; Kubitzki
2014) and protologs. Morphological terminology
followed Harris & Harris (2001) and authors’
nomenclature, IPNI (2018). For the materials
examined, only one specimen per municipality
is cited.
Results and Discussion
Humiriaceae, in the state of Espírito Santo,
is represented by six species distributed in four
genera: Humiria, Humiriastrum, Sacoglottis and
Vantanea (Fig. 1). Among the genera registered,
Vantanea is the most diversified, with three
species [V. bahiaensis Cuactrecasas (263: 1990),
V. compacta (Schnizlein) Cuactrecasas (65: 1961)
and V. spiritu-sancti (Cuactrecasas) K. Wurdack
& C.E. Zartman (102: 2019)], the other genera

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Figure 1 – a-o. Morphological characters of Humiriaceae species occurring in Espírito Santo – a-c. Humiria
balsamifera – a. open flower showing the union of stamens; b. open flower showing the indument at the apex of
ovary; c. ovary after herborization; d-e. Humiriastrum mussunugense – d. flower without petals; e. androecium with
heterodynamous stamens; f-g. Sacoglottis matogrossensis – f. leaf blade; g. drupe; h-k. Vantanea bahiaensis – h.
leaf blade; i. detail of the glands at the base of leaf blade; j. open flower showing the gynoecium; k. drupe; l-n. V.
compacta – l. leaf blade; m. flower; n. gynoecium; o. V. spiritu-sancti – leaf blade and drupe. (a-c. R. Nichio-Amaral
29; d-e. M.M. Monteiro & A.O. Giaretta 72; f-g. J. Freitas et al. 238; h-k. A. Nepomuceno et al. 792; l-n. L.D.Thomaz
1622; o. L.F.T. Menezes 2279).
5 mm
5 mm
5 mm
1.5 mm
5 mm
3 mm
3 mm
3 mm
3 mm
6 cm
1 cm
3 cm
2 cm
5 cm
4 cm
abc
d
e
f
g
h
i
j
k
l
m
n
o

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are represented by one species each: Humiria
balsamifera Aublet (564: 1775), Humiriastrum
mussunungense Cuactrecasas (235: 1993) and
Sacoglottis matogrossensis Malme (9: 1928).
In the SpeciesLink database (2020), ten
species names are listed, however, four were
not corroborated in this taxonomic treatment,
namely: Humiriastrum dentatum (Casaretto)
Cuatrecasas (136: 1961), Humiriastrum glaziovii
(Urban) Cuatrecasas (137: 1961), Humiriastrum
spiritu-sancti Cuatrecasas (137: 1964) and
Vantanea paraensis Ducke (99: 1925). The
delimitation of H. dentatum is mentioned
in the comments under H. mussunugense.
Humiriastrum glaziovii was not included in this
work, since the only sample collected in Espírito
Santo is deposited in the herbarium US, and is
probably H. mussunugense. In addition, together
with H. dentatum and H. mussunungense, it
forms a taxonomic complex, being, according to
the specialized literature, delimited by tenuous
characteristics of the indument of the leaves and
branches of the inflorescence, making a true
diagnosis impossible through the examination
of images. Humiriastrum spiritu-sanctii has
undergone nomenclatural combination as
Vantanea spiritu-sanctii and Vantanea paraensis
was not confirmed for the state, as no samples
were found in the local herbaria, the only existing
one being deposited in the herbarium U and
without access to the image.
The geographic distribution of the species
is shown in Figure 2. The occurrence of species
in the vegetation types proposed by Garbin et
al. (2017) are shown in Table 1, and Figure 3. In
the dense rain forest, all the Humiriaceae species
recorded here were found, followed by pioneer
formations of restingas, with four species and
seasonal semideciduous forest, with one species
(Tab. 1). In open ombrophilous forests and
ecological refuges, no species were registered
(Tab. 1).
The most important characters for specific
delimitation of Humiriaceae of Espírito Santo are
the length (homodynamous or heterodynamous)
and number of stamens, as well as the number of
stamen series and anther morphology.
According to the most recent list of
angiosperm species published for Espírito Santo
(Dutra et al. 2015), Vantanea compacta is a new
record for the state. However, considering the data
available at BFG (2018), three species are new
records (Sacoglottis matogrossensis, Vantanea
bahiaensis and V. compacta).
It is worth mentioning that among the six
species registered here, four are endemic to Brazil,
of which three (Humiriastrum mussunugense,
Vantanea bahiaensis, and V. spiritu-sancti) are
solely registered for the Atlantic Forest. Vantanea
compacta, found in the Atlantic Forest, also occurs
in the Caatinga and Cerrado (BFG 2018). Among
these species, only Vantanea spiritu-sancti has
a conservation status - Critically Endangered
(CNCFlora 2020), the others do not yet have their
conservation status assessed.
Taxonomic treatment
Humiriaceae A. Juss. Fl. bras. Merid. (4th ed.)
2(13): 87, 1829.
Shrubs or trees. Stipules deciduous or
persistent. Leaves alternate, simple, sessile to
petiolate, chartaceous to coriaceous, obovate,
elliptical to ovate, base attenuate, decurrent,
obtuse to cuneate, apex obtuse, rounded, retuse-
mucronate, attenuate, acuminate to cuspidate,
margin entire, serrate to crenate. Inflorescences
racemes, cymes to panicles, terminal and axillary;
flowers bisexual, actinomorphic, dichlamydeous;
calyx 5-merous, sepals free, corolla 5-merous,
petals free, white, greenish to cream, stamens
10 to many, homodynamous or heterodynamous,
1–3-seriate, ovary superior, placentation pendulous,
style simple, stigma capitate. Drupes oblongoid,
globose, subglobose to ellipsoid. Seeds unseen.
Key to Humiriaceae of Espírito Santo
1. Leaf margin crenulate to serrate.
2. Stipules persistent; stamens homodynamous ...................................3. Sacoglottis matogrossensis
2’. Stipules deciduous; stamens heterodynamous ........................... 2. Humiriastrum mussunungense
1’. Leaf margins entire.
3. Stamens 20, homodynamous, uniseriate ................................................... 1. Humiria balsamifera
3’. Stamens ≥ 30 (except Vantanea spiritu-sancti, which has exactly 20 stamens, biseriate),
heterodynamous, 2–3-seriate.

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4. Leaf blades 4–8 cm wide, obovate to obovate-elliptical; ovary tomentose ....... 4. Vantanea bahiaensis
4’. Leaf blades 2–4 cm wide, elliptical, elliptical-obovate to oblong-elliptical; ovary hirsute or glabrous.
5. Leaf blades with apex acuminate to cuspidate, margin plane; petals glabrous; stamens 60, biseriate;
ovary hirsute .................................................................................................5. Vantanea compacta
5’. Leaf blades with rounded apex, margin revolute; petals hirsute; stamens 20, biseriate; ovary
glabrous ..................................................................................................6. Vantanea spiritu-sancti
1. Humiria balsamifera Aubl., Histoire des Plantes
de la Guiane Françoise 1: 564–566, t. 225. 1775.
Figs. 1a-c; 2a
Shrubs or trees, 1.5–8m tall. Branches
cylindrical, glabrous, lenticulate. Stipules
deciduous. Leaves alternate, sessile to short
petiolate; petiole 1–2 mm long, glabrous to
sparsely hirsute, cylindrical; leaf blade 4–8 × 3–5
cm, coriaceous, obovate, base attenuate, apex
obtuse to retuse-mucronate, glabrous on both
surfaces, margin entire, slightly revolute, midrib
slightly prominent adaxially, prominent abaxially,
secondary veins 7–9 pairs, impressed on both
surfaces. Inflorescences racemose, terminal and
axillary; peduncle 0.5–1.5 cm long, cylindrical,
sparsely hirsute to glabrous. Flowers with pedicels
2–4(–5) mm long, cylindrical, hirsute; sepals
3–6 × 3–4 mm, ovate, hirsute; petals 5–7 ×
1.5–2 mm, deciduous, lanceolate to narrow-ovate,
glabrescent; stamens 20, uniseriate, connate at
base, homodynamous, filaments 3–5 mm long,
glabrous; anthers ovoid, pilose at base; ovary 4–5
× 2.5–3 mm, ovoid, glabrous to pilose towards the
apex; style 3–4 mm long, cylindrical, glabrescent;
stigma capitate, glabrous. Drupes 5–10 × 2–5 mm,
obloid, glabrous, sepals and stigma persistent.
Seeds not seen.
Examined material: Aracruz, Coqueiral, 7.XI.1995,
fl. and fr., J.N. Neves 297 (VIES). Conceição da Barra,
Parque Estadual de Itaúnas - trilha da Borboleta,
1.V.2015, fl., R. Nichio-Amaral 29 (VIES). Guarapari,
Parque Estadual Paulo Cesar Vinha, 20.IX.2010, fl.,
A.S. Zandonadi & S.S. Dutra 8 (VIES). Jaguaré, Giral,
16.I.2009, fr., L. Kollmann & R. Lopes 11498 (MBML).
Linhares, Reserva Natural Vale, 18.VI.2010, fr., V.B.R.
Ferreira 108 (CVRD, SAMES). Presidente Kennedy,
Praia das Neves, 7.VIII.1990, fl. and fr., J.M.L Gomes
et al. 1315 (VIES). Santa Teresa, Santo Antônio,
9.II.1999, fl., L. Kollmann et. al. 1866 (MBML, CEPEC).
São Mateus, Bairro Liberdade, 27.IX.2008, fr., A.O.
Giaretta & M.M. Monteiro 339 (VIES). Sooretama,
Reserva Natural Vale, 13.V.2009, fl., G.D. Colleta et al.
290 (VIES, ESA). Vila Velha, Alagados do Vale, Vale
Encantado, 1.II.2015, fl., R.T. Valadares 1332 (VIES).
Vitória, Reserva Ecológica de Camburi, 23.XII.1997, fr.,
A.M. Assis & I. Weiler-Júnior 318 (VIES).
Species with a Neotropical distribution, from
Colombia to Argentina (Cuatrecasas 1961). In
Brazil, it occurs in all phytogeographic domains,
except in the Pantanal (BFG 2018). In Espírito
Santo, the species is mainly distributed in the
pioneer vegetation of Restinga, despite having
records in areas of higher altitude in vegetation
of Dense and Open Rainforest (Tab. 1). Humiria
balsamifera has ample morphological plasticity,
with 13 varieties of the species being classified,
in addition to the type variety (Cuatrecasas 1961).
However, here in this work we have not adopted
the classification of varieties, due to the tenuous
nature of the diagnostic morphological characters.
Considering the size of the leaves, the species is
similar to Humiria crassifolia Martius ex. Urban
(441: 1877), however, it can be distinguished by
presenting leaves with a short petiole (1–2 mm long
vs. petiole > 10 mm long) and sometimes leaves
pubescent along the primary vein (vs. leaves always
glabrous) (Cuatrecasas 1961).
2. Humiriastrum mussunungense Cuatrec.
Phytologia 75(3): 235-237. 1993. Figs. 1d-e; 2b
Trees 10–20 m tall. Branches cylindrical,
glabrous, lenticulate. Stipules deciduous. Leaves
alternate, petiolate; petiole 3–11 mm long,
glabrous, cylindrical; leaf blade 5–13 × 2.5–5.5 cm,
chartaceous to coriaceous, elliptical to elliptical-
ovate, base obtuse to cuneate, apex attenuate to
acuminate, glabrous on both surfaces, margin
crenulate to serrate, slightly revolute, midrib
impressed adaxially, slightly prominent abaxially,
secondary veins 8–9 pairs, impressed adaxially,
slightly prominent abaxially. Inflorescences
cymose-paniculate, axillary; peduncle 1–3 cm
long, cylindrical, hirsute. Flowers with pedicels
0.5–2 mm long, cylindrical, sparsely hirsute;
sepals 1–1.8 × 1–2 mm, orbicular, connate at
base, margin ciliate, sparsely pilose abaxially;
petals 2.5–4 × 1.3–2 mm, oblong to elliptical,
sparsely hirsute; stamens 20, uniseriate, connate
at base, heterodynamous, filaments 2.5–3.5 mm
long; anthers oblong-lanceolate, glabrous; ovary
1.5–1.8 × 1–1.5 mm, ovate, glabrous; style 0.5–1.2

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Figure 2 – a-f. Geographic distribution of Humiriaceae species occurring in Espírito Santo – a. Humiria balsamifera;
b. Humiriastrum mussunugense; c. Sacoglottis matogrossensis; d. Vantanea bahiaensis; e. V. compacta; f. V. spiritu-
sancti.
a b
c
e f
d

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mm long, cylindrical, glabrous; stigma capitate,
glabrous. Drupes 3.5–4.5 × 3–3.5 cm, subglobose
to ellipsoid, glabrescent, sepals and stigma not
persistent. Seeds unseen.
Examined material: Aracruz, Retiro, 18.X.1993, fl.,
O.J. Pereira et al. 5080 (VIES). Conceição da Barra,
Parque Estadual de Itaúnas, 29.IX.2009, fl. and fr., M.
Ribeiro & A.G. Oliveira 29 (SAMES). Guarapari, Setiba,
2.IX.1996, fr., J.M.L. Gomes 2210 (VIES). Linhares,
Reserva Florestal da CVRD, 27.VII.1991, fl., D.A.
Folli 1393 (CVRD). São Mateus, Bairro Liberdade,
27.IX.2008, fl., M.M. Monteiro & A.O. Giaretta 72
(SAMES).
Species endemic to Brazil, being restricted to
the domain of the Atlantic Forest, occurring, until
now, only in the state of Espírito Santo, where it is
registered from the pioneer vegetation of Restinga
and lowland rain forest (Cuatrecasas 1993; BFG
2018; Tab. 1). Humiriastrum mussunungense has a
tenuous delimitation with H. dentatum, which can
be distinguished by the inflorescence and leaf blade
characteristics. However, these characteristics
vary according to the geographic distribution of
individuals found in Espírito Santo. As this work
does not specifically refer to a greater taxonomic
delimitation between Humiriastrum species, we
decided to keep Humiriastrum mussunungense
occurring in Espírito Santo, while pointing out the
need for a taxonomic revision of the genus aiming
to provide new and strong morphological characters
for the delimitation of species.
3. Sacoglottis matogrossensis Malme, Arkiv för
Botanik 22A(7): 9, t. 3. 1928. Figs. 1f-g; 2c
Shrubs or trees, 3–8 m tall. Branches
cylindrical, glabrous, lenticulate. Stipules
persistent. Leaves alternate, petiolate; petiole 4–10
mm long, glabrous, cylindrical; leaf blade 5–12
× 3–5 cm, coriaceous, elliptical, base cuneate to
attenuate in young leaves, apex acuminate, glabrous
on both surfaces, margin serrate, slightly revolute,
midrib impressed adaxially, slightly prominent
abaxially, secondary veins 8–10 pairs, impressed
on both surfaces. Inflorescences racemose, terminal
and axillary; peduncle 1–2 cm long, cylindrical,
pubescent. Flowers with pedicels 1–3 mm long,
cylindrical, hirsute; sepals 2–3 × 2–2.5 mm,
orbicular to obovate, ciliate; petals 4–7 × 2–2.5
mm, deciduous, lanceolate, glabrous; stamens
10, uniseriate, connate at base, homodynamous,
filaments 4–6 mm long, glabrous; anthers ellipsoid,
glabrous; ovary 2–3 × 1–2 mm, ovoid, glabrous;
style 2–4 mm long, cylindrical, glabrous; stigma
capitate, glabrous. Drupes 3–4 × 3–4 cm, globose,
glabrous, sepals, filaments and stigma sometimes
persistent. Seeds unseen.
Material examined: Anchieta, 10.X.2009, fr., J.M.L.
Gomes et. al. 3463 (VIES). Aracruz, Barra do Riacho,
15.V.1990, fl. and fr., P.C. Vinha 1013 (CVRD, VIES).
Conceição da Barra, Parque Estadual de Itaúnas,
19.V.2014, fl. and fr., J. Freitas et al. 238 (VIES).
Linhares, Reserva Biológica de Comboios, 27.VII.1992,
fl., O.J. Pereira & J.M.L. Gomes2604 (VIES). Santa
Teresa, Reserva Biológicas Augusto Ruschi, 13.V.2003,
fl. and fr., R.R. Vervloet et al. 2399 (MBML).
Species with a Neotropical distribution, from
Colombia, Venezuela and Brazil, where it has wide
distribution, occurring in the phytogeographic
domains of the Amazon, Cerrado and Atlantic
Forest (Cuatrecasas 1961, BFG 2018). In Espírito
Santo, the species is distributed in pioneer
vegetation of Restinga and lowland rain forest
(Tab. 1). The species also has broad morphological
plasticity, being classified into varieties based
Species Vegetational types
PFR SSF OOF DRF ER
Humiria balsamifera ● ● -●-
Humiriastrum mussunungense ●- - ●-
Sacoglottis mattogrossensis ●- - ●-
Vantanea bahiaensis ●- - ●-
Vantanea compacta - - - ●-
Vantanea spiritu-sancti - - - ●-
Table 1 – Geographic distribution of Humiriaceae species occurring in Espírito Santo in the vegetation types proposed
by Garbin et al. (2017). PFR = pionners formations of restingas; SSF = seasonal semi-deciduous forest; OOF = open
ombrophilous forest; DRF = dense rain forest; ER = ecological refuge. ● = presence; - absent.

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on shape, however, Sacoglottis matogrossensis
is morphologically similar to S. cydonioides
Cuatrecasas (183: 1961) because they have
persistent bracts and globose drupes, however, they
can be distinguished by having prominent ribs (vs.
superficial ribs) (Cuatrecasas 1961).
4. Vantanea bahiaensis Cuatrec. Phytologia 68(4):
263-265. 1990. Figs. 1h-k; 2d
Shrubs or trees, 4–6 m tall. Branches
cylindrical, glabrous, lenticulate. Stipules
deciduous. Leaves alternate, petiolate; petiole
4–8(–10) mm long, glabrous, cylindrical; leaf
blade (4–)6–10 × 4–8 cm, coriaceous, obovate
to obovate-elliptical, base cuneate, apex rounded
to rarely cuspidate, glabrous on both surfaces,
margin entire, revolute, midrib slightly prominent
on both surfaces, secondary veins 8–12 pairs,
slightly prominent on both surfaces. Inflorescences
racemose, terminal, peduncle 4–5 cm long,
cylindrical, pubescent. Flowers with pedicels
1–2 mm long, cylindrical, pubescent; sepals
1.5–2 × 1–1.5 mm, orbicular to obovate, ciliate;
petals 3–4 × 1–2 mm, deciduous, lanceolate,
pubescent to glabrescent; stamens 30–40, biseriate,
heterodynamous; filaments 2–5 mm long, glabrous;
anthers ellipsoid, glabrous; ovary 2–3 × 1.5–2 mm,
ovoid, tomentose, style 1–1.5 mm long, cylindrical,
glabrous; stigma capitate, glabrous. Drupes 1–1.5
×1 cm, globose to ellipsoid, velutinous, sepals,
filaments and stigma sometimes persistent. Seeds
unseen.
Material examined: Conceição da Barra, Parque
Estadual de Itaúnas, trilha da Borboleta, 2.IV.2019, fl.
and fr., A. Nepomuceno et al. 792 (VIES). Guarapari,
Setiba, 2.IX.1996, fl. and fr., J.M.L. Gomes 2209 (VIES).
Linhares, Reserva Natural Vale, 13.X.2012, fl. V.B.R.
Ferreira 124 (CVRD, SAMES). São Mateus, 10.I.2008,
fl. and fr., A.O. Giaretta et al. 168 (VIES, SAMES). Vila
Velha, 21.XI.1983, fl., P.C. Vinha (VIES 262).
Species endemic to Brazil, being restricted to
the Atlantic Forest domain, occurring, until now,
in the states of Bahia and Espirito Santo, where it
is registered in pioneer vegetation of Restinga and
lowland Ombrophilous Forest (Cuatrecasas 1990;
BFG 2018; Tab. 1). Among the Brazilian species
of the genus, Vantanea bahiaensis has greater
morphological similarity to V. spiritu-sancti, and
can be recognized based on the shape of the leaf
blade and type of ovary induction.
5. Vantanea compacta Contributions from the
United States National Herbarium 35(2): 65.
1961. Figs. 1l-n; 2e
Shrubs or trees, 5–9 m tall. Branches
cylindrical, glabrous to hirsute, lenticulate. Stipules
deciduous. Leaves alternate, petiolate; petiole
2–3 cm long, glabrous to slightly puberulous,
cylindrical; leaf blade 5–7 × 2–4 cm, coriaceous,
elliptical to oblong-elliptical, base attenuate to
decurrent, apex acuminate to cuspidate, glabrous
adaxially, glabrescent abaxially, margin entire,
plane, midrib impressed adaxially, prominent
abaxially, secondary veins 6–8 pairs, impressed on
both surfaces. Inflorescences paniculate, terminal
and axillary; peduncle 4–5 cm long, cylindrical,
puberulous. Flowers with pedicels 1–1.5 cm long,
cylindrical, glabrous; sepals 2–2.5 × 1–1.5 mm,
obovate to wide-ovate, ciliate; petals 4–6 × 1.5–2
mm, deciduous, elliptical-lanceolate, glabrous;
stamens 60, 2-seriate, heterodynamous; filaments
3–5 mm long, glabrous; anthers ovoid, glabrous;
ovary 1–2 × 1–1.5 mm, ovoid, hirsute; style 3–4
mm long, cylindrical, glabrous; stigma capitate,
glabrous. Fruits not seen. Seeds not seen.
Material examined: Linhares, Reserva Natural Vale,
3.XI.2014, fl. and fr., D.A. Folli 7295 (CVRD, HUEFS).
Santa Teresa, Estação Biológica Santa Lúcia, 28.X.1992,
fl., L.D. Thomaz 1622 (MBML).
Species endemic to Brazil, where it occurs
in the phytogeographic domains of the Caatinga,
Cerrado and Atlantic Forest (Cuatrecasas 1961;
BFG 2018). In Espírito Santo it is registered in
vegetation of Dense Rainforest, from low to high
altitudes (Cuatrecasas 1961; BFG 2018; Tab. 1). In
addition to being part of the group of species of the
genus that has glabrous petals, Vantanea compacta
is the species of the genus that has the smallest leaf
blade measurements, these being fully glabrous
with an apiculate, acuminate apex.
6. Vantanea spiritu-sancti K. Wurdack & C.E.
Zartman, PhytoKeys 124: 2019. Figs. 1o; 2f
Trees 10–15 m tall. Branches cylindrical,
glabrous, lenticulate. Stipules deciduous. Leaves
alternate, petiolate; petiole 1–2.5 cm long, glabrous,
cylindrical; leaf blade 6–9 × 2–3.5 cm, coriaceous,
elliptical-obovate, base cuneate, apex rounded,
glabrous on both surfaces, margin entire, revolute,
midrib impressed on the lower surface, prominent
on the upper surface, secondary veins 10–12
pairs, impressed on both surfaces. Inflorescences
corymbose, terminal and axillary; peduncle 3–4
cm long, cylindrical, glabrescent. Flowers with
pedicels 1–2 cm long, cylindrical, hirsute; sepals
1–1.5 × 1 mm, ovate, ciliate; petals 3–4 × 1–1.5
mm, deciduous, lanceolate, hirsute; stamens 20,

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Figure 3 – a-f. Distribution of Humiriaceae species occurring in Espírito Santo in the vegetation types proposed by
Garbin et al. (2017) – a. Humiria balsamifera; b. Humiriastrum mussunugense; c. Sacoglottis matogrossensis; d.
Vantanea bahiaensis; e. V. compacta; f. V. spiritu-sancti.
a b
c
e f
d

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biseriate, heterodynamous; filaments 4–5 mm long,
glabrous; anthers ovoid, glabrous; ovary 2–3 × 1–2
mm, ovoid, glabrous; style 5–8(–10) mm long,
cylindrical, glabrous; stigma 5-lobate, glabrous.
Drupes 3–4 × 2–3 cm, ellipsoid, glabrous, filaments
sometimes persistent. Seeds unseen.
Material examined: Fundão, APA Goiapaba-Açu,
15.VII.1998, fl. and fr., L. Kohlmann et al. 223 (MBML).
Linhares, Reserva Natural Vale, 20.V.2016, fl. and fr.,
L.F.T. Menezes 2279 (SAMES). Santa Teresa, Reserva
Biológica Augusto Ruschi, 9.IV.2003, fl. and fr., R.R.
Vervloet et al. 2185 (MBML).
Species endemic to Brazil, being restricted to
the Atlantic Forest domain, occurring, until now, in
the states of Bahia and Espirito Santo, where it is
registered in vegetation of Dense Rainforest from
low to high altitudes (Cuatrecasas 1964; BFG 2018;
Wurdack & Zartman 2019; Tab. 1), with occurrence
records in three conservation units: Goiapaba-
Açu Environmental Proctetion Area, Vale Nature
Reserve and Augusto Ruschi Biological Reserva.
Vantanea spiritu-sancti was recently transferred
from the genus Humiriastrum to Vantanea, for
presenting anthers with two bisporangiate thecae
and palynological characteristics (Wurdack
& Zartman 2019). The species has a greater
morphological similarity with V. bahiaensis, whose
morphological delimitation was presented in the
comments of that species.
Acknowledgments
We thank the curators and technical assistants
of the herbaria CVRD, MBML, SAMES and
VIES. The last author also thanks the Fundação
de Amparo à Pesquisa e Inovação do Espírito
Santo, for financial support (FAPES Nº 18/2018,
TO 525/2018).
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List of material examined
Alves M 2305 (1). Amorim BS 1954 (3), 1977 (1), 2027 (4). Andrade IR 225 (1). Assis AM 318 (1), 2219 (3), 2844 (1) 3274 (4), 3476
(3). Belém RP 3836. Cândito ES 718 (1). Coelho R 44 (1). Colleta GD 289 (1), 290 (1). Cruz TA 71 (6). Silva AF 1986 (1). Lima
HC 2918 (4), 2976 (4). Faria JEQ 4242 (1). Farias GL 65 (4), 83 (1). Farney C 4605 (1). Felitto G 1208 (1) 1250 (1). Ferreira VBR
12 (4), 39 (1), 55 (1), 59 (4), 79 (1), 82 (1), 86 (1), 89 (1), 92 (1), (1), 124 (4), 128 (4), 162 (1). Folli DA 1121 (3), 1291 (1); 1377 (3),
1558 (1), 1582 (4), 1683 (4), 2095 (6), 3422 (1), 4547 (4), 4639 (6), 5268 (3), 5376 (1), 5389 (3), 5530 (1), 7295 (5). Freire GQ 47
(4). Freitas J 238 (3). Giaretta AO 168 (4), 232 (1), 339 (1); 490 (1), 531 (4), 540 (4), 1070 (1). Giordano LCS 2678 (6), 2679 (1),
2680 (1), 2681 (4). Gomes JML 1315 (1), 2209 (4), 3463 (3), 3498 (3), 4111 (1). Gomes SM 4 (1). Groppo M 997 (1). Hatschbach
G 58054 (1). Kollmann L 223 (6), 265 (6), 293 (6), 707 (6), 1282 (6), 1866 (1), 11498 (1). Kuhlmann L 4166 (3). Lima JCA 3 (1).
Lombardi JA 9647 (1). Lopes LCM 35 (4), 45 (1), 148 (1), 149 (4). Luber J 187 (1), 193 (1). Lucas E 897 (1), 929 (4), 984 (1). Luz
AA 342 (4). Maas PJM 9872 (4). Machado JO 290 (4). Martinelli G 11032 (4). Martins RFA 21 (1). Meireles J 568 (1), 570 (4),
612 (1). Menezez LFT 1654 (3), 1712 (1), 2279 (6). Monteiro MM 162 (4), 172 (1). Moraes PLR 4608 (1). Moraes QS 29 (4), 198
(1). Nascimento AL 5 (1). Nepomuceno A 792 (4), 793 (1). Neve JN 297 (1). Nichio-Amaral R 29 (1). Nunes NL 29 (1). Paciencia
MB 2335 (4). Peixoto AL 340 (1). Pereira OJ 139 (1), 232 (4), 270 (4), 271 (1), 949 (1), 1138 (1), 1718 (1), 1889 (4), 2604, (3), 2927
(1), 3000 (4), 3049 (4), 3092 (1), 3134 (3), 3161 (1), 3171 (4), 3295 (1), 3523 (1), 3792 (1), 3970 (1), 4071 (1), 4244 (1), 4360 (3),
4463 (1), 4568 (1), 4780 (1), 4861 (1), 5349 (1), 5940 (4), 6095 (4), 6104 (1), 6141 (3), 6318 (4), 7614 (1) 7752 (4). Pirani JR 2617
(1). Queiroz LP 2456 (1). Reis R 9 (6). Romão GO 1251 (4). Salim-Filho J 174 (1), 175 (1). Silva SS 16 (1), 17 (1), 21 (1), 24 (4),
37 (1). Siqueira GS 847 (6). Sobral M 3994 (4). Souza S 7 (1), 26 (1), 40 (1). Souza WO 284 (1). Spada J 8 (6), 233 (5). Stehmann
JR 4643 (4), 4666 (1). Thomaz LD 1339 (6), 1622 (5). Valadares RT 1332 (1). Valdemarin KS 889 (4). Vervloet RR 2156 (6), 2185
(6), 2399 (3). Vinha PC 1013 (3), 1245 (1). Weinberg B 311 (1), 580 (1). Zandonadi AS 7 (1), 8 (1).