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Rodriguésia 73: e01862019. 2022
http://rodriguesia.jbrj.gov.br
DOI: http://doi.org/10.1590/2175-7860202273001
Abstract
The present work, which is part of a series of taxonomic monographs, aims to provide useful taxonomic features
for distinguishing species of Cactaceae from Itaúnas State Park-Brazil. After performing field expeditions
and observations, and analysis of herbaria vouchers, a total of five species of Cactaceae is herein recorded:
Brasiliopuntia brasiliensis (Least Concern), Cereus fernambucensis subsp. fernambucensis (Least Concern),
Melocactus violaceus subsp. violaceus (Vulnerable), Pereskia aculeata (Least Concern), and Pilosocereus
arrabidae (Near Threatened). Leaf (when present) and cladodes morphology, and presence or absence of
cephalium are the most useful characters to specific delimitation. Key identification, descriptions, photographs,
taxonomic comments, geographic distribution and habitat data are herein provided.
Key words: Atlantic Forest, cacti, Caryophylalles, Restinga, taxonomy.
Resumo
O presente trabalho, que faz parte de uma série de monografias taxonômicas, visa fornecer características
taxonômicas úteis para distinguir espécies de Cactaceae do Parque Estadual de Itaúnas-Brasil. Após a
realização de expedições e observações de campo e análise de vouchers herbários, um total de cinco espécies
de Cactaceae foram registrados: Brasiliopuntia brasiliensis (Pouco Preocupante), Cereus fernambucensis
subsp. fernambucensis (Pouco Preocupante), Melocactus violaceus subsp. violaceus (Vulnerável), Pereskia
aculeata (Pouco Preocupante) e Pilosocereus arrabidae (Quase Ameaçado). Morfologia da folha (quando
presente) e dos cladódios, e presença ou ausência de cefálio são os caracteres mais úteis para as delimitações
específicas. Chave de identificação, descrições, fotografias, comentários taxonômicos, distribuição geográfica
e dados de habitat são fornecidos aqui.
Palavras-chave: Mata Atlântica, cactos, Caryophylalles, Restinga, taxonomia.
Flora of Espírito Santo, Brazil
Flora of Itaúnas State Park, Espírito Santo, Brazil: Cactaceae
Álvaro Nepomuceno1,2,5, Victor Santos Miranda1,3 & Anderson Alves-Araújo1,4
1 Universidade Federal do Espírito Santo - UFES/CEUNES, Lab. Sistemática e Genética Vegetal, São Mateus, ES, Brasil.
2 ORCID: <https://orcid.org/0000-0002-4643-8177>.
3 ORCID: <https://orcid.org/0000-0003-2829-5587>.
4 ORCID: <https://orcid.org/0000-0001-5810-5145>.
5 Author for correspondence: alvaronepomuceno567@gmail.com
Introduction
Caryophyllales has 13 families sharing,
among other synapomorphies, the presence of
vessels with simple perforations and similar
aminoacid sequences of the citrochrome c (Judd
et al. 2009; APG IV 2016). In the suborder
Caryophyllineae, which is considered monophyletic
based on morphological and molecular studies
(Judd et al. 2009), Cactaceae and Portulacaceae are
phylogenetically related by sharing loss of internal
phloem and CAM photosynthetic metabolism
(Judd et al. 2009; Ocampo & Columbs 2010;
Hernández-Hernández et al. 2011). Cactaceae,
one of the most representative and iconic families
of the order, has approximately 1,400 species and
100 genera distributed exclusively in the American
continent, with Mexico, USA, and Northeastern
and Southeastern Brazil bearing its main centers
of diversity (Barthlott 1983; Valiente-Banuet et al.
1996; Dávila-Aranda et al. 2002; Casas et al. 2014).
Nepomuceno A, Miranda VS & Alves-Araújo A
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Cactaceae species are characterized mainly
by the reduction of branches, leaves (usually)
modified in spines, succulent and photosynthetic
stems, superficial and often thorny roots (Duque
1980; Wallace & Gibson 2002; Edwards &
Donoghue 2006; Judd et al. 2009). Due to their
features, many species might be used as human
food resource, forage, timber, medicinal and
ornamental (Ruedas et al. 2006; Santos et al.
2007; Lucena et al. 2013, 2014; Nunes et al.
2015).
In Brazil, Cactaceae is represented by
approximately 270 species (193 endemic) and
39 genera (14 endemic), which Pilosocereus
Byles & Rowley, Rhipsalis Gaertn. (29 spp.,
each) and Parodia Speg. (27 spp.) are the most
taxonomically diverse genera (BFG 2018). The
highest species richness can be found in arid and
semi-arid environments, such as those occurring
in Northeastern Brazil (Taylor & Zappi 2004).
However, many species can be also found in
moist environments such as the Atlantic Forest
where 102 species belonging to 21 genera are
recorded (BFG 2018). Amongst Brazilian states,
Rio de Janeiro is the one which bears the highest
number of Cactaceae taxa recorded (57 spp.)
(BFG 2018).
Identification of Cactaceae samples in
herbaria is difficult due to the different methods
of collection and drying specimens. Main
reproductive characters of Cactaceae are lost
during sampling and drying processes, especially
those from fruits. In addition, the number of
thorns in the areolas can also be modified
depending on processes mentioned above.
Under these considerations, the present
work aims to provide a taxonomic treatment for
Cactaceae occurring in the State Park of Itaúnas,
contributing to the knowledge of the regional and
state floras, and delimitation among the species
of the group.
Materials and Methods
Itaúnas State Park (ISP) (18°25’S, 39°42’W)
(Fig. 1) is located in the municipality of Conceição
da Barra, northern coast of Espírito Santo, and has
an area of 3,481 ha (Souza et al. 2016). The ISP
is fully inserted in the geological zone of the
trays and has several natural ecosystems, such
as restingas (predominant vegetation), tabuleiro
forests, dunes, wetlands and mangroves (Souza
et al. 2016).
Specimens at VIES and SAMES herbaria
(acronyms according to Thiers, continuously
updated) were analyzed, and fieldwork
expeditions were performed from August/2018
to May/2019. Samples were processed according
to Peixoto & Maia (2013), incorporated mainly
to VIES herbarium and duplicates sent to
MBML and SAMES herbaria. Identification
and geographical distribution comments were
performed using specialized literature (Zappi
1994; Taylor & Zappi 2004; Hunt et al. 2006;
Zappi et al. 2007; Menezes et al. 2013; Gonzaga
et al. 2014a,b, 2017).
Terminology for morphological characters
followed Harris & Harris (2001) and Taylor
& Zappi (2004). Scientific names and their
authorship are in agreement with IPNI (2019).
Habit and habitat data were based on voucher
tags information and field observations.
This work is part of the series of taxonomic
and floristic works carried out for Itaúnas State
Park, among them Souza et al. (2016, 2017) and
others which will be published soon.
Results and Discussion
For the Espírito Santo, 46 species of
Cactaceae are recorded, of which 22 occur in
the restingas of the state, of these, five (22%)
species are recorded in the ISP (BFG 2018). In
addition, of the 13 species of Cactaceae endemic
to Brazil that occur in the capixaba restingas,
four (23%) are registered for study area (BFG
2018).
Initially, Cactaceae at the ISP was
represented by six species (Souza et al.
2016), however, after the taxonomic treatment
presented here, five species are recognized for
the family: Brasiliopuntia brasiliensis (Willd.)
A. Berger, Cereus fernambucensis Lem. subsp.
fernambucensis, Melocactus violaceus Pfeiff.
subsp. violaceus, Pereskia aculeata Mill.
and Pilosocereus arrabidae (Lem.) Byles &
Rowley, being this the first taxonomic treatment
considering Cactaceae from Espírito Santo state.
Main useful morphological characters for species
delimitation were the leaf cladodes morphology
and presence or absence of cephalium.
According to the Goettsch et al.
(2015), Brasiliopuntia brasiliensis, Cereus
fernambucensis subsp. fernambucensis and
Pereskia aculeata are assigned as Least Concern
(LC), Pilosocereus arrabidae as Near Threatened
(NT), and Melocactus violaceus subsp. violaceus
as Vulnerable (VU).
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Figure 1 – a-f. Environments found in the Itaúnas State Park – a. Alméscar of Trail; b. Borboleta of Trail; c-d. Buraco
do Bicho of Trail; e. Pescador of Trail; f. Restinga of Trail.
ab
c
e f
d
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Taxonomic treatment
Identication key for the Cactaceae species of ItaúnasState Park
1. Leaveswelldevelopedandfunctionalpresent;pedicellateowersandgatheredinorescences .........
................................................................................................................................ 4. Pereskia aculeata
1’. Leavesreducedortransformedintospines;sessileowersandlonely.
2. Arborescent; glochids present, cladodes dimorphics, reduced leaves (in young shoots) ...............
.......................................................................................................... 1. Brasiliopuntia brasiliensis
2’. Shrubs; glochids absent, cladodes monomorphics, leaves transformed into spines.
3. Subglobosecladodes;presentcephalium,owers1.5–2.5×0.6–1.3cm,berry10–16×8–10
mm......................................................................... 3. Melocactus violaceus subsp. violaceus
3’. Cylindricalcladodes;absentcephalium,owers6–20×4–5cm,berry3–7×3.5–5.8cm.
4. Articulated cladodes, areoles well-spaced along the ribs ................................................
...........................................................2. Cereus fernambucensis subsp. fernambucensis
4’. Non-articulated cladodes, areoles congested along the ribs ............................................
................................................................................................5. Pilosocereus arrabidae
1. Brasiliopuntia brasiliensis (Willd.) A. Berger.
Fig. 2a-b
Arborescent4–5mtall.Cladodescylindrical,
branched, articulated, ribs absent, intermediate
segments 22–54 cm long, cylindrical, terminal
segments5–12×3–5cm,compresses,obovate,
deciduous, distal areolas 10–30 mm between
each other, spinescent, spines 1–2, 5–20 mm
long, glabrous. Glochids present. Leaves reduced
(young branches) or absent (mature branches).
Flowers appearing at the apex of the cladodes.
Flowers3–5×4–6cm,sessile,neartoterminal
segments, filaments and anthers glabrous,
ovary ovate, style 1.2–1.5 mm long, stigmatic
branches 3–6, 3–6 mm long. Berry 3–6 cm
diam., spinescent, globose, yellow-greenish
pericarp,smooth, areolated. Seeds 8–10 × 7–9
mm, subgloboids.
Examined material: Trilha da Restinga, 7.VII.2012,
fr., L.F.T. Menezes et al. 2037 (SAMES); 12.II.2014,
fr., J.O. Machado et al. 200 (VIES); 2.IV.2019, fr., A.
Nepomuceno et al. 795 (VIES).
Additional examined material: BRASIL. ESPÍRITO
SANTO: Conceição da Barra, Área 126 da Aracruz
Celulose, 26.II.1992, fr., O.J. Pereira et al. 2879
(VIES); Área 213 da Aracruz Celulose, 25.II.1992, fl.,
O.J. Pereira et al. 3099 (VIES).
Brasiliopuntia (K. Schum) A. Berger is
a monospecific genus belonging to the tribe
Opuntieae of the subfamily Opuntioideae,
occurring from Bolivia to Paraguay and
Argentina (Taylor & Zappi 2004; Hunt et al.
2006). In Brazil, B. brasiliensis occurs in all
phytogeographical domains, in Deciduous
Seasonal Forest, Semideciduous Seasonal Forest,
Ombrophilous Forest, Mixed Ombrophylous
Forest, restinga and rocky outcrops (BFG 2018).
It is characterized by having dimorphic branches,
compressed at the top and cylindrical at the base.
The species is categorized as Least Concern (LC)
according to Goettsch et al. (2015). In the study
area, it is considered as rare where it can be found
only on the edges of restinga forest.
2. Cereus fernambucensis Lem. subsp.
fernambucensis. Fig. 2c-d
Shrubs 1–2 m tall. Cladodes cylindrical,
branched, articulate, 4–5 ribbed, ribs 1–2 cm
high, oblique transverse folds, distal areolas
7–22mmbetweeneachother,spinescent,spines
4–8, 5–30 mm long. Glochids absent. Leaves
transformed in spines. Absent cephalium. Flowers
15–20 cm long, sessile, filaments and anthers
glabrous, ovary ovate, style 16–18 cm long,
stigmatic branches 12, 5–10 mm long. Berry
5–7 × 3.5–5.5 cm, glabrous, ovoid, pericarp
pinkishtoreddish,smooth.Seeds2–3×1.8–2
mm, suborbicular.
Examined material: Trilha da Borboleta, 2.VI.2019,
fl., A. Nepomuceno et al. 788, 789 (VIES); Trilha do
Pescador, 21.I.2013, fl., R. Coelho 255 (SAMES);
7.II.2013, fl., A.O. Giaretta 448 (VIES); 15.VIII.2013,
fl., J.O.Machado et al. 133 (VIES).
Additional examined material: BRASIL. ESPÍRITO
SANTO: Conceição da Barra, Área 126 da Aracruz
Celulose, 3.XII.1992, fl., O.J. Pereira et al. 4343
(VIES); Área 213 da Aracruz Celulose, 4.XI.1992, fl.,
O.J. Pereira et al. 4102 (VIES).
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Rodriguésia 73: e01862019. 2022
Figure 2 – a-j. Morphological characteristics of native Cactaceae from Itaúnas State Park – a-b. Brasiliopuntia
brasiliensis – a. berry; b. floral bud; c-d. Cereus fernambucensis subsp. fernambucensis – c. cladode with flower; d.
cross section of cladode; e-f. Melocactus violaceus subsp. violaceus – e. apical cephalium; f. flowers; g-h. Pereskia
aculeata – g. flower; h. berry; i-j. Pilosocereus arrabidae – i. cladode; j. cross section of cladode. Photographs: a-c.
SGV files; d-e. Rafael Mathielo; f. SGV files; g-h. Maria Ana Farinaccio; i. SGV files; j. Rafael Mathielo.
ab c
ef
g
h i j
d
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Cereus Mill. belongs to the Cereeae tribe of
the subfamily Cactoideae and has ca. of 20 species
distributed throughout South America (Taylor
& Zappi 2004). Cereus fernambucensis subsp.
fernambucensis is endemic to Brazil and can be
found Caatinga and Atlantic forest domains, in
restingas and rocky outcrops (BFG 2018). The
species differs from C. fernambucensis subsp.
sericifer (Ritter) N.P. Taylor & Zappi for presenting
red fruits (vs. yellow) (Taylor & Zappi 2004). In
addition, it is assigned as Least Concern (LC)
according to Goettsch et al. (2015). It can be found
throughout study area, especially in open areas and
forest edges.
3. Melocactus violaceus Pfeiff. subsp. violaceus.
Fig. 2e-f
Shrubs solitary 8–13 cm tall. Cladodes
subglobose,non-branched,9–13ribbed,ribs2–3.5
cm high, oblique transverse folds, distal areolas
5–10mm between eachother,spinescent,spines
10–12,15–25mmlong.Glochidsabsent.Leaves
transformed in spines. Apical cephalium with white
trichomes and spinning brown bristles. Flowers
1.5–2.5×0.6–1.3cm;sessile,ovaryovate.Berry
10–16×8–10mm,glabrous,claviforme,pericarp
pinktooff-whiteatthebase,smooth.Seeds1–1.5
×0.8–1mm,ellipsoids.
Examined material: Trilha da Borboleta, 28.III.2009,
fr., M.M. Monteiro 150 (SAMES), 14.IV.2014, fl., J.O.
Machado et al. 307 (VIES).
Additional examined material: BRASIL. ESPÍRITO
SANTO: Conceição da Barra, Área 213 da Aracruz
Celulose, 25.II.1992, fl., O.J. Pereira 3099 (VIES); São
Mateus, Bairro Litorâneo, 28.VIII.2008, fl., A.O.Giaretta
et al. 322 (SAMES).
Melocactus (L.) Link. & Otto belongs to
the Cereeae tribe of the subfamily Cactoideae
and has 34 species distributed from Mexico to
the east of Brazil, with limit distribution in Rio
de Janeiro (Taylor & Zappi 2004). Melocactus
violaceus is endemic in Brazil, occurring in the
phytogeographic domains of Caatinga and Mata
Atlântica, in carrasco and restinga vegetation
(BFG 2018). It is distinguished from Melocactus
violaceus subsp. margaritaceus N.P. Taylor for
presenting fruits pink to purple (vs. white) and
Melocactus violaceus subsp. ritteri N.P. Taylor
due to occur in the coastal region of Brazil, mainly
in restinga (vs. occurrence restricted to the rocky
outcrops of the interior of the Bahia state). The
species is classified as vulnerable (VU) according
to Goettsch et al. (2015). In the study area, the
species is often found in open areas.
4. Pereskia aculeata Mill. Fig. 2g-h
Climbing shrub. Cladodes cylindrical,
branched,not articulated,ribsabsent,spines 1–10
central,10–30mmlong,6–20radial,7–40mmlong,
cylindrical. Leaves well-developed and functional,
alternate,petiole3–5mmlong,glabrous,lamina2.5–
7× 0.5–3cm,elliptictonarrow-elliptic, glabrous.
Glochids absent. Inflorescences in panicles terminal
and axillary.Flowers 2.5–5 × 2–4 cm, pedicels
5–20mmlong,ovaryovate,style10–11mmlong,
stigmaticbranches4–7,3–5mmlong.Berry1–2cm
diam, glabrous, globose, pericarp yellowish, smooth,
spines 3–8 mm long, glabrous, cylindrical, bracts
8–15 × 1–4 mm, narrow-elliptic, glabrous. Seeds
4.5–5×4–4.2mm,obovatetoelliptical.
Examined material: Trilha da Restinga, 11.IV.2010, fr.,
M. Ribeiro 123 (SAMES, VIES); 22.III.2014, fl., B.S.
Amorim et al. 1966 (VIES); 19.V.2014, fl., N.T.L.Pena
96 (VIES); 2.VI.2019, A. Nepomuceno et al. 794 (VIES).
Additional examined material: BRASIL. ESPÍRITO
SANTO: São Mateus, Bairro da Liberdade, 6.III.2008,
fr., O.J. Pereira (SAMES); 5.IV.2008, fr., M.B. Faria et
al. 157 (SAMES).
Pereskia Mill. belongs to the subfamily
Pereskioideae and has 17 species distributed
throughout the neotropics (Taylor & Zappi 2004).
In Brazil, Pereskia aculeata occurs in the various
phytophysiognomies of the Caatinga, Cerrado,
and Atlantic Forest domains (BFG 2018). It can
be distinguished from other species by presenting
climbing habit. The species is classified as of Least
Concern (LC) according to Goettsch et al. (2015).
In the study area, the species is rare, being found in
the canopy and at the edges of forest.
5. Pilosocereus arrabidae (Lem.) Byles &
Romley. Fig. 2i-j
Shrubs ca. 2m tall. Cladodes cylindrical,
branched, non-articulated, 6 ribbed, ribs 1.5–2.5
mm high, oblique transverse folds, distal areolas
10–15mmapart,spinescent,spines4–10,5–15mm
long, glabrous. Glochids absent. Leaves transformed
in spines.Absent cephalium. Flowers 6–7 × 4–5
cm, sessile, filaments and anthers glabrous, ovary
obovate,stylet45–50mmlong,stigmaticbranches
9,6–10mmlong.Berry3–5×5–5.8cm,glabrous,
subgloboid, green pericarp to light red, smooth to
slightlyrough.Seeds1.5–2×1–1.3mm,ellipsoids.
Examined material: Trilha da Borboleta, 7.II.2009,
fl., A.O. Giaretta 448 (SAMES); 27.IX.2009, fl., A.O.
Giaretta 638 (VIES); 21.I.2013, fr., R. Coelho 254
(SAMES); 2.IV.2019, fl. and fr., A. Nepomuceno et al. 786
(VIES). Trilha da Restinga, 2.IV.2019, fr., A. Nepomuceno
et al. 796 (VIES).
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Rodriguésia 73: e01862019. 2022
Additional examined material: BRASIL. ESPÍRITO
SANTO: Conceição da Barra, Área 126 da Aracruz
Celulose, 3.XII.1992, fl., O.J. Pereira et al. 4342 (VIES);
Área 213 da Aracruz Celulose, 25.III.1992, fl., O.J.
Pereira et al. 3081 (VIES).
Pilosocereus Byles & G.D. Rowley belongs
to the Cereeae tribe of the subfamily Cactoideae
and has 37 species distributed from Mexico to
Paraguay (Taylor & Zappi 2004). Pilosocereus
arrabidae is endemic to the Brazilian Atlantic
Forest domain, in restinga vegetation and
rocky outcrops (BFG 2018). According to
Taylor & Zappi (2004), it is morphologically
related to Pilosocereus brasiliensis (Britton &
Rose) Backeb. subsp. brasiliensis, a sympatric
species in Espírito Santo state, especially due
to morphology of the areolas and the color of
the epidermis. However, it can be distinguished
by presenting acute floral buds (vs. obtuse in
Pilosocereus brasiliensis subsp. brasiliensis).
The species is classified as almost threatened
(NT) (Goettsch et al. 2015). It can be found
throughout study area, especially in open areas
and forest edges.
Acknowledgments
We are indebted to the organization which
funded our research, FAPES process n° 525/2018
capixaba researcher. The authors thank Rafael
Mathielo and Maria Ana Farinaccio, for the
photographs; and Mariana de Andrade Wagner,
for the English review.
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Received in October 03, 2019. Accepted in April 22, 2020.
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