Flora of Espírito Santo: Droseraceae

Abstract

We present a taxonomic treatment of the species of Droseraceae in Espírito Santo state, Brazil. Droseraceae is represented in the state by two species: Drosera intermedia, found in marshlands in the restingas (pioneer formations), and Drosera latifolia, which occurs in the campos de altitude (ecological refuges) and on granitic inselbergs in areas of ombrophilous forest in the mountainous region of Espírito Santo. We provide an identification key, morphological descriptions, and commentaries on taxonomy, ecology, conservation, and geographical distribution.

Full text

Rodriguésia 73: e00312021. 2022
http://rodriguesia.jbrj.gov.br
DOI: http://dx.doi.org/10.1590/2175-7860202273016
Abstract
We present a taxonomic treatment of the species of Droseraceae in Espírito Santo state, Brazil. Droseraceae
is represented in the state by two species: Drosera intermedia, found in marshlands in the restingas (pioneer
formations), and Drosera latifolia, which occurs in the campos de altitude (ecological refuges) and on
granitic inselbergs in areas of ombrophilous forest in the mountainous region of Espírito Santo. We provide
an identification key, morphological descriptions, and commentaries on taxonomy, ecology, conservation,
and geographical distribution.
Key words: Atlantic Forest, carnivorous plants, Drosera, inselbergs, restingas.
Resumo
Apresentamos o tratamento taxonômico das espécies de Droseraceae para o Espírito Santo. Droseraceae está
representada no estado por duas espécies: Drosera intermedia, encontrada em locais brejosos nas restingas
(formações pioneiras), e Drosera latifolia, que ocorre nos campos de altitude (refúgios ecológicos) e em
inselbergues graníticos inseridos em áreas de floresta ombrófila da região serrana do Espírito Santo. Fornecemos
chave de identificação, descrições morfológicas, comentários taxonômicos, ecológicos, de conservação e de
distribuição geográfica.
Palavras-chave: Mata Atlântica, plantas carnívoras, Drosera, inselbergs, restingas.
Flora of Espírito Santo, Brazil
Flora of Espírito Santo: Droseraceae
Paulo Minatel Gonella1,5, Sabrina Vasconcelos Caram2,3 & Valquíria Ferreira Dutra2,4
1 Universidade Federal de São João del-Rei, Campus Sete Lagoas, Depto. Ciências Exatas e Biológicas, Sete Lagoas, MG, Brasil. ORCID: <https://orcid.
org/0000-0001-8332-5326>.
2 Universidade Federal do Espírito Santo, Depto. Ciências Biológicas, Vitória, ES, Brasil.
3 ORCID: <https://orcid.org/0000-0002-3004-8182>.
4 ORCID: <https://orcid.org/0000-0003-1547-1377>.
5 Author for correspondence: pmgonella@gmail.com
Introduction
Droseraceae (non-core Caryophyllales,
Eudicots) is a family of carnivorous herbs
composed of three genera and around 250 species
(Fleischmann et al. 2018). Dionaea and Aldrovanda
are monotypic and do not occur naturally in Brazil
but are cultivated in the country by enthusiasts. On
the other hand, Drosera is a cosmopolitan genus
that contributes most species to the family, with
centers of diversity in Australia, Brazil, and South
Africa. Brazil, in particular, is home to 31 species,
one nothospecies, and two varieties, of which 19
species are endemic to the country (Gonella 2020;
Flora do Brasil 2020).
The genus is characteristic of open formations
and is often associated with quartzitic and
oligotrophic soils that are, at least, seasonally wet
(Juniper et al. 1989; Fleischmann et al. 2018).
Drosera is a characteristic element of the endemic
flora of the Brazilian montane grasslands called
campos rupestres (Colli-Silva et al. 2019) and is
usually absent from forest formations, except for a
few species that can be found occurring in riverine
forests (Gonella 2020).
The state of Espírito Santo is entirely located
within the Atlantic Forest Domain, with an original
cover of ombrophilous and seasonal semideciduous
forests, in addition to pioneer formations such as
vegetation in the coastal plains (restingas) and
ecological refuges for higher-altitude flora in the
montane grasslands of Serra do Caparaó (Garbin et
al. 2017). Despite having some of the greatest
floristic diversity of any Brazilian state (Dutra et
al. 2015; BFG 2018), only 12.6% of its original

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vegetation cover remained (Fundação SOS Mata
Atlântica & INPE 2019), highlighting the relevance
and urgency of floristic inventories, taxonomic
treatments, and conservation efforts.
This study is part of the Flora of Espírito
Santo Project and aims at presenting a taxonomic
treatment for the Droseraceae occurring in
the state and to provide an identification key
and descriptions, along with commentaries on
taxonomy, ecology, conservation, and geographical
distribution.
Material and Methods
Descriptions and phenological data were based
on specimens studied in the field and herbarium
specimens, complemented by information in the
literature. Specimens held at the herbaria HB,
MBML, MO, NY, R, RB, US, and VIES were
studied in person, while one specimen from the GH
herbarium was analyzed at the institution’s virtual
herbarium, available at <https://kiki.huh.harvard.
edu/databases/specimen_index.html> (acronyms
according to Thiers, continuously updated). The
morphological terminology used in the species
descriptions follows Gonella et al. (2014, 2015),
Rivadavia et al. (2014), and Fleischmann et al.
(2018).
Following a recent reassessment of the
Red List conservation status of the species in
Espírito Santo (Fraga et al. 2019), the Droseraceae
species discussed in this paper had their regional
conservation status newly assessed based on new
data on their geographical distribution in the
state. The assessment follows the IUCN (2012a)
categories and criteria, including the IUCN (2012b)
guidelines for regional assessments. We used the
conservation assessment tool GeoCAT (available at
<http://geocat.kew.org/>) to calculate the extent of
occurrence (EOO) and area of occupancy (AOO)
as described in Bachman et al. (2011) using the
standard IUCN cell size of 4 km2 for the AOO
estimation.
Results and Discussion
In Espírito Santo, Droseraceae are represented
only by two species of the genus Drosera. Drosera
intermedia Hayne in Dreves & Hayne (1798: 18)
occurs in wet areas of the restingas (Fig. 1). In
contrast, D. latifolia (Eichler 1872: 395) Gonella
& Rivadavia in Gonella et al. (2014: 20) is a
rupicolous species found in montane grasslands
and on granitic rock outcrops (inselbergs; Fig. 1).
Three Drosera species have been previously
reported for Espírito Santo. Valadares et al. (2020)
recorded D. intermedia in an area of marshland in
the restinga. Silva & Giulietti (1997) and Correa
& Silva (2005) reported D. villosa A.Saint-
Hilaire (1824: 2607), though these records were
reclassified as D. latifolia by Gonella et al. (2014).
A record of D. montana A.Saint-Hilaire (1824:
260) was listed by Dutra et al. (2015) based on the
voucher Ynês Mexia 40251 (GH, MO), but that is
also D. latifolia.
Drosera montana is recorded from the Minas
Gerais side of the Serra do Caparaó (Rivadavia
et al. 2014; Moreira et al. 2020). However, no
specimens have been found on the Espírito Santo
side thus far.
Droseraceae Salisb.
Annual or perennial carnivorous herbs.
Stem inconspicuous or elongated. Leaves simple,
stipules present or absent, vascularized glandular
Figure 1 – Geographic distribution of Drosera species
in the state of Espírito Santo. ● = Drosera intermedia;
▲ = Drosera latifolia.

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emergences present on the adaxial lamina
surface, or lamina modified into a bilobed snap
trap with sensitive trichomes over the adaxial
surface. Inflorescence cymose, multi-flowered
(rarely uniflorous). Flowers 5-merous (rarely
4-merous), actinomorphic; stamens 5(–many),
anthers longitudinally dehiscent; ovary superior,
3–5-carpellar; styles 1–5 (usually 3), usually
bifurcated at the base (rarely entire); ovary with
parietal placentation (rarely basal), pluriovulated.
Fruit a loculicidal capsule; seeds numerous.
Drosera L. (description based solely on the species
occurring in Espírito Santo).
Terrestrial or rupicolous herbs. Leaves
rosetted or spirally arranged, stipulate, petiolate,
red to green in color, with sessile glands; stipules
intrapetiolar, membranaceous, with apex laciniate;
petiole linear, glabrous or eglandular-pilose at least
abaxially; lamina oblong, lanceolate or spatulate,
with vascularized glandular emergences (tentacles)
on the adaxial surface, abaxial surface glabrous or
eglandular-pilose. Inflorescence a scorpioid cyme,
multiflorous (rarely uniflorous), 1–few per plant,
with sessile glands; scape glabrous or eglandular
and glandular-pilose. Flowers ephemeral, anthesis
lasting for a few hours; sepals fused at the base,
glabrous or glandular-pilose; petals white or pink;
stamens 5; ovary 1-locular; styles bifurcated at the
base. Petals, sepals and stamens persistent in fruit.
Seeds with papillose or reticulate testa.
Drosera is the largest genus in the family,
comprising about 250 species distributed
worldwide and concentrated in the Southern
Hemisphere. The main center of diversity of the
genus is in the southwest of Western Australia,
followed by northern Australia, the Cape Province
of South Africa, and the mountains of east-central
Brazil (Fleischmann et al. 2018). The genus is
immediately identified by the presence of glandular
emergences (tentacles) over the adaxial leaf lamina
surface (Figs. 2-4). In Espírito Santo, Drosera is
represented by two species, which are described
below.
Identication key to the species of Drosera from the Espírito Santo state
1. Leaves, scape and sepals glabrous. Lamina 5.2–8.1 mm long, spatulate. Sepals 3–3.5 mm long; petals
white. Seeds ovoid, testa surface papillose ..........................................................1. Drosera intermedia
1’. Leaves eglandular-pilose, at least on the abaxial surface, scape and sepals glandular and eglandular-
pilose. Lamina 8.4–9.75 mm long, lanceolate to oblong. Sepals 5–5.5 mm long; petals pink. Seeds
fusiform, testa surface reticulate .............................................................................. 2. Drosera latifolia
1. Drosera intermedia Hayne in Dreves & Hayne,
Bot. Bilderb. 3: 18, t. 3B. 1798. Figs. 1; 2a-d; 3
Perennial rosetted herb. Subglabrous
indumentum consisting solely of minute sessile
glands, red in color (black in dry specimens), on
both leaf surfaces, scapes and sepals abaxially.
Leaves persistent when old, reddish; petiole 15–30
mm long; lamina 5.2–8.1 mm long, spatulate, apex
rounded, adaxial surface covered with numerous
red, carnivorous, capitate tentacles, 0.1–3.4 mm
long, and minute sessile glands, abaxial surface
covered with minute sessile glands; stipules
2.6–4.1 mm long, rectangular, membranaceous,
the apical divided into several laciniate segments.
Scapes 2–4 per plant, 9–14 cm long (including
floriferous part), flattened, base curved, green
to red in color; inflorescence a scorpioid cyme,
bearing 6–10 flowers; sepals 5, 3–3.5 mm long,
lanceolate, minute sessile glands, apex acute;
petals 5, 4.5–5 mm long, whitish; stamens 5,
3.5–4 mm long, anthers 0.53–0.74 mm long; ovary
3-carpellate, fused, ca. 2.5 mm long, globose,
3-lobed in outline; styles 3, forked at the base; fruit
a dry dehiscent capsule, ca. 6 mm long, globose,
3-valvate; seeds ovoid, 0.4–0.82 mm long, testa
papillose.
Specimens examined: Conceição da Barra,
Parque Estadual de Itaúnas, 18.5932oS, 39.7322oW,
24.VIII.2002, O.J. Pereira et al. 6997 (VIES). Guarapari,
Parque Estadual Paulo César Vinha, 20.5994oS,
40.4108oW, 6.VIII.2020, J.C. Guarnier et al. 467 (VIES);
18.VIII.2020, fl. and fr., S.V. Caram et al. 01 (VIES).
Vitória, área da C.V.R.D., 20.3194oS, 40.3377oW,
15.VIII.2007, fl., O.J. Pereira & G. Lübe 7517 (VIES).
Vila Velha, PEPCN, 20.3297oS, 40.2924oW, 29.IX.2007,
fl., R.T. Valadares et al. 578 (HURB).
Drosera intermedia has a wide distribution,
occurring in central and western Europe, Asia
Minor, East Africa, and North, Central, and
South America, including Brazil (Fleischmann
& Gonella 2020; Gonella 2020). The species was

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recently recorded in Espírito Santo for the first
time (Valadares et al. 2020), expanding its known
distribution in Brazil, where it has also been
recorded in the states of Rio de Janeiro, Bahia,
Amapá, and Roraima (Gonella 2020). In Espírito
Santo, the species occurs in the restingas (Fig. 1),
in areas of herbaceous marshland. It has been found
in the protected areas of the Parque Estadual Paulo
César Vinha (PEPCV) and the Parque Estadual de
Itaúnas. In the wet areas of the PEPCV, which are
a structurally dynamic environment, D. intermedia
is always found associated with flat and stabilized
sedimentary zones (humid fields), protected
from natural and dynamic drainage channels that
constantly modify the structure of the vegetation
(Valadares et al. 2020). It was collected in flower
in August and September and with fruits in August.
The species is characterized by glabrous
leaves (only presenting sessile glands; Fig. 3),
spatulate (Figs. 2b; 3b), with a petiole narrower and
Figure 2 – a-d. Drosera intermedia – a. habit; b. detail of the leaf; c. flower; d. fruits (all photos taken at PEPCV).
e-f. Drosera latifolia – e. habit; f. detail of the leaf (all photos taken at PN Caparaó). (a-d. by VF Dutra; e. by HL
Silva; f. by AV Scatigna).
a
b
c
ef
d

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Figure 3 – a-c. Drosera intermedia – a. habit; b. leaf lamina, adaxial surface; c. tentacle head. (All based on Caram
et al. 01).
1.5 cm
1.5 cm
0.02 cm
a
b
c
at least three times longer than the lamina (Figs. 2b;
3a), stipules fimbriate from the base, inflorescence
and sepals glabrous, white petals (Fig. 2c), and
ovoid seeds with a papillose testa.
Drosera intermedia was not assessed for the
Espírito Santo Red List (Fraga et al. 2019) as no
records had been identified in the state at the time
of its elaboration. While the species is globally

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classified as Least Concern (Cross et al. 2020), its
occurrence in the state is restricted to an AOO of
16 km2 (criterion B2; IUCN 2012a). In addition,
it is limited to fewer than five known locations
(sub-criterion B2a), and its habitat is in a state of
continuous decline (sub-criterion B2biii) since
the state’s coast is considered to be under high
anthropic pressure due to pollution, fires, and urban
expansion over the restingas (Maciel 1990; MMA
2002; Fraga et al. 2019). Therefore, we assess the
species as Endangered (EN) in Espírito Santo.
2. Drosera latifolia (Eichler) Gonella & Rivadavia,
Phytotaxa 156: 21. 2014. Figs. 1; 2e-f; 4
Perennial rosetted herb. Indumentum
consisting of white eglandular trichomes (brown
in dry specimens), present on the abaxial leaf
surface and petiole; glandular capitate trichomes
- the stalk light red and translucent, the head
dark red - present on scapes and sepals abaxially;
minute sessile glands red in color (black in dry
specimens), on both leaf surfaces, scapes, pedicels,
bracts and sepals abaxially. Leaves persistent,
green to red; petiole 4.6–5.3(–10) mm long;
lamina 8.4–9.75(–25) mm long, lanceolate to
oblong, apex rounded, adaxial surface covered
with numerous red, carnivorous, capitate tentacles,
2.28–3.54 mm long, and minute sessile glands,
abaxial surface with eglandular trichomes, and
minute sessile glands; stipules 2.48–3.51 mm
long, square, membranaceous, the apical divided
into several laciniate segments. Scapes 1–3 per
plant, 9–18(–34.7) cm long (including floriferous
part), cylindrical, base erect, green in color;
inflorescence a scorpioid cyme, often bifurcating,
bearing 2–9(–27) flowers; sepals 5, 5–5.5 mm long,
lanceolate, with glandular capitate trichomes, and
few eglandular trichomes, apex acute or obtuse;
petals 5, 7–7.5 mm long, pink; stamens 5, 2.5–3.5
mm long, anthers ca. 1 mm long; ovary 3-carpellate,
fused, 2–3 mm in diam., globose, 3-lobed in outline;
styles 3, forked at the base; fruit a dry dehiscent
capsule, ca. 3 mm long, globose, 3-valvate; seeds
fusiform, ca. 0.8 mm long, testa reticulate.
Specimens examined: Alegre, Serra Caparaó, Pico da
Bandeira, 20.4340oS, 41.7966oW, 3.III.1959, fl., H.S.
Irwin 2804 (NY, US). Castelo, Forno Grande, 8.XII.1948,
A.C. Brade 19249 (RB). Domingos Martins, topo da
Pedra Azul, 19.XI.1975, Kautsky 517 (HB). Ibitirama,
Parque Nacional do Caparaó, entre o Acampamento
Macieira e o Pico da Bandeira, 18.I.2019, fl., H.L. Silva
& L.J. Lieven 257 (VIES). São Roque do Canaã, Alto
Misterioso, 19.7969oS, 40.7752oW, 11.V.2007, fr., C.
Esgario et al. 171 (MBML); 19.7972oS, 40.7750oW,
15.XI.2004, fl., A.P. Fontana et al. 1072 (MBML). Serra
da Caparaó, 25.XI.1929, fl., Y. Mexia 40251 (GH, MO).
Drosera latifolia is endemic to Brazil,
occurring in the mountain ranges of Serra do
Mar, Serra da Mantiqueira, and the Espinhaço
Range in the states of Santa Catarina, Paraná, São
Paulo, Minas Gerais, Rio de Janeiro, and Espírito
Santo (Gonella et al. 2014). In Espírito Santo, the
species is rupicolous (Fig. 2e), occurring in areas
of ecological refuges and ombrophilous forests
(Fig. 1). It is found on the summits of the state’s
larger granitic inselbergs, which are considered to
be a transition between the sugar loaf inselbergs
and campos de altitude (montane grasslands) by de
Paula et al. (2020), such as in the Parque Estadual
da Pedra Azul, Parque Estadual do Forno Grande,
and Alto Misterioso, and the campos de altitude
of the Parque Nacional do Caparaó. According to
Gonella et al. (2014), the individuals collected at
Caparaó represent the smallest sized morphotype
of D. latifolia, as well as the one that occurs at the
highest elevation (up to 2,550 m). The remaining
populations in the state represent the typical
morphotype described by Gonella et al. (2014).
The species was previously recorded in the floristic
inventory of Alto Misterioso (Esgario et al. 2009)
and Parque Nacional do Caparaó (Moreira et al.
2020) as D. villosa. Drosera latifolia was recorded
with flowers in January, March, and November and
with fruits in May.
The species is characterized by its lanceolate
to oblong leaves (Figs. 2f; 4), a petiole shorter
than or equaling the lamina, eglandular trichomes
present on the abaxial leaf surface and eventually
on the adaxial petiole surface, glandular-pilose
inflorescence and sepals, pink petals, and fusiform
seeds with reticulate testa.
Drosera latifolia is listed as Data Deficient
in the Espírito Santo Red List (Fraga et al. 2019)
and globally assessed as Least Concern (Cross et al.
2020). The new information on its distribution in the
state presented here allows its regional assessment
to be updated to Near Threatened (NT) in the state
of Espírito Santo. While the species has an AOO of
16 km2 (criterion B2) and a naturally fragmented
distribution due to its occurrence in mountaintops,
with fewer than five locations (sub-criterion B2a),
no continuing decline has been observed to affect
the subpopulations (sub-criterion B2b). Three of the
four known locations in the state are found within
protected areas, except for the Alto Misterioso, a
priority conservation area (Esgario et al. 2009).

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Figure 4 – a-b. Drosera latifolia – a. habit; b. leaf, adaxial surface. (All based on Irwin 2804).
0.8 cm
0.3 cm
a
b
Acknowledgments
The authors are grateful to the curators of the
herbaria they visited. They also thank Lidyanne
Aona, for sending images of the specimens at
HURB; André Vito Scatigna and Hiago Lourenço
da Silva, for in situ photographs of D. latifolia;

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and Andreas Fleischmann and Rodrigo Valadares,
for valuable suggestions for the improvement of
the manuscript.
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This is an open-access article distributed under the terms of the Creative Commons Attribution License.