Rodriguésia 73: e02432020.

2022
http://rodriguesia.jbrj.gov.br
DOI: http://dx.doi.org/10.1590/2175-7860202273027

Flora of Espírito Santo, Brazil

Angiosperm checklist and conservation of one the richest restingas
in southeastern Brazil

Julia Cristina Guarnier1,2,8, Valquíria Ferreira Dutra1,3, Rodrigo Theófilo Valadares1,4,

Hiago Lourenço da Silva1,5, Vinícius de Castro Freitas1,6 & Renata Henrique Mota1,7

Abstract
The Paulo César Vinha State Park (PEPCV), an important conservation unit in the state of Espírito Santo,
has one of the richest angiosperm communities of any restinga area in Brazil, making it a priority area for
biodiversity conservation. In this study, we perform an inventory of the PEPCV flora through an exploratory
collection over the park area and compilation of data from botanical collections available in virtual herbaria.
A total of 634 species were listed, distributed into 367 genera and 111 families of angiosperms, representing
43.75% of the species documented in the restingas of Espírito Santo. The richest families were Fabaceae (54
spp.), Cyperaceae (40 spp.), Orchidaceae (40 spp.), Myrtaceae (36 spp.), and Poaceae (30 spp.). Six species
were recorded for the first time in the park, ten are endemic to the Espírito Santo, and about 10% of the species
are threatened with extinction. We provide data on the habit, species distribution in phytophysiognomies,
and endangered species. Finally, we discussed the importance of broad knowledge about the park’s plants,
exposing points to conserve specific areas with high antopic rate, and plans to preserve the biodiversity of
this ecosystem.
Key words: angiosperms, Atlantic Forest, Espírito Santo, Paulo César Vinha State Park, phytophysiognomies.
Resumo
O Parque Estadual Paulo César Vinha (PEPCV) é uma importante Unidade de Conservação do estado do
Espírito Santo, e está entre as áreas de restinga mais ricas em espécies de angiospermas do Brasil, sendo
considerado uma área prioritária para conservação da biodiversidade. Neste estudo inventariamos a flora do
PEPCV, por meio de coletas exploratórias ao longo da área do parque e compilação de dados de coleções
botânicas disponíveis em herbários virtuais. Foram listadas 634 espécies, reunidas em 367 gêneros e 111
famílias de angiospermas para o parque, o que representa 45,5% das espécies citadas para as restingas
capixabas. As famílias mais ricas em espécies foram Fabaceae (54 spp.), Cyperaceae (40 spp.), Orchidaceae
(40 spp.), Myrtaceae (36 spp.) e Poaceae (30 spp.). Seis espécies são novos registros para a flora do parque,
10 são endêmicas do estado do Espírito Santo e cerca de 10% das espécies são consideradas ameaçadas de
extinção. São apresentados dados sobre o hábito, a distribuição das espécies nas fitofisionomias e as espécies
ameaçadas. Por fim, discutimos a importância de um amplo conhecimento sobre as plantas do parque, expondo
pontos para a conservação de áreas específicas com altos índices de antropização e planos de preservação da
biodiversidade desse ecossistema.
Palavras-chave: angiospermas, Floresta Atlântica, Espírito Santo, Parque Estadual Paulo César Vinha,
fitofisionomias.

See supplementary material at <https://doi.org/10.6084/m9.figshare.17192903.v1>

1
Universidade Federal do Espírito Santo - UFES, Centro de Ciências Humanas e Naturais, Depto. Ciências Biológicas, Lab. Sistemática Vegetal, Vitória, ES, Brasil.
2
ORCID: <https://orcid.org/0000-0003-3486-8265>. 3 ORCID: <https://orcid.org/0000-0003-1547-1377>. 4 ORCID: <https://orcid.org/0000-0002-6528-2278>.
5
ORCID: <https://orcid.org/0000-0003-1330-9085>. 6 ORCID: <https://orcid.org/0000-0001-9189-9522>. 7 ORCID: <https://orcid.org/0000-0002-6900-8911>.
8
Author for correspondence: [email protected]
2 de 12 Guarnier JC et al.

Introduction created to preserve the restinga ecosystem in the

The Atlantic Forest is recognized as a state and is home to vegetation with little anthropic
global biodiversity hotspot, with high biological interference (Chagas et al. 2014). In addition, it
richness and a large number of threatened species represents the main restinga area in the south of
(Mittermeier et al. 2004). The biome is made up Espírito Santo and represents one of the richest
of distinct phytoecological regions, distributed restingas in terms of angiosperm species along the
along an extensive latitudinal range with extremely Brazilian coast (Guarnier 2019).
heterogeneous geomorphological and climatic Although PEPCV is recognized as a priority
characteristics (Azevedo 1962; Veloso et al. 1991). area for conservation, continued impacts persist,
Its coastal area has seen varied and extensive such as illegally set fires (Espírito Santo 2010;
economic uses since Brazil’s colonial period (Dean Narciso 2012; Brasil 2018a). These environmental
1995), resulting in a fragmented natural landscape variations, in addition to the ecosystem services
of only 11.73% of its original coverage (Ribeiro et provided by the restinga environment, underpin
al. 2009; Fundação SOS Mata Atlântica & INPE decisions to direct resources to the PEPCV and
2019). local environmental authorities (Espírito Santo
Despite a well-documented literature on the 2009). In particular, floristic surveys are important
reforestation of the Atlantic Forest (Atlantic Forest for land managers and help to identify and conserve
Restoration Pact 2009), the recovery of some of its remnants with unique vegetation cover (Chaves
ecosystems - such as the restinga - has presented et al. 2013). Online platforms are increasingly
a challenge. Restinga vegetation occurs on coastal being used to assist with these floristic surveys
sandy deposits, which extend along the Brazilian and provide information on species richness within
coast (Cerqueira 2000). This vegetation occurs on a Brazilian conservation units (Moreira et al. 2020).
quaternary plain (IBGE 2012) and is remarkable for In spite of the volume of research that has been
the richness of its plant communities, which grow carried out in PEPCV (e.g., Behar & Viegas 1992;
on nutrient-poor, sandy sediments that are usually Martins et al. 1999; Assis et al. 2003, 2004; Fraga
of marine origin and rarely fluvial (Pereira 2003). & Peixoto 2004; Silva & Piassi 2010; Valadares et
In Espírito Santo state, the restinga was al. 2010; Gomes & Silva 2013; Chagas et al. 2014;
originally present along 411 km of the coastline Peterle et al. 2015), no study has yet been able to
on the state (Albino et al. 2016), its continuous present an exhaustive list of the park’s angiosperms.
nationwide decline is especially pronounced in this Therefore, the purpose of this study is
region (e.g., Pereira 2007; Brasil 2018a; Espírito to provide an updated floristic list of PEPCV,
Santo 2013). The coast has a geomorphological with information on the occurrence of
peculiarity that directly influences the restingas phytophysiognomies and conservation status.
ecosystem of the state. In the northern region, we By doing this, we aim to assist researchers and
observe more extensive restingas and are composed environmental authorities in their efforts to enact
of a small variety of habitats - a characteristic policies aimed at the conservation of restingas.
derived from the influence of the Tabuleiro Forest
located on the Northeast Coast. In the southern Material and Methods
region, below the Rio Doce delta, we observe more Study area
preserved restingas with more variety of habitats The PEPCV was created by decree nº 2.993-N
- the result of the influence of the Precambrian in 1990, with initial denomination of Setiba’s Park.
Crystalline Complex located on the Southeast Through the Law nº 4.903 in 1994, became known
Coast (Silveira 1968). The vegetation is composed as Paulo César Vinha State Park. Is situated on a
of different phytophysiognomies made up of coastal plain with an area of about 1,500 ha, located
established herbaceous, shrub, or tree communities, within the Environmental Protection Area of
with the makeup depending on the level of the water Setiba in the Guarapari municipality, metropolitan
table, among other factors (Pereira 2003; Magnago region of Grande Vitória, on the southern coast of
et al. 2011a). Espírito Santo (Fig. 1). The area is located between
The Paulo César Vinha State Park (PEPCV) 20°32’13” and 20°37’41” S and between 40°22’58”
in Espírito Santo contains most of the restinga and 40°25’39”W (CEPEMAR 2007). According to
phytophysiognomies identified in the literature the Köppen classification (Alvares et al. 2014), the
(Pereira 2003). This was the first conservation unit region’s climate is classified as Aw tropical, with an

Rodriguésia 73: e02432020. 2022

Angiosperm checklist from a restinga area 3 de 12

annual average temperature between 24 °C and 26 (2003). The mapping of typologies identified was
°C and average annual precipitation of 1,160 mm prepared using QGIS v 2.18 software (2016).
(INCAPER 2019). The vegetation is predominantly PEPCV also has a rocky outcrop adjacent to the
composed of restinga, and eight plant formations intermittent mouth of the Caraís Lagoon, which is
can be identified (Fig. 2) according to Pereira its main water body.

Collection and preparation

of the database
Fertile samples (flowers and/or fruits) of
angiosperms were collected between May 2016
and August 2020 using the walking method
(Filgueiras et al. 1994). The collected material
was processed according to the usual methods
of specimen preservation (Fidalgo & Bononi
1989), and then deposited at the VIES Herbarium
in the Federal University of Espírito Santo,
Vitória, Brazil. The identification of the collected
specimens was performed using dichotomous
keys, by the comparison with specimens deposited
in the VIES Herbarium collection, consultation
with the INCT-HVFF (<http://inct.splink.org.
br/>) and REFLORA (2020) virtual herbaria, as
well as by consulting with specialists on different
botanical families.
We also consulted virtual herbarium
collections to help prepare our database of
the specimens collected in the PEPCV. Our
search included the key terms “Parque de
Setiba”, “Lagoa do Milho”, and “Lagoa de
Caraís” to include materials collected before the
current nomenclature of the park and without
geographical coordinates. The data from this
research included specimens from the following
herbariums: ALCB, BHCB, BOTU, CEN,
CEPEC, CESJ, EAC, FLOR, HRCB, HUCS,
HUEFS, HURB, IAC, ICN, JPB, MBM, MBML,
MO, NY, OUPR, RB, RBR, RFA, SAMES, SP,
SPF, SPSF, TEPB, UEC, UFP, UPCB, VIC,
and VIES (Thiers, continuously updated). We
considered dried specimens with identification by
specialists to be reliably determined for inclusion
in the database.
The third source of data included material
from articles with floristic, phytosociological,
and taxonomic surveys carried out in the PEPCV
Figure 1 – Location of the Paulo César Vinha State area: Martins et al. 1999; Assis et al. 2003, 2004;
Park, Guarapari, Espírito Santo, southeastern Brazil, Valadares et al. 2010, 2020; Chagas et al. 2014;
showing the phytophysiognomies present, according Peterle et al. 2015). All names were validated by
to Pereira (2003). The non-floodable herbaceous consulting BFG (2018). The information from the
formation, although not represented due to the scale, three data acquisition sources was organized in a
occurs along the entire length of the park following floristic list following the classification proposed
the beach line. by APG IV (2016).

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4 de 12 Guarnier JC et al.

a b

c d

e f

Figure 2 – a-f. Phytophysiognomies observed at the Paulo César Vinha State Park, Guarapari, Espírito Santo,
southeastern Brazil – a,b. flooded herbaceous; c. open non-floodable shrub; d. open floodable shrub; e. floodable
forest; f. non-floodable forest.
Rodriguésia 73: e02432020. 2022
Angiosperm checklist from a restinga area 5 de 12

The species habit assessment was carried out supplementary material <https://doi.org/10.6084/
following the design proposed in the Technical m9.figshare.17192903.v1>).
Manual of Brazilian Vegetation (IBGE 2012). Forest phytophysiognomies (floodable
Data were obtained by observing specimens in and non-floodable) showed a greater number
the field and from the labels of material consulted of species habits than the shrub and herbaceous
in the herbaria. phytophysiognomies. Only the flooded forest
Finally, data on the geographic distribution formation had lower numbers. Herbaceous plants
of the species were obtained from BFG (2018). predominated on the rocky outcrops, though most
The species’ conservation status at national and of the species habits evaluated were also present.
state level was consulted in the Red Book of Our study found records of six species
Brazilian Flora (Martinelli & Moraes 2013) and that had not been previously reported in the
the List of Endangered Species in Espírito Santo PEPCV, as follows: Ageratum conyzoides L.
(Fraga 2019; Fraga et al. 2019), respectively. (Asteraceae), Lemna minuta Kunth. (Araceae),
Lepidaploa coulonioides (H.Rob.) H.Rob.
Results (Asteraceae), Peltaea obsita (Mart. ex Colla)
A total of 634 species were found, Krapov (Malvaceae), Rourea gardneriana Planch.
distributed into 367 genera and 111 families of & Cristóbal (Connaraceae), and Stachytarpheta
angiosperms in the PEPCV (Tab. S1, available on angustifolia (Mill.) Vahl (Verbenaceae).
supplementary material <https://doi.org/10.6084/ Ten species endemics to Espírito Santo
m9.figshare.17192903.v1>). The following ten were identified [Aechmea fosteriana L.B. Sm.,
families had the highest number of species: Fabaceae A. roberto-seidelii E. Pereira, Cryptanthus
(54 spp.), Cyperaceae (40 spp.), Orchidaceae (40 maritimus L.B. Sm, Neoregelia macrosepala L.B.
spp.), Myrtaceae (36 spp.), Poaceae (30 spp.), Sm. (Bromeliaceae), Clusia spiritu-sanctensis
Mariz & Weinberg (Clusiaceae), Rhynchospora
Rubiaceae (27 spp.), Bromeliaceae (25 spp.),
p l u s q u a m ro b u s t a L u c e ñ o & M . M a r t i n s
Apocynaceae (23 spp.), Asteraceae (22 spp.), and
(Cyperaceae), Ocotea arenicola L.C.S. Assis &
Melastomataceae (18 spp.). Together, these families
Mello-Silva (Lauraceae), Rhodostemonodaphne
make up approximately 50% of the species richness
capixabensis Baitello & Coe-Teix. (Lauraceae),
of the PEPCV.
Callianthe inaequalis (Link &Otto) Donnel.
The most representative genera in numbers
(Malvaceae), and Dichanthelium peristypum
of taxa were Eugenia (12 spp.), Cyperus (9 spp.),
(Zuloaga & Morrone) Zuloaga (Poaceae)]. Among
Rhynchospora (9 spp.), Myrcia (8 spp.), Aechmea
them, two species are restricted to restingas of
(8 spp.), Solanum (7 spp.), Chamaecrista (7
this state: Rhynchospora plusquamrobusta e
spp.), Paspalum (6 spp.), Eleocharis (6 spp.), and
Rhodostemonodaphne capixabensis.
Miconia (6 spp).
A total of 64 threatened species were found
The most representative species habits
(Tab. S2, available on supplementary material
were herbs (202 spp.), followed by trees (157
spp.), shrubs (101 spp.), climbers/liana (77 spp.),
subshrubs (45 spp.), epiphytes (34 spp.), palm
(7spp.), hemiparasite (6spp.) and hemiepiphytes
(5 spp.) (Figs. 3-4).
The phytophysiognomies more richness in
species are the non-floodable forest formation
(263 spp.), followed by the flooded and floodable
herbaceous formations (161 spp.), floodable forest
formation (101 spp.), non-floodable open shrub
(90 spp.), floodable open shrub (69 spp.), rocky
outcrops (30 spp.), non-floodable herbaceous (22
spp.), and flooded forest (17 spp.) (Fig. 5). The sum
of the species values in the formations is higher
than the total species that occur in the PEPCV Figure 3 – Percentage of species habits in the Paulo
due to the fact that some species occur in different César Vinha State Park, Guarapari, Espírito Santo,
phytophysiognomies (Tab. S1, available on southeastern Brazil.
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6 de 12 Guarnier JC et al.

a b

c d

e f

Figure 4 – a-f. Species with different habits sampled at the Paulo César Vinha State Park, Guarapari, Espírito Santo,
southeastern Brazil – a. Ouratea cuspidata (tree); b. Actinocephalus ramosus (herb); c. Phyllanthus klotzschianus
(subshrub); d. Marcetia taxifolia (shrub); e. Tillandsia stricta (epiphyte); f. Passiflora alata (climbers/liana).
Rodriguésia 73: e02432020. 2022
Angiosperm checklist from a restinga area 7 de 12

<https://doi.org/10.6084/m9.figshare.17192903. The peculiarity of angiosperms list presented

v1>), corresponding to 10.4% of the species present here can be explained in two ways. The first is due
in the PEPCV. Of these, 26 species are present in to the diversity of phytophysiognomies present in
the Red Book of Brazilian Flora (Martinelli & the area, which offers a greater variety of niches
Moraes 2013), with half at the vulnerable state than other restingas on the coast of Espírito
(VU) and the other half endangered (EN). At the Santo (e.g., restinga de Camburi; Pereira & Assis
state level, 55 species are present in the List of 2000). The second factor involves the geographic
Endangered Species of Espírito Santo (Fraga et al. positioning of the PEPCV. Espírito Santo is
2019b), with 6% critically endangered (CR), 34% geomorphologically unique as it is located between
EN, and 60% VU. two coastal regions (Silveira 1968) the Eastern
Most threatened species occur in the non- coast is made up of the Barreiras Formation, to
floodable forest formation (31 spp.), followed by the north of the Doce River delta; to the south of
the non-flooded open shrub formation (14 spp.). the Doce River delta, the Southeast Coast is cut by
The remaining phytophysiognomies account for the presence of the escarpments of the Precambrian
less than 15% of all threatened species (Fig. 6). Crystalline Complex.
The plants of the restinga are heavily affected
Discussion by neighboring ecosystems (Matias & Nunes
According to BFG (2018), the restinga in 2000), which has an effect on species distributions.
Espírito Santo has 1,447 species of angiosperms. The flora of the PEPCV is influenced by the
This study indicates that PEPCV contains the nearby forests of the Crystalline Complex, with a
richest restinga in terms of Angiosperms species predominance of the Atlantic Forest species (Assis
statewide, with 43.75% of all known angiosperm et al. 2004). However, statistical tests that include
species in the state’s restinga (Fig. 7). The PEPCV medium-altitude forests in the Southeast Coast
is followed by the Itaúnas State Park in Conceição are necessary to support the influence hypothesis.
da Barra at the extreme northern end of the state, This study shows that forest formations
which has 562 species of angiosperms, accounting have the highest richness of species and life habits
for 40.1% of the known angiosperm species in the PEPCV. Some authors (Freiberg 1994;
in the restingas of Espírito Santo (Souza et al. Parker 1995) argue that this richness in forest
2016). The significant contribution of the families formations occurs due to the greater stratification
Fabaceae, Myrtaceae, Poaceae, Bromeliaceae, in comparison to other phytophysiognomies,
Rubiaceae, and Asteraceae to the species richness which causes microclimate differences due to
of restingas has been documented in other studies the heterogeneity of strata and provides a greater
carried out in different sites in northeastern number of niches for plant species. However, there
(Queiroz et al. 2012; Oliveira et al. 2014; Santos- is an evident decrease in richness between non-
Filho et al. 2015), southeastern (Martins et al. floodable forest formations and those with a higher
2007; Araújo et al. 2009), and southern (Scherer degree of flooding. This result has been associated
et al. 2005) Brazil. with species selection caused by seasonal water

Figure 5 – Percentage distribution of flora species

among the phytophysiognomies of the Paulo Figure 6 – Distribution of threatened species among
César Vinha State Park, Guarapari, Espírito Santo, the phytophysiognomies of the Paulo César Vinha State
southeastern Brazil. Park, Guarapari, Espírito Santo, southeastern Brazil.
Rodriguésia 73: e02432020. 2022
8 de 12 Guarnier JC et al.
saturation in flooded restinga formations (Magnago A.Gray, and Schinus terebinthifolia Raddi, are
et al. 2010, 2013). routinely found in the literature of the restinga
Some of the species listed here are flora in Espírito Santo (Pereira 1990; Magnago et
characteristic of forest formations within the al. 2011b; Bove et al. 2003; Kuster et al. 2019).
PEPCV. Assis et al. (2003; 2004) cite Alchornea Our findings on the species richness of
triplinervia (Spreng.) Mull. Arg., Jacaranda flooded and floodable herbaceous formations
puberula Cham., Myrciaria floribunda O.Berg, significantly exceeded previous estimates (Pereira
Protium heptaphyllum (Aubl.) Marchand, 1990). The present results show that Cyperaceae
and Tapirira guianensis Aubl. as the most and Poaceae were the most representative of these
representative taxa of non-floodable forest formations, in line with previously findings. The
formations in the park. Other species listed here representativeness of these families in the PEPCV
for flooded forests, Bactris setosa Mart., Geonoma can be linked to two important factors: the great
schottiana Mart., and Symphonia globulifera L.f.), extent of herbaceous formations in the park area
are also characteristic of these formations in other (Fig. 1) and the capacity of the species of these
restinga areas of the Espírito Santo (Magnago et families to colonize open environments with
al. 2011b). flood pulses (Bove et al. 2003; Valadares et al.
Some authors indicate floristic differences 2020). Previous studies also indicated the strong
between flooded and non-flooded open shrub contribution of these families to species richness in
formations (Kuster et al. 2019). Although the similar areas (Araújo & Henriques 1984; Sá 1992;
water table outcrop has not been observed in Araújo et al. 1998). Although the data may indicate
the flooded areas mapped here, species like a strong effect caused by the taxonomic study
Abildgaardia baeothryon A.St.-Hil. found in of Cyperaceae in PEPCV (Martins et al. 1999),
areas between thickets were associated with taxonomic efforts that include Poaceae in the
stretches with observed humidity. Species like future are expected to further increase the richness
this can be indicators for such areas, supporting of flooded and floodable herbaceous formations.
the hypothesis that proximity to the water table It is a general rule that non-floodable
is a regulatory factor in the development of some herbaceous formation has lower richness species
species (Magnago et al. 2011a). Species inherent than other restinga phytophysiognomies (Araújo
to these formations, such as Allagoptera arenaria & Henriques 1984; Thomaz & Monteiro 1993;
(Gomes) Kuntze, Byrsonima sericea DC., Cupania Magnago et al. 2011b; Bove et al. 2003; Souza et
emarginata Cambess., Paullinia weinmanniifolia al. 2016). All species related to this formation are

Figure 7 – Comparison of the most representative families in terms of species richness cited for restingas, as follows:
Restingas’s ES (BFG 2018); PEPCV - Paulo César Vinha State Park (present study); PEI - Itaúnas State Park (Souza
et al. 2016); PN - Praia das Neves (Braz et al. 2013).
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Angiosperm checklist from a restinga area 9 de 12

resilient and widely distributed along the Brazilian Acknowledgments

coast (BFG 2018). However, some species that The authors are grateful to the Instituto
are present in lower frequencies (e.g., Scaevola Estadual de Meio Ambiente e Recursos Hídricos
plumieri (L.) Vahl and Sporobolus virginicus (L.) (IEMA), for aiding with data collection. We offer
Kunth) can serve as indicators of native vegetation our special thanks to the team at the Paulo César
on the beach, as a consequence of conservation of Vinha State Park (PEPCV), who provided us with
the area. access to the area and a positive relationship for
The threatened angiosperm checklist of future work. We are deeply grateful to researchers
the PEPCV has a representative percentage at the VIES herbarium, who contributed their time
of species, even though it is a fully protected and helped to refine the manuscript: our thanks to
Conservation Unit inserted in a biodiversity Aline Pitol Chagas, José Manoel Lúcio Gomes,
hotspot (Mittermeier et al. 2004). The park has and Luana Silva Braucks Calazans, his detailed
been designated a “Sítio-BAZE” (MMA 2018) reading in the manuscript enriched the research
since it shelters several species threatened and our scientific perspective.
with extinction within a declining ecosystem.
In the park, even historically poorly studied
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Area Editor: Dr. Rafael Pinto

Received in December 28, 2020. Accepted in May 18, 2021.
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