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CAVE FAUNA OF BRAZIL
1
ORGANIZERS
Robson de Almeida Zampaulo
Xavier Prous
FAUNA
CAVE
OF BRAZIL
CAVE FAUNA OF BRAZIL
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CAVE FAUNA OF BRAZIL
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CAVE FAUNA OF BRAZIL
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Belo Horizonte • aug 2023
FAUNA
CAVE
OF BRAZIL
ORGANIZERS
Robson de Almeida Zampaulo
Xavier Prous
CAVE FAUNA OF BRAZIL
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Gruta da Lapa Doce – Iraquara (BA)
Ataliba Coelho
CAVE FAUNA OF BRAZIL
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CAVE FAUNA OF BRAZIL
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Caverna Terra Ronca – São Domingos (GO)
Ataliba Coelho
CAVE FAUNA OF BRAZIL
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CAVE FAUNA OF BRAZIL
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Gruta Lago Azul – Bonito (MS)
Daniel Menin
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Organizers
ROBSON DE ALMEIDA ZAMPAULO
XAVIER PROUS
Designer
FLÁVIA GUIMARÃES
Edition
Cover photo – Lapa da Onça – Itacarambi (MG)
DANIEL MENIN
Sleeve mosaic photos
LUCAS MENDES RABELO
ROBSON DE ALMEIDA ZAMPAULO
RODRIGO LOPES FERREIRA
Illustrations and maps
MARCOS BRITO
Translation
ERIK R. WILD
Technical review
ALINE DA SILVA REIS
Orthographic review
EDDA CABRAL MARTINS DA COSTA
Standards review ABNT
CARINA REJANE MARTINS CRB6/2336
RACHEL BRAGANÇA DE CARVALHO MOTA CRB6/2838
authors
ADRIANO MARQUES DE SOUZA
Bios Réplicas
adrianoms15@yahoo.com.br
ALESSANDRO DAMASCENO MARQUES
Universidade do Vale do Rio dos Sinos -
Unisinos
alessandrodm@outlook.com
ALINE DA SILVA REIS
Observatório Espeleológico - OE
alineambiental@yahoo.com.br
ALMIR ROGÉRIO PEPATO
Universidade Federal de Minas Gerais -
UFMG
apepato@gmail.com
AMAZONAS CHAGAS-JR
Universidade Federal de Mato Grosso -
UFMT
amazonaschagas@gmail.com
ANA MARIA LEAL-ZANCHET
Pesquisadora do Conselho Nacional de
Desenvolvimento Cientíco e Tecnológico
- CNPQ
lealzanchet@gmail.com
ANDREY JOSÉ DE ANDRADE
Universidade Federal do Paraná - UFPR
andreyandrade@ufpr.br
ANGÉLICO ASENJO
Instituto Tecnológico Vale - ITV
pukara8@gmail.com
ANTONIO DOMINGOS BRESCOVIT
Instituto Butantan
antonio.brescovit@butantan.gov.br
BRENDA KAROLINA GOMES
ALMEIDA
Universidade Federal de Minas Gerais -
UFMG
g.almeidabrenda@gmail.com
CAIO CÉSAR PIRES DE PAULA
Biology Centre CAS
piresdepaula@yahoo.com.br
CLÁUDIO A. R. DE SOUZA
Instituto Butantan
claudio.souza@meioambiente.mg.gov.br
CRISTIANO SAMPAIO COSTA
Universidade Federal de Mato Grosso -
UFMT
csampaioc@gmail.com
DANILO PACHECO CORDEIRO
Instituto Nacional da Mata Atlântica -
INMA
d.pacheco.c@gmail.com
DANTE BATISTA RIBEIRO
Universidade Federal de Minas Gerais -
UFMG
dante.dantebatista@gmail.com
DIEGO DE MEDEIROS BENTO
Instituto Chico Mendes de Conservação
da Biodiversidade - ICMBio
Centro Nacional de Pesquisa e
Conservação de Cavernas - CECAV
diego.bento@icmbio.gov.br
DIEGO MONTEIRO VON SCHIMONSKY
Laboratório de Estudos Subterrâneos - LES
Universidade Federal de São Carlos, São
Carlos - UFSCar
dmvschimonsky@gmail.com
DOUGLAS ZEPPELINI
Universidade Estadual da Paraíba - UEPB
zeppelini@daad-alumni.de
ERICA CRISTINA PACÍFICO DE ASSIS
Museu de Zoologia da Universidade de
São Paulo - MZUSP
ericapacico81@gmail.com
ESTEVAM CIPRIANO ARAÚJO DE LIMA
Museu Nacional, Universidade Federal do
Rio de Janeiro - UFRJ
estevam.araujo@gmail.com
EUNICE APARECIDA BIANCHI GALATI
Universidade de São Paulo - USP
egalati@usp.br
FELIPE F. F. MOREIRA
Fundação Oswaldo Cruz - Fiocruz
ppmeiameiameia@gmail.com
GIOVANNA MONTICELLI CARDOSO
Centro de Estudos em Biologia
Subterrânea - CEBS
Universidade Federal de Lavras - UFLA
gmcardoso.bio@gmail.com
This work was produced with speleological compensation resources
established in the Licença de Instalação e Operação 001/2018 of the Projeto
Expansão Oeste Cava da Divisa – Brucutu Mine, Processo Administrativo
00022/1995/070/2017 and in compliance with Condicionante nº 7 do Termo
de Compromisso de Compensação Espeleológica (TCCE) established between
Vale and the Superintendência de Projetos Prioritários (SUPPRI) da Secretaria
de Estado de Meio-Ambiente e Desenvolvimento Sustentável do Estado de
Minas Gerais (SEMAD).
CAVE FAUNA OF BRAZIL
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GISELE LOPES NUNES
Instituto Tecnológico Vale - ITV
gisele.nunes@itv.org
GIULLIA ROSSI
Universidade Estadual Paulista - UNESP
giulliarossi@hotmail.com
GUILHERME OLIVEIRA
Instituto Tecnológico Vale - ITV
guilherme.oliveira@vale.com
HÉLCIO R. GIL-SANTANA
Fundação Oswaldo Cruz - Fiocruz
helciogil@uol.com.br
IGOR CIZAUSKAS
Organização de Apoio à Pesquisa da
Biodiversidade - OAPBio
icizauskas@gmail.com
IVANKLIN SOARES CAMPOS-FILHO
Department of Biological Sciences
Lefkosia (Nicosia) - Cyprus
ivanklin.lho@gmail.com
JADER OLIVEIRA
Universidade de São Paulo - USP
Universidade Estadual Paulista Julio de
Mesquita Filho - UNESP
jdr.oliveira@hotmail.com
JOSÉ PAULO LEITE GUADANUCCI
Universidade Estadual Paulista - UNESP
joseguadanucci@gmail.com
JULIO CÉSAR DO CARMO VAZ SANTOS
Centro de Estudos em Biologia Subterrânea -
CEBS
Universidade Federal de Lavras - UFLA
Julio.santos3@estudante.ua.br
JÚLIO CÉSAR ROCHA COSTA
Instituto Chico Mendes de Conservação da
Biodiversidade - ICMBio
Centro Nacional de Pesquisa e Conservação de
Cavernas - CECAV
julio.costa@icmbio.gov.br
KLEBER MAKOTO MISE
Grupo de Estudos Espeleológicos do Paraná -
GEEP - Açungui
klebermise@yahoo.com.br
LEONARDO SILVA MARUJO
Acadêmico de Ciências Biológicas e Ilustrador
cientíco
leomarujo.ornito@gmail.com
LEOPOLDO FERREIRA DE OLIVEIRA BERNARDI
Universidade Federal de Lavras - UFLA
leopoldobernardi@gmail.com
LETÍCIA MARIA VIEIRA
Departamento de Entomologia, Universidade
Federal de Lavras - UFLA
leticia.vieira@ua.br
LIVIA CORDEIRO MEDEIROS
Ciclo Azul Meio Ambiente e Sustentabilidade
liv.biosubt@gmail.com
LUCAS DENADAI DE CAMPOS
Universidade de São Paulo - USP
lcdenadai@gmail.com
LUDSON NEVES DE ÁZARA
Museu Nacional, Universidade Federal do Rio
de Janeiro - UFRJ
ludsonazara@yahoo.com.br
LUIZ F. MORETTI INIESTA
Universidade de São Paulo - USP
Instituto Butantan
luiz-moretti@hotmail.com
LUIZ RICARDO L. SIMONE
Museu de Zoologia da Universidade de São
Paulo - MZUSP
lrsimone@usp.br
MARCIO PEREZ BOLFARINI
Laboratório de Estudos Subterrâneos - LES
Universidade Federal de São Carlos - UFSCar
biobolf@gmail.com
MARCONI SOUZA-SILVA
Centro de Estudos em Biologia Subterrânea -
CEBS
Universidade Federal de Lavras - UFLA
marconisilva@ua.br
MARCOS RYOTARO HARA
Universidade de São Paulo - USP
harvestmaniac@gmail.com
MARIA ELINA BICHUETTE
Laboratório de Estudos Subterrâneos - LES
Universidade Federal de São Carlos - UFSCar
lina.cave@gmail.com
MARIANE SOARES RIBEIRO PEREIRA
Vale S.A.
mariane.ribeiro@vale.com
MARINA HENRIQUES LAGE DUARTE
Pontifícia Universidade Católica de Minas
Gerais - PUC MINAS
marinabioacustica@hotmail.com
MATHEUS HENRIQUE SIMÕES
Vale S.A.
matheus.simoes@vale.com
MAURÍCIO CARLOS MARTINS DE ANDRADE
Instituto Chico Mendes de Conservação da
Biodiversidade - ICMBio
Centro Nacional de Pesquisa e Conservação de
Cavernas - CECAV
mauricio.icmbio@gmail.com
MAYARA HIDAKA COVIZZE
Instituto Butantan
mayara.hidaka@butantan.gov.br
MAYSA FERNANDA VILLELA REZENDE
SOUZA
Centro de Estudos em Biologia Subterrânea -
CEBS
Universidade Federal de Lavras - UFLA
mvillelabio@yahoo.com.br
PAULA CABRAL ETEROVICK
Zoologisches Institut - Braunschweig -
Germany
pceterovick@gmail.com
PEDRO GUILHERME BARRIOS SOUZA DIAS
Museu Nacional do Rio de Janeiro - Museu
MNRJ
pedrogdias@gmail.com
RAFAELA BASTOS-PEREIRA
Centro de Estudo em Biologia Subterrânea -
CEBS
Universidade Federal de Lavras - UFLA
rafaelabastospereira@gmail.com
RENATA NEVES BIANCALANA
Universidade Federal do Pará - UFPA
Museu Paraense Emílio Goeldi - MPEG
renata.biancalana@gmail.com
ROBSON DE ALMEIDA ZAMPAULO
Gerência de Espeleologia e Tecnologia – Vale
Grupo de Estudos Ambientais da Serra do Mar
– GESMAR
Observatório Espeleológico - OE
rzampaulo@yahoo.com.br
RODRIGO LOPES FERREIRA
Centro de Estudos em Biologia Subterrânea -
CEBS
Universidade Federal de Lavras - UFLA
drops@ua.br
RODRIGO S. BOUZAN
Universidade de São Paulo - USP
Instituto Butantan
rodrigobouzan@outlook.com
RONNIE ALVES
Instituto Tecnológico Vale - ITV
ronnie.alves@itv.org
SAMUEL CHAGAS BERNARDES
Museum für Naturkunde
samuel.geneticafarm@gmail.com
SAMUEL GEREMIAS DOS SANTOS COSTA
Universidade Federal de Minas Gerais - UFMG
estoupa.bob@gmail.com
SANTELMO VASCONCELOS
Instituto Tecnológico Vale - ITV
santelmo.vasconcelos@itv.org
TANISE GRACIELE DE OLIVEIRA
VASCONCELOS
Universidade do Vale do Rio dos Sinos -
Unisinos
tanisevasconcelos@gmail.com
THAIS GIOVANNINI PELLEGRINI
Departamento de Entomologia, Universidade
Federal de Lavras - UFLA
thais.g.pellegrini@gmail.com
THIAGO DOS SANTOS
Bios Réplicas
thiagoherpeto@gmail.com
VALÉRIA DA CUNHA TAVARES
Universidade Federal da Paraíba - UFPB
Instituto Tecnológico Vale - ITV
val.c.tavares@gmail.com
CAVE FAUNA OF BRAZIL
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Thecadactylus rapicauda
Robson de Almeida Zampaulo
Biological research in caves, of dierent lithologies, has provided signicant advances in the knowledge of the biodiversity
and functioning of subterranean ecosystems. Such advances are crucial for the evolution of environmental analysis relevant to
speleology, whether in the context of environmental licensing or that of scientic research in general.
In this sense, Vale, in partnership with dozens of researchers from dierent institutions, promoted the organization of this book,
which includes the main taxonomic groups and a signicant part of the diversity of known species of subterranean fauna in Brazil.
Throughout this precious work, it is possible to perceive the evolution of the theme in recent years, expressed by the dedication of
everyone in the construction of each chapter and in the search for excellence portrayed in high quality photographs in scenarios
of extreme ecological value and, many times, even untouched, which reect the valuable biodiversity existing in the subterranean
universe.
The results of years of research resulting from speleological monitoring and taxonomic studies, as well as cave relevance analysis
work, are reported here. The result of a highly specialized eort that shares knowledge about the subterranean fauna, combined
with beautiful images of several species in their natural environment (many of which are endemic, rare and unique), this wonderful
book allows us to “dive” into the subterranean world and get to know a little more of the life present in these environments devoid
of light and with few resources for the survival of these extremely adapted living beings.
In this way, and to perpetuate and disseminate the knowledge achieved through the arduous eort of scientists, consultants
and private initiatives, Vale, through its technical sta, promotes with this work another step towards advancing the training of
new professionals and specialists in Subterranean Biology. Undoubtedly, we have important information here that will certainly
contribute to environmental licensing processes and to the evolution of this science in the country, corroborating the commitment
to promote eective actions for environmental preservation and sustainable mining.
Iuri Viana Brandi
Speleology and Technology Manager – Ferrous Value Chain
Rodrigo Dutra Amaral
Executive Manager of Environmental Licensing, Studies and Speleology – Ferrous Value Chains
presentation
CAVE FAUNA OF BRAZIL
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Giupponia chagasi
Rodrigo Lopes Ferreira
The publication of the work “Essai sur les problèmes biospéologiques”, in 1907, by the Romanian zoologist Emil G. Racovitza, is
considered an important milestone in the establishment of world Biospeleology (from the Greek bios = life; spelaion = cave; logos
= study). The term Subterranean Biology is currently most common and can be dened as the science that is dedicated to the
study of organisms that live in the subterranean environment, as well as the interactions between species and the relationships
between organisms and the environment. This is because the subterranean environment is complex and vast, with caves being our
main “window”, but only one of the environments that make up this mosaic of ecosystems. Thus, the subterranean environment
comprises the set of interconnected subsoil natural cavities, of variable sizes, formed by large networks of heterogeneous spaces
that can be lled by air or water.
Approximately 22 thousand caves have been cataloged in the Brazilian territory to date, with the potential for more than one
million, according to the most recent estimates. At least 591 species of fauna have been described from holotypes collected in these
environments, of which 283 are considered troglobites, that is, endemic species that have evolved and have specic adaptations to
survive in these unique ecosystems. However, there are still hundreds of species already known and waiting for scientic description,
in addition to an incalculable number of species to be discovered.
In this context, we have gathered in 22 chapters the most up-to-date information on the main groups of cave fauna in Brazil, in
addition to ve initial chapters that bring historical aspects, legislation, ecosystem dynamics and areas of science still little explored,
such as molecular biology and microbiology. Consolidating all the knowledge available in this work was an intricate task, in view
of the signicant increase in the number of professionals who are specializing and dedicating themselves to the study of our
subterranean biodiversity, which, consequently, has resulted in a signicant increase in publications on new species. The production
capacity of our researchers is indisputable, and they certainly have a notable prominence internationally. Therefore, it is essential to
invest nancial resources in the training of specialists and in the implementation and maintenance of the infrastructure of current
and new research centers, as well as in the maintenance of our valuable scientic collections. Only then will we be able to continue
producing high quality knowledge about our caves and other subterranean environments.
Thus, we hope that this work will be an important reference for all those who are dedicated to the study of cave fauna in Brazil,
whether in the context of academic research, environmental licensing or activities with speleology groups. We also hope that it will
provide all readers a beautiful journey into a small part of our wonderful caves and the life that coexists there.
Robson de Almeida Zampaulo
Speleology and Technology Management – Vale
Serra do Mar Environmental Studies Group – GESMAR
Speleological Observatory - OE
preface
CAVE FAUNA OF BRAZIL
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Caverna B05 - Doresópolis (MG)
André Bernardes Machado
CAVE FAUNA OF BRAZIL
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Zelurus zikani
Robson de Almeida Zampaulo
341
16
Hemiptera:
Heteroptera
HÉLCIO R. GIL-SANTANA
Laboratório de Diptera, Instituto Oswaldo Cruz, Fundação Oswaldo Cruz - Fiocruz
JADER OLIVEIRA
Laboratório de Entomologia em Saúde Pública, Departamento de Epidemiologia,
Faculdade de Saúde Pública, Universidade de São Paulo - USP
Laboratório de Parasitologia, Faculdade de Ciências Farmacêuticas
Universidade Estadual Paulista Julio de Mesquita Filho - UNESP
FELIPE F. F. MOREIRA
Laboratório de Biodiversidade Entomológica,
Instituto Oswaldo Cruz, Fundação Oswaldo Cruz - Fiocruz
ROBSON DE ALMEIDA ZAMPAULO
Gerência de Espeleologia e Tecnologia – Vale
Grupo de Estudos Ambientais da Serra do Mar – GESMAR
Observatório Espeleológico - OE
CAVE FAUNA OF BRAZIL
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General introduction
Heteroptera is a suborder of the order Hemiptera (Insecta)
whose representatives are popularly called “percevejos” or
“true bugs”. Among insects with incomplete metamorphosis
(hemimetabolous), it is the group with the greatest number of
species. It is estimated that around 37,000 to 45,000 species of
heteropterans have already been described (SCHAEFER; PANIZZI,
2000; SCHUH; WEIRAUCH, 2020) and approximately 25,000 have
yet to be discovered (SCHAEFER; PANIZZI, 2000). Approximately
5,450 species of Heteroptera have been cataloged for Brazil
(Catálogo Taxonômico da Fauna do Brasil – CTFB, available
online: http://fauna.jbrj.gov.br/fauna/).
The etymology of Heteroptera, derived from the Greek
terms hetero (= dierent) and pteron (= wing), alludes to
the structure of the forewings, the hemelytra, which have
a coriaceous anterior half and a membranous posterior
half (GRAZIA; FERNANDES, 2012). Although the forewing is
completely coriaceous or membranous in some species, most
representatives of the suborder have the hemelytra type
(COSTA LIMA, 1940). A prominent feature of the group shared
with other Hemiptera is the structure of the mouthparts in the
form of a segmented labium, which contains piercing-sucking
mouthparts. These may have been one of the factors behind
the evolutionary success of hemipterans in general (GRAZIA et
al., 2012). The presence of a connexivum, i.e., attened lateral
margins of the abdomen, standing out laterally to a greater or
lesser extent (COSTA LIMA, 1940), is another peculiarity of most
heteropterans that facilitates their recognition in relation to
other insects.
Metamorphosis from egg to adult occurs as in other
hemimetabolous insects. After emerging from the eggs, the
immatures (nymphs), pass, in general, through ve instars.
Nymphs, although structurally similar to adults, have neither
genitalia nor developed wings and can also dier from them
in color. The wings are only functional and fully developed in
adults, while in nymphs they develop while externally protected
by alar thecae, which appear in the third nymphal stage and
gradually increase in size until the last stage (COSTA LIMA, 1940;
GRAZIA; FERNANDES, 2012).
Heteropterans are terrestrial, semi-aquatic or aquatic
insects, ranging in length from 0.5 to 110 mm (GRAZIA;
FERNANDES, 2012). They are mainly phytophagous, with
several species of economic importance for causing damage
to cultivated plants, or predatory, basically of other arthropods,
while species of a few families are hematophagous, including
Triatominae (Reduviidae) (“barbeiros” or “kissing bugs”), vectors
of Chagas disease. Heteroptera is currently divided into seven
infraorders — Enicocephalomorpha, Dipsocoromorpha,
Gerromorpha, Nepomorpha, Leptodomorpha, Cimicomorpha
and Pentatomomorpha (GRAZIA; FERNANDES, 2012; PANIZZI;
GRAZIA, 2015; SCHUH; WEIRAUCH, 2020) — with 57 families
occurring in Brazil (CTFB).
While no other major group of insects is considered to utilize
such a wide variety of habitats as successfully as heteropterans
(SCHUH; WEIRAUCH, 2020), including caves (e.g., KŁYS; LIS, 2013;
PAPE, 2013; WYGODZINSKY, 1966), little has been recorded
about their biology and ecology in this environment in Brazil
(e.g., OLIVEIRA et al., 2008; PINTO-DA-ROCHA, 1995; RESENDE et
al., 2016; TRAJANO, 2000). One of the few exceptions is Ferreira
et al. (2016), which presents data on the diversity of Zelurus
Hahn, 1826, recorded in caves in Brazil, as well as an assessment
of the distribution of dierent species of this genus.
Here, we present a synthesis of current knowledge about
the main families of Heteroptera recorded in Brazilian caves,
including recent contributions. We hope that the knowledge
presented here will serve as a starting point for all those
interested in studying this group.
Main groups found in Brazilian caves
Terrestrial heteropterans
ANTHOCORIDAE
Anthocorids are very small (tiny), dark, attened, oval or
elongated insects. Although they are predators, they can feed
on pollen and other plant parts (CARPINTERO, 2015). Schuh and
CAVE FAUNA OF BRAZIL
343
Weirauch (2020) presented an overview of the family worldwide,
including a key to its seven tribes, whereas Carpintero (2002)
published the rst catalog dedicated to Neotropical species. Lattin
(2000) reviewed the species that occur in managed ecosystems.
Apparently, they are important in the biological control of pests
in agroecosystems (BORROR et al., 1989). There are about 100
species registered for Brazil (GRAZIA; FERNANDES, 2012). Pinto-
da-Rocha (1995) and Loyola et al. (2007) reported them in caves.
CYDNIDAE
Cydnids are called percevejos-cavadores or digger bugs.
They are ovoid in shape, small to medium in size and vary in
color from black to brown. They have a attened head and
legs with strong, long spines; the tibiae and tarsi are modied
in species with fossorial habits, which feed on roots. Some
cydnids live above ground, in vegetation, feeding on seeds or
plant tissues that fall (SCHAEFER, 2009; SCHWERTNER; NARDI,
2015; SCHUH; WEIRAUCH, 2020). Cydnids have a worldwide
distribution, being well represented in tropical and temperate
regions. The family includes over 750 described species in 93
genera and is divided into six subfamilies. Cydnids are poorly
known for the Neotropics. Froeschner (1981) presented a key
to the South American fauna of the subfamily Cydninae, the
largest and most diverse of the family, with 90 genera and 300
species. Only data on unidentied cydnids has been presented
in many articles about Brazilian caves, especially those found
associated with guano from fruit bats (GNASPINI NETTO 1989;
PELLEGRINI; FERREIRA, 2012; PINTO-DA-ROCHA, 1995; SIMÕES
et al., 2015; SOUZA-SILVA; FERREIRA, 2015; SOUZA et al., 2020).
In general, such species are abundant and found on the soil
surface near roots and guano, as well as under blocks. Dias et al.
(2018) sequenced the mitochondrial genome of a troglophilous
species found in iron formation caves in the Carajás region.
DIPSOCORIDAE
The family Dipsocoridae comprises the genera Alpagut
Kiyak, 1995 (Palearctic), Cryptostemma Herrich-Schäer,
1835 (worldwide distributed) and Pachycoleus Fieber, 1860
(Palearctic), with about 30 described species. According to
Štys (1995), Pachycoleus, which has often been considered a
subgenus of Cryptostemma, is represented in the Neotropics by
many undescribed species. Species are dicult to distinguish,
in part due to poorly-illustrated original descriptions that
lack details about important genital features (WEIRAUCH;
FERNANDES, 2015); the exceptionally well-illustrated description
by Wygodzinsky (1952) of C. haywardi Wygodzinsky, 1952 is
an exception. Given this context, Pinto da Rocha (1995) and
Cordeiro et al. (2014) reported the occurrence of the family in
caves but did not advance in species identication.
ENICOCEPHALIDAE
This family includes most of the described genera and species
of Enicocephalomorpha and most of the specimens deposited in
collections. The species are small (from 2 to 15mm in length) and,
in general, range in color from uniform yellow or brown to opaque
black (with exceptions) (ŠTYS, 1995). Enicocephalidae comprises
about 300 described species worldwide, which are classied into
47 genera and ve subfamilies (ŠTYS, 1995, 2002). Štys (2008)
indicated that at least 30 genera and hundreds of new species
remain undescribed so far (FERNANDES; WEIRAUCH, 2015). Štys
(1995) provided a key to the subfamilies of Enicocephalidae. The
family has already been reported in caves by Bernardi et al. (2012),
Pinto da Rocha (1995) and Silva et al. (2020), however, none of the
specimens were identied to the species level.
LYGAEIDAE
Lygaeids range 1.2–12 mm in length and vary in shape and
color, with patterns ranging from brown to black, in some cases
exhibiting aposematic coloration. Baranowski and Slater (2005)
published a key to the 10 subfamilies. A total of 120 species and
58 genera are registered for Brazil (GRAZIA; FERNANDES, 2012).
In a survey of six caves in Parque Estadual do Sumidouro (Pedro
Leopoldo, Minas Gerais), Iniesta et al. (2012) reported the family
with a sample of six individuals.
CAVE FAUNA OF BRAZIL
344
REDUVIIDAE
Reduviidae is the second largest family of Heteroptera,
with approximately 7,000 species described worldwide
(MALDONADO, 1990; SCHUH; WEIRAUCH, 2020; WEIRAUCH et
al., 2014). Of this total, about 11% (~800) are cataloged for Brazil
(CTFB). With the exception of hematophagous Triatominae,
reduviids are considered predators of other arthropods. They
are highly variable in size, body conformation and appendages,
from some species of Empicoris Wollf, 1811 (Emesinae), with
delicate appendages and reaching only 3 mm in total length,
to several species with robust bodies, reaching 40 mm in length
(SCHUH; WEIRAUCH, 2020; WYGODZINSKY, 1966). Diagnostic
features for Reduviidae include: eyes usually large; ocelli usually
present; labium usually curved and robust, but may be straight
and/or thin, with three visible segments (except Hammacerinae
with four); prosternum with stridulatory sulcus (stridulitrum);
hemelytral membrane usually with two or three elongated cells;
and spongy fossa present at the ventral end of the anterior and
middle tibiae in many taxa (GIL-SANTANA et al., 2015; SCHUH;
WEIRAUCH, 2020). Among these characteristics, the stridulatory
sulcus has been considered diagnostic of Reduviidae for a long
time (e.g., COSTA LIMA, 1940), being particularly useful for
non-specialists to separate them from heteropterans of other
families (HRG-S, pers. obs.).
Currently, 24 subfamilies of Reduviidae are recognized as
valid (SCHUH; WEIRAUCH, 2020), of which 20 are represented in
the Neotropical region (GIL-SANTANA et al., 2015) and 18 in Brazil
(CTFB). The most recent keys for subfamilies of Reduviidae are
those of Forero (2004), covering the Neotropical region; that of
Grazia and Fernandes (2012) for Brazil; and those by Weirauch et
al. (2014) and Schuh and Weirauch (2020), for those around the
world. The last two, because of the accompanying illustrations,
facilitate the understanding of the dichotomies. Consultation of
Gil-Santana et al. (2015a) allows access to identication keys for
the genera of several subfamilies or respective information, and
to summaries about the taxonomy and biology of Neotropical
reduviids. Maldonado (1990) is the general taxonomic catalog
of Reduviidae, which covers the entire world fauna, while CTFB
allows the consultation of updated faunal data for Brazil. Among
the reduviids found in caves in Brazil, the largest number of
species belong to the subfamilies Emesinae and Reduviinae.
Representatives of Ectrichodiinae, Triatominae and Saicinae
have also been recorded in Brazilian caves, as shown below:
Ectrichodiinae
Twenty-four genera are currently recognized for
Ectrichodiinae in the Americas, including more than a hundred
species (GIL-SANTANA et al., 2015, 2020a; FORTHMAN;
WEIRAUCH, 2017), among which 16 genera and just over 60
species have been registered for Brazil (CTFB). The genera found
in Brazil can be identied by the key presented by Gil-Santana
et al. (2015) or the most up-to-date version by Gil-Santana et
al. (2020a). Representatives of Ectrichodiinae can be easily
recognized by having a bid scutellum (e.g., Figure 1B) and a
completely or mostly glabrous integument (e.g., Figure 1A-
C), with many species showing aposematic, brightly colored
patterns (e.g., Figure 1A). The wings of females of many species
are reduced in size (e.g., Figure 1A), to varying degrees, or absent.
The scarce biological data suggest that species of Ectrichodiinae
are specialized predators of millipedes (Gil-Santana et al., 2015).
The genera with the greatest number of species, including the
most common ones, are Brontostoma Kirkaldy, 1904, Pothea
Amyot & Serville, 1843 and Rhiginia Stål, 1859 (CTFB; COSTA
LIMA, 1940a). Also belonging to these genera are the species
already found in caves, whose record was made by the junior
author (RAZ) and are presented here for the rst time: B.
trux (Stål, 1859) (Figure 1A; the female in this gure presents
coloration with a very pronounced variation, yet we consider it
to be of this species); another specimen of B. trux with a peculiar
coloration also collected by RAZ; P. furtadoi Gil-Santana & Costa,
2005 (Figure 1B) and R. atripennis (Walker, 1873). It should be
noted that all these species are primarily known from open
environments (e.g., GIL-SANTANA et al., 2013; GIL-SANTANA,
2014; HRG-S pers. obs.).
Emesinae
The elongated and relatively thin body and legs, together
with the appearance of the raptorial forelegs, which resemble
those of mantids (“louva-a-Deus” or “praying mantis”),
characteristics presented by most emesines, make it easy to
dierentiate them from other reduviids (Figure 1D-L). It is also
noteworthy that the emesines are one example of heteropterans
with an entirely membranous forewing, without the structure of
a hemelytron. Almost 90 species, included in 17 genera, have
been recorded in Brazil (CTFB). Many species of Emesinae are
occasionally or frequently found in caves (WYGODZINSKY,
1966), with the degree of adaptation and the frequency or
exclusivity to this habitat being variable. Mayemesa lapinhaensis
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Figure 1. A) Brontostoma cf. trux (Stål, 1859), female; B) Pothea furtadoi Gil-Santana & Costa, 2005, female; C) nymph of unidentied species of Ectrichodiinae;
D-F) Phasmatocoris catarinae Gil-Santana, 2015, D) adult, E) couple in copula, F) nymph; G-I) Phasmatocoris galvaoi Gil-Santana, 2015, G) adult, H-I) adult performing
predation, I – detail; J-K) Ploiaria sp., adult at rest, J) dorsal view, K) dorsolateral view; L) Emesinae sp., lateral view. Photos: (Robson de A. Zampaulo).
A
D
G
J
B
E
H
K
C
F
I
L
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(Wygodzinsky, 1950), for example, originally known only from
caves, (WYGODZINSKY, 1966), was later found both in completely
dissociated environments (GIL-SANTANA et al., 1999) and again
in caves (HRG-S, personal observation). An updated summary
of the world literature, including a discussion of ecological and
biological knowledge of emesines found or living in caves, was
presented by Pape (2013). Species of Dohrnemesa Wygodzinsky,
1945, Phasmatocoris Breddin, 1904 (Figure 1D-I) and Ploiaria
Scopoli, 1786 (Figure 1J-K) are frequent in Brazilian caves,
with some species of the rst two genera being described
and, until now, known only from this environment, such as
Ph. catarinae Gil-Santana, 2015 and Ph. galvaoi Gil-Santana,
2015 (e.g. GIL-SANTANA et al., 2007; GIL-SANTANA 2015; GIL-
SANTANA; FERREIRA, 2016, 2017), while there are at least two
common, and possibly new, species of Ploiaria in caves awaiting
identication (HRG-S, personal observation) (e.g., Figure 1J-K).
In general, immatures of dierent stages of emesines can be
found associated with litter on the oor of caves in entrance and
dysphotic zones, where they probably nd food in abundance.
In turn, adults or last instars are generally more frequently
observed along the walls and ceilings of caves, especially in
predominantly dry areas. These reduviids are often recorded
feeding on dipterans (e.g., Psychodidae), coleopteran larvae,
spiders and immature crickets, among other small arthropods.
Emesines can also be found associated with spiders of the
genera Loxosceles Heineken & Lowe, 1832 and Mesabolivar
González-Sponga, 1998, and may exhibit kleptoparasitic
behavior (RESENDE et al., 2016).
Reduviinae
Reduviinae includes nine genera that occur in Brazil, with
about a hundred species (CTFB; GIL-SANTANA et al., 2015),
of which Zelurus is the most diverse, with almost 90 species
recorded in the country. Ferreira et al. (2016) presented a
synthesis of the numerous taxonomic references on species
of Zelurus, which is one of the most ubiquitous cave taxa in
Brazil. These authors presented a list of ten species of Zelurus
recorded in Brazilian caves, among which Z. diasi (Costa Lima,
1940) (Figures 2A-B), Z. sipolisi (Fallou, 1889) (Figure 2C), Z.
zikani (Costa Lima 1940) (Figures 2D-E) and Z. festivus (Stål,
1859) (Figure 2F) are illustrated in this chapter. We also add
the occurrence of Z. circumcinctus (Hahn, 1825) and Z. pintoi
(Costa Lima, 1940), updating the number of species recorded
in Brazilian caves to 12.
It is worth noting that the known nymphs of this genus
have cryptic coloration in relation to the substrate, often being
covered by soil particles or even other environmental structures
(e.g., lichens) (Figures 9A-D), which increases this camouage.
On the other hand, the adults seem to mimic wasps of the
genus Pepsis Fabricius, 1804 (Hymenoptera: Pompilidae), both
due to the color and shape of the body (HOGUE, 1993) and the
movement they perform when ying (COSTA LIMA, 1940b). They
are generalist predators, being nocturnal and quite aggressive
(GROSSI et al., 2012; SCHUH; WEIRAUCH, 2020).
In general, nymphs are found in large numbers in the entrance
zone of predominantly dry caves, while adults are relatively rare
in these environments. One of the plausible hypotheses is that
caves function as sites for reproduction and the development of
immature forms, since selective pressures dierent from those
of the epigean environment (e.g., lower predator abundance,
lack of light and specic microclimatic conditions) could favor
the reproductive success of countless species of this genus.
Adult forms of Zelurus sp. were recorded as predators of
harvestmen (Goniosoma sp., Arachnida, Opiliones, Gonyleptidae)
in almost all caves surveyed by Machado et al. (2003), while
Serracutisoma spelaeum (Mello-Leitão, 1933; cited as Goniosoma
spelaeum) was observed as prey of Zelurus travassosi (Costa
Lima, 1940). This last species was considered the most important
predator of the trogloxene S. spelaeum in the state of São Paulo
and of the troglophile Daguerreia inermis Soares & Soares, 1947
(Opiliones, Gonyleptidae) in the state of Paraná (MACHADO et
al., 2003). The immature forms, on the other hand, have a large
repertoire of prey, and can feed on crickets, coleopteran larvae,
dipterans and spiders, among others.
Saicinae
There are seven genera and 22 species of Saicinae in Brazil
(CTFB; GIL-SANTANA et al., 2015, 2020b). Little is known about
the biology of its representatives, which are usually collected
by light attraction (SCHUH; WEIRAUCH, 2020). Biological
data for the group in the Americas were summarized by Gil-
Santana et al. (2010), while Gil-Santana et al., (2020b), based
on nymphs and adults found in a single cave in the state of
Tocantins, described a new genus and species of Saicinae,
Quasitagalis Gil-Santana, Oliveira & Zampaulo, 2020, and Q.
afonsoi Gil-Santana, Oliveira & Zampaulo, 2020, recording for
the first time a species of this subfamily as a cave dweller in
the Americas.
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Triatominae
Triatomines stand out among reduviids for feeding on
vertebrate blood in all their life stages and for being vectors of
the protozoan Trypanosoma cruzi (Chagas, 1909), the etiological
agent of Chagas disease, a pathology that is still of signicant
public health importance in Latin America (MONTEIRO et al.,
2018). Several species of Triatominae have been collected or
found in caves (LENT; WYGODZINSKY, 1979). The etymology of
Cavernicola Barber, 1937, reects the specialization for living
in caves of the species for which the genus was described, C.
pilosa Barber, 1937, which has a wide geographical distribution
in South America and is found mainly in caves inhabited by bats
(OLIVEIRA et al., 2008; GALVÃO; GONÇALVES, 2014; GALVÃO,
2020). Panstrongylus geniculatus (Latreille, 1811) (Figures 2J-L)
is one of the relatively common triatomine species and has the
widest geographical distribution in South and Central America,
where it is found in a variety of habitats, including caves with
bat colonies (LENT; WYGODZINSKY, 1979) or caves in general
(GALVÃO; GONÇALVES, 2014). Another species that occurs
in Brazil and has already been recorded in caves is Eratyrus
mucronatus Stål, 1859 (GALVÃO; GONÇALVES, 2014; GALVÃO,
2020).
Reduviids of other subfamilies
In addition to representatives of the subfamilies listed above,
which are the most frequent, we have recorded the occurrence
of other subfamilies of Reduviidae in caves (HRG-S personal
observation), such as Apiomerus sp. (nymph) (Harpactorinae;
Apiomerini) and Vescia sp. (Vesciinae) (Figure 2 I).
THYREOCORIDAE
Members of Thyreocoridae (= Corimelaenidae) are known
as black bugs. The family is divided into Thyreocorinae
and Corimelaeninae, the latter being represented in the
Neotropical Region. They are relatively small and dark in
color, with an oval-elongated body and a large scutellum
(GRAZIA; FERNANDES, 2012). Galgupha Amyot & Serville, 1843
is the most diverse Neotropical genus, with 155 species in 15
subgenera. About 160 species of the family have been recorded
in the Neotropics, but their biology is poorly understood. Black
bugs are phytophagous, feeding on developing or mature
owers and fruits (MATESCO; GRAZIA, 2015). Despite occurring
in many crops, they are considered minor pests in orchards
and are not mentioned among heteropterans of economic
importance (SCHAEFER; PANIZZI, 2000). Simões et al. (2015)
found a species in caves in the state of Minas Gerais, while
studying the how physical attributes of caves inuence the
structure and composition of cave invertebrate communities.
TINGIDAE
Tingids, known as “lace bugs”, are delicate and small
insects with a growth of the pronotum and forewings having
a lacy appearance. The family is divided into three subfamilies
(Tinginae, Cantacaderinae and Vianaidinae), all represented
in the Neotropical Region (SCHUH et al., 2006, GUIDOTI et
al., 2015). There are currently about 300 genera and 2,500
species in this family, with 70 genera and 600 species in the
Neotropics. Few species have had their biology studied in this
region, such as Corythaica cyathicollis (Costa, 1864) (KOGAN,
1960) and Leptopharsa heveae Drake & Poor, 1935 (Cividanes
et al., 2004), both of economic importance. There is only one
record for caves, namely fth-instar nymphs and males of
Thaumamannia vanderdrifti Van Doesburg, 1977, in caves in
the iron formation of the Floresta Nacional de Carajás, in the
municipality of Parauapebas, state of Pará (GUIDOTI et al.,
2014). Although there are other records for the family in the
state of Minas Gerais (unpublished data), such specimens are
considered relatively rare in Brazilian caves in general.
Aquatic and semiaquatic heteropterans
Aquatic and semiaquatic true bugs are much less diverse
than their terrestrial relatives but correspond to almost 5,000
species distributed in the infraorders Leptopodomorpha (ca.
380), Gerromorpha (ca. 2,100) and Nepomorpha (ca. 2,300)
(POLHEMUS; POLHEMUS, 2008). The leptopodomorphan
fauna of Brazil is poorly known, with scarce records of only
10 species of the family Saldidae (CTFB), none of which were
based on specimens collected in caves. Records in these
environments in Brazil are also relatively rare for Gerromorpha
and Nepomorpha, being more abundant in the case of the
former. Pinto-da-Rocha (1995) summarized records from caves
in Brazil for the families Gerridae, Veliidae (Gerromorpha),
Belostomatidae and Naucoridae (Nepomorpha), but none
identied to species.
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Figure 2. A-B) Zelurus diasi (Costa Lima, 1940), adult, A) dorsal view, B) detail, lateral view; C) Zelurus sipolisi (Fallou, 1889), adult, frontal view; D-E) Zelurus zikani
(Costa Lima, 1940), D) adult, dorsal view, E) couple in copula; F) Zelurus festivus (Stål, 1859); G) Zelurus cf. avipennis (Mayr, 1865), dorsolateral view; H) Zelurus sp., dorsal
view; I) Vescia sp., dorsal view; J-L) Panstrongylus geniculatus (Latreille, 1811), adults, J) dorsal view, K) lateral view, L) nymph, partially covered with soil detritus. Photos:
(Robson de A. Zampaulo).
A
D
G
J
B
E
H
K
C
F
I
L
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GERROMORPHA
Gerromorphans are semi-aquatic true bugs, most of which
spend almost their entire active life on the surface or shores
of freshwater bodies. Some species, however, are secondarily
terrestrial or occupy coastal marine habitats, or even the
open ocean (MOREIRA 2015). The infraorder is divided into
eight families, of which Gerridae, Hebridae, Hydrometridae,
Mesoveliidae and Veliidae are recorded from Brazil. A total of
244 species have been recorded from the country thus far, of
which 15 have been reported in caves (CTFB).
Brachymetra albinervus (Amyot & Serville, 1843) (Gerridae:
Charmatometrinae) (Figure 3A), a widely distributed species in
the Neotropics, was recorded in Caverna dos Troncos, Parque
Nacional Cavernas do Peruaçu, state of Minas Gerais (MONTE;
BICHUETTE, 2020), and in Duto do Quebó, municipality of Nobres,
state of Mato Grosso (MAGALHÃES et al., 2021). The species
is very common and abundant, and generally lives in sandy-
bottomed streams, occasionally being found in and around
caves. Cylindrostethus palmaris Drake & Harris, 1934 (Gerridae:
Cylindrostethinae) (Figures 3B-C) has habitat preferences similar
to those of B. albinervus and was recently recorded in Gruta
do Catão, municipality of São Desidério, state of Bahia, and in
the previously mentioned Duto do Quebó (MAGALHÃES et al.,
2021). It is the most common species of Cylindrostethus in South
America and the only one that has winged adults, which may be
related to its wide distribution
The family Hydrometridae includes two monotypic genera
that were recently described based on troglobitic specimens.
Both were tentatively allocated to the subfamily Hydromerinae
based on the apical articulation between antennomeres I and
II. Cephalometra pallida Polhemus & Ferreira, 2018 (Figure
7D) occurs in Gruta da Natividade, state of Tocantins, while
Spelaeometra gruta Polhemus & Ferreira, 2018 (Figures 7B-C)
was collected in four caves of the Parque Nacional Cavernas do
Peruaçu (POLHEMUS; FERREIRA, 2018).
Mesovelia amoena Uhler, 1894 (Mesoveliidae: Mesoveliinae)
(Figure 4A) occurs throughout most of the Americas and has
even been introduced to Hawaii (DAMGAARD et al., 2012). The
species was recorded once in caves in Brazil, based on material
from São Desidério, as well as the aforementioned C. palmaris
(CORDEIRO; MOREIRA, 2015).
The greatest number of species of Gerromorpha recorded
in caves in Brazil is in the family Veliidae. Microvelia ayacuchana
Drake & Maldonado, 1952 (Veliidae: Microveliinae) (Figure 4B)
was recorded based on material collected in pools in a cave in
the municipality of Canaã dos Carajás, state of Pará (MOREIRA;
CAMPOS, 2012). Individuals of this species generally live on
the banks in pools beside waterfalls, running to land when
disturbed. Microvelia ioana Drake & Hottes, 1952 (Figures 6E-F),
endemic to the Atlantic Forest, was recorded in Gruta do Saco
Grande, Florianópolis, state of Santa Catarina (MAGALHÃES et
al., 2021).
Rhagovelia kararao Floriano & Moreira, 2015 (Veliidae:
Rhagoveliinae) (Figures 4C-D) was described based on
material collected in two caves, one in Altamira and another
in Vitória do Xingu, state of Pará, and has not been recorded
again. Specimens of this genus are usually collected in
open lotic environments with moderate to strong currents,
but the above species was found in slow flowing waters in
sandstone caves (FLORIANO; MOREIRA 2015). Rhagovelia
robusta Gould, 1931 (Figures 3F-G) and R. whitei (Breddin,
1898) (Figures 6A-B), more widely distributed in south-
central South America, were recorded in caves in the states
of Mato Grosso and Minas Gerais (CORDEIRO; MOREIRA,
2015; MAGALHÃES et al., 2021; TAYLOR; FERREIRA, 2012),
but they are much more common in open environments.
Rhagovelia henryi Polhemus, 1997 (Figures 3D-E), in turn,
is found exclusively in the Atlantic Forest and, recently,
was recorded for the first time in caves, based on material
collected in the municipality of Iporanga, state of São Paulo
(MAGALHÃES et al., 2021).
Two species of the genus Paravelia Breddin, 1898 (Veliidae:
Veliinae) have, thus far, only been found in caves. Paravelia
cunhai Rodrigues & Moreira, 2016 (Figures 5A-B) was collected
in this type of environment in the municipalities of Canaã dos
Carajás and Vitória do Xingu, Pará (Rodrigues & Moreira 2016a),
while P. digitata Rodrigues & Moreira, 2016 (Figures 6C -D)
was collected in a series of caves in Bahia and Rio Grande do
Norte (MAGALHÃES et al., 2021; RODRIGUES; MOREIRA, 2016b).
The morphology of these animals, however, does not indicate
restriction to subterranean environments. Paravelia polhemusi
Rodrigues, Moreira, Nieser, Chen & Melo, 2014 (Figure 5C) and
P. splendoris (Drake & Harris, 1933) (Figure 5D), in turn, have
already been recorded both in caves and in open environments
in Brazil (DIAS-SILVA et al., 2013; RODRIGUES et al., 2014;
RODRIGUES; MOREIRA, 2016b; RODRIGUES; ÁLVAREZ-ARANGO,
2019; FRANCO et al., 2020).
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Figure 3. A) Brachymetra albinervus, apterous female, dorsal view; B) Cylindrostethus palmaris, macropterous male dorsal view; C) Cylindrostethus
palmaris, macropterous male, abdomen apex, ventral view; D) Rhagovelia henryi, apterous male, dorsal view; E) Rhagovelia henryi, apterous
male, ventral view; F) Rhagovelia robusta, apterous male, dorsal view; G) Rhagovelia robusta, apterous male, ventral view. Originally published in
Magalhães et al., (2021).
A
B C
D
E
FG
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Figure 4. A) Mesovelia amoena, apterous female, dorsal view;
B) Microvelia ayacuchana, apterous male, dorsal view; C) Rhagovelia
kararao, apterous male, dorsal view; D) Rhagovelia kararao, apterous
male, ventral view. Scale bars: 1 mm and only applies to gures A and
B. Originally published in Rodrigues et al. (2021) (A, B) and Floriano &
Moreira (2015) (C, D).
A
C
B
D
NEPOMORPHA
Nepomorphans, or truly aquatic true bugs, live submerged
in freshwater bodies most of their lives, with representatives of
two families being secondarily aerial (BARBOSA; RODRIGUES,
2015). Depending on the author, the infraorder is divided into 11
or 13 families, of which Belostomatidae, Corixidae (sensu lato),
Gelastocoridae, Helotrephidae, Micronectidae, Naucoridae,
Nepidae, Notonectidae, Ochteridae, Pleidae and Potamocoridae
occur in Brazil. In all, 310 species have been recorded in the
country (CTFB), but only three were reported based on material
collected in caves.
Nerthra terrestris (Kevan, 1948) (Gelastocoridae: Nerthrinae)
can be found on the banks of water bodies or away from them
(NIESER, 1975). In Brazil, the species was recorded once at Abrigo
da Gravura, municipality of Altamira, state of Pará (CORDEIRO;
MOREIRA, 2015). Maculambrysus stali (La Rivers, 1962) and
Australambrysus teutonius (La Rivers, 1951) (Naucoridae:
Ambrysinae), in turn, are commonly collected in stony streams in
open areas, with moderate to strong currents, but have already
been recorded in caves in Pará and Minas Gerais, respectively
(CORDEIRO; MOREIRA, 2015; TAYLOR; FERREIRA, 2012).
Main troglomorphisms
Among the species known from Brazil thus far (Figure 8),
only two — Cephalometra pallida and Spelaeometra gruta
(Gerromorpha: Hydrometridae) — present morphological
adaptations that indicate some degree of restriction to
subterranean environments, being considered eectively
troglobites. Specializations, such as reduction of wings,
depigmentation, reduction of ocular structures and elongation
of body structures, were observed in these species.
Pape (2013) discusses that advanced troglomorphisms are
apparently uncommon for Emesinae (Reduviidae), and so far only
two troglobitic species of the genus Collartida Villiers, 1949 (C.
anophthalma Espanol & Ribes, 1983 and C. tanausu Ribes, Oromi &
Ribes, 1998), described from lava tubes of the Canary Islands, and
a species of the genus Nesidiolestes Kirkaldy, 1902 (N. ana Gagné
& Howarth, 1974), described from lava tube caves of Hawaii, have
been considered troglobites. However, in this same work, the
author mentions that, of the 44 species of Emesinae with records
in caves, 24 are known only for this type of environment, which
may indicate that some eectively subterranean species may
not present the classic troglomorphisms for the majority of cave
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Figure 5. A) Paravelia cunhai, macropterous male, dorsal view; B) Paravelia cunhai, macropterous male, ventral view; C)
Paravelia splendoris, brachypterous male, dorsal view; D) Paravelia polhemusi, macropterous male, dorsal view. Scale bar: 1 mm
and only applies to gures A and B. Originally published in Rodrigues & Moreira (2016a) (A, B), Rodrigues et al. (2014) (C) and
Rodrigues Álvarez-Arango (2019) (D).
A
C
B
D
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Figure 6. A) Rhagovelia whitei, apterous male, dorsal view; B) Rhagovelia whitei, apterous male, ventral view; C) Paravelia digitata, macropterous
male, dorsal view; D) Paravelia digitata, macropterous male, lateral view; E) Microvelia ioana, apterous female, dorsal view; F) Microvelia ioana,
apterous male, dorsal view. Originally published in Magalhães et al., (2021).
groups, as observed by Wygodzinsky, (1966), Gil-Santana et al.
(2007), Pape (2013) and Chłond et al. (2018). This condition may
indicate an underestimation of the number of troglobitic species
known for Brazil.
Main methods of collection
and conservation
Collection methods for terrestrial heteropterans in epigean
environments include the use of entomological umbrellas, light
traps, baited traps, pitfall traps and extractors (Berlese, Berlese-
Tüllgren and Winkler), in addition to manual collection on
various substrates such as vegetation, soil, logs or under blocks,
for example. As for other invertebrates, the most ecient and
commonly used collection method in caves is the active search.
Among the most ubiquitous groups, the reduviids of the genus
Zelurus are often found associated with oor sediment during
the young phase where, most of the time, they use this substrate
to cover their bodies as a camouage strategy (Figures 9A-D).
Adult specimens are much less frequent (compared to
immature forms), being found associated with the walls and
ceilings of caves. For the other groups, young emesines are
recurrent among litter, but they can also be found associated
with bare oors and cave walls. Specimens of the families
Enicocephalidae and Tingidae are more frequent near the soil,
litter and roots and under blocks. Specimens of the family
Cydnidae are often found associated with guano deposits,
especially those produced by fruit and hematophagous bats.
The other families can be found in dierent substrates, especially
in the entrance and penumbra zones.
A
D
B
E
C
F
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Figure 7. A) Live specimen of Spelaeometra sp.; B) Spelaeometra gruta, apterous female, dorsal view ; C) Spelaeometra gruta, apterous male, dorsal view; D) Cephalometra
pallida, apterous male, lateral view. Photos: A (Rodrigo L. Ferreira); B-C (Originally published in Polhemus & Ferreira, 2018).
A
D
B C
CAVE FAUNA OF BRAZIL
355
Figure 8. Map with type localities of troglobitic hemipteran species (Heteroptera) described for Brazil.
CAVE FAUNA OF BRAZIL
356
In relation to aquatic (Nepomorpha) and semi-aquatic
(Gerromorpha) heteropterans (NIESER; MELO, 1997), in practice,
the vast majority of accessible caves are dry, as they are
currently in a senile condition, and therefore these groups are
poorly represented in caves in general. However, these groups
can be found in caves with the presence of upwellings, rivers,
lakes (associated with the base level or not), travertine dams or
any other type of water deposit (including puddles), whether
perennial or intermittent. Individuals of Gerromorpha live
on the surface lm or banks of water bodies (SCHUH; SLATER,
1995), which facilitates their visual identication and sampling.
Representatives of Nepomorpha, on the other hand, tend to
have closer relationships with bottom substrates because they
are benthic (HUNGERFORD, 1948), and so more specic methods
must be employed.
In general, there are few works that have evaluated the
diversity of aquatic invertebrates in caves in Brazil and that have
used particular methods for sampling this faunal compartment,
making these groups completely underestimated in the country
(e.g., PELLEGRINI et al., 2020; PELLEGRINI et al., 2018; ZEPON et
al., 2019). Sampling in small puddles of water and travertine
dams can be easily carried out with the aid of pipettes and small
manual nets. However, in larger drainages or lakes, sampling
should include the dierent submerged substrates, such as
roots, sediments, gravel, rocks and organic matter deposited at
the bottom of these water bodies. For this, sieves, nets, Surber-
type collectors or “D” nets (rapiché), all with ne mesh, are
recommended. In more specic studies, the removal of samples
Figure 9. A-D. Nymphs of Zelurus sp.; A) Specimen without camouage; B) Specimen camouaged with sediment; C) Specimen camouaged
with lichens; D) Specimen camouaged with sediment of an iron ore cave. Photos: (Robson de A. Zampaulo).
A
C
B
D
CAVE FAUNA OF BRAZIL
357
of sediment and organic matter for processing in loco with the
aid of plastic trays or later in the laboratory is recommended. A
detailed description of the processing of these samples in the
laboratory can be obtained in Silveira et al. (2004). Conservation
of collected material should use 70% alcohol for general
taxonomic studies and absolute alcohol and storage at -20ºC
to avoid the degradation of the genetic material for molecular
studies.
Main Brazilian collections
For Heteroptera in general, and particularly the families
Miridae and Reduviidae, the collection with the largest
number of types and specimens in Brazil was that of the Museu
Nacional, Universidade Federal do Rio de Janeiro (UFRJ), until 02
September 2018 when it was completely destroyed in the great
re that hit this institution (ESCOBAR, 2018). Next, for Reduviidae,
it is worth mentioning the collections on the campus of Instituto
Oswaldo Cruz in Manguinhos, Rio de Janeiro. The Coleção de
Triatomíneos of Instituto Oswaldo Cruz (CTIOC) is not only in the
largest collection of triatomines in the world, but also includes
numerous types and specimens of predatory Reduviidae,
particularly species of Zelurus referred to in numerous articles
by Herman Lent and Pedro (Petr) Wolfgang Wygodzinsky on this
group (MALDONADO, 1990); while the Coleção Entomológica of
the same institute (CEIOC), highlights the Reduviidae material
studied by Ângelo Moreira da Costa Lima, separated into its
own collection (Coleção Costa Lima), which includes specimens
and types, recently cataloged by Almeida et al. (2014) and
Rodrigues et al. (2017), and other specimens referred to in their
articles, such as his review of Zelurus (as Spiniger) (COSTA LIMA,
1940b). Other collections with fewer type specimens, such as
the collection of the Museu de Zoologia da Universidade de São
Paulo (MZUSP; recent catalog by CARRENHO et al., 2020), or with
less numerous collections but including types or specimens
from caves, described more recently, such as the Coleção de
Invertebrados Subterrâneos of the Universidade Federal de
Lavras (ISLA/UFLA) Minas Gerais, also deserve to be mentioned.
For aquatic heteropterans from caves, most of the material
currently collected is deposited at the Museu de Zoologia
da Universidade de São Paulo (MZUSP), the Coleção de
Invertebrados Subterrâneos of the Centros de Estudos em
Biologia Subterrâneas of the Universidade Federal de Lavras
(CEBS /UFLA), the Coleção Entomológica of the Instituto Oswaldo
Cruz (IOC) and the Laboratório de Estudos Subterrâneos of the
Universidade Federal de São Carlos (LES/UFSCar).
Acknowledgements
We would like to thank everyone who contributed to the
collection of specimens of cave-dwelling heteropterans and
information that has allowed a better understanding of the
presence of these animals in these unique ecosystems. Jader
Oliveira and Felipe F. F. Moreira would like to thank the Fundação
de Amparo of the Pesquisa do Estado de São Paulo (FAPESP),
the Fundação Carlos Chagas Filho de Amparo of the Pesquisa
do Estado do Rio de Janeiro (processes #E-26/201.362/2021
and #E -26/203,250/2021) and the Conselho Nacional de
Desenvolvimento Cientíco e Tecnológico (CNPQ; process
#301942/2019-6) for funding received.
ALMEIDA, M. D.; RODRIGUES, J. M. S.; MOREIRA, F. F. F.; JURBERG, J.
Lista dos exemplares tipo de Heteroptera (Insecta: Hemiptera)
da Coleção Costa Lima, Instituto Oswaldo Cruz, EntomoBrasilis,
Rio de Janeiro, Brasil, v. 7, n. 2, p. 151-158, 2014.
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