Armadillo burrows: a meeting point for biodiversity in the Pantanal Wetland

Abstract

Dasypus novemcinctus and Euphractus sexcinctus are common armadillo species in the Brazilian Pantanal. Due to their
digging habits, they alter the surroundings providing environmental alternatives that can be used by other organisms. We
therefore aimed to identify the animal species that utilize the burrows of these armadillos in the Pantanal Wetland and
to investigate how they use them. This study was carried out by camera trapping, targeting 27 armadillo burrows in the
Pantanal. We obtained a total of 594 records of 34 species of mammals, birds, and reptiles. The way wildlife utilized
the burrows was categorized into pass, approach, entrance, and interior. Records showed that Carnivora was the richest
order recorded, mainly Nasua nasua, Cerdocyon thous, and Leopardus pardalis. We found that Thrichomys fosteri and
Clyomys laticeps were the only species using the burrows as shelter or refuges. Our data show that D. novemcinctus and
E. sexcinctus burrows play an important ecological role in the Pantanal wetland, acting as meeting points for several species
of local fauna.

Full text

SHORT COMMUNICATION
Mammalian Biology
https://doi.org/10.1007/s42991-024-00466-8
Introduction
Physical modications of natural environments by verte-
brates and invertebrates can benet dierent species (Jones
et al. 1994). Knowing which and how some species favor
others provide arguments for biodiversity conservation that
can inuence policy decisions. The burrows of fossorial
animals play important roles in several ecological processes
by altering vegetation, promoting sediment movement and
increasing nutrient availability, thus aecting forest dynam-
ics and creating new environments for other organisms
(Sawyer et al. 2012; Fleming et al. 2014; Rodrigues et al.
2020). Due to these characteristics, they can be considered
ecosystem engineers (Jones et al. 1994; Reichman and
Seabloom 2002; Desbiez and Kluyber 2013).
Armadillos are semi-fossorial mammals that are widely
distributed in Americas (Redford and Wetzel 1985). Of the
25 species of known armadillos (Abreu et al. 2023; Barthe
et al. 2024), seven occur in the Brazilian Pantanal: Cabas-
sous tatouay, Cabassous squamicaudis, Dasypus novem-
cinctus, Dasypus septemcinctus, Euphractus sexcinctus,
Priodontes maximus, and Tolypeutes matacus (Alho et al.
Handling editor: Heiko G. Rödel.
Heitor Miraglia Herrera
herrera@ucdb.br
1 Programa de Pós-Graduação em Ciências Ambientais e
Sustentabilidade Agropecuária, Universidade Católica Dom
Bosco, Campo Grande, Mato Grosso do Sul, Brazil
2 Programa de Pós-Graduação em Ecologia e Conservação,
Universidade Federal do Mato Grosso do Sul, Campo
Grande, Mato Grosso do Sul, Brazil
3 Laboratório de Entomologia em Saúde Pública,
Departamento de Epidemiologia, Faculdade de Saúde
Pública, Universidade de São Paulo, São Paulo, São Paulo,
Brazil
4 Departamento de Biologia & CESAM, Universidade de
Aveiro, Campus de Santiago, Aveiro, Portugal
5 Quinta de Prados, Departamento de Biologia, CoLAB
ForestWISE – Laboratório Colaborativo para a Gestão
Integrada da Floresta e do Fogo, Campus Universitário de
Santiago Aveiro, Santiago Aveiro, Portugal
6 Instituto Homem Pantaneiro, Corumbá, Mato Grosso do Sul,
Brazil
Abstract
Dasypus novemcinctus and Euphractus sexcinctus are common armadillo species in the Brazilian Pantanal. Due to their
digging habits, they alter the surroundings providing environmental alternatives that can be used by other organisms. We
therefore aimed to identify the animal species that utilize the burrows of these armadillos in the Pantanal Wetland and
to investigate how they use them. This study was carried out by camera trapping, targeting 27 armadillo burrows in the
Pantanal. We obtained a total of 594 records of 34 species of mammals, birds, and reptiles. The way wildlife utilized
the burrows was categorized into pass, approach, entrance, and interior. Records showed that Carnivora was the richest
order recorded, mainly Nasua nasua, Cerdocyon thous, and Leopardus pardalis. We found that Thrichomys fosteri and
Clyomys laticeps were the only species using the burrows as shelter or refuges. Our data show that D. novemcinctus and
E. sexcinctus burrows play an important ecological role in the Pantanal wetland, acting as meeting points for several spe-
cies of local fauna.
Keywords Dasypus novemcinctus · Euphractus sexcinctus · Camera traps · Floodplain · Burrow utilization
Received: 19 August 2024 / Accepted: 17 October 2024
© The Author(s) under exclusive licence to Deutsche Gesellschaft für Säugetierkunde 2024
Armadillo burrows: a meeting point for biodiversity in the Pantanal
Wetland
Filipe MartinsSantos1· Nayara YoshieSano1,2 · William Oliveirade Assis1· Leonardo FrançaNascimento1·
Jaderde Oliveira3· CarlosFonseca4,5 · Amadeu M.V.M.Soares4· Heitor MiragliaHerrera1,2 ·
Grasiela Edithde OliveiraPorrio6
1 3

F. M. Santos et al.
2011; Feijó and Anacleto 2021). The most common species
in the Pantanal biome are D. novemcinctus and E. sexcinctus
(Maccarini et al. 2015). While D. novemcinctus is usually
associated with densely forested habitat (McDonough et al.
2000; Goulart et al. 2009), E. sexcinctus inhabits open veg-
etation and forest edges (Maccarini et al. 2015).
Many animals have been reported using armadillo bur-
rows, including insects, lizards, snakes, frogs, and mammals
(Taber 1945; Clark 1951; Vitt and Caldwell 1993; Dias-
Lima et al. 2003; Platt et al. 2004; Desbiez and Kluyber
2013; Aya-Cuero et al. 2017; Rodrigues et al. 2020). Des-
biez and Kluyber (2013) proposed that the frequency and
time of interaction of 24 vertebrate species with the burrows
of the giant armadillo, P. maximus, occur at dierent scales
in the Pantanal biome. However, there are no studies that
evaluate the uses of the most abundant armadillo burrows
and the local fauna. Therefore, our goal was to verify which
organisms are utilizing the burrows of D. novemcinctus and
E. sexcinctus, recording species richness as well as the dif-
ferent types of uses.
Materials and methods
Data collection
The study was carried out in an area of 10 km2 of a Pri-
vate Natural Heritage Reserve, located in the Pantanal sub-
region of Nhecolândia (19º 03’ 54.62” S; 56º 45’ 53.05”
W). The eld work was carried out from November 2015
to January 2016; and from June to August 2016. We used
27 camera traps (Bushnell®, Overland Park, Kansas, USA),
in ve transects (~ 250 m each one) separated by 500 m
from each other at the study site. Two or three cameras were
installed along each transect, about 50–70 m from each
other, 10–15 cm above the ground in front of armadillo bur-
rows with evidence of activity such as recently removed soil
and fresh armadillo footprints. Camera traps operated 24 h/
day for 71 days with ve seconds interval between pictures,
with a total eort of 927 camera-days. We did not use baits
to attract animals, and equipment was checked in 20-day
intervals to change batteries and to download pictures. Bur-
rows of armadillos were identied to be from D. novem-
cinctus and E. sexcinctus based on their morphometric
characteristics, but we could not determine which burrow
was from each species mainly due to the characteristics of
the region’s sandy soil (Rodrigues et al. 2020).
After the camera trapping sampling, we searched for
invertebrates and other taxa inside the monitored burrows
through excavations using shovels and hoes. Soil samples
were sieved with two dierent meshes (4 and 1 mm, respec-
tively) for a detailed search for invertebrates, which were
later identied by external morphological characters by spe-
cialists. Reptiles and amphibians were identied in the eld.
Data analysis
The data obtained through camera trapping were screened,
and pictures of the same species at the same camera trap-
ping station within a period of 1 h were excluded to guar-
antee independence between records (Silveira et al. 2003).
A species accumulation curve was obtained by means of
1,000 randomizations of dierent sample sizes using the
package vegan (Oksanen et al. 2018) in R 4.2.1 software (R
Core Team 2022). We classied the vertebrates utilizing the
armadillo burrows using the categories proposed by DeGre-
gorio et al. (2022): (1) pass, when the animal passed in front
of the burrow without showing any type of interaction; (2)
approach, animal interacted with accumulated sand, bare
soil or litter piled in front of the burrow for less than ve
seconds; (3) entrance, animal interacted with the burrow
entrance by foraging, resting, looking for prey, entering the
burrow, and/or remaining visible for the camera for more
than ve seconds; (4) interior, when the animal entered the
burrow and left the eld of view of the camera trap.
We used a network structure together with the modular-
ity metric (Almeida-Neto and Ulrich 2011; Santos and Sano
2022) to visualize the distribution of species records and the
dierent categories of uses. For greater robustness of the
network analyses, we used species with ≥ 10 camera trap
records. The signicance of the modularity was estimated
through Monte Carlo procedures based on comparisons with
randomized distributions generated with null models. We
generated 1,000 randomized matrices based on the original
weighted matrix using the algorithm proposed by Vázquez
et al. (2007) and Pinheiro et al. (2019). Network structure
was considered to signicantly deviate from the null model
when p < 0.05. All analyses were performed using the R
4.2.1 software (R Core Team 2022).
Results
A total of 27 active burrows of armadillos were sampled, but
we were unable to characterize which species were using
the holes based on their morphometric characteristics. We
obtained 594 independent records of dierent mammals,
birds, and reptiles distributed in 15 orders and 34 species
(Table 1; Online Resource 1, 2, and 3). The accumulation
curve of camera trapping data showed an asymptotic ten-
dency (Online Resource 4). For mammals, we observed
seven species of Carnivora (Cerdocyon thous, Eira bar-
bara, Herpailurus yagouaroundi, Leopardus pardalis,
Puma concolor, Nasua nasua, and Procyon cancrivorus),
1 3

Armadillo burrows: a meeting point for biodiversity in the Pantanal Wetland
ve Cetartiodactyla (Mazama americana, Subulo gouazou-
bira, Dicotyles tajacu, Tayassu pecari and the exotic feral
Sus scrofa), four Rodentia (Clyomys laticeps, Dasyprocta
azarae, Oecomys mamorae, and Thrichomys fosteri), two
Cingulata (E. sexcinctus and D. novemcinctus), two Pilosa
(Myrmecophaga tridactyla and Tamandua tetradactyla),
one Didelphimorphia (Monodelphis domestica), and one
Perissodactyla (Tapirus terrestris) (Table 1).
Concerning birds, we observed three species of Galli-
formes (Aburria cumanensis, Crax fasciolata, and Ortalis
canicollis), three Passeriformes (Cyanocorax cyanomelas,
Synallaxis albilora, and Turdus leucomelas), and one spe-
cies each of Cariamiformes (Cariama cristata), Columbi-
formes (Leptotila verreauxi), Falconiformes (Micrastur
semitorquatus), and Tinamiformes (Crypturellus undulatus).
We also registered two species of reptiles: one Squamata
(Tupinambis teguixin) and one Testudinata (Geochelone
carbonaria) (Table 1; Online Resource 1, 2, and 3). The
rodent species T. fosteri, D. azarae, and C. laticeps had the
highest number of records, and the species G. carbonaria,
M. tridactyla, M. semitorquatus, and T. teguixin were the
least recorded (Table 1). The armadillo species recorded in
Table 1 Utilization records of vertebrate species revealed by camera traps in front of burrows of Dasypus novemcinctus and Euphractus sexcinctus
in the Southern Pantanal wetland from November 2015 to August 2016. The records were classied as pass, when the animal passed in front of
the burrow without presenting any type of interaction; Approach, animal interacted with accumulated sand, bare soil or litter piled in front of the
burrow for less than ve seconds; entrance, animal interacted with the burrow opening by foraging, resting, looking for prey, entering the entrance
of the burrow, and/or remaining visible for the camera for more than ve seconds; Interior, when the animal entered the burrow and left the eld
of view of the camera trap
Class Order Family Species (No. of records) Burrow utilization
Passage Approach Entrance Interior
Mammals Carnivora Canidae Cerdocyon thous (n = 8) 3 3 2 0
Felidae Herpailurus yagouaroundi (n = 2) 2 0 0 0
Leopardus pardalis (n = 6) 4 1 1 0
Puma concolor (n = 2) 1 1 0 0
Mustelidae Eira barbara (n = 3) 2 1 0 0
Procyonidae Nasua nasua (n = 24) 911 4 0
Procyon cancrivorus (n = 3) 1 1 1 0
Cetartiodactyla Cervidae Mazama americana (n = 2) 2 0 0 0
Subulo gouazoubira (n = 13) 13 0 0 0
Suidae Sus scrofa feral (n = 30) 24 0 6 0
Tayassuidae Dicotyles tajacu (n = 53) 44 8 1 0
Tayassu pecari (n = 32) 28 4 0 0
Cingulata Dasypodidae Dasypus novemcinctus (n = 10) 1 4 3 2
Euphractus sexcinctus (n = 48) 19 17 9 3
Didelphimorphia Didelphidae Monodelphis domestica (n = 11 ) 72 2 0
Perissodactyla Tapiridae Tapirus terrestris (n = 14) 14 0 0 0
Pilosa Myrmecophagidae Myrmecophaga tridactyla (n = 1) 1 0 0 0
Tamandua tetradactyla (n = 10) 72 1 0
Rodentia Cricetidae Oecomys mamorae (n = 4) 1 2 0 1
Dasyproctidae Dasyprocta azarae (n = 72) 53 13 6 0
Echimyidae Clyomys laticeps (n = 64) 1 18 29 16
Thrichomys fosteri (n = 81) 0 13 54 14
Birds Cariamiformes Cariamidae Cariama cristata (n = 4) 4 0 0 0
Columbiformes Columbidae Leptotila verreauxi (n = 56) 54 2 0 0
Falconiformes Falconidae Micrastur semitorquatus (n = 1) 0 0 1 0
Galliformes Cracidae Aburria cumanensis (n = 4) 3 0 1 0
Crax fasciolata (n = 10) 8 0 2 0
Ortalis canicollis (n = 2) 1 0 0 0
Passeriformes Corvidae Cyanocorax cyanomelas (n = 2) 1 1 0 0
Furnariidae Synallaxis albilora (n = 3) 3 0 0 0
Turdidae Turdus leucomelas (n = 2) 2 0 0 0
Tinamiformes Tinamidae Crypturellus undulatus (n = 18) 12 3 3 0
Reptiles Squamata Teiidae Tupinambis teguixin (n = 1) 1 0 0 0
Testudinata Testudinidae Geochelone carbonaria (n = 1) 1 0 0 0
1 3

F. M. Santos et al.
the armadillos D. novemcinctus and E. sexcinctus and the
medium-sized mammal N. nasua (Module 2 [Color Green],
Fig. 1). The remaining species (D. tajacu, T. pecari, S.
scrofa feral, S. gouazoubira, T. terrestris, T. tetradactyla,
M. domestica, D. azarae, C. fasciolata, L. verreauxi, and
C. undulatus) were only related with pass (Module 3 [Color
Red], Fig. 1). Regarding vertebrate species that had < 10
records (Table 1), we highlight C. thous and L. pardalis,
with approach and entrance with burrows (Fig. 2), as well
our study were E. sexcinctus and D. novemcinctus. During
the excavations, we found eight taxa of invertebrates and
small vertebrates inside 51% of monitored burrows (Online
Resource 5).
The network data from the 16 vertebrate species with
≥ 10 records (Table 1) showed signicant modular patterns
with three established modules (Qw modularity = 0.33,
P = 0). The small rodents C. laticeps and T. fosteri were
associated with the entrance and interior (Module 1 [Color
Blue], Fig. 1). Three species were associated with approach,
Fig. 1 Network demonstrating the association between species of ver-
tebrates and their use of armadillo burrows sampled in the Southern
Pantanal wetland between November 2015 to August 2016. Only spe-
cies with > 10 records were considered in this analysis, revealing three
modules highlighted by shadows: Module 1, in blue, included Interior
and Entrance categories; Module 2, in green, refers to Approach cat-
egory; and Module 3, in red, refers to Pass category. The thickness of
the line is related to the times that category was recorded
1 3

Armadillo burrows: a meeting point for biodiversity in the Pantanal Wetland
of medium-sized armadillos to start a new gallery. The asso-
ciations of small rodents with armadillo burrows were also
observed in the Cerrado region (Vieira et al. 2005), and in
southeastern Brazil (Bueno et al. 2004).
Our results revealed that the carnivorous L. pardalis and
C. thous showed some interesting uses of approach and
entrance (Fig. 2). As predators, they may be able to smell
the presence of common prey, such as the small rodents
observed here entering the armadillo burrows, as reported
by Bianchi et al. (2014). The presence of potential prey may
also explain why the top predator bird M. semitorquatus
was observed interacting with the burrows. We recorded
ten species of birds, but only three of them (C. undulatus,
L. verreauxi, and C. fasciolata) had more than ten records.
Leptotila verreauxi and C. undulatus exhibited a dierent
pattern than other animals: dozens of photos were taken
in sequence but discarded to obtain a record. This could
suggest that they are using the burrows for foraging small
as birds and the gregarious T. pecari that searched for food
around the burrows (Online Resource 6).
Discussion
Our data indicate that E. sexcinctus and D. novemcinctus
create new microenvironments that act as meeting points for
the local fauna in the Pantanal wetland. We observed that
the two most abundant small rodent species in the Panta-
nal, T. fosteri and C. laticeps (Herrera et al. 2007; Sano et
al. 2021; Pessanha et al. 2023), entered the burrows. There
is no evidence that T. fosteri makes burrows, but there are
many reports of this species using holes and tree hollows
for protection (Antunes et al. 2016; Menezes et al. 2018). In
the case of C. laticeps, this semifossorial species is known
to dig its own gallery (Lacher and Alho 1989; Antunes et al.
2016, 2022). We hypothesize that C. laticeps uses burrows
Fig. 2 Sequences showing the use of armadillo burrows by rodents
(prey) and carnivores (predators) in the Southern Pantanal wetland,
Brazil, November 2015 to August 2016. A1-3: Leopardus pardalis
staying near a burrow where a Clyomys laticeps recently entered B1-3:
Thrichomys fosteri entering the burrow, and C1-3: Cerdocyon thous
smelling the B1-3 burrow just after the Thrichomys fosteri entered
1 3

F. M. Santos et al.
Funding This work was supported by the following Brazilian re-
search agencies: Coordenação de Aperfeiçoamento de Nível Superior
(CAPES – Finance code 001), Conselho Nacional de Desenvolvimen-
to Cientíco e Tecnológico (CNPq). FMS received a fellowship from
CAPES (88887.369261/2019–00). HMH received a fellowship from
CNPq (311769/2023-3). NYS received a fellowship from Project Rede
Pantanal from the Ministry of Science, Technology, and Innovations of
Brazil (FINEP Grant No. 01.20.0201.00).
Declarations
Conict of interest One of the authors, Carlos Fonseca, is a subject
editor of Mammalian Biology.
Statement of animal ethics This study was authorized by Brazilian
Laws (SISBIO 49662-1) and the Ethics Committee of Animal Use
(CEUA-UCDB 019/2015).
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Armadillo burrows have been reported as shelters in dif-
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the present study.
In the last few years, the Pantanal region has been suer-
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(Garcia et al. 2021; Tomas et al. 2021; Libonati et al. 2022).
Small vertebrates represented more than 80% of the verte-
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(Tomas et al. 2021) and some small rodents were, surpris-
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al. 2022). This suggests that the (armadillo) burrows in the
Pantanal region can also act as protection for some small
vertebrates during wildres, which are predicted to be more
frequent each year (Libonati et al. 2022). A similar phenom-
enon also occurs with wombats in Australia (Linley et al.
2024; Old et al. 2018), where 48 vertebrate species were
record using wombat burrows as shelter during gigares
(Linley et al. 2024). Therefore, as observed for the giant
armadillo, the role played by smaller armadillo species in
maintaining biodiversity and ecological dynamics in the
Pantanal is evident.
Supplementary Information The online version contains
supplementary material available at https://doi.org/10.1007/s42991-
024-00466-8.
Acknowledgements We thank Gabriel de Oliveira Freitas Freitas
(https://icterusecoturismo.com) for the identication of Leptotila ver-
reauxi, Micrastur semitorquatus, Synallaxis albilora and Turdus leu-
comelas. We also thank team of Insana Huna (www.insanahuna.com)
research group and the sta of Alegria farm.
Author contributions Filipe Martins Santos, Grasiela Edith de Olivei-
ra Porrio, and Heitor Miraglia Herrera contributed to the study con-
ception and design. Equipment and eld work logistics funding were
partially provided by Carlos Fonseca and Amadeus Soares. Material
preparation, data collection, and analysis were performed by Grasiela
Edith de Oliveira Porrio, William Oliveira de Assis, Leonardo França
Nascimento, Nayara Yoshie Sano, Jader de Oliveira and Filipe Martins
Santos. The rst draft of the manuscript was written by Grasiela Edith
de Oliveira Porfírio, William Oliveira de Assis, and Filipe Martins
Santos. All authors commented on previous versions of the manuscript
and read and approved the nal manuscript.
1 3

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