Mammalia 2023; aop
Short Note
Hernani F. M. Oliveira*, Nícholas F. Camargo, Humberto C. Nappo, Isac M. Venâncio, Tânia A. Queiroz,
Joseana L. Freitas, Diainara S. Figueiredo, Pedro R. Alencar, Ticiane L. Costa and Guarino R. Colli
Unprecedented habitat use by an arboreal
Neotropical marsupial (Didelphimorphia:
Didelphidae) in the Cerrado
https://doi.org/10.1515/mammalia-2023-0040
Received April 4, 2023; accepted October 26, 2023;
published online November 29, 2023
Abstract: The brown-eared woolly opossum (Caluromys
lanatus) is an elusive frugivorous Neotropical canopy
specialist marsupial considered primarily arboreal, but here
we report the first records of individuals from this species
captured on the ground in the Cerrado. We hypothesize two
main reasons to explain this behavior: first related to canopy
openness, which would force the animals to the ground to
keep moving across the landscape; and the second related
with search for food on the ground during periods of food
scarcity due to increased canopy gaps. Our findings can have
important implications for the species conservation and
understanding forest ecological dynamics.
Keywords: canopy; ground; mammal; savanna; vertical
stratification
Forest canopies remain one of the last ecological frontiers to
be explored and understood by science (Nakamura et al.
2017). There is generally much more information about the
ecology of ground and understory animals than about can-
opy specialists (Nakamura et al. 2017). This situation is
partially related not only to their elusiveness but also due to
the challenges of accessing and studying animal ecology in
canopies, which in some forests can reach over 40 m (Lang
et al. 2022; Nakamura et al. 2017; Santana et al. 2017). How-
ever, different mammals previously regarded as primarily
*Corresponding author: Hernani F. M. Oliveira, Coleção Herpetológica
da Universidade de Brasília, Departamento de Zoologia, Universidade de
Brasília, Brasília 70910-900, Brazil, E-mail: oliveiradebioh@gmail.com.
https://orcid.org/0000-0001-7040-8317
Nícholas F. Camargo and Humberto C. Nappo, Departamento de
Ecologia, Universidade de Brasília, Brasília, DF, Brazil
Isac M. Venâncio, Tânia A. Queiroz, Joseana L. Freitas, Diainara S.
Figueiredo, Pedro R. Alencar, Ticiane L. Costa and Guarino R. Colli,
Coleção Herpetológica da Universidade de Brasília, Departamento de
Zoologia, Universidade de Brasília, Brasília 70910-900, Brazil
arboreal have also occasionally been recorded on the ground
(Mourthé et al. 2007; Pauli et al. 2014). The vertical stratifi-
cation patterns of these animals may vary according to
different factors, such as habitat structure, food resource
availability, and predation (Camargo et al. 2018; Rader and
Krockenberger 2006). For instance, in the Brazilian savanna
(i.e., Cerrado), the arboreal rodent Rhipidomys macrurus is
more associated with the canopy in less complex forests to
avoid predators (Camargo et al. 2018). Therefore, unusual
records of arboreal mammals on the ground may represent
relevant information to expand our understanding of their
ecology and different aspects of forest dynamics (Barnett
et al. 2012; Pauli et al. 2014).
Cerrado is mainly formed by open vegetation forma-
tions and is highly influenced by edaphic factors and the
frequency and strength of fires (Hoffmann and Moreira
2002; Ribeiro and Walter 1998). The latter has an essential
role in shaping the vegetation structure within the Cerrado
by increasing tree mortality and changing tree density
across the landscape (Miranda et al. 2002). However, not all
vegetation formations are equally susceptible to fire within
the Cerrado (Hoffman et al. 2012). Even though gallery forests
do not usually burn due to the lower temperatures and sun
exposure, and high tree cover (Hoffman et al. 2012; Miranda
et al. 2010), there have been reports of fires within gallery
forests, with further consequences for its associated fauna
(Oliveira and Aguiar 2015). Moreover, gallery forests within
the Cerrado present a unique ecological opportunity for
arboreal animals to feed and move, with approximately 10 %
(25 out of the 249 spp.) of Cerrado mammals presenting some
degree of arboreality (Paglia et al. 2012). However, fires can
reduce canopy cover in vegetation formations of the Cerrado
(Miranda et al. 2002), with potentially cascading conse-
quences for the movement patterns and food availability of
arboreal mammals using these habitats.
The brown-eared woolly opossum (Caluromys lanatus)
is an elusive arboreal Neotropical frugivorous marsupial
with a wide distribution ranging from east of the Andes in
Colombia throughout northeastern Argentina and
2 H.F.M. Oliveira et al.: Arboreal marsupial recorded in pitfall traps
southeastern Brazil (Astúa 2015). This marsupial heavily
feeds on fruits, but it may also feed on invertebrates, ver-
tebrates, and nectar (Camargo et al. 2017; Casela and Cáceres
2006; Gribel 1988). C. lanatus is a canopy specialist (Camargo
et al. 2018; Lambert et al. 2005; Vieira and Camargo 2012),
with the genus presenting adaptive convergence with pri-
mates regarding behavior, locomotion, posture, and foot
morphology, such as the use of terminal branches, walking
gait, and a diagonal-sequence footfall pattern during locomo-
tion (Cartmill et al. 2002; Lemelin et al. 2003; Schmitt and
Lemelin 2002, 2004). To our knowledge, it has never been
recorded at ground level in the Cerrado, but records for other
biomes, such as the Amazon have been reported (Santos-Filho
et al. 2015). Moreover, studies with considerable sampling ef-
forts in the canopy usually have only recorded a few in-
dividuals, indicating low-density populations (Carmignotto and
Aires 2011; Camargo et al. 2018; Lambert et al. 2005). Thus, this
species’ ecology and natural history have been scarcely re-
ported (Camargo et al. 2017; Casela and Cáceres 2006; Lambert
et al. 2005).
Here, we report five records of C. lanatus individuals in
pitfall traps inside gallery forests of the Brazilian Cerrado
that burned in 2005 and 2011, representing the first records
of this species on the ground within this biome. Although
C. lanatus have some morphological similarities with its
congener species Caluromys philander, it differs from it in
relation its conspicuous tail hair pattern, which is covered by
hair until the half part of its dorsal side and one fifth on its
ventral side (Gardner 2007; Rossi et al. 2012). In C. philander,
the tale is covered with hairs in similar extension on the
dorsal and ventral side, which is close to one third of its total
length (Gardner 2007; Rossi et al. 2012). Moreover, we pro-
pose potential hypotheses explaining why this species use
ground level for its movement and foraging.
Cerrado has a highly seasonal climate with a well-
defined warm-wet season from October to April when close
to 90 % of the annual precipitation of 1100–1600 mm occurs,
and a cool-dry season from May to September (Miranda et al.
1993). All captures were made inside gallery forests in Dis-
trito Federal in central Brazil at two adjacent protected
areas: Reserva Ecológica do IBGE (RECOR) (15°56′41″S, 47°53′
07″W) (1300 ha) and Estação Ecológica do Jardim Botânico de
Brasília (EEJBB) (15°53′09″S, 47°50′34″W) (4518 ha) (Figure 1).
Representing 5 % of the Cerrado distribution, gallery forests
form corridors along water courses and wetlands (Ribeiro
and Walter 1998). This formation is characterized by 20–
30 m trees with a canopy cover ranging between 70 % and
95 % (Ribeiro and Walter 1998). Gallery forests represent
some of the few forest formations in the Cerrado, which is
characterized mainly by open formations that cover over
70 % of its total distribution (Ribeiro and Walter 1998). The
gallery forest at the RECOR burned during non-prescribed
fires in 2005 and 2011, which affected most of this protected
area (Caetano and Colli 2021; Mendonça et al. 2015). Fires can
cause significant tree mortality and the opening of vegeta-
tion clearings in gallery forests (Kellman and Meave 2007;
Santiago et al. 2005). Consequently, this locality presents a
lower canopy cover compared to other non-burned gallery
forests in the region located in other protected areas.
Forty permanent pitfall traps (20 in each site) have been
checked twice a week inside these two areas since 2000 to
monitor the dynamics of lizard populations and commu-
nities. Each trap consists of four PVC pipe sections (50 cm
depth, 20 cm diameter – 15 litter buckets) buried vertically in
the ground and connected by 5 m long metal fences forming
a Y shape (Figure 2). Traps are separated by a distance of at
least 20 m.
On three occasions during 2022, we found C. lanatus
(three individuals) inside pitfall traps in the gallery forest of
RECOR (Figure 3), one during the end (March 29) and two at
the beginning of the wet season (November 8 and December
13). We also recorded two other individuals from this species
in the pitfalls of the same area: one on June 14, 2017, and
another on November 23, 2018, totaling five records of
C. lanatus in pitfall traps of our study sites. However, we
could not track back the site and buckets of the last two
individuals mentioned. Since our monitoring focused on the
herpetofauna, these individuals were only noticed and
recorded anecdotally, and even more captures may have
occurred in previous years. All recorded individuals were
released after capture, but at least one individual, after being
released from the pitfall quickly climbed back to the canopy,
reinforcing the pattern of high arboreal activity of this
marsupial species despite the unusual ground records.
Our records of C. lanatus (three within a single year) in
pitfall traps indicate a relatively high exploitation intensity by
the marsupial at ground level. For three years (2012–2014)
(after the fires during 2005/2011), Camargo et al. (2018) sampled
eight forest sites within the same protected area of our study,
including also one of our sampling sites at EEJBB, with Sher-
man live traps on the ground, understory, and canopy (43,200
trap-nights). They captured 13 individuals of C. lanatus, all of
them in the canopy. Moreover, only very few other studies
within the species range has ever recorded the species on the
ground (Fleck and Harder 1995; Santos-Filho et al. 2015). Pitfall
traps are designed to capture ground-dwelling animals, but
might occasionally trap some arboreal mammal species
(Ardente et al. 2017); thus, repeatedly capturing such an arbo-
real species at ground level is worth noting.
A possible explanation is that these animals were forced
to climb down the trees to move through the area. Usually,
this would not be necessary for a canopy specialist; however,
 H.F.M. Oliveira et al.: Arboreal marsupial recorded in pitfall traps 3

Figure 1: Locations of the two gallery forests in the Brazilian Cerrado where we captured the brown-eared woolly opossum (Caluromys lanatus) for the
first time in pitfall traps. The white circle in the left indicates the gallery forest at the Reserva Ecológica do IBGE (RECOR), and the white circle on the right
indicates the gallery forest at the Estação Ecológica do Jardim Botânico de Brasília (EEJBB).

the gallery forest in the RECOR is characterized by a
discontinuous canopy, a lasting consequence of a wildfire
back in 2011. Thus, this forest seems to present a discon-
nected ecological corridor for the movement of this species,
which is a limiting factor for the movement of arboreal
mammal species (Cudney-Valenzuela et al. 2023). Even low-
intensity fires can significantly impact the structure of
tropical forests, affecting the locomotion of arboreal mam-
mals and overall gallery forest ecological dynamics (Mar-
ques et al. 2022; Widyastuti et al. 2022). Hence, animals
willing to move through the forest must climb down to the
ground at some point, allowing them to be caught in our
pitfall traps. The behavior of walking on the ground has been
reported for other arboreal mammal species, such as
4 H.F.M. Oliveira et al.: Arboreal marsupial recorded in pitfall traps
Figure 2: Pitfall traps that were used to capture brown-eared woolly opossum (Caluromys lanatus) for the first time on the ground in the Brazilian
Cerrado.
arboreal sugar gliders from Australia (van der Ree et al. 2004)
and primates of the Atlantic Forest (Dib et al. 1997; Mourthé
et al. 2007), which have increased ground activity in places
where there are forest gaps. Similarly, anthropogenic forest
disturbances increase the terrestrial activity of Bornean
orangutans (Pongo pygmaeus) according to the degree of forest
disturbance and canopy gap size (Ancrenaz et al. 2014).
Terrestrial activity has also been reported among
arboreal frugivorous Neotropical primates (pithecines) due
to food scarcity in the canopy, which forces individuals to
look for food on the ground (Barnett et al. 2012). Although
fruit production in Cerrado gallery forests is highly seasonal
(Funch et al. 2002; Ragusa-Netto 2008), the decreased canopy
cover and the simplification of the vertical structure may
have further accentuated the low food availability (Cudney-
Valenzuela et al. 2023), forcing animals to climb down the
trees to access alternative food resources. This strategy
would also minimize intra- and interspecific competition in
the canopy, as seen for other arboreal mammals (Eppley
et al. 2022; Mourthé et al. 2007). Alternatively, these in-
dividuals could have been attracted to the traps while
already foraging on the ground since the pitfalls usually trap
invertebrates, one of their potential food items.
It is important to note that this unusual behavior can
significantly affect survival within this ecological context.
Terrestrial activity in arboreal mammals is linked with
increased predatory risk and higher chances of contracting
new parasites (Kawabe and Mano 1972; Mourthé et al. 2007).
Feral dogs, for example, attack arboreal mammals moving
on the ground within many Neotropical protected areas
(Chaves et al. 2022; Galetti and Sazima 2006; Lessa et al. 2016).
Thus, among many other terrestrial predator species
(including native ones), they could represent an essential
new pressure for these individuals. Indeed, feral dogs are
often recorded in the protected area of our study (e.g., Pet-
razzini and Aguiar 2021). Furthermore, higher infection
rates in arboreal animals by different pathogens during
locomotion on the ground (Zommers et al. 2013) may impact

Figure 3: Photos of two of the five individuals of the brown-eared woolly
opossum (Caluromys lanatus) captured in pitfall traps in gallery forests of
the Brazilian Cerrado.
their health and survival, with further consequences for
population dynamics.
The relatively frequent observations of C. lanatus
exploiting the ground level across multiple years suggest
that this marsupial, despite its high arboreal specialization,
presents certain behavioral plasticity in the vertical strata
use of forests. On the other hand, such ground-level activity
of C. lanatus might be highly context-dependent, here linked
with the intensity and frequency of wildfires that have
reduced forest canopy cover. Indeed, other multiple-year
studies evaluating the assemblage of small mammals in
other gallery forests within our protected area did not
document it, and it has only been rarely documented else-
where. However, it is essential to note that C. lanatus plas-
ticity in vertical space use can significantly affect the species’
survival in different deforestation scenarios and have rele-
vant implications for forest dynamics (e.g., Havmøller et al.
2021; Lefevre and Rodd 2009). Thus, more studies on the
natural history and ecology of this rare and elusive
Neotropical marsupial are warranted.
H.F.M. Oliveira et al.: Arboreal marsupial recorded in pitfall traps 5
Acknowledgments: The authors are grateful to the admin-
istration of RECOR and EEJBB and the team at Coleção Her-
petológica da Universidade de Brasília, responsible for
maintaining and checking the traps.
Research ethics: The authors declare that they have
complied with the guidelines of the Care and Use
Committee of the American Society of Mammalogists
[Sikes, R.S. and Animal Care and Use Committee of the
American Society of Mammalogists (2016). Guidelines of the
American Society of Mammalogists for the use of wild
mammals in research and education. J. Mammal. 97 (3): 663–
688].
Author contributions: Conceptualization: H.F.M. Oliveira,
N.F. Camargo, H.C. Nappo, G.R. Colli; methodology, formal
analysis and investigation: H.F.M. Oliveira, N.F. Camargo,
G.R. Colli; writing – original draft preparation: H.F.M.
Oliveira, N.F. Camargo, H.C. Nappo, G.R. Colli; writing –
review and editing: H.F.M. Oliveira, N.F. Camargo, H.C.
Nappo, I.M. Venâncio, T.A. Queiroz, J.L. Freitas, D.S.
Figueiredo, P.R. Alencar, T.L. Costa, G.R. Colli; funding
acquisition: H.F.M. Oliveira, G.R. Colli; resources: G.R. Colli;
supervision: G.R. Colli.
Competing interests: The authors declare no competing
financial or non-financial interests regarding the
publication of the current article.
Research funding: The authors thank the Coordenação de
Apoio à Formação de Pessoal de Nível Superior (CAPES),
Conselho Nacional do Desenvolvimento Científico e
Tecnológico (CNPq), and Fundação de Apoio à Pesquisa do
Distrito Federal (FAPDF) for financial support.
Data availability: All data is available in the manuscript.
References
Ancrenaz, M., Sollmann, R., Meijaard, E., Hearn, A.J., Ross, J., Samejima, H.,
Loken, B., Cheyne, S.M., Stark, D.J., Gardner, P.C., et al. (2014). Coming
down from the trees: is terrestrial activity in Bornean orangutans
natural or disturbance driven? Sci. Rep. 4: 4024–4029.
Ardente, N.C., Ferreguetti, A.C., Gettinger, D., Leal, P., Martins-Hatano, F.,
and Bergallo, H.G. (2017). Differencial efficiency of two sampling
methods in capturing non-volant small mammals in an area in eastern
Amazonia. Acta Amaz 47: 123–132.
Astúa, D. (2015). Order didelphimorphia. In: Wilson, D.E. and
Mittermeier, R.A. (Eds.), Handbook of the mammals of the world,
Monotremes and marsupials. Lynx Edicions, Barcelona, pp. 70–187.
Barnett, A.A., Boyle, S.A., Norconk, M.M., Palminteri, S., Santos, R.R., Veiga,
L.M., Alvim, T.H.G., Bowler, M., Chism, J., Fiore, A.D., et al. (2012).
Terrestrial activity in pitheciins (Cacajao, Chiropotes, and Pithecia). Am.
J. Primatol. 74: 1106–1127.
Caetano, G.H.D. and Colli, G.R. (2021). Climate instability causing the decline
of a Neotropical savanna lizard population (Squamata: Tropiduridae).
Biol. J. Linn. Soc. 134: 474–485.
6 H.F.M. Oliveira et al.: Arboreal marsupial recorded in pitfall traps
Camargo, N.F., Sano, N.Y., and Vieira, E.M. (2017). Predation upon small
mammals by Caluromys lanatus (didelphimorphia: didelphidae) and
Callithrix penicillata (primates: Callitrichidae) in the Brazilian savanna.
Mammalia 81: 207–210.
Camargo, N.F.D., Sano, N.Y., and Vieira, E.M. (2018). Forest vertical
complexity affects alpha and beta diversity of small mammals.
J. Mammal. 99: 1444–1454.
Carmignotto, A.P. and Aires, C.C. (2011). Mamíferos não voadores
(Mammalia) da Estação Ecológica Serra Geral do Tocantins. Biota
Neotrop. 11: 313–328.
Cartmill, M., Lemelin, P., and Schmitt, D. (2002). Support polygons and
symmetrical gaits in mammals. J. Linn. Soc. 136: 401–420.
Casela, J. and Cáceres, N.C. (2006). Diet of four small mammal species from
Atlantic forest patches in South Brazil. Neotrop. Biol. 1: 5–11.
Chaves, Ó.M., Júnior, J.S., Buss, G., Hirano, Z.M., Jardim, M.M.A., Amaral,
E.L.S., Godoy, J.C., Peruchi, A.R., Michel, T., and Bicca-Marques, J.C.
(2022). Wildlife is imperiled in peri-urban landscapes: threats to
arboreal mammals. Sci. Total Environ. 821: 152883.
Cudney‐Valenzuela, S.J., Arroyo‐Rodríguez, V., Morante‐Filho, J.C., Toledo‐
Aceves, T., and Andresen, E. (2023). Tropical forest loss impoverishes
arboreal mammal assemblages by increasing tree canopy openness.
Ecol. Appl. 33: 1–12.
Dib, L.R.T., Oliva, A.S., and Strier, K.B. (1997). Terrestrial travel in muriquis
(Brachyteles arachnoides) across a forest clearing at the Estação
Biológica de Caratinga, Minas Gerais, Brazil. Neotrop. Primates 5:
8–9.
Eppley, T.M., Hoeks, S., Chapman, C.A., Ganzhorn, J.U., Hall, K., Owen, M.A.,
Adams, D.B., Allgas, N., Amato, K.R., Andriamahaihavana, M., et al.
(2022). Factors influencing terrestriality in primates of the Americas
and Madagascar. Proc. Natl. Acad. Sci. U. S. A. 119: e2121105119.
Fleck, D.W. and Harder, J.D. (1995). Ecology of marsupials in two Amazonian
rain forests in northeastern Peru. J. Mammal. 76: 809–818.
Funch, L.S., Punch, R., and Barroso, G.M. (2002). Phenology of gallery and
montane forest in the Chapada diamantina, Bahia, Brazil. Biotropica
34: 40–50.
Galetti, M. and Sazima, I. (2006). Impact of feral dogs in an urban Atlantic
Forest fragment in southeastern Brazil. Nat. Conserv. 4: 146–151.
Gardner, A.L. (2007). Genus Caluromys. In: Gardner, A.L. (Ed.). Mammals of
South America: marsupials, xenarthrans, shrews, and bats. The University
of Chicago Press, Chicago, pp. 3–11.
Gribel, R. (1988). Visits of Caluromys lanatus (Didelphidae) to flowers of
Pseudobombax tomentosum (Bombacaceae): a probable case of
pollination by marsupials in Central Brazil. Biotropica 20: 344–347.
Havmoller, L.W., Loftus, J.C., Havmoller, R.W., Alavi, S.E., Caillaud, D., Grote,
M.N., Hirsch, B.T., Tórrez-Herrera, L.L., Kays, R., and Crofoot, M.C.
(2021). Arboreal monkeys facilitate foraging of terrestrial frugivores.
Biotropica 53: 1685–1697.
Hoffman, W.A. and Moreira, A.G. (2002). The role of fire in population
dynamics of woody plants. In: Oliveira, P.S. and Marquis, R.J. (Eds.), The
cerrados of Brazil: ecology and natural history of a Neotropical savanna.
Columbia University Press, New York, pp. 159–177.
Hoffmann, W.A., Jaconis, S.Y., Mckinley, K.L., Geiger, E.L., Gotsch, S.G., and
Franco, A.C. (2012). Fuels or microclimate? Understanding the drivers
of fire feedbacks at savanna–forest boundaries. Austral. Ecol. 37:
634–643.
Kellman, M. and Meave, J. (2007). Fire in the tropical gallery forests of Belize.
J. Biogeogr. 24: 23–34.
Kawabe, M. and Mano, T. (1972). Ecology and behavior of the wild proboscis
monkey, Nasalis larvatus (Wurmb), in Sabah, Malaysia. Primates 13:
213–227.
Lambert, T.D., Malcolm, J.R., and Zimmerman, B.L. (2005). Variation in small
mammal species richness by trap height and trap type in southeastern
Amazonia. J. Mammal. 86: 982–990.
Lang, N., Jetz, W., Schindler, K., and Wegner, J.D. (2022). A high-resolution
canopy height model of the Earth, ArXiv preprint arXiv 2204: 08322.
Lefevre, K.L. and Rodd, F.H. (2009). How human disturbance of tropical
rainforest can influence avian fruit removal. Oikos 118: 1405–1415.
Lemelin, P., Schmitt, D., and Cartmill, M. (2003). Footfall patterns and
interlimb co‐ordination in opossums (Family Didelphidae): evidence
for the evolution of diagonal‐sequence walking gaits in primates.
J. Zool. 260: 423–429.
Lessa, I., Guimarães, T.C.S., de Godoy, H.B., Cunha, A., and Vieira, E.M.
(2016). Domestic dogs in protected areas: a threat to Brazilian
mammals? Nat. Conserv. 14: 46–56.
Marques, N.C., Machado, R.B., Aguiar, L.M., Mendonça-Galvão, L., Tidon, R.,
Vieira, E.M., Marini-Filho, O.J., and Bustamante, M. (2022). Drivers of
change in tropical protected areas: long-term monitoring of a
Brazilian biodiversity hotspot. Perspect. Ecol. Conserv. 20: 69–78.
Mendonça, A.F., Armond, T., Camargo, A.C.L., Camargo, N.F., Ribeiro, J.F.,
Zangrandi, P.L., and Vieira, E.M. (2015). Effects of an extensive fire on
arboreal small mammal populations in a neotropical savanna
woodland. J. Mammal. 96: 368–379.
Miranda, A.C., Miranda, H.S., Dias, I.D.F.O., and de Souza Dias, B.F. (1993).
Soil and air temperatures during provocated cerrado fires in central
Brazil. J. Trop. Ecol. 9: 313–320.
Miranda, H.S., Bustamante, M.M., and Miranda, A.C. (2002). The fire factor.
In: Oliveira, P.S. and Marquis, R.J. (Eds.), The cerrados of Brazil.
Columbia University Press, New York, pp. 51–68.
Miranda, H.S., Neto, W.N., and Neves, B.M.C. (2010). Caracterização das
queimadas de Cerrado. In: Miranda, H.S. (Ed.). Efeitos do regime do
fogo sobre a estrutura de comunidades de cerrado: resultados do Projeto
Fogo. IBAMA, Brasília, pp. 23–34.
Mourthé, Í.M., Guedes, D., Fidelis, J., Boubli, J.P., Mendes, S.L., and Strier,
K.B. (2007). Ground use by northern muriquis (Brachyteles
hypoxanthus). Am. J. Primatol. 69: 706–712.
Nakamura, A., Kitching, R.L., Cao, M., Creedy, T.J., Fayle, T.M., Freiberg, M.,
Hewitt, C.N., Itioka, T., Koh, L.P., Ma, K., et al. (2017). Forests and their
canopies: achievements and horizons in canopy science. Trends Ecol.
Evol. 32: 438–451.
Oliveira, H.F.M. and Aguiar, L.M.S. (2015). The response of bats (Mammalia:
Chiroptera) to an incidental fire on a gallery forest at a Neotropical
savanna. Biota Neotrop. 15: 1–5.
Paglia, A.P., Da Fonseca, G.A., Rylands, A.B., Herrmann, G., Aguiar, L.M.,
Chiarello, A.G., Leite, Y.L.R., Costa, L.P., Siciliano, S., Kierulff, M.C.M., et al.
(2012). Lista anotada dos mamíferos do Brasil 2a edição/annotated
Checklist of Brazilian mammals. Occas. Papers Conserv. Biol. 6: 1–82.
Pauli, J.N., Mendoza, J.E., Steffan, S.A., Carey, C.C., Weimer, P.J., and Peery,
M.Z. (2014). A syndrome of mutualism reinforces the lifestyle of a
sloth. Proc. Royal Soc. B 281: 20133006.
Petrazzini, P.B. and Aguiar, L.M.S. (2021). You say goodbye, and I say hello: the
giant anteater (Myrmecophaga tridactyla) activity pattern in response to
temperature and human presence. Mastozool. Neotrop. 28: 469–479.
Rader, R. and Krockenberger, A. (2006). Does resource availability govern
vertical stratification of small mammals in an Australian lowland
tropical rainforest? Wildl. Res. 33: 571–576.
Ragusa-Netto, J. (2008). Toco toucan feeding ecology and local abundance in a
habitat mosaic in the Brazilian cerrado. Ornitol. Neotrop. 19: 345–359.
Ribeiro, J.F. and Walter, B.M.T. (1998). Fitofisionomias do bioma cerrado. In:
Sano, S.M. and Almeida, S.P. (Eds.), Cerrado: ambiente e flora.
Embrapa, Brasília, pp. 89–167.
 H.F.M. Oliveira et al.: Arboreal marsupial recorded in pitfall traps
Rossi, R.V., Brandão, M.V., Carmignotto, A.P., Miranda, C.L., and Gerem, J.
(2012). Diversidade morfológica e taxonômica de marsupiais
didelfídeos, com ênfase nas espécies brasileiras. In: Cáceres, N.C.
(Ed.). Os marsupiais do Brasil: biologia, ecologia e evolução. Editora
UFMS, Campo Grande, pp. 23–72.
Santana, R.A.S.D., Dias-Júnior, C.Q., Vale, R.S.D., Tóta, J., and Fitzjarrald, D.R.
(2017). Observing and modeling the vertical wind profile at multiple
sites in and above the Amazon rain forest canopy. Adv. Meteorol. 2017:
1–8, https://doi.org/10.1155/2017/5436157.
Santiago, J., Junior, M.C.S., and Lima, L.C. (2005). Fitossociologia da
regeneração arbórea na mata de galeria do Pitoco (INGE-DF), seis
anos após fogo acidental. Sci. Flor. 67: 64–77.
Santos-Filho, M.D., Lázari, P.R.D., Sousa, C.P.F.D., and Canale, G.R. (2015).
Trap efficiency evaluation for small mammals in the southern Amazon.
Acta Amazon 45: 187–194.
Schmitt, D. and Lemelin, P. (2002). Origins of primate locomotion: gait
mechanics of the woolly opossum. Am. J. Phys. Anthropol. 118: 231–238.
7
Schmitt, D. and Lemelin, P. (2004). Locomotor mechanics of the slender loris
(Loris tardigradus). J. Human Evol. 47: 85–94.
van der Ree, R., Bennett, A., and Gilmore, D. (2004). Gap-crossing by gliding
marsupials: thresholds for use of isolated woodland patches in an
agricultural landscape. Biol. Conserv. 115: 241–249.
Vieira, E.M. and Camargo, N.F. (2012). Padrões do uso vertical do habitat por
marsupiais brasileiros. In: Cáceres, N.C. and Monteiro-Filho, E.L.A.
(Eds.), Os marsupiais do Brasil: biologia, ecologia e evolução. Editora
UFMS, Campo Grande, pp. 347–364.
Widyastuti, K., Reuillon, R., Chapron, P., Abdussalam, W., Nasir, D., Harrison,
M.E., Morrogh-Bernard, H., Imron, M.A., and Berger, U. (2022).
Assessing the impact of forest structure disturbances on the arboreal
movement and energetics of orangutans: an agent-based modeling
approach. Front. Ecol. Evol. 10: 983337.
Zommers, Z., Macdonald, D.W., Johnson, P.J., and Gillespie, T.R. (2013).
Impact of human activities on chimpanzee ground use and parasitism
(Pan troglodytes). Conserv. Lett. 6: 264–273.