Mammalia 2017; aop

Short Note

Edson Fiedler de Abreu-Júnior*, Jacob Daniel Charters and Alexandre Reis Percequillo
The Giant Tree Rat, Toromys grandis (Wagner, 1845): new record
with range extension and comments on its morphology, biology
and conservation
https://doi.org/10.1515/mammalia-2017-0042 hypotheses regarding Toromys (Upham and Patterson
Received April 13, 2017; accepted October 4, 2017 2015, Fabre et al. 2017) recovered it as a sister-group of Pat-
tonomys, a sister-clade to the genera: Makalata (Husson,
Abstract: We present data on a new record of Toromys
1978), Phyllomys (Lund, 1839) and Echimys F. (Cuvier,
grandis, a species not sampled for almost 60  years. The
1809). Toromys is characterized by large size [average
species is known from the newly-collected specimen and
head and body length ranging from 237  mm in Toromys
206  specimens housed in museums worldwide. The gut
rhipidurus (Thomas, 1928) to 307  mm in Toromys grandis
morphology, consisting of very long intestines and well-
(Wagner, 1845)], a well-haired tail (representing 80–90%
developed caecum, suggest that T. grandis is a potential
of the head and body length), body covered with stiff seti-
herbivore also adapted to omnivory. Our record at the
form and aristiform hairs, forefeet and hindfeet with small
Reserva de Desenvolvimento Sustentável (RDS) Piagaçu-
tubercular rugosities between the plantar and palmar
Purus extends the range of T. grandis approximately 150
pads, and short rounded ears. In regards to craniodental
km southwestward and represents the second record of
morphology, Toromys is distinguished by the nasal bones
the species within a protected area.
constricted medially, the squamosotympanic fenestra as a
Keywords: Amazon Forest; arboreal echimyid; digestive narrow horizontal slit, the supraorbital ridges expanded
tract; geographical distribution; protected area. into a shelf, and the protoloph and metaloph of molari-
forms often connected by a slender mure (Iack-Ximenes
et al. 2005, Emmons et al. 2015).
The giant tree rat or giant toro, Toromys grandis, is a
The genus Toromys (Rodentia, Echimyidae) was described
nocturnal arboreal rat that inhabits lowland rainforests,
by Iack-Ximenes, de Vivo and Percequillo (2005), based on
being restricted to floodplain forests (Emmons and Feer
the species Loncheres grandis Wagner, 1845, which had
1997, Iack-Ximenes et al. 2005). It is known from 16 locali-
previously been moved between various genera during the
ties, mostly in the central Amazon region, along both sides
last century (see generic synonymy in Iack-Ximenes et al.
of lower Rio Amazonas, in the lower Rio Solimões, and in
2005). Emmons et al. (2015) later added Echimys rhipidu-
the mid-lower Rio Tapajos (Iack-Ximenes et al. 2005). Little
rus Thomas, 1928 to the Toromys genus; another species of
is known about the systematics, biology and conservation
uncertain position. Emmons (2005) had previously united
status of this species. Most available information, includ-
E. rhipidurus and L. grandis within a “grandis group” based
ing geographic distribution, is from 57 voucher specimens
on morphological affinities. The most recent phylogenetic
of T. grandis reported by Iack-Ximenes et al. (2005), which
represented most of the available specimens within Bra-
*Corresponding author: Edson Fiedler de Abreu-Júnior, Laboratório zilian and European collections.
de Mamíferos, Departamento de Ciências Biológicas, Escola Here, we report on a new specimen of Toromys grandis
Superior de Agricultura “Luiz de Queiroz”, Universidade de São that is apparently the first individual collected since
Paulo, Av. Pádua Dias 11, Caixa Postal 9, 13418-900, Piracicaba,
1958. This compelled us to review the information from
São Paulo, Brazil, e-mail: edson.abreu@usp.br.
http://orcid.org/0000-0003-4547-6160 148 specimens not analyzed by Iack-Ximenes et al. (2005)
Jacob Daniel Charters: Laboratório de Ecologia Isotópica, Centro – housed in one Brazilian and three North-­ American
de Energia Nuclear na Agricultura, Universidade de São Paulo, museums – providing a new map of geographical distribu-
Av. Centenário 303, Caixa-Postal 96, 13416-000, Piracicaba, tion with a range extension for the species. Based on the
São Paulo, Brazil
new specimen, we comment on its morphology, biology
Alexandre Reis Percequillo: Laboratório de Mamíferos,
Departamento de Ciências Biológicas, Escola Superior de
and conservation.
Agricultura “Luiz de Queiroz”, Universidade de São Paulo, Av. Pádua We made a short-term collecting expedition (sensu Voss
Dias 11, Caixa Postal 9, 13418-900, Piracicaba, São Paulo, Brazil and Emmons 1996) to the Reserva de Desenvolvimento
2      E.F. de Abreu-Júnior et al.: New data on Toromys grandis (Rodentia: Echimyidae)
Sustentável (RDS) Piagaçu-Purus, on the lower Rio Purus,
Amazonas, from February 6 to March 6, 2017. We selected
three sampling areas surrounding the communities of
Itapuru (−4.290751 S, −61.806055 W; right bank of Rio
Purus), Caua-Cuiuanã (−4.234045 S, −61.734642 W; left
bank of Rio Purus) and Uixi (−4.435613 S, −62.306623 W;
left bank of Rio Purus). Within these areas we sampled
along small rivers in seasonally inundated floodplain
forests (“várzea”) and in non-flooded upland forests
(“terra firme”). In each area we spent about 8 days under-
taking diurnal and nocturnal searches for small mammals
(Rodentia and Didelphimorphia). To capture the speci-
mens we used a 28 gauge shotgun and a 5.5  mm caliber
air gun. The specimens were collected and prepared fol-
lowing the recommendations of the Animal Care and
Use Committee of the American Society of Mammalogists
(Sikes and The Animal Care and Use Committee 2016) and
will be deposited at Coleção do Laboratório de Mamíferos
da Escola Superior de Agricultura “Luiz de Queiroz”, Uni-
versidade de São Paulo (LMUSP). Collections were made
under permits issued by the “Instituto Chico Mendes de
Conservação da Biodiversidade” (SISBIO 8499-1) and by
the “Departamento de Mudanças Climáticas e Gestão de
Unidades de Conservação da Secretaria Estadual de Meio
Ambiente do Amazonas” (DEMUC/SEMA 09/2017).
We examined the specimen of Toromys grandis col-
lected in the lower Rio Purus, as well as specimens housed
at the Museu de Zoologia da Universidade de São Paulo
(MZUSP) and the Museu Paraense Emílio Goeldi (MPEG).
To compile collecting information we searched VertNet
database (http://vertnet.org/) for T. grandis and in online
catalogs of the major North-American scientific collec-
tions. To identify (external and cranial morphology) the
specimen we followed Emmons (2005) and Iack-Ximenes
et al. (2005). The analysis of the digestive tract morphol-
ogy followed Emmons (1981) and Leite (2003); meaning
that we compared the size of the segments of the digestive
tract of different species of arboreal echimyids using the
mean relative size, which is expressed in percent of head
and body length.
During the sampling effort we collected 24 specimens
of rodents and marsupials, including one Toromys grandis
(cataloged with collector’s field number EFA21). This
specimen was collected near the confluence of Igarapé do
Caua with Rio Purus, at the left bank of Rio Purus, on Feb-
ruary 17, 2017, during a nocturnal search, at around 10:00
pm. The area is characterized by seasonally flooded forest
with many palm trees and patches of bamboo. On this
occasion we sighted two other specimens of T. grandis,
however, when we approached them, they both dived into
the water and swam under the surface. This appears to be
an anti-predator behavior as documented in other rodent
species (see Emmons and Feer 1997) and was witnessed
with other echimyid rodents within the area (e.g. Isothrix
spp. and Makalata spp.). We also collected at precisely the
same place, within minutes, a specimen of the Amazon
Bamboo Rat, Dactylomys dactylinus (Desmarest 1817),
showing that these two echimyid species co-occur in
this habitat. Diurnal sampling uncovered no specimen of
Toromys. Only specimens of genus Isothrix (Wagner, 1845)
were observed during the day censuses.
The specimen EFA21 is an adult female that we iden-
tified as Toromys grandis as it has both quantitative and
qualitative traits with specimens diagnosed by Iack-
Ximenes et  al. (2005), and also with specimens from
MZUSP and MPEG. Our specimen is large with: head and
body length = 280  mm; tail length = 253  mm; hind foot
with claws = 50  mm; hind foot without claws = 45  mm;
ears length = 23  mm; and body mass = 795 g. Externally,
the dorsal pelage is harsh, with stiff, black and gold hairs
(long and thin); the head is predominantly black; the
lateral pelage is similar to dorsum, but paler. The ventral
pelage is grayish, with a silver tinge. The dorsal surface of
manus and pes are covered by short black hairs; the claws
are robust and covered by short ungual black hairs. The
tail is densely and entirely covered by short black hairs,
except for the basal portion, which is covered by black
and golden hairs (Figure 1). On the skull, the nasal bones
are constricted medially; the frontal with lateral wings are
well-developed and form a roof over the orbital region;
the postorbital process of the zygoma is formed by jugal
and squamosal; the buccinator and masticator foramina
are separated from each other; the transverse canal is well
developed; the external auditory meatus is separated from
squamosal bone by a thin ridge of petrosal bone (Figure 2).
This specimen was pregnant with two fetuses, one
in each ramus of the uterus. The fetuses were in the
final stages of development and measured 65 and 70 mm
head and body length, 63 and 61 tail length, respectively,
and 17.5 mm (both) hind foot length including the claws
(Figure 3A). We measured and compared the gut morphol-
ogy of specimen EFA21  with information available for
four other species of arboreal echimyids. The segments of
the digestive tract of Toromys grandis are larger than the
segments of all other species reported in the literature
(Figure 3B; Table 1). The small intestine and the caecum
of T. grandis are about twice the relative length of the
comparable regions of Phyllomys pattoni (Emmons, Leite,
Kock, and Costa, 2002) (so far the species with the longest
known small intestine; Leite 2003) and Dactylomys dacty-
linus (so far the species with the longest known caecum;
Leite 2003). The large intestine is also very long, about one
 E.F. de Abreu-Júnior et al.: New data on Toromys grandis (Rodentia: Echimyidae)      3
Figure 1: External morphology of Toromys grandis (EFA21; head and
body length = 280 mm and tail length = 253 mm).
Dorsal view (left) and ventral view (right).
third longer than that of D. dactylinus. The stomach of T.
grandis is very similar to that of D. dactylinus in morphol-
ogy (simple) and size (see Emmons 1981). Long and exten-
sively folded intestines and well-developed caecum are
indications of a primarily herbivorous diet (Leite 2003).
However, Emmons (1981) highlights that a long small
intestine is more associated with an omnivorous diet and
a long large intestine is correlated with an herbivorous
diet. Both intestines were extremely long in the Giant Tree
Rat, suggesting that this species is specialized to a her-
bivorous diet but also well adapted to an omnivorous diet.
The stomach contents of this individual included a highly
digested pulp (not identifiable), with very small fragments
of arthropod exoskeleton and very small seeds.
Iack-Ximenes et  al. (2005) examined 57  specimens
of  Toromys grandis deposited in nine scientific col-
lections  in Brazil (MZUSP, n = 16; Museu Nacional da
­Universidade do Rio de Janeiro – MN, n = 25) and Europe
(The Natural History Museum – BMNH, n = 3; Museum
Figure 2: Skull and mandible of Toromys grandis (EFA21; greatest
skull length = 63.99 mm).
Skull on dorsal, ventral, and lateral view, and mandible on lateral
view.
für Naturkunde – MNK, n = 1; Naturhistorika Riksmu-
seet  – NRM, n = 10; and Naturhistorisches Museum
– NMW, n = 1), none of which were collected after the
1950s (G.  Iack-Ximenes, personal communication, 2017).
Our online search for specimens of T. grandis within
Brazilian and North-American collections allowed us to
find: 14 specimens at MPEG, collected from 1909 to 1956
– only three specimens included information on the col-
lector (two collected by I.O.C. and one by M. Moreira);
116 specimens at the American Museum of Natural History
(AMNH), all collected by the Olalla brothers between 1930
and 1931; 15  specimens at the Museum of Comparative
Zoology, Harvard University (MCZ), collected by Alfonso
M. Olalla from 1932 to 1933; and three specimens at The
Field Museum of Natural History (FMNH), one collected
by E. Snethlage in 1912 and two by Alfonso M. Olalla in
1958. Our survey in other collections found no speci-
men of this species (e.g. Museum of Vertebrate Zoology,
4      E.F. de Abreu-Júnior et al.: New data on Toromys grandis (Rodentia: Echimyidae)

University of California; Smithsonian Institution National

Figure 3: Fetus (A) and digestive tract (B) of Toromys grandis (EFA21).
See Table 1 for the mean relative size of segments of digestive tract.
Museum of Natural History). Therefore, there are at least
206 specimens of T. grandis available in museums world-
wide, as assessed by Iack-Ximenes et  al. (2005) and the
current authors, plus our newly collected specimen. Apart
from the collected specimen we also saw two other indi-
viduals on the same night, within 3  h, suggesting that
this species is relatively common locally. Moreover, locals
reported that this species, known as “Rato Toró” (distinc-
tively from other arboreal echimyids, such as Isothrix spp.
and Makalata spp., that are known as “Ratos Corós”), is
indeed relatively common in flooded forests (“igapós”).
These records of geographical distribution of
Toromys grandis add seven more localities from museum
specimens (previously cited), plus our new record from
the lower Rio Purus, for a total of 24 localities (Figure 4).
Our record on the Purus River extends the distributional
range of T. grandis approximately 150 km towards the
southwest. We also have corrected some problems with
the map of known collection localities provided by Iack-
Ximenes et al. (2005; figure 2, p. 93), in which all locali-
ties were mislabeled: the locality numbers on the map
do not correspond to the locality names on the gazetteer
(Iack-Ximenes et al. 2005: appendix II, p. 111), following
the Ornithological Gazetteer of Brazil (Paynter Jr. and
Traylor Jr. 1991). As a consequence of this mislabeling,
some localities on the list of specimens examined (Iack-
Ximenes et  al. 2005: appendix I, p. 109) are not on the
map (Iack-Ximenes et al. 2005: figure 2, p. 93), and there
are some localities in that map that do not correspond to
collection localities of T. grandis.
According to Cassola (2016) there are no records of
Toromys grandis in conservation units. However, this is

Table 1: Mean relative size (expressed as percentage of head and body length) of segments of digestive tracts of arboreal echimyids.

  Phyllomys   Mesomys   Dactylomys dactylinus   Kannabateomys   Toromys grandis

pattoni (n = 1) hispidus (n = 2) (n = 2) amblyonyx (n = 3) (n = 1)

Head and body length (mm)  230   188   290   273   280
Small intestine
 Length   370   344   159   174   643
 Maximum width   2   3   5   NA   2.5
Large intestine
 Length   226   140   247   172   314
 Maximum width   4   2   7   NA   6
Caecum
 Length   57   48   66   37   125
 Maximum width   9   8   13   NA   10
Stomach length   NA   17   24   NA   23
Reference   Leite (2003)   Emmons (1981)   Emmons (1981)   Silva (1993)   Present report

Measurements were taken in specimens fixed with formalin and preserved in 70% ethanol, except for K. amblyonyx, which was measured
before fixation. NA, not available.
 E.F. de Abreu-Júnior et al.: New data on Toromys grandis (Rodentia: Echimyidae)      5

Figure 4: Updated map of geographical distribution of Toromys grandis.

Dots: localities reported by Iack-Ximenes et al. (2005) (Brazil: Amazonas: 1 – Lago do Batista, Rio Amazonas [−3.283333, −58.266667]; 2
– Manaqueri, Rio Solimões [−3.483333, −60.516667]; 3 – Silves [−2.833333, −58.216667]; 4 – Urucurituba, Rio Amazonas [−2.766667,
−57.816667]; Pará: 5 – Belterra, Santarém [−2.633333, −54.933333]; 6 – Bravo, north bank of Rio Amazonas [−1.9, −55.166667]; 7 –
Fazenda Recreio, Ilha Caviana [−0.166667, −50.166667]; 8 – Fazenda Paraíso, Faro, Rio Amazonas [−2.083333, −56.766667]; 9 – Lago
Cuiteua, north bank of Rio Amazonas [−1.816667, −54.966667]; 10 – Paraná do Bom Jardim, Paissandú, Nhamundá [−2.033333, −56.2];
11 – Paraná de Faro [−2.166667, −56.733333]; 12 – Santarém [−2.433333, −54.7]; 13 – Igarapé Açú, left bank of Rio Tapajós, Santarém
[−3.733333, −55.516667]; 14 – Fazenda São Pedro, Monte Alegre [−2.007778, −54.069167]; 15 – Óbidos [−1.916667, −55.516667]; 16 –
Santa Rita, Rio Amazonas [−2.033333, −55.3]). Squares: new localities from museum specimens (Brazil: Amazonas: 17 – Parintins, Serra
do Parintins, south bank of Rio Amazonas [−2.6, −56.733333]; Pará: 18 – Faro, north bank of Rio Amazonas [−2.183333, −56.733333]; 19
– Porto de Moz, Villarinho de Monte, Rio Xingu [−1.616667, −52.016667]; 20 – Rio Amazonas, Boca do Igarapé Piaba [−1.916667, −55.55];
21 – Rio Amazonas, Lago Jauary [−1.883333, −55.533333]; 22 – Rio Arapiuns, Santarém [−2.3, −55]; 23 – Rio Nhamunda, Fazenda Paraiso
[−2.18333, −55.5]). Star: present record at lower Purus River (Brazil: Amazonas: 24 – Confluency of Igarapé do Caua with Rio Purus, left bank
of Rio Purus, Comunidade do Caua-Cuiuanã [−4.243572, −61.762304]).

incorrect, as a record from Ilha Caviana, Pará, is in the
“Área de Proteção Ambiental do Arquipélago do Marajó”.
Our specimen likewise comes from a conservation unit at
the RDS Piagaçu-Purus, making it more significant. All
other sampling localities are from non-protected areas
(see Figure 4), although some records are in close proxim-
ity to conservation unit boundaries.
Iack-Ximenes et al. (2005) suggested that the narrow
geographical distribution of Toromys grandis is probably
a sampling bias and further surveys may demonstrate the
presence of this species along other tributaries of the Rio
Amazonas. We only partially agree with this, since during
the past 6 years we have made short- and long-term field
surveys along six rivers of the Amazon basin (Abacaxis,
Iça, Japurá, Jufari, Madeira and Xingu) collecting over
1200  specimens of non-volant small mammals, and no
specimens of T. grandis were sighted or collected, despite
employing the same sampling methods. Moreover, no
published surveys for the Amazon region in the last
decades have recorded this species (e.g. George et al. 1988
in the Rio Tapajos, Patton et  al. 2000 in the Rio Juruá,
Voss et al. 2001 in the Paracou region, da Silva et al. 2007
in the Rio Madeira, Gettinger et  al. 2012 in the Floresta
Nacional de Carajás, Rossi et al. 2017 in seven sites in the
northern State of Pará, and Abreu-Júnior et al. 2016, 2017
in upper Rio Acre and Rio Jufari, respectively). Therefore,
6      E.F. de Abreu-Júnior et al.: New data on Toromys grandis (Rodentia: Echimyidae)
considering the size of the Amazon basin, we hypothesize
that T. grandis occupies a small geographic range.
Our record from the lower Purus represents an impor-
tant rediscovery of a poorly known species of echimyid
rodent from the Amazon forest. It also highlights the
importance of continuous survey in this biome, whether
on short- or long-term approaches. Our study shows this
species to be nocturnal and potentially herbivorous, and
can occur in close syntopy with Dactylomys dactylinus.
This raises questions about the niche partitioning of these
two species in the unique and highly dynamic seasonally
inundated floodplain forests of the Amazon.
Acknowledgments: We are grateful for the financial sup-
port provided by the American Society of Mammalo-
gists through the Latin American Student Field Research
Award and by the Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) through the PhD fellow-
ship, both to E.F. Abreu-Júnior; to Cristiano Neves de
Oliveira and Maria do Carmo for logistical and bureau-
cratic support at RDS Piagaçu-Purus; to people who
hosted us and helped us during the field work: Padre,
Noca, Barbozinha and Bengala; to our field work assis-
tants: Faris, Berge, Prefeito, and Bengala; to Eduardo
Von Mühlen and Heloisa Brum to introducing the region
to us and for the logistical support in the field; to Jero-
nymo Dalapicolla and Ana Carolina Pavan for checking
specimens housed at the MPEG and AMNH, respectively.
We are also thankful to Gilson E. Iack Ximenes for shar-
ing with us valuable information on this species and to
Louise H. Emmons for critically reviewing the manuscript
and providing important suggestions, both on the Eng-
lish language and scientific contents.
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