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: [email protected].
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, an anti-predator behavior as documented in other rodent
Amazonas, from February 6 to March 6, 2017. We selected species (see Emmons and Feer 1997) and was witnessed
three sampling areas surrounding the communities of with other echimyid rodents within the area (e.g. Isothrix
Itapuru (−4.290751 S, −61.806055 W; right bank of Rio spp. and Makalata spp.). We also collected at precisely the
Purus), Caua-Cuiuanã (−4.234045 S, −61.734642 W; left same place, within minutes, a specimen of the Amazon
bank of Rio Purus) and Uixi (−4.435613 S, −62.306623 W; Bamboo Rat, Dactylomys dactylinus (Desmarest 1817),
left bank of Rio Purus). Within these areas we sampled showing that these two echimyid species co-occur in
along small rivers in seasonally inundated floodplain this habitat. Diurnal sampling uncovered no specimen of
forests (“várzea”) and in non-flooded upland forests Toromys. Only specimens of genus Isothrix (Wagner, 1845)
(“terra firme”). In each area we spent about 8 days under- were observed during the day censuses.
taking diurnal and nocturnal searches for small mammals The specimen EFA21 is an adult female that we iden-
(Rodentia and Didelphimorphia). To capture the speci- tified as Toromys grandis as it has both quantitative and
mens we used a 28 gauge shotgun and a 5.5  mm caliber qualitative traits with specimens diagnosed by Iack-
air gun. The specimens were collected and prepared fol- Ximenes et  al. (2005), and also with specimens from
lowing the recommendations of the Animal Care and MZUSP and MPEG. Our specimen is large with: head and
Use Committee of the American Society of Mammalogists body length = 280  mm; tail length = 253  mm; hind foot
(Sikes and The Animal Care and Use Committee 2016) and with claws = 50  mm; hind foot without claws = 45  mm;
will be deposited at Coleção do Laboratório de Mamíferos ears length = 23  mm; and body mass = 795 g. Externally,
da Escola Superior de Agricultura “Luiz de Queiroz”, Uni- the dorsal pelage is harsh, with stiff, black and gold hairs
versidade de São Paulo (LMUSP). Collections were made (long and thin); the head is predominantly black; the
under permits issued by the “Instituto Chico Mendes de lateral pelage is similar to dorsum, but paler. The ventral
Conservação da Biodiversidade” (SISBIO 8499-1) and by pelage is grayish, with a silver tinge. The dorsal surface of
the “Departamento de Mudanças Climáticas e Gestão de manus and pes are covered by short black hairs; the claws
Unidades de Conservação da Secretaria Estadual de Meio are robust and covered by short ungual black hairs. The
Ambiente do Amazonas” (DEMUC/SEMA 09/2017). tail is densely and entirely covered by short black hairs,
We examined the specimen of Toromys grandis col- except for the basal portion, which is covered by black
lected in the lower Rio Purus, as well as specimens housed and golden hairs (Figure 1). On the skull, the nasal bones
at the Museu de Zoologia da Universidade de São Paulo are constricted medially; the frontal with lateral wings are
(MZUSP) and the Museu Paraense Emílio Goeldi (MPEG). well-developed and form a roof over the orbital region;
To compile collecting information we searched VertNet the postorbital process of the zygoma is formed by jugal
database (http://vertnet.org/) for T. grandis and in online and squamosal; the buccinator and masticator foramina
catalogs of the major North-American scientific collec- are separated from each other; the transverse canal is well
tions. To identify (external and cranial morphology) the developed; the external auditory meatus is separated from
specimen we followed Emmons (2005) and Iack-Ximenes squamosal bone by a thin ridge of petrosal bone (Figure 2).
et al. (2005). The analysis of the digestive tract morphol- This specimen was pregnant with two fetuses, one
ogy followed Emmons (1981) and Leite (2003); meaning in each ramus of the uterus. The fetuses were in the
that we compared the size of the segments of the digestive final stages of development and measured 65 and 70 mm
tract of different species of arboreal echimyids using the head and body length, 63 and 61 tail length, respectively,
mean relative size, which is expressed in percent of head and 17.5 mm (both) hind foot length including the claws
and body length. (Figure 3A). We measured and compared the gut morphol-
During the sampling effort we collected 24 specimens ogy of specimen EFA21  with information available for
of rodents and marsupials, including one Toromys grandis four other species of arboreal echimyids. The segments of
(cataloged with collector’s field number EFA21). This the digestive tract of Toromys grandis are larger than the
specimen was collected near the confluence of Igarapé do segments of all other species reported in the literature
Caua with Rio Purus, at the left bank of Rio Purus, on Feb- (Figure 3B; Table 1). The small intestine and the caecum
ruary 17, 2017, during a nocturnal search, at around 10:00 of T. grandis are about twice the relative length of the
pm. The area is characterized by seasonally flooded forest comparable regions of Phyllomys pattoni (Emmons, Leite,
with many palm trees and patches of bamboo. On this Kock, and Costa, 2002) (so far the species with the longest
occasion we sighted two other specimens of T. grandis, known small intestine; Leite 2003) and Dactylomys dacty-
however, when we approached them, they both dived into linus (so far the species with the longest known caecum;
the water and swam under the surface. This appears to be 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). Figure 2: Skull and mandible of Toromys grandis (EFA21; greatest
Dorsal view (left) and ventral view (right). skull length = 63.99 mm).
Skull on dorsal, ventral, and lateral view, and mandible on lateral
view.

third longer than that of D. dactylinus. The stomach of T.

grandis is very similar to that of D. dactylinus in morphol- für Naturkunde – MNK, n = 1; Naturhistorika Riksmu-
ogy (simple) and size (see Emmons 1981). Long and exten- seet  – NRM, n = 10; and Naturhistorisches Museum
sively folded intestines and well-developed caecum are – NMW, n = 1), none of which were collected after the
indications of a primarily herbivorous diet (Leite 2003). 1950s (G.  Iack-Ximenes, personal communication, 2017).
However, Emmons (1981) highlights that a long small Our online search for specimens of T. grandis within
intestine is more associated with an omnivorous diet and Brazilian and North-American collections allowed us to
a long large intestine is correlated with an herbivorous find: 14 specimens at MPEG, collected from 1909 to 1956
diet. Both intestines were extremely long in the Giant Tree – only three specimens included information on the col-
Rat, suggesting that this species is specialized to a her- lector (two collected by I.O.C. and one by M. Moreira);
bivorous diet but also well adapted to an omnivorous diet. 116 specimens at the American Museum of Natural History
The stomach contents of this individual included a highly (AMNH), all collected by the Olalla brothers between 1930
digested pulp (not identifiable), with very small fragments and 1931; 15  specimens at the Museum of Comparative
of arthropod exoskeleton and very small seeds. Zoology, Harvard University (MCZ), collected by Alfonso
Iack-Ximenes et  al. (2005) examined 57  specimens M. Olalla from 1932 to 1933; and three specimens at The
of  Toromys grandis deposited in nine scientific col- Field Museum of Natural History (FMNH), one collected
lections  in Brazil (MZUSP, n = 16; Museu Nacional da by E. Snethlage in 1912 and two by Alfonso M. Olalla in
­Universidade do Rio de Janeiro – MN, n = 25) and Europe 1958. Our survey in other collections found no speci-
(The Natural History Museum – BMNH, n = 3; Museum 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

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
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. 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 surveys along six rivers of the Amazon basin (Abacaxis,
“Área de Proteção Ambiental do Arquipélago do Marajó”. Iça, Japurá, Jufari, Madeira and Xingu) collecting over
Our specimen likewise comes from a conservation unit at 1200  specimens of non-volant small mammals, and no
the RDS Piagaçu-Purus, making it more significant. All specimens of T. grandis were sighted or collected, despite
other sampling localities are from non-protected areas employing the same sampling methods. Moreover, no
(see Figure 4), although some records are in close proxim- published surveys for the Amazon region in the last
ity to conservation unit boundaries. decades have recorded this species (e.g. George et al. 1988
Iack-Ximenes et al. (2005) suggested that the narrow in the Rio Tapajos, Patton et  al. 2000 in the Rio Juruá,
geographical distribution of Toromys grandis is probably Voss et al. 2001 in the Paracou region, da Silva et al. 2007
a sampling bias and further surveys may demonstrate the in the Rio Madeira, Gettinger et  al. 2012 in the Floresta
presence of this species along other tributaries of the Rio Nacional de Carajás, Rossi et al. 2017 in seven sites in the
Amazonas. We only partially agree with this, since during northern State of Pará, and Abreu-Júnior et al. 2016, 2017
the past 6 years we have made short- and long-term field 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 da Silva, M.N.F., M.C. Arteaga, C.G. Bantel, D.M. Rossoni, R.N.
that T. grandis occupies a small geographic range. Leite, P.S. Pinheiro, F. Röhe and E. Eler. 2007. Mamíferos de
pequeno porte (Mammalia: Rodentia and Didelphimorphia).
Our record from the lower Purus represents an impor-
In: (L.R. Py-Daniel, C.P. Deus, A.L. Henriques, D.M. Pimpão
tant rediscovery of a poorly known species of echimyid and O.M. Ribeiro, eds.) Biodiversidade do Médio Madeira:
rodent from the Amazon forest. It also highlights the Bases científicas para propostas de conservação. Instituto
importance of continuous survey in this biome, whether Nacional de Pesquisas da Amazônia, Manaus, AM.
on short- or long-term approaches. Our study shows this pp. 179–194.
Emmons, L.H. 1981. Morphological, Ecological, and Behavioral
species to be nocturnal and potentially herbivorous, and
Adaptations for Arboreal Browsing in Dactylomys dactylinus
can occur in close syntopy with Dactylomys dactylinus.
(Rodentia, Echimyidae). J. Mamma. 62: 183–189.
This raises questions about the niche partitioning of these Emmons, L.H. 2005. A revision of the genera of arboreal Echimyidae
two species in the unique and highly dynamic seasonally (Rodentia, Echimyidae, Echimyinae); with description of two
inundated floodplain forests of the Amazon. new genera. Zoology 133: 247–310.
Emmons, L.H. and F. Feer. 1997. Neotropical rainforest mammals: a
field guide. 2nd ed. The University Chicago Press, Chicago, IL.
Acknowledgments: We are grateful for the financial sup- pp. 307.
port provided by the American Society of Mammalo- Emmons, L.H., Y.L.R. Leite and J.L. Patton. 2015. Genus Toromys
Iack-Ximenes, de Vivo, and Percequillo, 2005. In: (J.L. Patton,
gists through the Latin American Student Field Research
U.F.J. Pardiñas and G. D’Elía, eds.) Mammals of South America,
Award and by the Conselho Nacional de Desenvolvimento volume 2, Rodents. The University of Chicago Press, Chicago
Científico e Tecnológico (CNPq) through the PhD fellow- and London. pp. 929–931.
ship, both to E.F. Abreu-Júnior; to Cristiano Neves de Fabre, P.H., N.S. Upham, L.H. Emmons, F. Justy, Y.L.R. Leite, A.C.
Oliveira and Maria do Carmo for logistical and bureau- Loss, L. Orlando, M.K. Tilak, B.D. Patterson and E.J.P. Douzery.
cratic support at RDS Piagaçu-Purus; to people who 2017. Mitogenomic phylogeny, diversification, and bioge-
ography of South American spiny rats. Mol. Biol. Evol. 34:
hosted us and helped us during the field work: Padre,
613–633.
Noca, Barbozinha and Bengala; to our field work assis- George, T.K., S.A. Marques, M. de Vivo, L.C. Branch, N. Gomes and
tants: Faris, Berge, Prefeito, and Bengala; to Eduardo R. Rodrigues. 1988. Levantamento de mamíferos do Parna –
Von Mühlen and Heloisa Brum to introducing the region Tapajós. Brasil Florestal 63: 33–41.
to us and for the logistical support in the field; to Jero- Gettinger, D., N. Ardente and F. Martins-Hatano. 2012. Peque-
nos mamíferos não-voadores (Roedores e Marsupiais). In:
nymo Dalapicolla and Ana Carolina Pavan for checking
(F.D. Martins, A.F. Castilho, J. Campos, F.M. Hatano and S.G.
specimens housed at the MPEG and AMNH, respectively. Rolim, eds.) Fauna da Floresta Nacional de Carajás, estudos
We are also thankful to Gilson E. Iack Ximenes for shar- sobre vertebrados terrestres. Nitro Imagens, São Paulo, SP.
ing with us valuable information on this species and to pp. 145–161.
Louise H. Emmons for critically reviewing the manuscript Iack-Ximenes, G.E., M. de Vivo and A.R. Percequillo. 2005. A new
and providing important suggestions, both on the Eng- genus for Loncheres grandis Wagner, 1845, with taxonomic
comments on other arboreal echimyids (Rodentia, Echimyidae).
lish language and scientific contents.
Arquivos do Museu Nacional 63: 89–112.
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Rats, Genus Phyllomys (Rodentia, Echimyidae), with descrip-
tion of two new species. University of California Publications
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