Agrellos 2012

Zootaxa 3220: 1–28 (2012) ISSN 1175-5326 (print edition)
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Copyright © 2012 · Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
The taxonomic status of the Castelo dos Sonhos Hantavirus reservoir,

Oligoryzomys utiaritensis Allen 1916 (Rodentia: Cricetidae: Sigmodontinae)
RODRIGO AGRELLOS*1, CIBELE R. BONVICINO*1,2, ELIZABETH SALBÉ T. ROSA3, APARECIDO A. R.

MARQUES4, PAULO S. D’ANDREA1 & MARCELO WEKSLER5,6
1
Instituto Oswaldo Cruz, Fundação Oswaldo Cruz (FIOCRUZ), Avenida Brasil, 4365
Manguinhos, 21040-900, Rio de Janeiro, Brazil
2
Divisão de Genética, Instituto Nacional do Câncer (INCA), Rua André Cavalcanti, 37, Centro 20231-050, Rio de Janeiro, Brazil.
3
Instituto Evandro Chagas, Seção de Arbovirologia e Febres Hemorrágicas, Rodovia BR-316, Km 0, 67030-000, Ananindeua, Pará,
Brazil.
4
Secretaria de Saúde do Estado de Mato Grosso, Cuiabá, Mato Grosso, Brazil
5
Depto. de Vertebrados, Museu Nacional - Universidade Federal do Rio de Janeiro,
Quinta da Boa Vista, São Cristóvão, 20940-040, Rio de Janeiro, Brazil.
6
Corresponding author; E-mail: marcelo.weksler@gmail.com
* Both authors contributed equally to this paper
Abstract
Species of the genus Oligoryzomys are commonly found accross Latin America, and several of them play important roles
as natural reservoirs of Hantaviruses. Here we demonstrate that O. utiaritensis, the natural reservoir of hantavirus Castelo
dos Sonhos in northwestwern Brazil and previously considered a junior synonym of O. nigripes or O. eliurus, is a valid
species. Morphology, morphometry, karyotyping, and phylogenetic reconstructions based on nuclear (intron 7 of the beta-
fibrinogen gene) and mitochondrial (cytochrome b) DNA show that O. utiaritensis differs from O. nigripes and from other
forms of the genus, including the recently described taxon O. moojeni. Oligoryzomys utiaritensis differs in external (whit-
ish ventral pelage and tail weakly bicolored) and cranial (incisive foramina never extending posteriorly the alveolus line
of M1) characters from sympatric species. It has the highest diploid number (2n=72) within Oligoryzomys, and is charac-
terized by three putative synapomorphies in cytochrome b, and one in intron 7 of beta fibrinogen. We also point to the
assignment of Oligoryzomys eliurus as a junior synonym of O. nigripes. Finally, we present phylogenetic analyses of in-
trageneric relationships showing that O. utiaritensis is a member of a clade containing Amazonian and Cerrado taxa, in-
cluding O. moojeni, O. rupestris, and O. delicatus.
Keywords: pygmy rice rat, hantaviruses, South America, phylogeny, cytocrome b, intron 7 beta fibrinogen, karyotype,
morphometrics
Introduction
Oligoryzomys Bangs is a speciose and widespread rodent genus found in all Neotropical countries, from Mexico to
Argentina. Eighteeen Oligoryzomys species are currently recognized (Musser & Carleton, 2005; Weksler & Bonvicino,
2005), nine of which are putatively distributed in Brazil: Oligoryzomys chacoensis (Myers & Carleton), O. flave-
scens (Waterhouse), O. fornesi (Massoia), O. fulvescens (Saussure), O. microtis (Allen), O. moojeni Weksler &
Bonvicino, O. nigripes (Olfers), O. stramineus Bonvicino & Weksler, and O. rupestris Weksler & Bonvicino.
Through Brazil, Oligoryzomys species occupy a variety of habitats, from humid environments like the Atlantic For-
est, Pantanal, and Amazonia to drier environments like the Cerrado and Caatinga (Carleton & Musser, 1989), and from
the sea level to 3,000 m above sea level.
Oligoryzomys species inhabit several areas currently undergoing massive development in the “agricultural
frontier” of Central and Northern Brazil, where cases of Hantavirus pulmonary syndrome (HPS) have been
reported (Oliveira et al., 2009; Oliveira et al., 2011; Rosa et al., 2005; Rosa et al., 2010; Suzuki et al,. 2004). Four
Accepted by P. Gaubert: 27 Jan. 2012; published: 2 Mar. 2012 1

Brazilian Oligoryzomys species are known hantavirus reservoirs: O. nigripes, the host of Juquitiba-like Virus
(Oliveira et al., 2009; Suzuki et al., 2004 ), O. flavescens, host of Central Plata Virus (Delfraro et al., 2003), O.
fornesi, host of Anajatuba Virus (Rosa et al., 2005), and O. microtis, host of Rio Marmoré Virus (Ritcher et al.,
2010). A fifth form of Brazilian hantavirus, the Castelo dos Sonhos Virus, has been found in Mato Grosso and Pará
states (Medeiros et al., 2010) from a reservoir belonging to the genus Oligoryzomys, the specific identity of which
is still uncertain. This new virus is found in a region of intense agricultural activity in the ecotone between Amazo-
nia and Cerrado, and the crucial role of Oligoryzomys in Hantavirus transmission makes the understanding of its
taxonomy, distribution, evolutionary history, and ecology necessary. However, despite recent phylogenetic studies
of the genus (González-Ittig et al., 2010; Hanson et al., 2011; Miranda et al., 2009; Palma et al., 2010; Rivera et
al., 2007; Rogers et al., 2009), Oligoryzomys systematics is still highly controversial due to the high phenotypic
similarity between species, which makes diagnosis and species identification difficult (see Carleton & Musser,
1989 and Weksler & Bonvicino, 2005 for historical accounts of the generic taxonomy).
In the present study, we report morphologic, karyologic, and molecular (mitochondrial and nuclear) data on the
host species of the Castelo dos Sonhos Hantavirus. Some specimens herein analyzed were collected ca. 50 km from
the type locality of Oligoryzomys utiaritensis, a species described by Allen (1916) based on specimens collected by
the Roosevelt-Rondon Expeditions in Utiariti, Rio Papagaio, Mato Grosso (Allen, 1916). Almost a century after its
original description, the identity of O. utiaritensis is still controversial. In the summary report on the genus Oligo-
ryzomys by Carleton & Musser (1989), O. utiaritensis was assigned as a probable junior synonym of O. eliurus
Wagner (itself considered sometimes a synonym of O. nigripes; see discussion), a position still sustained by
Musser & Carleton (2005) in the latest compendium of mammals of the world (Wilson & Reeder, 2005). We herein
present evidence that O. utiaritensis is a valid species and provide an emended diagnosis and re-description of this
enigmatic taxon. We also carry out a phylogenetic analysis of this genus, and discuss the taxonomic identication of
some GenBank nucleotide sequences that had been studied by molecular-only approaches.
Material and methods
Specimens. We examined 282 Oligoryzomys specimens deposited in the following collections: Museu Nacional,
Universidade Federal do Rio de Janeiro (MN), Rio de Janeiro, Brazil; American Museum of Natural History
(AMNH), New York, USA — including the type series of O. utiaritensis and O. mattogrossae (Allen); and Museu
de Zoologia da Universidade de São Paulo (MZUSP), São Paulo, Brazil. The holotype of Hesperomys eliurus at the
Natural History Museum (NHMV), Vienna, Austria was also analyzed, through photographs and personal exami-
nation by Dr. A. Langguth. Additional information on collection locality data (Figure 1) and museum acronyms
and numbers are presented in Appendix 1 (see also Bonvicino & Weksler 1998, and Weksler & Bonvicino 2005,
for previously analyzed specimens of other Oligoryzomys species). Four of the recently collected specimens were
positive carriers of Castelo dos Sonhos hantavirus (Rosa et al., 2011): MN75063 (field number CNP-SVS35),
MN75064 (CNP-SVS103), MN74939 (SVS328), MN74965 (CNP-SVS39).
The terminology and illustration of characters herein analyzed were reported by Carleton (1980), Reig (1977),
Voss (1988), Carleton & Musser (1989), Voss (1993), Voss & Carleton (1993), Steppan (1995) and Weksler (2006).
External and cranial measurements. The following external dimensions were measured (in millimeters) in
specimens collected by us or obtained from original specimen tags: head and body length (HBL), tail length (LT),
ear length (Ear), hind foot length with claw (HF) and body mass (Wt). Whenever Total Length had been originally
reported in specimen tags, HBL was estimated by subtracting tail length (TL) from total length. Cranial measure-
ments were taken with digital calipers to the nearest 0.01 mm. For morphometric analyses, we employed 12 cranial
dimensions following Bonvicino & Weksler (1998): condylo-incisive length (CIL), length of diastema (LD), pala-
tal bridge (PB), length of maxillary molars (LM), breadth of first maxillary molar (BM1), external alveolar breadth
(M1M), length of incisive foramen (LIF), breadth of incisive foramen (BIF), rostrum breadth (BRO), orbital length
(ORL), zygomatic breadth (ZB), and breadth of zygomatic plate (BZP). These variables were chosen because they
provided consistent values by different investigators (i.e., did not display inter-researcher error; unpublished data).
Statistical analyses. Morphometric analyses of skull characters were performed for adult specimens (i.e.,
specimens with all teeth erupted and with at least minimal wear; Oliveira et al., 1998); males and females were
grouped due to lack of sexual dimorphism (t-tests, p<0.05; not shown). Analysis of Variance (ANOVA) with the
2 · Zootaxa 3220 © 2012 Magnolia Press AGRELLOS ET AL.

post hoc test Fisher’s Least Significant Difference (LSD; Sokal & Rohlf, 1994) and Canonical Discriminant Anal-
ysis with estimation of Squared Mahalanobis Distances (Strauss, 2010) using logarithmic-transformed data were
carried out for identifying patterns of morphometric variation and for comparing O. utiaritensis with O. nigripes
and the closely related species O. moojeni (see phylogenetic results). Statistical analyses were performed with
STATISTICA 7.0 (StatSoft Inc., 2004).
FIGURE 1. Map showing the localities of O. utiaritensis (square), O. moojeni (triangles), and O. nigripes (circles). Open sym-
bols are type localities. Localities of O. nigripes are based on newly karyotyped specimens (Appendix 1), Weksler & Bonvicino
(2005), Paresque et al. (2007), and Miranda et al. (2009). Localities of O. utiaritensis and O. moojeni are: Brazil, Pará state (1)
Castelo dos Sonhos; Mato Grosso state (2) Peixoto de Azevedo (3) Feliz Natal, (4) Sapezal (5) Utiariti, Rio Papagaio, (6)
Campo Novo do Parecis; Tocantins state (7) Lajeado, (8) Porto Nacional, (9) Dianópolis, (10) Novo Jardim; Goiás state (11)
Minaçu, (12) Cavalcante (type locality of O. moojeni), (13) Colinas do Sul, (14) Uruaçu, (15) Niquelância, (16) Sítio D’Aba-
dia, and (17) Mimoso de Goiás. A= PN Ybicuí, Paraguay, type locality of O. nigripes, B= Itataré, São Paulo state, Brazil, type
locality of O. eliurus, C= Conceição do Mato Dentro, Minas Gerais state, Brazil. Gray area corresponds to the Cerrado morpho-
climatic domain.
Karyotypic analysis. We karyotyped 25 specimens from four populations of Oligoryzomys utiaritensis, three
specimens of O. nigripes, and six specimens of O. moojeni (Appendix 1). Chromosome preparations were obtained
with short-term bone marrow cultures incubated for two hours (around 37oC) in 15 ml Falcon tubes containing ster-
ile medium (80% RPMI, 20% fetal calf serum, 5 μmg/ml of ethidium bromide and 10-6 M colchicine). Chromo-
somes were ordered according to morphology and decreasing size, fundamental numbers referring to the autosome
complement.
Molecular data. DNA was isolated from livers preserved in 100% ethanol following the standard phenol-
chloroform protocol (Sambrook & Russell, 2001). The full-length cytochrome b gene (cyt-b) was amplified by
TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS Zootaxa 3220 © 2012 Magnolia Press · 3

standard PCR procedures with the primer pair L14724 (Irwin et al., 1991) and Citb-Rev (Casado et al., 2010).
Amplifications were performed in 50 μL reactions with Platinum® Taq Polymerase (Invitrogen™) and concentra-
tions of primers and templates as recommended by the manufacturer. Reactions were run for 35 cycles at 94°C for
30 s, 58°C for 30 s, and extension at 72°C for 90 s, with initial denaturation at 94°C for 2 min and final extension at
72°C for 7 min. We also amplified intron 7 of the nuclear β-fibrinogen gene (i7FGB) with the primer pair β17-
mammL and βfib-mammU as reported by Matocq et al. (2007). Amplifications were performed with 30 cycles of
denaturation at 94°C for 1 min, 60°C for 30 s, and extension at 72°C for 1 min, with an initial denaturation at 94°C
for 5 min and final extension at 72°C for 7 min.
Amplicons were purified with GFX® PCR DNA and Gel Band Purification Kit (GE™ Healthcare) and
sequenced both with the primers used in the PCR amplification and two additional internal primers for cyt-b,
namely MEU1 (Bonvicino & Moreira, 2001) and MVZ16 (Smith & Patton, 1993). Sequencing reactions were run
in an ABI3130xl (Applied Biosystems™) platform and electropherograms were manually checked and aligned
using BioEdit 8.0 (Hall, 1999) and Chromas 1.45 (McCarthy, 1998). Resulting sequences (n = 32 for cyt-b and n =
52 for i7FGB; Appendix 2) were deposited in GenBank (cyt-b: JQ013746-JQ013777; i7FGB: JQ282842-
JQ282893).
Additional cyt-b data from other 43 GenBank Oligoryzomys specimens (Appendix 2) were used for phyloge-
netics reconstructions. Outrgoup taxa included Oreoryzomys balneator (EU579510), Microryzomys minutus
(AF108698), Oryzomys palustris (GU126539) and Neacomys spinosus (EU579504) for constructing the cyt-b tree,
and Oryzomys palustris (AY274219), Holochilus sciureus (AY274218) and Sigmodon hirsutus (EU652893) for the
i7FGB tree. Analyses were performed seperately for each gene because datasets had different sets of OTUs; final
alignments consisted of 1,140 bp for cytochrome b and 729 pb for i7FGB.
Phylogenetic analyses. Maximum likelihood (Felsenstein, 1981) and Bayesian analysis (Huelsenbeck et al.,
2001) were carried out for phylogenetic reconstructions. Maximum-likelihood trees were estimated with RaxML-
HPC 7.2.8 (Stamatakis, 2006) at the CIPRES Science Gateway (Miller et al., 2010). The GTR model of nucleotide
substitution (Rodríguez et al., 1990) corrected for site-specific rate heterogeneity using gamma distribution with
four classes (Yang, 1994) was used for cyt-b, while the K81 model of nucleotide substitution (Kimura, 1981) cor-
rected for site-specific rate heterogeneity using gamma distribution with four classes (Yang, 1994) was used for
i7FGB; both models were selected using the Akaike Information Criterion (AIC) in jModelTest 0.1.1 (Posada,
2008). Nodal bootstrap values (Felsenstein, 1985) for the likelihood analyses were calculated using 1,000 pseu-
doreplicates. Bayesian analyses were performed using Markov chain Monte Carlo (MCMC) sampling as imple-
mented in MrBayes 3.1.2 (Ronquist & Huelsenbeck, 2003), also at the CIPRES Science Gateway (Miller et al.,
2010). Uniform interval priors were assumed for all parameters except base composition for which we assumed a
Dirichlet prior. We performed four independent runs of 10,000,000 generations each, with 2 heated chains sam-
pling for trees and parameters every 10,000 generations. The first 10,000 generations were discarded as burn-in,
and the remaining trees were used to estimate posterior probabilities for each node. All analyses were checked for
convergence by plotting the log-likelihood values against generation time for each run with Tracer 1.4 (Rambaut &
Drummond, 2007). All parameters had an effective sample size (ESS) over 500.
In order to assess genetic divergence between closely related taxa, Kimura´s two-parameter (K2P; Tamura &
Kumar, 2002) distances for cyt-b were estimated with MEGA 4 (Tamura et al., 2007). Since K2P includes some
assumptions that do not fit those expected for intronic sequences (Prychitko & Moore, 2000), we used Tamura
three-parameter (T3P; Tamura, 1992) for i7FGB. The presence of open reading frames and indels, as well as pair-
wise ts/tv ratio and pairwise K2P genetic distance within 1st, 2nd and 3rd codon positions of cyt-b were also estimated
with MEGA 4.
Based on phylogenetic results (see below), median joining network (Bandelt & Forster, 1999) was used for
analysing O. moojeni and O. utiaritensis cyt-b haplotypes with NETWORK 4.6 (Foster, 2010). In this analysis,
only 800 bp were considered in view of the available sequence data in GenBank.
Results
Morphological analysis. The distinctive features of the integumental and cranial anatomy shared by the holotype
(Figures 2 and 3) and the recently collected O. utiaritensis specimens were: (i) whitish ventral pelage, with white

hair tips and gray base; (ii) limits between ventral and lateral pelage well-defined; (iii) unicolored or weakly bicol-
ored tail; and (iv) long incisive foramina, the posterior borders reaching or almost reaching the alveolus of the first
upper molars, but never extending posteriorly. Although each of these features have been found in other Oligoryo-
mys species (table 1; also see below), this combination of characters was unique among known Oligoryzomys spe-
cies of Northern Mato Grosso and Southern Para and Amazonas states (O. microtis, O. moojeni, and O. fornesi). In
addition, the examination of the holotype of O. mattogrossae, which is also from Utiariti, revealed that this is a dif-
ferent species (the taxonomic status of O. mattogrossae is under analysis and will be reported in a subsequent
paper).
FIGURE 2. Dorsal, lateral, and ventral views of the skin of the holotype of Oligoryzomys utiaritensis (AMNH 37541).
Table 2 shows summary statistics of external and skull measurements of O. utiaritensis, O. moojeni, O.
nigripes, and the O. utiaritensis holotype (AMNH37541). Analyses of variance between O. utiaritensis, O.
nigripes, and O. moojeni showed significant differences (p<0.05) in 11 characters (CIL: F=13.63, p<0.001; M1M:
F=3.56, p=0.03; LD: F=8.51, p<0.001; PB: F=8.54, p<0.001; LIF: F=16.51, p<0.001; BIF: F=8.99, p<0.001; LM:
F=56.33, p<0.001; BM1: F=12.90, p<0.001; ORL: F=10.50, p<0.001; ZB: F=13.42, p<0.001; BZP: F=12.82,
p<0.001). Fisher’s LSD test showed O. utiaritensis to be significantly different (p <0.05) from O. nigripes in five
variables (PB, LM, BM1, ZB and BZP) and from O. moojeni in seven (CIL, LD, LIF, BIF, LM, ORL and ZB).
Canonical Discriminant Analysis discriminated O. utiaritensis, O. moojeni and O. nigripes (Figure 4). Six
specimens (MN13386, 13416, 13433, 13440, 13464, and 13475) from Minas Gerais State, previously identified as
O. utiaritensis by Avila-Pires (1960), fell within the O. nigripes range (Figure 4).

TABLE 1. Morphological comparison among Brazilian Oligoryzomys species. Included here are head-and-body length, tail length,
molar series length (all in mm), dorsal and ventral body coloration; information are, whenever possible from holotypes (or neotypes);
when type information is not available, the data are marked with *; color information from direct examination of holotypes (see Ridg-
way, 1912, for color chart). Abbreviations: AMNH – American Museum of Natural History; BMNH—British Museum of Natural His-
tory; UMMZ—University of Michigan Museum of Zoology; MN—Museu Nacional / UFRJ; CEM—Colecion Elio Massoia. Data for
molar series of O. flavescens (mean and range) are from Weksler & Bonvicino (2005). Data for body and tail measurements of O. del-
icatus (mean and range) are from AMNH specimens from Venezuela.
Species / Head- Tail Molar Dorsal color Ventral color

Holotype Body series
O. utiaritensis
100 120 3.45 grizzled yellowish brown Whitish with gray base
AMNH 37541
O. nigripes
100 119 3.59 dark brown to dark yellow Whitish with gray base
UMMZ 133872 (neotype)
O. stramineus
103 116 3.65 pale grayish yellow Whitish with gray base
MN 34439
O. chacoensis Clay, heavily lined with
104 149 3.45 Whitish with gray base
UMMZ 125524 black
O. rupestris grizzled yellowish brown,
84 105 3.29 Whitish with gray base
MN 50286 gray head
O. microtis
93 90 3.08 dull yellowish brown Pure White1
AMNH 37090
O. moojeni grizzled reddish- to
81 115 3.21 Cream with gray base
MN 50309 yellowish-brown
O. flavescens 3.2* Ochraceous Buffy with
95 105 bright brownish orange
BMNH 55.12.24.162 (3.0–3.5) gray base
O. delicatus 84* 94*
3.0 Yellowish brown Cream with gray base
AMNH 7317/5925 (77–89) (84–106)
O. messorius Dull whitish, with trace
82 95 3.1 Grizzled greyish fawn
BMNH 1.6.4.97 of buffy
O. mattogrossae Ochraceous Buffy with
95 115 3.30 rufous yellowish brown
AMNH 37542 gray base
O. fornesi Ochraceous Buffy with
82 105 3.3 chestnut yellowish brown
CEM3561 gray base
1. Other O. microtis, including topotypes, are tinged with buff
Karyotypic analyses. In 25 specimens of Oligoryzomys utiaritensis from three localities (see Appendix 1), a
diploid number (2n) of 72 chromosomes and an autosome fundamental number (FNa) of 76 was found (Figure 5a).
The autosome complement was composed of three small-sized, biarmed pairs and 32 acrocentric pairs, three of
which were large-sized and 29 pairs varying in size from medium to small. The X chromosome was a large-sized
submetacentric.
Six O. moojeni showed 2n = 70 and FNa = 74 (Figure 5b). The autosome chromosome complement was com-
posed by three biarmed, small-sized pairs and 31 acrocentric pairs, one of which was very large, two other were
large and 28 varying in size from medium to small. The X chromosome was a large-sized submetacentric and the Y
chromosome a small-sized chromosome.
Oligoryzomys nigripes showed 2n = 62 and FNa = 82 (Figure 5c). The autosome chromosome complement
was composed by 11 biarmed pairs, varying in size from large to small, and 19 acrocentric pairs, varying in size
from medium to small. The X chromosome is a large metacentric. The karyotype of Oligoryzomys utiaritensis dif-
fered from that of O. nigripes in diploid number (10 more autosome pairs), fundamental autosome number and in
morphology of the chromosomes. Furthermore, comparisons with other Oligoryzomys karyotypes (Table 3)
showed that the karyotype 2n=72, NFa=76 is unique among Oligoryzomys species.

FIGURE 3. Dorsal, ventral, and lateral views of the skull of the holotype of Oligoryzomys utiaritensis (AMNH 37541). Bar =
1 cm.

TABLE 2. External and skull measuremens (in millimeters) of the type of O. utiaritensis (AMNH 37541) and descriptive statistics for
O. utiaritensis, O. moojeni, and O. nigripes. Values are given as average ± standard deviation (minimum-maximum) sample size. See
material and methods for acronyms.
Holotype of O. utiaritensis O. utiaritensis O. moojeni O. nigripes

AMNH 37541
HBL 100 91.3 ± 6.5 (78-104) 28 87.3 ± 6.0 (79-97) 12 93.0 ± 7.8 (80-115) 47
LT 120 118.8 ± 7.5 (103-135) 28 115.1 ± 11.0 (93-132) 12 122.1 ± 13.0 (89-180) 47
HF 22 22.5 ± 2.0 (19-26) 28 23.0 ± 1.5 (21-25) 13 23.9 ± 2.4 (18-29) 49
Ear 14 14.9 ± 1.2 (13-18) 28 15.5 ± 1.1 (13-17) 13 16.2 ± 1.7 (13-20) 46
Wt - 23.6 ± 3.8 (14-32) 27 20.0 ± 8.2 (10-35) 7 21.6 ± 5.3 (9-32) 41
CIL 22.3 22.1 ± 0.9 (19.8-23.5) 42 20.9 ± 1.2 (19.1-22.6) 18 22.3 ± 1.2 (19.9-24.5) 84
LD 5.7 6.2 ± 0.3 (5.4-6.8) 42 5.8 ± 0.4 (5.2-6.4) 18 6.1 ± 0.4 (5.3-7.2) 92
PB 4.5 4.2 ± 0.2 (3.9-4.6) 42 4.3 ± 0.2 (3.8-4.6) 18 4.4 ± 0.3 (3.8-5.2) 92
LM 3.45 3.4 ± 0.1 (3.1-3.6) 42 3.4 ± 0.1 (3.2-3.7) 18 3.6 ± 0.2 (3.2-4.0) 90
BM1 1.1 1.0 ± 0.0 (0.9-1.1) 42 1.0 ± 0.0 (1.0-1.1) 18 1.1 ± 0.1 (0.9-1.2) 92
M1M 4.3 4.4 ± 0.2 (4.1-4.8) 42 4.4 ± 0.1 (4.1-4.7) 18 4.5 ± 0.2 (3.9-5.0) 92
LIF 4.1 4.5 ± 0.3 (3.8-5.1) 42 4.0 ± 0.3 (3.5-4.6) 17 4.5 ± 0.4 (3.7-5.3) 92
BIF 1.6 1.7 ± 0.1 (1.5-2.0) 42 1.6 ± 0.1 (1.4-1.8) 16 1.7 ± 0.1 (1.3-2.0) 93
BRO 4.5 4.5 ± 0.3 (4.0-5.0) 42 4.4 ± 0.3 (3.7-4.9) 18 4.5 ± 0.3 (3.6-5.3) 91
ORL 8.5 8.5 ± 0.3 (8.0-9.1) 41 8.1 ± 0.4 (7.4-8.6) 18 8.4 ± 0.4 (7.4-9.2) 92
ZB 12.7 12.4 ± 0.6 (11.0-13.7) 40 12.2 ± 0.6 (11.3-13.1) 15 12.9 ± 0.7 (11.3-14.9) 87
BZP 2.3 2.3 ± 0.2 (1.9-2.6) 42 2.4 ± 0.2 (2.1-2.7) 18 2.5 ± 0.2 (1.9-2.9) 92
FIGURE 4. Plot of two Canonical Functions (CF) of the discriminant analysis between Oligoryzomys utiaritensis, O. moojeni,
and O. nigripes. Eigenvalues are 2.66 (CF1) and 0.59 (CF2). Numbered symbols refer to holotypes of O. utiaritensis (1;
AMNH 37541) and O. moojeni (4; MN 50309); to paraphyletic specimens of O. moojeni in cyt-b phylogeny (2 and 3; MN
36426 and 50307); and to specimens identified as O. utiaritensis by Avila-Pires (1960; 5 to 10; MN 13386, 13416, 13433,
13440, 13464 and 13475).

FIGURE 5. Karyotypes with giemsa coloration of (A) O. utiaritensis (male MN 74937), (B) O. moojeni (male CRB 1018) and
(C) O. nigripes (female CRB 1440).
Phylogenetic analyses. Maximum likelihood (ML) and Bayesian inference (BI) analyses based on cyto-
chrome b data showed similar topology. Oligoryzomys was monophyletic, with bootstrap support (bs) of 100% in
ML and posterior probability (pp) of 1.0 in BI (Figure 6). In both analyses, O. microtis was found as sister group to
all other Oligoryzomys, followed by clades of O. fornesi and O. costaricensis, but these basal nodes did not show
high supports. The remaining species were divided into four clades: (O. vegetus, O. fulvescens); (O. chacoensis,

(O.nigripes, O. stramineus)); (O magellanicus, O.longicaudatus, O. flavescens); and ((O. messorius, ((O. destruc-
tor, O. rupestris), (O. delicatus, (Oligozyzomys sp., (O. moojeni, O. utiaritensis)))). Most of these clades showed
moderate/high nodal support, but the relationships between these clades were poorly supported (Figure 6).
All species represented by more than one sample were shown to be monophyletic with high support in cyt-b
topologies, except O. moojeni and O. longicaudatus (Figure 6). O. moojeni was not monophyletic because three
specimens (MN 36426, MN 50307, and MN 67087) were grouped in a clade with O. utiaritensis (bs=84%; pp=1),
while O. magellanicus grouped within the O. longicaudatus specimens. Both ML and BI trees showed O. utiariten-
sis and O. moojeni in a clade with moderate nodal support in ML (bs=66%) and high in BI (pp=1.0). The mono-
phyly of O. utiaritensis was highly supported (bs=99%; pp=1.0).
TABLE 3. Karyotypic data for Oligoryzomys. 2n= diploid number, FNa = autosome fundamental number. The collecting countries of
four karyotyped, undetermined Oligoryzomys species are given between parentheses after the reference.
Taxon 2n FNa References

O. rupestris 44-46 52-53 Silva & Yonenaga-Yassuda (1997); Weksler & Bonvicino (2005)
O. stramineus 52 68-70 Andrades-Miranda et al. (2001); Bonvicino & Weksler (1998); Maia et al.
(1983); Weksler & Bonvicino (2005)
O. magellanicus 54 66 Gallardo & Patterson (1985)
O. costaricensis 54 68 Carleton & Musser (1995); Gardner & Patton (1976)
O. messorius 56 58 Andrades-Miranda et al. (2001)
O. longicaudatus 56 66 Gallardo & Patterson (1985)
O. chacoensis 58 74 Myers & Carleton (1981)
a
Oligoryzomys sp. 58 74 Espinosa & Reig (1991)
(Argentina)
O. andinus 60 70 Gardner & Patton (1976)
Oligoryzomys sp. 60 72 Kiblisky (1969)
(Venezuela)
O. fulvescens 60 74 Carleton & Musser (1995); Haiduk et al. (1979)
O. destructor 60 76 Gardner & Patton (1976)
O. nigripes 61-62 78-82 Almeida & Yonenaga-Yassuda (1991); Andrades-Miranda et al. (2001);
Bonvicino & Weksler (1998); Bonvicino et al. (2001); Brum-Zorrila et al.
(1988); Espinosa & Reig (1991); Myers & Carleton (1981); Paresque et al.
(2007); Yonenaga et al. (1976)
O. fornesi 62 64 Andrades-Miranda et al. (2001); Bonvicino & Weksler (1998); Myers &
Carleton (1981); Svartman (1989)
O. delicatus 62 74, 76 Gardner & Patton (1976)
O. microtis 64 66 Aniskin & Volobouev (1999); Gardner & Paton (1976); Patton et al. (2000)
O. flavescens 64-68 66-70 Andrades-Miranda et al. (2001); Aniskin & Volobouev (1999); Brum-Zorrila
et al. (1988); Espinoza & Reig (1991); Myers & Carleton (1981); Sbalqueiro
et al. (1991); Vidal-Rioja et al. (1988)
Oligoryzomys sp. 66 74 Andrades-Miranda et al. (2001)
(Brazil, Amapá)
Oligoryzomys sp. (Peru) 68 74, 76 Gardner & Patton (1976)
O. moojeni 70 74-76 Andrades-Miranda et al. (2001), Lima et al. (2003); Weksler & Bonvicino
(2005)
O. utiaritensis 72 76 This study
a
Despite sharing the same 2n and FNa with O. chacoensis, this karyotype differs in the morphology of biarmed chromosomes.

FIGURE 6. Phylogenetic relationships among species of Oligoryzomys based on Maximum likelihood (ML) analysis of the
cytochrome b gene. Nodal support values shown above branches are ML bootstrap values (>50%) and Bayesian posterior prob-
ability (>0.50). Model of substitution selected was GTR+G.

Analysis of the i7FGB dataset included 52 Oligoryzomys from eight different species, including 18 specimens
of O. utiaritensis and type specimens of O. moojeni (Appendix 2). ML and BI trees showed similar topologies,
with Oligoryzomys as monophyletic (Figure 7) with high nodal support (bs = 99% and pp = 1). In the i7FGB anal-
yses, O. microtis grouped with O. fornesi in the most basal offshoot (bs = 68% and pp = 0.97), while in cyt-b these
two species were paraphyletic. The remaining species grouped in two clades: the first composed by (O. stramineus,
O. nigripes) with bs = 80% and pp = 1.0, and the second by (O. flavescens, (O. utiaritensis, (O. rupestris, O.
moojeni))) with bs = 56% and pp = 0.94. Within the last clade, O. rupestris and O. moojeni were sister taxa with
high nodal support (bs=90% and pp=1), while in cyt-b O. moojeni and O. utiaritensis were sister taxa. All the spe-
cies represented by more than one sample were shown to be monophyletic with high support in both analyses (Fig-
ure 7), including O. moojeni and O. utiaritensis (in the cyt-b tree, O. moojeni was not monophyletic).
FIGURE 7. Phylogenetic relationships among species of Oligoryzomys based on Maximum likelihood (ML) analysis of the
intron 7 of beta-fibrinogen gene. Nodal support values shown above branches are ML bootstrap values (>50%) and Bayesian
posterior probability (>0.50). Model of substitution selected was K81+G.

TABLE 4. Genetic distances for Oligoryzomys species (in %): Kimura 2-parameter distances for cyt-b (lower diagonal), and Tamura three-parameter
distances for i7FGB (upper diagonal). Gray cells indicate distance within species (only for cyt-b). Specimens used in this analysis are listed in Appendix
2; - = nuclear DNA sequences not available for these species .
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
1.O. microtis 2.2 0.8 - - - - - 1.6 - 2.9 1.6 - 2.2 - - - 1.7 1.7
2.O. fornesi 12.6 0.3 - - - - - 1.4 - 2.8 1.4 - 2.3 - - - 1.7 1.5
3.O. costaricensis 13.0 11.3 2.0 - - - - - - - - - - - - - - -
4.O. vegetus 10.9 11.0 8.9 0.4 - - - - - - - - - - - - - -
TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS

5 .O. fulvescens 12.1 10.5 10.7 8.1 1.6 - - - - - - - - - - - - -
6 .O. longicaudatus 11.7 11.9 9.2 10.4 9.7 0.6 - - - - - - - - - - - -
7. O. magellanicus 10.6 11.9 8.0 9.4 8.5 1.3 - - - - - - - - - - - -
8. O. flavescens 12.0 12.0 9.5 10.3 11.0 4.7 4.5 3.9 - 3.2 1.8 - 2.2 - - - 2.0 1.5
9. O. chacoensis 12.3 12.2 10.6 9.7 10.1 9.4 8.9 9.5 0.5 - - - - - - - - -
10. O. stramineus 11.6 11.9 9.7 8.5 9.2 8.4 7.5 8.2 6.9 0.3 1.9 - 3.0 - - - 2.8 2.6
11. O. nigripes 11.8 12.5 9.9 9.3 9.6 8.2 6.9 9.8 7.8 5.7 1.3 - 2.2 - - - 1.9 1.5
12. O. messorius 12.5 12.6 9.4 9.7 10.7 9.5 8.1 9.1 9.3 8.9 9.8 - - - - - - -
13. O. rupestris 12.5 13.6 10.2 10.2 10.2 9.3 8.7 10.6 10.9 9.6 9.1 10.3 0.6 - - - 1.1 2.0
14. O. destructor 13.3 12.1 10.5 11.4 12.2 10.8 10.1 10.9 11.5 9.7 11.0 10.5 9.3 0.1 - - - -
15. O. delicatus 11.1 13.5 9.9 9.4 9.3 9.0 7.9 9.8 10.5 9.3 8.9 9.0 7.8 10.1 4.9 - - -
16. Oligoryzomys sp. 12.5 12.8 10.2 9.9 10.4 11.0 10.3 11.0 10.8 10.6 10.2 10.3 8.5 10.0 7.6 0.6 - -
17. O. moojeni 9.7 10.8 8.9 8.3 9.7 8.6 7.1 8.8 8.8 7.6 8.3 8.8 8.4 9.0 6.6 8.1 2.1 1.2
18. O. utiaritensis 10.5 12.0 10.6 9.9 10.5 9.4 9.0 10.2 11.2 9.0 9.7 10.8 8.6 9.6 5.9 7.3 6.4 0.5
Zootaxa 3220 © 2012 Magnolia Press ·

13
K2P distances (Table 4) between cyt-b haplotypes of different species varied from 1.3% (between O. longicau-
datus and O. magellanicus) to 13.6% (between O. fornesi and O. rupestris). Oligoryzomys utiaritensis differed
from O. nigripes by 9.7% and from its sister species O. moojeni by 6.4%. The overall K2P distance between Oligo-
ryzomys species was 8.2%. Average K2P intra-specific distances (Table 4) varied from 0.1% in O. rupestris to
4.9% in O. delicatus; the average K2P being 0.5% within O. utiaritensis and 2.1% within O. nigripes.
T3P distances for i7FGB (Table 4) among different species varied between 0.8% (O. microtis – O. fornesi) and
3.2% (O. flavescens – O. stramineus). Oligoryzomys utiaritensis differed from O. nigripes by 1.5%, and from O.
moojeni by 1.2%. The overall average T3P distance between Oligoryzomys species was 1.9%.
In order to clarify the relation between O. moojeni and O. utiaritensis, a median joining network was con-
structed with cyt-b haplotypes (Figure 8). Oligoryzomys utiaritensis haplotypes differed from those of O. moojeni
by at least 16 mutations. The Oligoryzomys moojeni haplotypes of specimens grouping with O. utiaritensis in the
ML and BI topologies (haplogroup 2 in Figure 8) differed from the other O. moojeni haplotypes (haplogroup 1) by
at least 15 mutations.
FIGURE 8. Median joining network of cytochrome b haplotypes from O. moojeni (white circles) and O. utiaritensis (gray cir-
cles). Haplotype circles are proportional to their frequency. Dashes indicate mutations among haplotypes and dark circles indi-
cate estimated haplotypes that were not sampled. The arrow shows the position of root, based on ML analysis (see Fig. 6).
Discussion
Taxonomy of Oligoryzomys utiaritensis. Our findings indicated that the Oligoryzomys specimens that are natural
reservoirs of the Castelo dos Sonhos hantavirus from Northern Mato Grosso and Southeastern Pará belong to a dif-
ferent species from all others currently recognized in the genus (Musser & Carleton, 2005). Their karyotype (2n =
72; FNa = 76), herein described for the first time, differed from all other Oligoryzomys karyotypes (Table 3). The
identification of these specimens as O. utiaritensis was based on (i) geographic proximity – less than 50km –
between their site of capture respective to the type locality of O. utiaritensis in Utiariti (Mato Grosso, Brazil); (ii)
their extremely similar integumental and cranial morphology to the O. utiaritensis holotype (AMNH37541; Fig-
ures 2 and 3); and (iii) their placement with the O. utiaritensis holotype in discriminant analysis (Figure 4).
Oligoryzomys utiaritensis was described by Allen (1916) from specimens collected by the Roosevelt-Rondon
Expedition in Utiariti, Rio Papagaio, Mato Grosso. Except for a few mentions in checklists (e.g., Gyldenstolpe,
1932; Moojen, 1952; Cabrera, 1961), the only reports of O. utiaritensis in the literature accounted for specimens
from Gradaús, southeast Pará state (Carvalho, 1960) and from Conceição do Mato Dentro (Avila-Pires, 1960), Car-
atinga (Botelho & Williams, 1980; Botelho et al., 1981; Linardi et al., 1984), and Tiradentes (Lopes et al., 1989), in
Minas Gerais state (Figure 1). Oligoryzomys utiaritensis was considered a junior synonym of O. eliurus by Carle-
ton & Musser (1989) in their latest summary of this genus, and by Musser & Carleton (2005). Myers & Carleton
(1981) were the first to provide evidence in favor of this taxonomic arrangement, pointing to the close morphologic
similarity of Oligoryzomys utiaritensis and O. eliurus. The elucidation of the taxonomic status of O. utiaritensis
therefore requires clarifying the taxonomic status of O. eliurus, a taxon that has been placed under O. nigripes in

several reports (González-Ittig, et al., 2010; Massoia, 1973; Myers & Carleton, 1981; Paresque et al., 2007; Wek-
sler & Bonvicino, 2005). Here we synthetize the arguments in favor of the synonimization of O. eliurus under O.
nigripes, based on karyotypic and molecular, and corroborated by our morphological assessment of the O. eliurus
holotype.
Hesperomys eliurus was described by Wagner (1845) based on specimens collected by Natterer from Itararé, in
southern São Paulo state, in the Atlantic Forest domain (Figure 1). The ecological features of these two regions are
very different because Itararé is in an area of mixed vegetation, including semideciduous forest and natural grass-
lands, locally known as “campos”, on the southeast to the Cerrado-Atlantic Forest ecotonal belt across Paraná and
São Paulo (Scaramuzza, 2006). Conversely, Utiariti is located in the northern margin of the Cerrado, along its tran-
sition with the Amazonian forest.
Extensive work on Oligoryzomys karyology in southeastern São Paulo state and adjacent regions in Paraná
reported only two karyomorphs: 2n=62 and FN=80-82 in larger animals with whitish venter identified as O.
nigripes, and 2n=64 and FN=66-68 in smaller animals with yellowish venter identified as O. flavescens (Almeida
& Yonenaga, 1991; Andrades-Miranda et al., 2001; Bonvicino & Weksler, 1998; Bonvicino et al., 2001; Paresque
et al., 2007; Sbalqueiro et al., 1991; Weksler & Bonvicino, 2005; Yonenaga et al., 1976). The former karyomorph
shared the same phenotype (i.e., large body size and whithish venter) with the O. eliurus holotype (NHM 423) and
with the neotype of O. nigripes (UMMZ 133872; Myers and Carleton, 1981). The Oligoryzomys karyotype (2n=62
and FN=64) reported by Andrades-Miranda et al. (2001) from Niquelândia, Minaçú, Uruaçú, Ipameri, Caldas
Novas and Corumbaíba (all in Goiás state, Brazil), and identified as O. eliurus has never been reported from the
region of the type locality of this species, and therefore probably belongs to a third species, O. fornesi (sensu Myers
& Carleton, 1981), a smaller species with yellow venter (Bonvicino & Weksler, 1998).
Molecular data, altough sparse, also corroborate our arguments. Specimens with large body size and whitish
venter from southern and southwestern Brazil, including Paraná and São Paulo states, were all included in the O.
nigripes clade (Miranda et al., 2009 and this study). The specimen from São Paulo state (Guariba) considered to be
O. eliurus in the molecular analysis of Palma et al. (2010) should also be identified as O. nigripes1.
Given this series of evidence, we believe that Oligoryzomys eliurus should be regarded as a junior synonym of
O. nigripes; we therefore question whether O. utiaritensis might also be a junior synonym of O. nigripes or a valid
species. Although both species are morphologically similar, karyotypic data cleary demonstrate that O. utiaritensis
and O. nigripes are different species: O. utiaritensis showed 2n=72 and FNa=76, unlike O. nigripes with 2n=62,
FNa=78-82. Moreover, analyses of mitochondrial (Figure 6) and nuclear (Figure 7) markers indicated that O.
nigripes and O. utiaritensis were not sister taxa. O. nigripes is member of a clade with O. stramineus and O. cha-
coensis that shows a strong nodal support, similarly to recent analyses based on cyt-b (Palma et al., 2010; Rogers et
al., 2010; González-Ittig et al., 2010; Miranda et al., 2009) and IRBP (Weksler, 2003; Miranda et al., 2009). In
contrast, O. utiaritensis was placed in a clade with O. moojeni, O. rupestris, O. delicatus (Allen & Chapman), O.
destructor (Tschudi), O. messorius (Thomas) and an unknown species of northern South America, Oligoryzomys
sp. Finally, sequence divergence between the O. nigripes and O. utiaritensis was around 9.7% for cyt-b and 1.5%
for i7FGB.
Oligoryzomys moojeni, from Central Brazil (states of Goiás and Tocantins), is apparently the closest taxon to
O. utiaritensis. These two species have a similar karyotype and are phygenetically closely related, despite their
morphological differences (especially in integumental characteristics; see below). Karyotypic data suggest a close
relationship, but is also indicative that they are different species; the O. utiaritensis karyotype (2n=72, FNa=76) is
very similar to the O. moojeni karyotype (2n=70, FNa=74; Weksler & Bonvicino, 2005) but differed by one pre-
sumed tandem fusion/fission accounting for a large-sized pair and one small-sized pair in O. utiaritensis corre-
sponding to one very large pair of O. moojeni (Figure 4). Tandem fusions/fissions, as mechanisms of karyotypic
evolution, have been documented in several mammalian taxa although these rearrangements are not present as sta-
ble polymorphisms (Robinson & Elder, 1993), suggesting that abnormal disjunction in heterozygotes results either
in rapid fixation or loss from populations (Elder & Hsu, 1988). This scenario suggested that tandem fussions/fis-
sions might be efficient as post-mating isolating mechanisms preventing the presence of fertile hybrids.
1. Oligoryzomys nigripes also includes O. delticola (Thomas) as junior synonym, as demonstrated by molecular analysis of the D-
loop and cyt-b mitochondrial markers (Francés & D'Elía, 2006; see also Weksler & Bonvicino, 2005). Thus, the Oligoryzomys
specimen (GD 569 = UMMZ 176262) from Lunarejo (Rivera, Uruguay), identified as O. delticola by Palma et al. (2010), should
be identified as O. nigripes, as demonstrated by Francés & D'Elía (2006).

The cyt-b tree did not show O. moojeni as a monophyletic lineage because three specimens identified by kary-
otype and morphology as O. moojeni (MN 36426, MN 50307, and MN 67087) were paraphyletic with respect to O.
utiaritensis (fig. 6; haplogroup 2 in figure 8). On the other hand, the two species were identified as separate mono-
phyletic lineages in the i7FGB topology, and not even as sister taxa. How could these discordant results be
explained? Sample contamination could be discarded because DNA extraction, amplification and sequencing of
haplogroup 2 specimens were carried out in different laboratories. Pseudogene (nuclear transposition of mitochon-
drial DNA – Numt; Lopez et al., 1994) amplification also seemed unlikely because all haplogroup 2 specimens did
not have indels, and showed open reading frames. In comparison with the others haplogroups, it also had a high ts/
tv ratio and the expected pattern of pairwise differences between 1st, 2nd and 3rd codon positions (Triant &
DeWoody, 2007; data not shown). Genetic introgression between O. utiaritensis and O. moojeni is also unlikely in
view of fixed karyotypic differences and lack of intermediate karyomorphs between O. utiaritensis and O. moojeni
(all three specimens of haplogroup 2 in the cyt-b topology showed a typical O. moojeni karyotype). Besides, the
known distribution of O. moojeni is distant at least 650 km from the places of collection of O. utiaritensis. Finally,
a shallow coalescence, or retention of ancestral polymorphisms (Maddison, 1997), in O. moojeni seem unlikely
because the lineages are reciprocally monophyletic in the analyses with the slower-evolving i7FGB nuclear
sequence.
Network analysis (Figure 8) indicated that the total genetic diversity of O. moojeni has not been fully sampled,
as shown by the long branches separating the two haplogroups that contain individuals from the same populations
in Goiás. Additional work is thus required to study the phylogeographic pattern of these Oligoryzomys species of
the transitional zone of the Cerrado-Amazon of Brazil. Oligoryzomys utiaritensis and O. moojeni are apparently
distributed in neighboring river basins, the former in the Tapajós and Xingu basins, while the latter is found in the
Araguaia and Tocantins basins. The gap between their distributions could either be real or a result of lack of proper
sampling between river basins.
We thus conclude that our data corroborate the valid taxonomic status of O. utiaritensis, a finding that is rele-
vant for Government agencies, such as the Brazilian Ministry of Health, in view that 2% of O. utiaritensis collected
in Campo Novo do Parecis, Mato Grosso state, were seropositive for hantavirus (Nunes et al., 2008). This species
is the reservoir of hantavirus Castelo dos Sonhos (Rosa et al., 2008; Rosa et al., 2011) and its correct taxonomic
identification is extremely important for the planning of public policies of hantavirus control. The redescription of
O. utiaritensis is herein provided.
Systematics
Oligoryzomys utiaritensis J. A. Allen, 1916

(Figures 2, 3, 4)
Holotype. AMNH 37541, adult female (Figures 2 and 3), collected by Leo E. Miller on January 30th, 1914; mea-
surements of holotype are provided in Table 2.
Type Locality. Brazil, Mato Grosso state, Rio Papagaio, Sapezal municipality, Utiariti; geographical coordi-
nates 12°59’07’’S, 55°36’23’’W (taken by GPS).
Geographic Distribution. The six known collecting localities of O. utiaritensis (Figure 1, Appendix 1) are
distributed across the northwest of Mato Grosso and southwest of Pará. The area includes the Chapada dos Parecis,
a massive plateau in northwestern Mato Grosso that marks the transition between the Cerrado and Amazonian
domains in Central Brazil. All localities are restricted to the Rio Tapajós and Rio Xingu watershed. Carvalho
(1960) did not specify where the specimens collected at Gradaús (Pará state) and identified by him as O. utiariten-
sis were housed – thus their identity was not confirmed; nevertheless, this locality, nowadays at the municipality of
São Félix do Xingu (06°38' S, 51°59' W), is near the known range of the species and therefore could represent a
seventh locality for O. utiaritensis.
External and cranial Measurements. See Table 2.
Diagnosis. A medium-sized Oligoryzomys species characterized by: (1) grizzled yellowish-brown dorsal pel-
age, contrasting with the whitish ventral pelage and tail weakly bicolored; (2) long incisive foramina, the posterior
borders reaching or almost reaching the alveolus of the first upper molars, but never extending posteriorly; (3) the

highest diploid number (2n=72) among Oligoryzomys species; and (4) three putative synapomorphies in cyto-
chrome b, and one in intron 7 of beta fibrinogen (alignments available from the authors).
External Characters. Adult dorsal pelage grizzled yellowish-brown, between Antique Brown and Dresden
Brown (Ridgway, 1912), composed of long guard hairs and slightly shorter overhairs with a sub-apical brown-yel-
lowish band. Lateral color lighter than in dorsum and with a clearly defined limit with the whitish ventral pelage.
Ventral hairs white at their upper half and gray at their basal half. Internal surface of pinnae with brown-yellowish
hairs; external surface brownish. Dorsal surface of feet covered with withish hairs, sometimes washed with cream
color; short tufts of white ungual hairs at bases of claws on dII–dV. Tail longer than combined length of head and
body, sparsely haired, and covered with more or less conspicuous epidermal scales, lacking a long tuft of terminal
hairs and weakly bicolored, dorsal surface dark gray and ventral surface light gray (some specimens are unicolored
in the distal half of the tail). Superciliary, genal, and mystacial vibrissae not extend beyond ears. Juveniles with
grayish dorsum, with whitish ventral surface. Presence of eight mammae in inguinal, abdominal, postaxial, and
pectoral positions.
Cranial Characters. Delicate skull, medium and narrow rostrum, but slightly wider than interorbital region;
interorbital region hourglass-shaped. Braincase without supraorbital and postorbital ridges and with weakly devel-
oped lambdoidal ridge. Interparietal bone as broad as parietal. Relatively large zygomatic plate with deep zygo-
matic notch. Jugal bone absent, resulting in zygomatic process of squamosal in contact with the zygomatic process
of maxillary. Incisive foramina with almost parallel margins, the posterior borders reaching or almost reaching the
alveolus of the first upper molars, but never extending posteriorly. Palate with single large posterolateral palatal
pits not recessed in palatine fossa. Palatal bridge broad and long. Bony roof of mesopterygoid fossa perforated by
large sphenopalatine vacuities. Width of parapterygoid plate slightly greater than width of mesopterygoid fossa.
Alisphenoid strut absent (buccinator-masticatory foramen and accessory foramen ovale confluent), alisphenoid
canal with large anterior opening. Stapedial foramen and the posterior opening of the alisphenoid canal large, but
squamosal–alisphenoid groove and sphenofrontal foramen absent (= carotid circulatory pattern 2; Voss 1988). Pos-
terior suspensory process of the squamosal absent. Large subsquamosal fenestra, slightly smaller than postglenoid
foramen. Periotic exposed posteromedially between ectotympanic and basioccipital, extending anteriorly to carotid
canal. Mastoid perforated by conspicuous posterodorsal fenestra. In mandible, capsular process of lower incisor
alveolus well developed in most adults; superior and inferior masseteric ridges converging anteriorly as open chev-
ron below m1.
Dental Characters. Upper and lower incisors opisthodont; molars pentalophodont. Superior molar rows paral-
lel. The first upper molar (M1) anterocone is divided into anterolabial and anterolingual conules by a weak antero-
median flexus only in young animals; animals with moderate wear lack anteromedian flexus; the anteroloph is well
developed, separated from anterocone by a persistent anteroflexus; mesolophs are present on all upper molars in
most specimens, but absent in specimens with moderate or heavy wear; the paracone is connected by an enamel
bridge to the anterior moiety of protocone, and in some specimens to the mesoloph by the paralophule. M2 mor-
phology is similar to M1 except in the anterior region: M1 has an anterocone, while M2 only has an anteroloph.
Third upper molar (M3) is reduced, and has a single posterior cup, which we equate to the hypocone; hypoflexus is
diminutive. Anteroconid of first lower molar (m1) without anteromedian flexid; the anterolabial cingulum is pres-
ent on all lower molars; the anterolophid is present on m1 but absent on m2 and m3; an ectolophid is absent on m1
and m2; the mesolophid is distinct on unworn m1 and m2, sometimes joined to the entoconid; the posteroflexid is
present on m3.
Karyotype. Karyotypic analyses of 25 specimens of Oligoryzomys utiaritensis showed 2n=72, FNa=76 (Fig-
ure 4). Autosome complement comprising three pairs of small-sized biarmed chromosomes and 32 acrocentric
pairs, three large-sized and 29 varying in size from medium to small. X chromosome large submetacentric and
small Y chromosome.
Habitat. Specimens of Oligoryzomys utiaritensis were recently captured between 110 and 570 m of altitude, in
secondary semideciduous forest in the limits of corn and soy plantations, in eucalypt plantations with herbaceous
vegetation, and in very altered vegetation (semi-deciduous forest) near plantations. No information about the vege-
tation of the type locality was provided in the description of O. utiaritensis (Allen, 1916), but the vegetation along
the Papagaio River, including in the Utiariti region, is gallery forest.
Comparisons. Oligoryzomys utiaritensis differs from all other Oligoryzomys species by its unique karyotype.
In addition, O. utiaritensis differs from other Oligoryzomys that occur in Brazil by a combination of other charac-

ters including (1) whitish ventral pelage, (2) a sharply defined limit between lateral and ventral pelage color, in
comparison to a buffy venter without defined limit between lateral and ventral coloration in adult specimens of O.
moojeni, O. fornesi and O. flavescens (see table 1); (3) slightly bicolored tail, contrary to unicolored tail in O.
nigripes and O. rupestris (other species of the genus also have slightly bicolored tails); (4) a large size (adult HBL
> 91 mm in average) comparable to O. nigripes, O. stramineus, and O. chacoensis, and opposed to small size spe-
cies O. rupestris, O. microtis, O. moojeni, O. flavescens, O. delicatus, O. messorius, O. fornesi (HBL < 90 in aver-
age).
In addition, we present here more detailed comparison between O. utiaritensis and O. moojeni in view of their
closer geographic distribution and their molecular and karyotypic similarities. In Oligoryzomys utiaritensis, the
dorsum is yellowish-brown (unlike the reddish-brown dorsum of O. moojeni), with a clearly defined limit between
dorso-lateral and ventral pelage (unlike the poorly defined limit in O. moojeni, forming a gradient between lateral
and ventral coloration). In O. utiaritensis, the rostrum is slightly wider than the interorbital constriction (unlike O.
moojeni in which the rostrum and interorbital constriction are of similar width), and the subsquamosal fenestra is
almost of the same size of postglenoid foramen (unlike O. moojeni with a smaller subsquamosal fenestra about half
the size of the postglenoid foramen).
Etymology. Oligoryzomys utiaritensis was named by J.A. Allen based on the type locality Utiariti (spelled
Utiarity in old maps and accounts), referring to a Amerindian village in Sapezal municipality.
Specimens examined. See Appendix 1.
Remarks. The specimens referred as O. utiaritensis from Conceição do Mato Dentro (Avila-Pires, 1960) and
Caratinga (Botelho & Williams, 1980; Botelho et al., 1981; Linardi et al., 1984) are herein identified as O.
nigripes, based on morphometric (Figure 4) and morphological analyses of the voucher material deposited at the
Museu Nacional (see voucher numbers below in Appendix 1). Myers and Carleton (1981) discussed the composite
nature of the paratype of O. utiaritensis (AMNH 37540); our examination of this specimen was in agreement with
their assessment: skull and skin do not belong to the same animal, the skin being from a larger animal than the
skull. We thus remove this specimen from the type series of O. utiaritensis.
Oligoryzomys phylogenetics. Our phylogenetic results were generally coincident with recent studies of Oligoryzo-
mys (Francés & D'Elía, 2006; González-Ittig et al., 2010; Miranda et al., 2009; Palma et al., 2005, 2010; Richter et
al., 2010; Rivera et al., 2007; Rogers et al., 2009). Analyses of an additional marker, the nuclear i7FGB gene, pro-
vided independent corroboration, usually with high nodal support (especially for the Bayesian analysis), of several
clades found with mitochondrial cyt-b.
We could observe that O. utiaritensis was found in one the major lineages recovered by the cyt-b analyses with
6 other species: ((O. messorius, ((O. destructor, O. rupestris), (O. delicatus, (Oligozyzomys sp., (O. moojeni, O.
utiaritensis)))); in the i7FGB analyses, O. rupestris, O. moojeni, and O. utiaritensis also clustered together (the
other species were not available for these analyses).
Our phylogenetic reconstructions were also the first to include O. rupestris, which appeared as sister group to
O. destructor with respect to the (O. moojeni, O. utiaritensis) clade in the cyt-b phylogeny and as a sister lineage of
O. moojeni in the i7FGB phylogeny. Nevertheless, our results confirmed the need for analyzing additional loci to
reach a more complete understanding of Oligoryzomys phylogeny, mainly because analyses exclusively based on
cyt-b did not provide sufficient phylogenetic signal for a robust resolution of several interspecific relationships.
Acknowledgments
We appreciated the facilities provided by the owners of the Fazenda Caetitu in Sapezal (MT), Faz. Campo Belo,
Faz. Ponte Senada, Faz. Itamarati, Faz. Ouro Verde II and Faz. Alvorada in Campo Novo do Parecis (MT), and the
Secretaries of Health of Mato Grosso state, Sapeza and Campo Novo dos Parecis municipalities. The collaboration
in fieldwork by the field team of SVS (Secretaria de Vigilância em Saúde) and Laboratório de Biologia e Parasito-
logia de Mamíferos Silvestres Reservatórios, IOC, FIOCRUZ, was most useful. ICMBio (Instituto Chico Mendes
de Conservação da Natureza) granted license to collect the specimens. Work was supported by Conselho Nacional
de Desenvolvimento Científico e Tecnológico (CNPq) fellowships to CRB (302951/2007-5), PSD, and MW
(481286/2011-0) and Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) to CRB. We also

would like to thank the curators and staff from the AMNH (Robert Voss, Nancy Simmons, and Eileen Westwig)
and MNRJ (João O. Alves and Stella Franco) that gently allowed us to analyze the specimens under their care; Paul
Velazco for help in taking pictures of the holotype of O. utiaritensis; Mauro Elkoury and Marilia Lavocat for strong
efforts and facilities for the project development; to Héctor Seuánez for a critical review of an earlier version of the
manuscript; and to Philippe Gaubert and two reviewers for commentaries and suggestions that greatly improved
the manuscript.
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APPENDIX 1. List of localities and specimens of Oligoryzomys used in morphometric analyses (*), karyotyped (#), and
sequenced (σ for cyt-b and β for i7FGB). Italicized place names are municipalities. Geographic coordinates are provided
in parentheses after the locality or the municipality, with the source for these data. Species name are separated from the
locality and specimens by a dash. Numbered localities for O. utiaritensis and O. moojeni are plotted on the map (Figure
1). F: refers to females, M: refers to males. IBGE = Instituto Brasileiro de Geografia e Estatística. For additional ana-
lyzed specimens, see Weksler & Bonvicino (2005). Museum and collectors acronyms are: CRB (Cibele Rodrigues Bon-
vicino), LBCE (Laboratório de Biologia e Parasitologia de Mamíferos Silvestres Reservatórios, FIOCRUZ, Brazil), MN
(Museu Nacional, UFRJ, Brazil), NHM (Natural History Museum, Vienna, Austria), SVS (Serviço de Vigilância em
Saúde, Ministry of Health, Brazil), UFPB (Universidade Federal da Paraíba, Brazil), AMNH (American Museum of Nat-
ural History, USA) and USNM (United States Natural Museum, USA).
BRAZIL
Pará
1. Altamira, Castelo dos Sonhos (08º18’S 55º05’W, IBGE); O. utiaritensis– F: MN 75602, 75603*, 75609#*σ, 75611*,
75613*σβ, 75614*, SVS 278*; M: MN 75604*, 75605*, 75606# , 75608*#, 75610*, 75612*σβ
Mato Grosso
2. Peixoto de Azevedo (10º13’S 54º58’W, IBGE); O. utiaritensis– F: MN MN75600, 75601
3. Feliz Natal (12°23'S 54°55'W, IBGE) Aldeia Sobradinho; O. utiaritensis – F: SVS 819#, M: SVS 820
4. Sapezal, Fazenda Begolim (13º20’S 55º36’W, GPS); O. utiaritensis – F: MN 75591*, 75592*σβ, 75594*αβ, 75595*β,
75596*αβ, 75597*αβ, 75598σ; M: MN 75593*σβ
5. Sapezal, Rio Papagaio, Utiariti (12°59’S 5536’W, GPS); O. utiaritensis– F: AMNH 37541* (holotype of O.
utiaritensis); O. “mattogrossae” – M: AMNH 37542 (holotype of O. mattogrossae).
6. Campo Novo do Parecis (13º40’S 57º53’W, IBGE); O. utiaritensis – F: SVS 306σβ, 357σβ, CNP-SVS 5#, 265#, MN
75616*#σβ, 75623#*, 75624, 75625*#β ; M: CNP-SVS 263#, 264#, MN 75617*, 75618*#,75619*#σβ, 75620#σβ, 75621*,
75622*#β, 75626*#σβ, 75627#, 75628#, 75629#, 75630#, 75631*, 75632# ,75633#, 75634*#, 75635*, 75636*, 75637,
75638, 75639*, 75640*, 75641, 75642, 75643, 75644, 75645, 75646, 75647, 75648, 75649, 75650*, 75651, 75652,
75653*, 75654*, 75655,75656*, 75657, 75658*#, 75659#, 75660*, 75661, 75662, 75663, 75664, 75665, 75666,
75667, 75668, 75669
Tocantins
7. Lajeado, Fazenda Elizeu (9°55’S 48°17’W, Lima et al. 2003); O. moojeni – F: MN LJ35# – karyotyped by Lima et al.
(2003).
8. Porto Nacional, Chácara União (around 10°44’S 48°23’W, Lima et al. 2003); O. moojeni – F: ZUT 30# – karyotyped
by Lima et al. (2003).
9. Dianópolis (11°37’S 46°49’W, IBGE); O. moojeni – M: LBCE 12846#
10. Novo Jardim (11°49’S 46°37’W, IBGE); O. moojeni – M: LBCE 12012#, 12013#
18. São Sebastião do Tocantins (5°17'S 48°18'W), Fazenda Osara; O. microtis – F: CRB 1450β, 1451β, 1452β, M: CRB
1448 β
Goiás
11. Minaçu, Rio Tocantinzinho, 40km SW Minaçu (13°31’S 48°13’W, Miranda et al., 2009); O. moojeni – unsexed: MN
36832*#σ – karyotyped by Andrades-Miranda et al. (2001) and sequenced by Miranda et al. (2009).

12. Cavalcante, Fazenda Fiandeira (14°04’S 47°45’W, GPS); O. moojeni – F: MN 50310*#, 50312*#, 50313*#,
50315*#σβ, 50319*#σβσ, 50320*#σβ, 50321*#; M: MN 50287*#σβ,50307*#σβ, 50308*#, 50309*#σβ (holotype of O.
moojeni), 50311*#, 50314*#σ, 50318*#, 50377*# ; unsexed MN 50316*#, 50317*#σβ – all karyotyped by
Weksler&Bonvicino (2005).
13. Colinas do Sul, Rio Maranhão, 20km NW Colinas do Sul (14°09’S 48°04’W, Miranda et al., 2009); O. moojeni – M:
MN 36220*#, 36357*#, 36426*#; unsexed: MN: 36224#, 36234#, 36242#, 36345#, 36356#, 36367#, 36433#, 36438# –
all karyotyped by Andrades-Miranda et al. (2001; MN 36220, 36357, 36426 also sequenced by Miranda et al.
2009).
14. Uruaçu, Rio do Peixe, 40km NE Uruaçu (14°31’S 49°08’W, Miranda et al., 2009); O. moojeni – M: MN 37441*#σ;
unsexed: 37282#σ – all karyotyped by Andrades-Miranda et al. (2001) and sequenced by Miranda et al. (2009)
15. Niquelândia (14º28’S 48º27’W, IBGE); O. moojeni – unsexed: MN 36082#, 36083#, 36145#, 36147#, 36161# – all
karyotyped by Andrades-Miranda et al. (2001)
16. Sítio D'Abadia (14º48’S 46º15’W, GPS); O. moojeni – F: LBCE 11615#σβ, 11648#σβ
17. Mimoso de Goiás, Fazenda Cadoz (15°03’S 48°09’W, GPS); O. moojeni – F: MN 67087#σ
Alto Paraíso de Goiás (14º01’S 47º32’W, GPS); O. rupestris – F: MN 50286β, 50322σβ; M: MN 50326σβ
Teresina de Goiás (13°46'S 47°15'W, IBGE); O. stramineus – F: MN 46410σ / M: MN 34439β
Flores de Goiás (14°26'S 47°03'W, IBGE); O. nigripes – F: UFPB 1826*, 1827*, 1831*, 1833*, 1838*, 1839*, 1843*,
1846*, 1848*, 1850*, 1852*; M: UFPB 1830*, 1832*, 1834*, 1836*, 1840*, 1847*, 1849*, 1853*, 1854*
Bahia
Jaborandi (13°37'S 44°25' W, IBGE); O. fornesi – F: MN 62640β.
Minas Gerais
Conceição do Mato Dentro, Boca da Mata (19º10’S 43º33’W, IBGE); O. nigripes – F: MN 13428*, 13440*; M: MN
13386*, 13416*, 13433*, 13449*, 13454*, 13464*, 13475*
Espírito do Santo
Santa Teresa (19º56’S 40º35’W, IBGE); O. nigripes – F: SVS/ES 166*, 230*, 235*, 236*; M: SVS 163*#, 168*, 169*,
191*, 193*, 195*, 221*, 228*, 229*, 232*, 233*
São Paulo
Pedreira (22º44’S 46º54’W, IBGE); O. nigripes – F: CRB1422σ, 1424β; M: CRB 1209β, 1387β, 1436β; O. flavescens – M:
CRB1430σβ
Itararé (24º10’S 49º11’W, IBGE); O. nigripes– M: NHMV423 (holotype of Hesperomys eliurus)
Santa Catarina
Jaborá (27º10’S 51º44’W, IBGE); O. nigripes – F: LBCE 6910*#, 6911*#, 6949*#, 7344*, 7345*, 8096*, 8103*, 8115*,
8923*, 8928*, 8934*, 9721*, 9722*, MN 69934β, 69935β; M: LBCE 6953*#, 7326*#, 7329*, 7343*, 7346*, 8057*β,
8091*β, 8093*, 8098*, 8100*, 8108*, 8109*, 8160*β, 8926*, 8960*, 9607*, 9718*, 9786*, 9791*, 9841*
Itá (27°17'S 52°19'W, IBGE); O. nigripes – M: MN 62150β
Rio Grande do Sul
Aratiba (27°23'S 52°18'W, IBGE); O. nigripes – F: MN 62108β; M: MN 62113#β, 62155#β; unsexed: MN 62165β

Taim Ecological Station (32°29'S; 52°34'W, Miranda et al., 2009); O. nigripes – F: UFPB 616*, 617*; M: UFPB 613*,
618*
Rio de Janeiro
Teresópolis (22°24'S 42°57'W, IBGE); O. nigripes – M: MN 71984#β
VENEZUELA
Apure
Hato “El Frio”, aprox. 31 km W El Samán (7°55N 68°44’W); O. delicatus – M: AMNH 257241
Sucre
Finca Vuelta Larga, 9,7 km SE Guaraunos (10°33’N 63°07’W); O. delicatus – M: AMNH 257229, 257246
PARAGUAY
Paraguari Department
Sapucay (25°40'S 56°55'W, Weksler & Bonvicino, 2005); O. nigripes – F: USNM 121107*, 121403*, 121404*,
121406*, 121455*; M: USNM 121405*, 121456*
Tacuati (23°27'S 56°44'W, Bonvicino & Weksler, 1998); O. nigripes – F: USNM 293146*
Caaguazú Department
Caaguazú (25°28'S 56°00'W, Bonvicino & Weksler, 1998); O. nigripes – F: USNM 293144*, 292145*
Misiones Department
San Pablo, 20km de San Ignácio (26°52'S 57°03'W, Weksler & Bonvicino, 2005); O. nigripes – M: USNM 390106*
San Francisco, 36km NE de San Ignácio (26°52'S 57°03'W, Weksler & Bonvicino, 2005); O. nigripes – F: USNM
390109*; M: USNM 390107*
ARGENTINA
Chaco Province
Las Palmas (27°04'S 58°42'W, Weksler & Bonvicino, 2005); O. nigripes – M: USNM 236285*, 236286*, 236287*

APPENDIX 2. Specimen data, including museum and/or collector number, GenBank accession number and locality of
the specimens used in the molecular analysis. Reference codes are: 1 = Carroll et al. (2005), 2 = Miranda et al. (2009), 3
= Palma et al. (2005), 4 = Smith & Patton (1993), 5 = Rogers et al. (2009), 6=Percequillo et al. (2011), 7= our study,
8=Hanson et al. (2011), 9 =González-Ittig et al. (2010), 10=Palma et al. (2010), 11=Rocha et al. (2011). Acronyms of
Brazilian states are: AM (Amazonas), AP (Amapá), BA (Bahia), ES (Espírito Santo), GO (Goiás), MT (Mato Grosso),
PA (Pará), PR (Paraná), RJ (Rio de Janeiro), RR (Roraima), RS (Rio Grande do Sul), SC (Santa Catarina), SP (São
Paulo), TO (Tocantins). Museum and collector acronyms are: AMNH (American Museum of Natural History, USA),
ASNHC (Angelo State Natural History Collections, USA). BYU (Monte L. Bean Museum, Brigham Young University,
USA), CRB (Cibele Rodrigues Bonvicino), LB (Colección de Mamíferos del Centro Nacional Patagoónico, Argentina),
LBCE (Laboratório de Biologia e Parasitologia de Mamíferos Silvestres Reservatórios, FIOCRUZ, Brazil), LF (Luís
Flamarion), MN (Museu Nacional, UFRJ, Brazil), MVZ (Museum of Vertebrate Zoology, University of California,
USA), NK (Museum of Southwestern Biology, University of New Mexico, USA), SVS (Serviço de Vigilância em Saúde,
Ministry of Health, Brazil), TK (The Museum Texas Tech University, USA), UFPB (Universidade Federal da Paraíba,
Brazil). n/a = not applicable.
Voucher
Taxon Locality CYTB I7FGB Ref.
Number
O. microtis Bolivia, Santa Cruz BYU19014 AY439000 n/a 1
O. microtis Brazil, AM, Rio Juruá MVZ190401 HM594624 n/a 11
Brazil, TO, São Sebastião do
O. microtis CRB1448 n/a JQ282857 7
Tocantins
Tocantins
Tocantins
O. fornesi Brazil, GO, Minaçu MN36746 DQ826022 n/a 2
O. fornesi Brazil, GO, Minaçu MN36928 DQ826023 n/a 2
O. fornesi Brazil, BA, Jaborandi MN62640 n/a JQ282862 7
O. costaricensis Panama, Los Santos NK101588 EU192164 n/a 8
O. costaricensis Panama, Gamboa TK163369 GU393988 n/a 8
O. fulvescens Mexico, Chiapas ASNHC1665 EU294233 n/a 5
O. fulvescens Mexico, Chiapas ASNHC1666 EU294234 n/a 5
O. vegetus Costa Rica, Alajuela BYU15215 EU294251 n/a 5
O. vegetus Costa Rica, Alajuela BYU15217 EU294252 n/a 5
O. chacoensis Paraguay, Boqueron TK62932 EU258543 n/a 8
O. chacoensis Argentina, Salta Or22498 GU185904 n/a 9
O. stramineus Brazil, GO, Terezina de Goiás MN46410 JQ013747 n/a 7
O. stramineus Brazil, GO, Fazenda Regalito UFPB1827 DQ826027 n/a 2
O. stramineus Brazil, GO, Terezina de Goiás MN34439 n/a JQ282842 7
O. nigripes Brazil, ES, Monte Verde UFPB357 DQ826004 n/a 2
O. nigripes Brazil, GO, Caldas Novas MN37493 DQ825993 n/a 2
Brazil, PR, Parque Nacional de
O. nigripes JR78 DQ825988 n/a 2
Iguaçu
O. nigripes Brazil, RJ, Teresópolis MN71984 n/a JQ282868 7
O. nigripes Brazil, RS, Aratiba MN62108 n/a JQ282863 7
O. nigripes Brazil, RS, Mostardas MN37633 DQ826000 n/a 2
O. nigripes Brazil, RS, Rio Grande LF606 DQ825990 n/a 2
O. nigripes Brazil, RS, São Francisco de Paula MN37531 DQ825996 n/a 2

O. nigripes Brazil, RS, Sapiranga MN37565 DQ825997 n/a 2
O. nigripes Brazil, RS, Tenente Portela MN37501 DQ825994 n/a 2
O. nigripes Brazil, SC, Concórdia MN37685 DQ826002 n/a 2
O. nigripes Brazil, SC, Florianópolis MN37683 DQ826001 n/a 2
O. nigripes Brazil, SC, Itá MN62150 n/a JQ282865 7
O. nigripes Brazil, SC, Jaborá MN69934 n/a JQ282870 7
O. nigripes Brazil, SC, Jaborá MN69935 n/a JQ282872 7
O. nigripes Brazil, SC, Jaborá LBCE8057 n/a JQ282869 7
O. nigripes Brazil, SP, Guariba NK42266 EU192163 n/a 10
O. nigripes Brazil, SP, Pedreira CRB1209 n/a JQ282853 7
O. nigripes Brazil, SP, Pedreira CRB1422 GU126530 n/a 7
O. messorius Brazil, RR, Surumu MN37751 DQ826024 n/a 2
O. magellanicus Chile, Magallanes IPAT AY275705 n/a 3
O. longicaudatus Argentina, Neuquén LB012 AY275702 n/a 3
O. longicaudatus Argentina MVZ154463 GU393998 n/a 8
O. flavescens Bolivia, Oruro NK11547 AY452200 n/a 3
O. flavescens SP, Pedreira CRB1430 JQ013746 JQ282855 7
O. flavescens Brazil, GO, Rio Maranhão MN37699 DQ826013 n/a 2
O. delicatus Venezuela, Caño Delgadito TK138080 DQ227457 n/a 8
O. delicatus Venezuela, Finca Vuelta Larga AMNH257262 GU126529 n/a 6
O. rupestris Brazil, GO, Alto Paraíso MN50322 JQ013763 JQ282850 7
O. rupestris Brazil, GO, Alto Paraíso MN50326 JQ013764 JQ282851 7
O. rupestris GO, Alto Paraíso MN50286 n/a JQ282852 7
O. destructor Ecuador, Pichincha TEL1479 EU258544 n/a 5
O. destructor Ecuador, Pichincha TEL1481 GU393991 n/a 8
Oligoryzomys sp. Suriname, Nickerie TK17858 EU258546 n/a 8
Oligoryzomys sp. Brazil, AP, Tartarugalzinho MN37756 DQ826025 n/a 2
O. moojeni Brazil, GO, Cavalcante MN50287 JQ013772 JQ282845 7
O. moojeni Brazil, GO, Cavalcante MN50314 JQ013770 n/a 7
O. moojeni Brazil, GO, Mimoso de Goiás MN67087 JQ013777 n/a 7
O. moojeni Brazil, GO, Sítio D' Abadia LBCE11615 JQ013771 JQ282874 7
O. moojeni Brazil, GO, Sítio D' Abadia LBCE11648 JQ013773 JQ282875 7
O. moojeni Brazil, GO, Colinas do Sul MN36220 DQ826016. n/a 2

O. moojeni Brazil, GO, Colinas do Sul MN36357 DQ826017 n/a 2
O. moojeni Brazil, GO, Colinas do Sul MN36426 DQ826018 n/a 2
O. moojeni Brazil, GO, Minaçu MN 36832 DQ826019 n/a 2
O. moojeni Brazil, GO, Uruaçu MN37282 DQ826020 n/a 2
O. moojeni Brazil, GO, Uruaçu MN37441 DQ826021 n/a 2
O. utiaritensis Brazil, MT, Sapezal MN75592 JQ013753 JQ282876 7
O. utiaritensis Brazil, MT, Sapezal MN75595 n/a JQ282879 7
O. utiaritensis Brazil, PA, Castelo dos Sonhos MN75609 JQ013762 n/a 7
O. utiaritensis Brazil, PA, Castelo dos Sonhos MN75612 JQ013776 JQ282884 7
O. utiaritensis Brazil, PA, Castelo dos Sonhos MN75613 JQ013761 JQ282885 7
O. utiaritensis Brazil, MT, Campo Novo do Parecis MN75616 JQ013754 JQ282886 7
O. utiaritensis Brazil, MT, Campo Novo do Parecis SVS306 JQ013765 JQ282887 7
O. utiaritensis Brazil, MT, Campo Novo do Parecis SVS357 JQ013750 JQ282892 7

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Zootaxa 3220: 1–28 (2012) ISSN 1175-5326 (print edition)

The taxonomic status of the Castelo dos Sonhos Hantavirus reservoir,

RODRIGO AGRELLOS*1, CIBELE R. BONVICINO*1,2, ELIZABETH SALBÉ T. ROSA3, APARECIDO A. R.

Accepted by P. Gaubert: 27 Jan. 2012; published: 2 Mar. 2012 1

Material and methods

2 · Zootaxa 3220 © 2012 Magnolia Press AGRELLOS ET AL.

TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS Zootaxa 3220 © 2012 Magnolia Press · 3

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TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS Zootaxa 3220 © 2012 Magnolia Press · 5

Species / Head- Tail Molar Dorsal color Ventral color

1. Other O. microtis, including topotypes, are tinged with buff

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TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS Zootaxa 3220 © 2012 Magnolia Press · 7

Holotype of O. utiaritensis O. utiaritensis O. moojeni O. nigripes

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TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS Zootaxa 3220 © 2012 Magnolia Press · 9

Taxon 2n FNa References

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TAXONOMIC STATUS OF OLIGORYZOMYS UTIARITENSIS Zootaxa 3220 © 2012 Magnolia Press · 15

Oligoryzomys utiaritensis J. A. Allen, 1916

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RODRIGO AGRELLOS1, CIBELE R. BONVICINO1,2, ELIZABETH SALBÉ T. ROSA3, APARECIDO A. R.