Mammalian Biology

Zeitschrift fuÈr SaÈugetierkunde

www.elsevier.de/mammbiol

Original investigation
A new species of Wiedomys (Rodentia:
Sigmodontinae) from Brazilian Cerrado
By P. R. GONCËALVES, FRANCISCA C. ALMEIDA, and CIBELE R. BONVICINO
Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; The American Museum of Natural
History, New York, USA; Genetics Division, Instituto Nacional de CaÃncer and IOC ± FIOCRUZ, Rio de Janeiro, Brazil

Receipt of Ms. 06. 10. 2003

Acceptance of Ms. 15. 12. 2003

Abstract
On the basis of integrated analyses of karyologic, molecular and morphologic data of Wiedomys
samples distributed throughout north-eastern Brazil, we suggest the existence of more than one
evolutionary lineage within the genus and describe a new species apparently restricted to western
Bahia state, in the Cerrado morphoclimatic domain. This new taxon differs from W. pyrrhorhinus
by narrower incisive foramina, smaller molar rows, presence of an alisphenoid strut, and diploid
number of 60 chromosomes. Phylogenetic analyses of cytochrome b sequences of this new species
and samples of W. pyrrhorhinus from the Caatinga of Bahia state, depict the two taxa as reciprocally
monophyletic clades separated by 14.7±15.6% corrected genetic distance. This new species and
W. pyrrhorhinus are allopatric, the former occurring in the Cerrado morphoclimatic domain and the
latter endemic of the Brazilian semi arid Caatinga.

Key words: Wiedomys, new species, phylogeny, karyotype

Introduction
The Caatinga and Cerrado morphoclimatic cation of geographic limits of endemic ro-
domains occupy an extensive area in South dent species from Caatinga (Oliveira et al.
America, corresponding to approximately 2003). In this latter case, one of the poorly
2 650 000 km2, which together with the Cha- known rodent taxa apparently restricted to
co is commonly termed as the open diago- the Caatinga biome in northeastern Brasil
nal belt due to the predomination of xeric is the red nosed mouse genus Wiedomys
formations and grasslands dissected by (Oliveira et al. 2003), the unique extant re-
semideciduous forests (Eiten 1972; Reis presentative of the tribe Wiedomyini (Reig
1976). The knowledge of this region in- 1980).
creases with the descriptions of new rodent Recent phylogenetic studies have consid-
species from Cerrado (Bonvicino and ered this genus as an independent and basal
Weksler 1998; Bonvicino 2003; Langguth offshoot within the subfamily Sigmodonti-
and Bonvicino 2002) and by the clarifi- nae with apparently no closely related ex-

1616-5047/05/70/01-046 $ 30.00/0. Mamm. biol. 70 (2005) 1´ 46±60

 A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado 47

tant tribal groups (Steppan 1995; Smith and

Patton 1999). Despite the relatively wide
distribution of Wiedomys throughout north-
eastern Brazil (Thomas 1928; Pine 1980;
Oliveira et al. 2003), only W. pyrrhorhinus
(Wied, 1821) has been regarded as extant
species since the description of the genus
(Hershkovitz 1959). Few ecologic (Mares
et al. 1981; Streilen 1982 a) and karyologic
(Maia and Langguth 1987) studies were
carried out with W. pyrrhorhinus as a conse-
quence of its rarity and occasional occur-
rences in localities of the Caatinga (Strei-
len 1982 b). In the same way, morphologic
and genetic variability within and among
populations of W. pyrrhorhinus is poorly
known and the species diversity within this
genus may have been neglected.
Here we describe a new Wiedomys species
from the Cerrado morphoclimatic domain Fig. 1. Geographic location of ($) type-locality of
Wiedomys pyrrhorhinus and (*) samples analyzed in
on the basis of integrated karyologic, mole-
this study: Bahia state: (1) Tremedal, (2) Jaborandi,
cular (cytochrome b) and morphological an- (3) CaetiteÂ, (4) Seabra, (5) Canudos, (6) Feira de San-
alyses of samples from north-eastern Brazil. tana. Alagoas state: (7) QuebraÃngulo, (8) Santana do
Ipanema. Pernambuco state: (9) Caruaru, (10) Pes-
queira, (11) Bom Conselho (karyotyped by Maia and
Material and methods Langguth 1987) and (12) Cachoeirinha (karyotyped
by Maia and Langguth 1987). The shaded area deli-
We collected and karyotyped 5 specimens of mits the Caatinga biome as defined by IBGE (1965).
Wiedomys from 2 Brazilian localities in Bahia
state (Fig. 1): Jaborandi, Faz. SertaÄo do Formoso
(14°37¢ S, 45°51¢ W) (males MN 61657, MN 67023, MEU1 in an ABI Prismä 377 automatic DNA
and female MN 67022); and Caetite (13°50¢ S, sequencer following Smith and Patton (1993).
42°23¢ W) (males MN 63357 and MN 63420). All Sequences were manually aligned using the Se-
specimens were deposited in Museu Nacional quence Navigator software (Applied Biosystems,
(MN), Rio de Janeiro, Brazil. Inc. 1994) and deposited in GenBank. We also
Chromosome preparations from these individuals included sequence data available in GenBank
were obtained from short-term bone marrow cul- for one exemplar of Wiedomys pyrrhorhinus
tures. Cell suspensions were cultivated for two from Canudos (GenBank AF108685, Fig. 1, local-
hours at 37° C in RPMI 1640 culture medium sup- ity 5), Bahia state, and for selected species of
plemented with 20% fetal calf serum, ethidium the muroid rodents Oryzomys nitidus (Gen-
bromide (5 lg/ml) and colchicine (10±6 M). FNa Bank AF251529), Oryzomys lamia (AF181273),
refers to autosome fundamental number. Karyo- Eligmodontia puerelus (AF159289), Eligmodon-
logic data were compared with previously pub- tia morgani (AF108691), Thomasomys aureus
lished karyologic data of specimens from Bom (U03540), Thomasomys gracilis (AF108674) and
Conselho and Cachoeirinha, Pernambuco state Scotinomys teguina (AF109796). This last taxon
(localities 11 and 12 in Fig. 1, see Maia and Lang- was used as outgroup on the basis of previous
guth 1987). muroid phylogeny assessments (Smith and Pat-
DNA samples of the 3 specimens from Jaborandi ton 1999). Kimura 2 parameters model (K2p)
and 1 from Caetite were isolated from livers was used to calculate genetic distances between
preserved in ethanol following the procedures haplotypes and for constructing neighbour-join-
of Sambrook et al. (1989). Cytochrome b mt ing dendrograms using the software MEGA 2.1
DNA (ca. 1140 bp) was amplified with primers (Kumar et al. 2000). Maximum parsimony trees
MVZ 05 and MVZ 14 and sequenced with the were obtained through heuristic searches using
same primers, in addition to MVZ 16 and the tree-bisection-reconnection branch-swapping
48 P. R. GONCËALVES et al.

algorithm in PAUP 4.0b10 (Swofford 1999), with bridge (PB), length of incisive foramen (LIF),
all sites equally weighted. Confidence intervals breadth of incisive foramen (BIF), length of max-
for neighbour-joining and maximum parsimony illary molar row (LM), breadth of first maxillary
trees were obtained by Bootstrap analysis based molar (BM1), external alveolar breadth (M1M),
on 1000 replicates. cranial height (CH), rostrum length (RL), ros-
For morphologic comparisons, in addition to the trum breadth (BRO), least interorbital breadth
karyotyped specimens from Jaborandi and Cae- (LIB), zygomatic breadth (ZB), and breadth of
titeÂ, we included 61 specimens from the following braincase (BB). Only adult animals (with all mo-
Brazilian localities: Bahia state, Seabra (MN lar teeth erupted and functional) were included
18507, 18509±511, 18514, 18516±517, 18705, in morphometric comparisons. Principal compo-
18707±708); Feira de Santana (MN 18715±716, nent analyses were performed to explore patterns
18719±720, 18723, 18760±762, 18765±766); Ala- of size and shape variation within and among
goas state, Quebrangulo (MN 18549, 18776±777, samples using the software SYSTAT and Mat-
18779±781, 18784, 18786, 61603), Santana do Ipa- Lab 4.0 (MathWorks, Inc.) routines available at
nema (MN 18597, 18686, 18690, 18767±772, http://www.biol.ttu.edu/Faculty/FacPages/Strauss/
18774); Pernambuco state, Caruaru (MN 18468, Matlab/matlab.htm.
18470, 18472, 18661±663, 18666±668, 60736), Pes-
queira (MN 18588, 18675, 18685, 18743±744,
18747, 52229, 60742±746). Morphological descrip-
tions and comparisons were based on qualitative
Results
anatomical external and cranial characters, using
terminology referenced by Reig (1980) and Voss Karyologic analyses
(1988, 1993). Karyologic analyses from Caetite speci-
For morphometric characterisation we used exter- mens showed 2n = 62, FNa = 90 (Fig. 2).
nal measurements recorded on specimen tags, and
measured 16 cranial dimensions defined by Bon-
The autosomal complement is composed
vicino and Weksler (1998) with the aid of a digi- by 15 pairs of biarmed chromosomes vary-
tal calliper. Greatest skull length (GSL), condy- ing in size from large to small, and 15 pairs
lo-incisive length (CIL), breadth of the occipital of acrocentric chromosomes varying in size
condyles (BOC), length of diastema (LD), palatal from medium to small. The X chromosome

Fig. 2. Conventional Giemsa coloration of specimens of Wiedomys from (A) Jaborandi and (B) CaetiteÂ.
 A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado 49

is a medium sized acrocentric, and the Y Canudos samples (Fig. 3). In both analyses
chromosome a small sized metacentric. Oryzomys, Eligmodontia and Thomasomys
Karyologic analysis from Jaborandi speci- were depicted as monophyletic genera,
mens showed 2n = 60, FNa = 88 (Fig. 2). although their intergeneric relationships
The autosome complement is composed by are not fully resolved. The only exception
15 pairs of biarmed chromosomes one large, is the clustering of Thomasomys with Wied-
the largest of the chromosome complement, omys in both NJ and MP analyses, although
and 14 decreasing in size from medium to there is no bootstrap support in the last one.
small, and 14 acrocentric pairs varying in
size from medium to small. The X chromo-
some is a medium sized acrocentric, and Morphological variation
the Y chromosome a small acrocentric.
Considering the karyologic and molecular
distinctiveness of the Jaborandi sample
Molecular analyses (2n = 60, FNa = 88), we evaluated the hy-
pothesis that this population is morpholog-
The genetic distance estimates revealed re- ically distinct from the Caetite sample
latively low divergence levels between Cae- (2n = 62, FNa = 90) including 6 additional
tite and Canudos haplotypes (0.025 K2p) larger samples from Bahia, Alagoas and
and between Jaborandi haplotypes (0.013± Pernambuco states. In addition to the prin-
0.015 K2p) (Tab. 1). Conversely, higher val- cipal components analysis we evaluated the
ues of genetic divergence were found in craniometric diagnosis of the sample from
comparisons between Jaborandi haplotypes Jaborandi by inspecting bivariate plots por-
and either Caetite or Canudos haplotypes traying combinations between the different
(0.147±0.156 K2p). These values are higher craniometric characters.
than those found between selected species The principal component analysis summar-
of Oryzomys (0.079 K2p), Thomasomys ized the majority (62.9%) of the total cra-
(0.143 K2p) and Eligmodontia (0.123 K2p) niometric variance in the first component
(Tab. 1). (PC1) and only 7.5% in the second compo-
The neighbour-joining and maximum par- nent (Fig. 4 a). The PC1 rather represents
simony trees placed Wiedomys specimens the cranial variability related to variation in
in a well-supported monophyletic clade size due to the positive correlation of all
(100% bootstrap value), divided in two craniometric variables with this axis (GSL-
major lineages, one constituted by Jaboran- 0.2348; CH-0.1218; CIL-0.2489; ZB-0.2601;
di specimens and another by Caetite and BB-0.1318; LIB-0.1239; RL-0.3164; BRO-

Table 1. Pairwise Kimura 2-parameters genetic distances among cytochrome b haplotypes used in molecular
analyses.

Taxa 1 2 3 4 5 6 7 10 11 12

1. W. pyrrhorhinus ± CaetiteÂ
2. W. pyrrhorhinus ± Canudos 0.025
3. Wiedomys sp.n. ± Jaborandi 0.156 0.147
4. Wiedomys sp.n. ± Jaborandi 0.152 0.150 0.014
5. Wiedomys sp.n. ± Jaborandi 0.154 0.148 0.015 0.013
6. Oryzomys lamia 0.202 0.203 0.205 0.214 0.214
7. Oryzomys nitidus 0.201 0.205 0.211 0.218 0.222 0.079
10. Eligmodontia puerelus 0.218 0.216 0.242 0.248 0.248 0.182 0.185
11. Eligmodontia morgani 0.206 0.206 0.227 0.225 0.226 0.192 0.204 0.123
12. Thomasomys aureus 0.223 0.207 0.215 0.217 0.214 0.197 0.200 0.242 0.232
13. Thomasomys gracilis 0.209 0.190 0.221 0.218 0.218 0.179 0.187 0.235 0.219 0.143
50 P. R. GONCËALVES et al.

Fig. 3. Phylogenetic relationships among Wiedomys haplotypes and selected muroid species based on 1 140 bp
of the cytochrome b gene. (A) Neighbour-joining dendrogram and (B) single Maximum Parsimony tree obtained
(tree statistics: 944 steps, C. I. = 0.66, R. I. = 0.63). Numbers above nodes are bootstrap support values esti-
mated on 1000 replications.

0.2294; BOC-0.1262; BL-0.2380; BM1± foramina (Fig. 4 b; Tab. 2). In this combina-
0.1672; LM-0.1203; M1M-0.1824; PB-0.1998; tion, the Jaborandi specimen diverges from
LIF-0.2767; LD-0.3293; BIF-0.4344), while most samples displaying relatively shorter
PC2 depicts shape variation. Most samples maxillary molar row and narrower incisive
overlap along PC1 and PC2 suggesting foramina. One specimen from Caetite pre-
extensive intrapopulational size and shape sents the widest incisive foramina whereas
variation across samples. Specimens from the other specimen is superimposed within
Caetite generally have larger skulls while most samples presenting intermediate val-
the specimen from Jaborandi presents a re- ues for the two characters. The qualitative
latively smaller skull. The remaining sam- character variation related to the alisphe-
ples are intermediate in size and display noid foramina further contributed for the
some specimens with scores near Caetite or discrimination of the specimens from Jabor-
Jaborandi samples. andi in relation to the other samples. The
The inspection of bivariate plots revealed Jaborandi specimens present the buccina-
that the specimen from Jaborandi segre- tor-masticatory foramen separated from
gates best from the other samples in com- the foramen ovale accessorius by a distinct
parisons considering both length of maxil- bony alisphenoid strut whereas specimens
lary tooth row and breadth of incisive from the remaining samples lack this struc-
 A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado 51

Fig. 4. Bivariate plots of (A) individual scores of the first and second principal components, (B) measurements
of breadth of incisive foramina and maxillary tooth row length of specimens. Samples: (*) Feira de Santana, Ba-
hia state; (*) Seabra, Bahia state; (^) Pesqueira, Pernambuco state; (´) Caruaru, Pernambuco state; (&) San-
tana do Ipanema, Alagoas state; (^) Jaborandi, Bahia state; (~) QuebraÃngulo, Alagoas state; (~) CaetiteÂ, Ba-
hia state. See Material and methods for character abbreviations.

ture (Fig. 5). Samples from Pesqueira, San- (MN 61657) is represented by skin and
tana do Ipanema and Seabra were the only complete skull, M2 and M3 are not fully
polymorphic samples for this character, pre- erupted. The adult male (MN 67023) is re-
senting one out of 10 individuals bearing presented only by skin.
the alisphenoid strut. Nevertheless, these Etymology: Wiedomys pyrrhorhinus has
samples lie well apart from Jaborandi speci- been suggested as a Caatinga endemic and
men in comparisons of breadth of incisive the new form is apparently found outside
foramina and length of maxillary tooth row this biome, in a semi deciduous forest in
mentioned above. Therefore, the combina- western Bahia, an area enclosed by the Cer-
tion of these two craniometric characters rado morphoclimatic domain. Therefore,
with the conditions of the alisphenoid strut the epithet cerradensis refers to the mor-
consistently discriminates the Jaborandi phoclimatic domain where this species was
specimen from other samples of Wiedomys. found.
Diagnosis: A small sigmodontine rodent
with tail longer than head and body, pelage
Wiedomys cerradensis sp. nov.
of muzzle, ears, eye-ring and rump bright
Holotype: An adult female (MN 67022) re- orange, narrow incisive foramina and small
presented by skin, skull, mandible, partial molar rows, buccinator-masticatory and ac-
skeleton and ethanol-preserved liver tis- cessory oval foramina separated by a dis-
sues, field number CRB 1878 (Fig. 6). Col- tinct alisphenoid strut, and diploid number
lected at Fazenda SertaÄo do Formoso of 60 chromosomes.
(14°37¢888² S and 45°51¢293² W), Jaborandi, Measurements: See table 2.
state of Bahia, October 29. 1999, by C. R. Distribution: Known only from the type lo-
Bonvicino and J. F. Vilela in a semi decid- cality.
uous forest in Brazilian Cerrado. External characters: Dorsal pelage soft and
Paratypes: Two males, collected at the same dense with overall color grizzled-brown,
locality and date of the holotype. The young but with the rump, ears and muzzle dis-
 52
Table 2. Measurements (in mm) of the holotype of Wiedomys cerradensis (n. sp.), means and interval of variation (Max-Min) of selected samples of W. pyrrhorhinus from
northeastern Brazil. Char = variables, see material and methods from abbreviations, HB = head-body length, T = tail length, HF = hind foot length without claw, E = ear
length.

P. R. GONCËALVES et al.
n. sp. Wiedomys pyrrhorhinus

Char type Caetite Seabra Feira de Santana QuebraÃngulo Santana do Caruaru Pesqueira
Ipanema

HB 107 116.5 111 116 105.7 110 109 110.3

107±126 102±120 100±125 95±120 100±130 100±115 100±120
T 142 188.5 184 173 158 162 149.6 168.8
183±194 172±205 149±188 140±180 145±180 110±164 150±185
HF 25.5 25.5 26 27 25 24 24.4 25.8
25±26 24±28 26±30 24±27 22±25 22±26 25±29
E 19 21.5 21 21 19 21.5 19.6 19.7
20±23.5 20±23 19±23 18±20 19±24 17±21 18±20
GSL 27.85 32.26 29.93 29.82 29.21 28.38 27.88 28.92
32.05±32.47 28.22±32.1 27.39±30.99 26.87±30.79 26.6±30.61 25.7±29.29 27.05±30.58
CH 8.54 9.55 8.87 8.96 8.80 8.72 8.55 8.81
9.53±9.57 8.14±9.4 8.66±9.32 8.24±9.44 7.91±9.19 8.23±8.82 8.4±9.29
CIL 25.15 29.85 27.12 27.22 26.48 25.79 25.37 26.52
29.11±30.6 25.29±28.91 24.72±28.26 24.17±27.95 24.22±27.84 22.95±26.95 24.61±28.62
ZB 13.89 16.78 15.25 15.19 14.50 14.51 14.14 15.03
16.48±17.08 14.41±16.14 14.03±16.06 13.07±15.56 13.84±15.45 13.5±14.83 14.06±16.35
BB 11.61 13.1 11.99 11.95 11.86 11.77 11.39 12.04
13±13.2 11.5±12.33 11.59±12.2 11.22±12.29 11.05±12.27 11.28±11.74 11.38±13.14
LIB 4.44 4.65 4.54 4.6 4.37 4.34 4.40 4.41
4.6±4.69 4.26±4.79 4.28±4.95 3.97±4.61 4.22±4.75 4.06±4.62 4.22±4.7
RL 9.55 12.13 10.85 10.75 10.73 10.40 9.95 10.47
12.12±12.14 9.63±11.73 9.36±11.52 9.86±11.71 9.46±11.48 8.13±10.46 9.38±11.68
 n. sp. Wiedomys pyrrhorhinus

Char type Caetite Seabra Feira de Santana QuebraÃngulo Santana do Caruaru Pesqueira
Ipanema

BRO 4.67 5.14 4.85 4.81 4.78 4.61 4.62 4.81

A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado

5±5.29 4.46±5.23 4.56±5.14 4.26±5.19 4.17±5.03 4.44±4.84 4.33±5.67
BOC 6.04 6.67 6.48 6.56 6.33 6.48 6.27 6.74
6.67 6.23±7.05 6.38±6.79 5.86±6.84 6.24±6.73 6.04±6.54 6.24±7.23
BU 5.75 6.98 6.04 5.95 6.14 5.94 5.59 6.13
6.84±7.13 5.59±6.54 5.32±6.31 5.45±6.69 5.61±6.43 5.25±6.16 5.66±6.49
BM1 1.28 1.36 1.31 1.29 1.18 1.23 1.18 1.26
1.31±1.42 1.25±1.41 1.22±1.36 1.13±1.27 1.16±1.3 1.13±1.26 1.14±1.38
LM 4.35 4.85 4.90 4.76 4.62 4.70 4.65 4.77
4.8±4.9 4.62±5.45 4.45±5 4.46±4.87 4.41±4.8 4.31±5 4.4±5.27
M1M 5.1 5.67 5.40 5.39 5.10 5.24 5.03 5.28
5.65±5.69 5.13±5.67 5.15±5.73 4.43±5.43 5.06±5.42 4.79±5.4 5.08±5.73
PB 4.16 5.18 4.67 4.53 4.34 4.40 4.32 4.46
5.15±5.21 4.25±5.05 4.36±4.79 3.79±4.58 4.23±4.6 3.79±4.44 4.22±4.81
LIF 5.34 6.68 6.32 6.28 6.11 5.90 5.86 6.15
6.47±6.89 5.74±6.68 5.8±6.91 5.52±6.52 5.43±6.37 5.4±6.48 5.46±6.61
LD 6.24 7.89 7.10 7.17 6.84 6.59 6.45 6.84
7.7±8.08 6.74±7.65 6.61±7.77 5.84±7.46 6.05±7.41 5.53±6.95 6.46±7.4
BIF 1.78 2.45 2.36 2.25 2.22 2.07 2.02 2.16
2.24±2.66 2.21±2.62 2.09±2.62 1.9±2.51 1.83±2.45 1.88±2.36 1.83±2.45

53
54 P. R. GONCËALVES et al.

Fig. 5. Lateral view of the alisphenoid region showing the different conditions of the alisphenoid strut between
specimens from (A) Jaborandi and (B) CaetiteÂ. (ab) auditory bulla, (als) alisphenoid strut, (bmf) buccinator-mas-
ticatory foramen, (foa) foramen ovale accessorius. Scale bar = 1 mm.

tinctly brightened orange. Two types of is well delineated from laterals presenting
hairs of similar length (8±9 mm) are present entirely white hairs extending from the gu-
in dorsum, a rather homogeneously melanic lar to the inguinal region, including the ven-
type and a banded type with basal, subapi- tral region of forelimbs and medial region
cal and apical bands. In the head and dor- of hind limbs. Pinnae rounded and exter-
sum, banded hairs have a large plumbeous nally covered by entirely orange hairs,
basal band (7 mm), a very short orange co- which originate in the proximal helix and
lored subapical band (2 mm) and a more in the region just anterior to the base of
distal clay coloured apical band (4 mm). the ear. Eyes surrounded by short orange
Melanic hairs are entirely black in their dis- hairs forming a conspicuous orange eyering.
tal parts and mixed with banded hairs pro- Muzzle well furred by small orange hairs,
vide the grayish-brown tone of the dorsum. which extend to the caudal limit of the mys-
In the rump, banded hairs present apical tacial vibrissae array; mystacial vibrissae
and sub apical bands apparently fused in a are well developed (3±3.5 cm long), the
large (6±7 mm) rich orange layer, strength- longest extending beyond the distal limit of
ening the brightened orange tone of the pinnae; two supraciliary and one genal vi-
rump. Laterals of body are slightly grayer brissae also present and shorter (1.8±2 cm).
than dorsum due to the paler and wider Manus and pes dorsally furred by entirely
sub apical and apical bands. Ventral pelage orange hairs; silvery ungual tufts are pre-
 A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado 55

sent in the distal phalanxes of digits 2±5 in oped in one side of the skull, represented
manus and digits 1±5 in pes. Volar surface only by a thin bony projection. Paraptery-
of manus displays well-developed and fle- goid fossae shallow and large, wider than
shy metacarpal pads, the thenar being the mesopterygoid fossa at its caudal limit and
largest with an oblong shape and the hy- perforated by two small fontanelles. The
pothenar about the same size of the posterior openings of the oval foramina are
interdigital pads. Tail larger than combined reduced and unequally divided by a slender
head and body length; entirely dark brown strut of the aliphenoid. Hamular process
with its proximal ventral surface slightly slender, delimiting a small postglenoid fe-
paler; dorsal and ventral regions covered nestra and a large subesquamosal foramen.
by small black and white hairs averaging Ectotympanic (auditory bulla) notably in-
2 scales in length, which become longer to- flated and globular in its ventral surface
wards the tail tip, but do not form a distal and contacted to the squamosal by the teg-
hair tuft. The single adult female examined men tympani and hamular process; stape-
(MN 67022) displays 8 mammae: 2 inguinal, dial process well developed. Petrotympanic
2 abdominal, 2 thoracic, 2 pectoral. The sole fissure deep, large stapedial and spheno-
young specimen examined (MN 61657) dif- frontal foramina present, providing evi-
fers from adult individuals externally only dence of the carotid circulation pattern 1
by presenting more brownish dorsal colora- (Voss 1988). Mastoid fenestra quadrangular
tion throughout the flanks due to the richer and relatively large, occupying approxi-
orange colour of apical and sub apical mately 1/3 of the mastoid surface.
bands of banded hairs, thus, providing a less Mandible: Coronoid process small, not pro-
conspicuously brightened rump. jecting posterodorsally beyond the level of
Cranial characters: Skull with relative short the condyloid process and delimiting a
rostrum and laterally expanded braincase. small sigmoid notch. Angular notch deep.
Frontal bones with anteriorly convergent Condylar process well developed, extending
lateral outlines and sharply defined su- posterodorsally to the level of the angular
praorbital edges, but without distinct su- process. Inferior masseteric crest more pro-
praorbital crests. Suture between parietals nounced than superior masseteric crests.
and frontals V-shaped. Interparietal broad Dental characters: Upper incisors opistho-
and wide, not contacting the squamosal lat- dont with rounded yellow enameled bands
erally. Nasals short, strongly depressed in in anterior face. Molars brachyodont and
the median region of the fronto-nasal su- crested with alternated cusps. Upper series
ture. Zygomatic plate narrow not projected (M1±M3) parallel sided. M1 with rounded
anteriorly, with roughly vertical anterior anterior margin of procingulum. M2 squared
outline. Zygomatic notch moderately deep and M3 triangular with reduced metacone
and broad. Incisive foramina extending to and hypocone. Anteromedian flexus well de-
the line of M1 protoflexus, with 2/3 of the fined in M1, unequally dividing the antero-
septum formed by premaxillary bones. cone in a small anterolingual conule and
Palate short, not extending posteriorly be- larger anterolabial conule; anteroloph slen-
yond the line of M3 hypoflexus, presenting der in M1 and obsolescent in M2. Anterior
a pair of posterolateral palatal pits. Mesop- (M1) and median mures (M1, 2) diagonally
terygoid fossa narrow with U-shaped ante- oriented. Mesoloph distinct in M1 and M2,
rior notch and straight lateral margins. originated at the median mure and lingually
Sphenopalatine vacuities well developed fused to mesostyle. Protostyle and enteros-
occupying approximately 2/3 of the basi- tyle present as low stylar cusps. Mesoflexus
sphenoid roof. A distinct alisphenoid strut and posteroflexus reduced as enameled is-
separates the buccinator-masticatory and lands in M1 and M2. Procingulid of m1 nar-
accessory oval foramina. Even the young rower than breadth of m2, with anteroconid
specimen (MN61657) presented the ali- equally divided by a deep antermedian flex-
sphenoid strut although incompletely devel- id. Mesolophid absent. Protoflexid wide in
56 P. R. GONCËALVES et al.

Fig. 6. Dorsal, ventral and lateral view of the skull, lateral view of mandible and occlusal view of upper and lower
molar rows of Wiedomys cerradensis holotype (female MN 67022).
 A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado 57

m1 and deep in m2. Mesostylid present in with the description of the fossil Wiedomys
m1 and m2. Hypoflexid wide, presenting a marplatensis from the Upper Pliocene de-
well developed ectostylid in the lingual mar- posits of Argentina (Quintana 2002), the
gins of m1 and m2. Posteroflexid deep and paleontological evidence accumulated so
pronounced in m2 but reduced to a small en- far suggests that W. pyrrhorhinus may be an
amel island in m3. extant representative of an ancient group of
Comparisons: Wiedomys cerradensis differs sigmodontine rodents that was distributed
from W. pyrrhorhinus in karyotype (diploid throughout South America during Plio-
and fundamental numbers), and by present- Pleistocene times. Likewise, the molecular
ing an alisphenoid strut separating the buc- phylogenetic studies have confirmed the sta-
cinator-masticatory foramen from the acces- tus of Wiedomys as an independent and very
sory oval foramen combined with smaller divergent lineage within the Sigmodontinae
cranial and body dimensions (Tab. 2), of (Smith and Patton 1999). Despite the num-
which the most distinctive are the relative ber of discussions concerning the systema-
small molar row length (4.35 mm) and nar- tics and evolution of W. pyrrhorhinus, most
row incisive foramina (1.78 mm). Both spe- of them have been centered on its interge-
cies display the same colour pattern, with neric relationships and no attempt has been
rumps, ears and eye-rings brightened or- made to understand its intraspecific variabil-
ange, strongly contrasted to the grizzled- ity.
brown color of head and dorsum. W. cerra- The results obtained in this study on the
densis differs from the recently described basis of integrated karyologic, mitochon-
fossil species W. marplatensis (Quintana drial sequence and morphologic data are in-
2002) by displaying a much more simplified dicative of a higher diversity and complex-
m3, which lacks a distinct protoflexid and ity within the genus. The karyotypes here
has a posteroflexid reduced to an enamel is- described differed from those previously re-
land. ported from specimens collected in Ca-
choeirinha and Bom Conselho, Pernambu-
co state (2n = 62, FNa = 86; Maia and
Discussion Langguth 1987). The Jaborandi karyotype
(2n = 60, FNa = 88) departs to a greater ex-
Since its original description as Mus pyr- tent from the previously published 2n = 62,
rhorinos by Wied (1821) on the basis of a FNa = 86 karyotype by differing in both
specimen collected in southern Bahia, diploid and fundamental number, where-
W. pyrrhorhinus had been at times included as the karyotype from Caetite (2n = 62,
or related to a number of divergent sigmo- FNa = 90) differs only in fundamental num-
dontine genera such as Hesperomys (Bur- ber, having two additional pairs of biarmed
meister 1854), Oryzomys (Thomas 1886), chromosomes. Maia and Langguth (1987)
Rhipidomys (Trouessart 1898) and Tho- reported heteromorphism of pericentric in-
masomys (Thomas 1928; Gyldenstolpe version affecting pair number 16 in one kar-
1932), owing to its divergent set of cranial yotyped specimen from Pernambuco State.
and external features. Based on the unique- The distinctive morphology of this auto-
ness of character-states of this taxon, some pair (a submetacentric with very short
Hershkovitz (1959) described the monotyp- arm) made it easily recognizable in the two
ic genus Wiedomys and expressed his doubts karyotypes here described, which appar-
regarding its suprageneric and tribal affi- ently present no heteromorphism for this
nities. Finally, Reig (1980) formalized the pair. Both 2n = 60, FNa = 88 and 2n = 62,
uniqueness of Wiedomys as a very distinct FNa = 90 karyotypes described here lack
lineage within the Sigmodontinae, coining the large acrocentric pair 14, the largest of
the tribe Wiedomyini that included the fossil the chromosomal complement reported by
Cholomys pearsoni as the sole species re- Maia and Langguth (1987), which in the
lated to W. pyrrhorhinus. More recently, case of the 2n = 62, FNa = 90 karyotype
58 P. R. GONCËALVES et al.

may have evolved to one of its additional Jaborandi sample as a more divergent unit
biarmed chromosomes. The existence of within Wiedomys. In addition to karyologic
three different karyotypes within the genus divergence of Jaborandi specimens to the
raises the possibility of the existence of at 2n=62, FNa=90 and 2n=62, FNa=86 karyo-
least three previously unrecognised evolu- types, sequence divergence is high (14.7±
tionary units. We evaluated the significance 15.6%) in comparisons with either CaetiteÂ
of this variation to species limits within or Canudos haplotypes. These values are
Wiedomys by considering the additional higher than those found between species of
molecular and morphological evidence. other sigmodontine genera included in the
Morphological data show that despite the molecular analyses such as Oryzomys, Tho-
variation in chromosomal morphology be- masomys and Eligmodontia. Moreover, the
tween the 2n = 62, FNa = 86 (Maia and maximum parsimony and neighbour-joining
Langguth 1987) and 2n = 62, FNa = 90 analyses depicted the two karyotypes as
(CaetiteÂ) karyotypes, the samples from Per- two reciprocally monophyletic clades sug-
nambuco state (Pesqueira and Caruaru) gesting the existence of two divergent and
herein analysed overlapped extensively in relatively old evolutionary lineages within
craniometric characters with samples from the genus. The morphological analyses re-
central Bahia and Alagoas states. Moreover, vealed that the Jaborandi specimen displays
these samples share the absence of alisphe- a unique combination of traits related to
noid strut with the sole exception of one in- the alisphenoid, molar series length and in-
dividual in both Caruaru and Santana do cisive foramina width, which are useful for
Ipanema samples that present this structure. the taxonomic distinction of this new line-
The absence of any remarkable morphologic age within Wiedomys. Since the name Wied-
divergence among samples suggests that the omys pyrrhorhinus may be applied to the
2n = 62, FNa = 86 karyotype from Pernam- lineage represented by the central Bahia
buco state and the 2n = 62, FNa = 90 karyo- samples, the western sample from Jaborandi
type from Bahia state may represent a single represents an unnamed form, which we de-
species widely distributed along the semiar- scribed in the preceding sections as W. cer-
id Caatinga. Unfortunately, we do not have radensis.
sequence data available for samples from Wiedomys pyrrhorhinus has been consid-
Pernambuco state in order to provide more ered as one of the few endemic mammal
comprehensive comparisons. species of the Caatinga in a recent revision
The specimens from Caetite are the geo- of the geographic records of this species
graphically nearest relatives to the type spe- (Oliveira et al. 2003). The recognition of
cimen of W. pyrrhorhinus collected by the two evolutionary lineages within the genus
Prince Maximilian Wied-Neuwied in the Wiedomys, one presumably restricted to
caatingas along Riacho da Ressaca, at Tre- the Caatinga and the other apparently out-
medal municipality in Bahia state, thus, this er to this biome, raises an important evi-
name may be used to refer to samples from dence for the zoogeographic differentiation
the semiarid zone of Bahia, Alagoas and of the Caatinga fauna in relation to more
Pernambuco states analysed here. The mo- mesic surrounding biomes. Oliveira et al.
lecular analyses of the specimen from Ca- (2003) argued that the set of Caatinga
nudos and the 2n = 62, FNa = 90 sample endemic mammals, although reduced in
from Caetite revealed low genetic distances species number, comprises autochthonous
among their haplotypes (2.5% K2p), which and strikingly differentiated rodent genera
were grouped in a highly supported mono- favouring a hypothesis of an ancient origin
phyletic clade within the genus. These re- of the biome. In order to further evaluate
sults confirm the genetic cohesion of sam- the antiquity of the Caatinga we assessed
ples along central Bahia state. divergence dates between W. cerradensis
Karyologic, molecular and morphologic and the Caatinga endemic W. pyrrhorhinus,
comparisons were congruent in portraying correlating the amount of sequence diver-
 A new species of Wiedomys (Rodentia: Sigmodontinae) from Brazilian Cerrado 59

gence between these two lineages with time the genus (Quintana 2002). Therefore, the
considering the calibrated evolutionary rate molecular and fossil evidence available sug-
of 3rd position transversions of the cyto- gest that diversification within the genus
chrome b gene of sigmodontine rodents may have occurred during Pliocene times.
(2.3% sequence difference per million years
Smith and Patton 1993, 1999). Applying
the calibrated rate, the two lineages split at
approximately 3.4 millions of years before Acknowledgements
present, at the Upper Pliocene, corroborat- We are grateful to the owners of Fazenda SertaÄo
ing the hypothesis of an ancient origin of do Formoso for permission to collect animals at
the Caatinga rodent fauna proposed by their land, to S. Lindbergh, J. F. Vilela for their
Oliveira et al. (2003). The time estimates collaboration in fieldwork. To P. S. D'Andrea for
generated by the molecular analyses are specimens from Caetite and J. A. Oliveira for
also congruent with the fossil record of granting access to the Museu Nacional mammal
collection and for revising earlier versions of this
Wiedomys, which documents the first ap-
manuscript. To A. Langguth for the Zusamenfas-
pearance of the genus in the Upper Plio- sung. License to collect the specimens was
cene of the Buenos Aires province, Argen- granted by Brazilian Institute of Environment
tina, on the basis of the discovery of and Natural Resources (IBAMA).
W. marplatensis, the sole fossil species of

Zusammenfassung
Eine neue Art von Wiedomys (Rodentia: Sigmodontinae) aus dem Cerrado Brasiliens
Auf Grund von karyologischen, molekularen und morphologischen Daten beschreiben wir eine neue
Art von Wiedomys aus West Bahia Staat, in der Cerrado DomaÈne von Nord-Ost-Brasilien. Diese neue
Art unterscheidet sich von W. pyrrhorhinus durch schmaÈlere Foramina incisiva, kleinere Backenzahn-
reihe, einen Alisphenoid-Fortsatz und einen diploiden-Satz von 60 Chromosomen. Phylogenetische
Analysen von Cytochrom b Sequenzen der neuen Art und von W. pyrrhorhinus aus dem Caatinga
und aus Bahia weisen die beiden Taxa als Schwestergruppen aus, die durch eine korrigierte geneti-
sche Distanz von 14,7±15,6% getrennt sind. Die Wiedomys-Arten sind allopatrisch, eine kommt in
Cerrado und die andere in Caatinga vor.

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