andrades-miranda - chromosomes studies of seven species of oligoryzomys from brazil

8/8/2019 Andrades-Miranda - Chromosomes Studies of Seven Species of Oligoryzomys From Brazil

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ANDRADES-MIRANDA ET AL.CHROMOSOMES OF OLIGORYZOMYS November 2001 1081

FIG. 1.Collection sites (coordinates given in Appendix I): 1) Surumu, Roraima State (RR); 2)

Fazenda Sao Bento, Tartarugalzinho, Amapa State (AP); 3) including the localities: 40 km SW

Minacu, 55 km N Niquelandia, 20 km NW Colinas do Sul, and 40 km NE Uruacu; 4) Mamba, and

5) Ipameri, Caldas Novas, and Corumbaba, Goias State (GO); 6) Rio de Una, 10 km ESE Sao Jose,

Bahia State (BA); 7) Monte Verde, Esprito Santo State (ES); 8) Parque Nacional de Iguacu, Parana

State (PR); 9) Concordia, and 10) Costa de Dentro, Santa Catarina Island, Santa Catarina State (SC);

11) Parque Estadual do Turvo, Derrubadas, 12) Nonoai, 13) Aratiba, 14) Aracuri Ecological Station,

Muitos Capoes, 15) Iva, Tupancireta, 16) Caxias do Sul, 17) Tainhas, 18) Faxinal, Torres, 19) Sao

Francisco de Paula, 20) Riozinho, 21) Picada Verao and Alto Ferrabraz, Sapiranga, 22) Charqueadas,

23) including 2 localities: Rio Jacu, Eldorado do Sul, and Belem Novo, 24) Osorio, 25) Tramanda

Lagoon, Tramanda, 26) Quintao, 27) northern Tapes Bay, Tapes, 28) Capao do Leao, Mostardas,

and 29) Taim Ecological Station, Rio Grande, Rio Grande do Sul State (RS).

longing to the Sigmodontinae (Carleton and

Musser 1989), and studies have been per-

formed on some aspects of the genetic

structure of the populations and relation-

ships of a few species (Chiappero et al.

1997; Dickerman and Yates 1995; Myers et

al. 1995; Patton et al. 1996). Karyologic

studies have helped to clarify systematics ofthis complex group, which presents diploid

numbers (2n) from 44 found in the taxon

named as Oligoryzomys sp. 2 by Silva and

Yonenaga-Yassuda (1997) to 68 in the an-

imals considered as variant 1 of Oligory-

zomys longicaudatus by Gardner and Patton

(1976). We describe for the 1st time the

karyotypes of 3 species of this genus and 4

other species collected at 33 localities com-

prising the major part of the Brazilian range

of the genus.

MATERIALS AND METHODS

The sample included specimens of 7 taxa of

Oligoryzomys caught at 33 localities in an area

ranging from 01N to 32S in Brazil (Fig. 1; Ta-

ble 1). Skins and skulls of the individuals stud-

ied are deposited in the mammal collections of

the Museu Nacional, Rio de Janeiro (MN), and

the Universidade Federal da Paraiba, Joao Pes-


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1082 Vol. 82, No. 4JOURNAL OF MAMMALOGY

TABLE 1.Localities and number (n) of Oli-

goryzomys specimens analyzed.

Species Localitiesa n

O. stramineusO. cf. messorius

O. nigripes

O. eliurus

O. flavescens

O. microtis

Oligoryzomys sp.

41

511, 1329

3, 5

10, 12, 17, 22, 26

2

3

74

207

8

59

6

21

a Numbers correspond to those in Fig. 1.

TABLE 2.Diploid number (2n) and autosomal arm fundamental number (FN), X- and Y-chro-

mosome morphologies, and NOR-band patterns of Oligoryzomys.

Species 2n FN X Y

NOR

(pairs)

O. stramineus

O. cf. messorius

O. nigripes

O. eliurusO. flavescens

O. microtis

Oligoryzomys sp.

52

56

61, 62

626467

66

70

68

58

78, 8082

64, 666668

74

74

SMa

SM

M

A/STd

SM/M

A

A/ST

Mb

SM

Ac

A/SMA

A

A

1

13

2

25

a Submetacentric.b Metacentric.c Acrocentric.d Subtelocentric.

soa (UFPB). Voucher specimen numbers are

given in Appendix I.

Mitotic preparations were made employing the

technique of Baker et al. (1982). The C-, G-, and

NOR-banding were performed following themethods of Sumner (1972), Seabright (1971), and

Howell and Black (1980), respectively. The fluo-

rescent in situ hybridization process with the telo-

meric sequence (T2AG3)n was applied to speci-

mens of Oligoryzomys cf. messorius, O. eliurus,

and Oligoryzomys sp., with the all human telo-

meres probe (Oncor, Gaithersburg, Maryland)

stained with digoxygenin according to the man-

ufacturers protocol.

Eleven sets of microsatellite DNA primers de-

signed for Rattus (R12, R47, R65, R69, R75,

and R97Serikawa et al. 1992) and Mus (M13,

M23, and M49Love et al. 1990; and ATP and

KRTSantos et al. 1995) were amplified byheterologous polymerase chain reaction in O.

eliurus and O. nigripes, as described in Lima-

Rosa et al. (2000).

RESULTS AND DISCUSSION

The 7 species analyzed displayed diploid

numbers from 52 to 70. The diploid number

of 70 that we found in Oligoryzomys sp. is

the highest number reported to in the genus,

verified in 1 of the 3 undescribed karyo-

types we report in this paper (Table 2).

Oligoryzomys stramineus.Seven O.

stramineus (Tables 1 and 2) showed 2n

52, with 9 biarmed and 16 acrocentric au-

tosome pairs. The X chromosome was a

large submetacentric, between pairs 3 and

4 in size, and the Y chromosome was a me-

dium-sized metacentric. The C-bands (data

not shown) were centromeric in the major-

ity of the autosomes (but were absent in the

1st pair) and of the X chromosome. The Ychromosome was entirely heterochromatic.

The karyotype of O. stramineus could be

included in species group 3 (the first 24

pairs of similar size, according the classi-

fication we made of the Oligoryzomys spe-

cies based in the size and shape of its pair

1, see Table 3). That same Giemsa karyo-

type was reported by Bonvicino and Wek-

sler (1998:94, figure 2) in the type series of

O. stramineus, a new species that they de-

scribed. Mamba (Goias State; Fig. 1), from

which our sample came, was located inside

the distribution area proposed for this spe-cies (Cerrado of northern Goias and Minas

Gerais states and Caatinga of the states of

Paraiba and Pernambuco).


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TABLE 3.Karyotyped species of Oligoryzomys grouped according to the size and shape of pair

1 (2n diploid number; FN fundamental number of autosomal arms).

Species 2n FN Locality Referencesa

First pair 1.5 larger than pair 2; acrocentricOligoryzomys sp. 1

O. cf. messorius

O. longicaudatus

O. longicaudatusb

O. delicatusc

O. eliurus

O. aff. eliurus

O. fornesi

O. flavescens

O. flavescens

O. cf. flavescens

O. microtis

O. longicaudatus var. 1

Oligoryzomys sp.

46

56

56

56

60

62

62

6266

6066

6467

66, 68

66

68

70

52

58

64, 66

66

72

64, 66

64

6468

6670

6670

68, 70

74

74

74

Brazil

Brazil

Argentina

Chile

Venezuela

Brazil

Brazil

Paraguay

Uruguayd

Argentina

Brazil

Peru

Brazil

1

2

3

4 6

7

2

8

9

10, 11

2, 3, 1113

3

2

14

2

First pair 1.5 larger than pair 2; biarmed

Oligoryzomys sp. 2

O. magellanicuse

O. microtisf

44, 45

54

64

52, 53

66

66

Brazil

Chile

Peru

1

5, 6

1315

First 24 pairs of similar sizes

O. stramineus

O. fulvescens

O. fulvescens

O. chacoensis

O. cf. longicaudatus

O. andinus

O. destructorg

O. longicaudatus var. 3

O. nigripes/delticola

52

54

60

58

58

60

60

62

61, 62

68

68

74

74

74

70

76

74

7882

Brazil

Costa Rica

Mexico

Paraguay

Argentina

Peru

Peru

Venezuelah

16

14

17

9

3

14

14, 15

14

2, 3, 9, 11, 16, 18

a 1) Silva and Yonenaga-Yassuda (1997); 2) this work; 3) Espinosa and Reig (1991); 4) Gallardo and Gonzalez (1977); 5)Gallardo and Patterson (1985); 6) Gallardo and Palma (1990); 7) Kiblisky (1969); 8) Furtado (1981); 9) Myers and Carleton

(1981); 10) Brum (1965); 11) Brum-Zorrilla et al. (1988); 12) Sbalqueiro et al. (1991); 13) Aniskin and Volobouev (1999); 14)

Gardner and Patton (1976); 15) Musser and Carleton (1993); 16) Bonvicino and Weksler (1998); 17) Haiduk et al. (1979); 18)

Yonenaga et al. (1976).b Described as Oryzomys longicaudatus philippii.c Synonymized as Oligoryzomys fulvescens by Musser and Carleton (1993).d Brazil, Bolivia, and Argentina.e Described as Oryzomys longicaudatus magellanicus.f The karyotype was reported as Oryzomys longicaudatus variant 2.g The karyotype was reported as Oryzomys longicaudatus variant 4.h Brazil, Paraguay, Uruguay, and Argentina.

Oligoryzomys cf. messorius.Amazonian

specimens of O. cf. messorius (Fig. 1; Table

1) had 56 chromosomes, a karyotype hith-erto not described in the genus. The 1st pair

was a large acrocentric, 2 pairs were small

metacentrics, and the other 24 pairs were

medium to small acrocentrics, resulting in

a fundamental number (FN) 58 (FN is

used herein to designate only the number of

autosomal arms). The X chromosome was

a large submetacentric (between pairs 1 and

2 in size), and the Y chromosome was alarge metacentric (Fig. 2a). The C-bands

(heterochromatic material) were centromer-

ic in most of the chromosomes, including

the 1st pair. The short arm of the X chro-

mosome and the entire Y chromosome were

heterochromatic (Fig. 3a). One pair dis-


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FIG. 2.Giemsa-stained karyotypes of a) Oligoryzomys cf. messorius, diploid number (2n) 56,

fundamental number (FN)

58, male; b) O. eliurus, 2n

62, FN

66, female (in the square,acrocentric X and Y chromosomes); and c) O. microtis, 2n 66, FN 74.

played NOR-bands in the short arms (Table

2).

Oligoryzomys messorius was considered

as a synonym of O. fulvescens by Musser

and Carleton (1993). The species we stud-

ied in the Amazon presents 2n 56, FN

58 and belonged to species group type 1 of

the Oligoryzomys karyotypes, in which the


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FIG. 3.The C-bands of a) Oligoryzomys cf. messorius, male; b) O. eliurus, male (in the square,

pair of sex chromosomes of a female); c) O. microtis (the X and Y chromosomes are below the

letters); and d) Oligoryzomys sp.


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1st pair is 1.5 larger than pair 2 (Table 3).

However, O. fulvescens is chromosomally

quite distinct, presenting 2n 54, FN 68

in Costa Rica (Gardner and Patton 1976)

and 2n 60, FN 74 in Mexico (Haiduket al. 1979), and these karyotypes are in-

cluded in species group 3 (the first 24

pairs of similar size; see Table 3). The taxon

messorius was reported by Thomas (1901)

in the Kanuku Mountains, Guyana, about

150 km from the locality where we col-

lected our sample (Surumu, Roraima State;

Fig. 1). This geographic vicinity, allied to

the chromosome distinctiveness from ful-

vescens, suggests that the taxon of Surumu

may be cf. messorius.

Oligoryzomys nigripes.We found 2n

62 in specimens of O. nigripes from several

places in the states of Bahia, Goias, Esprito

Santo, Parana, Santa Catarina, and Rio

Grande do Sul (Fig. 1). That karyotype pre-

sented FN 78, 80, 81, and 82. That var-

iation was due to pericentric inversions oc-

curring in pairs 3 and 4 (acrocentrics or

submetacentrics, with all combinations of

heterozygous and homozygous individuals;

Tables 1 and 2). Pair 2 was always biarmed,

but an inversion made it either subtelocen-

tric or submetacentric. The X chromosome

was a large metacentric, submetacentric, orsubtelocentric, and the Y chromosome, also

polymorphic, was a small acrocentric, a

medium metacentric, or a large submeta-

centric. The 2n 61 karyotype correspond-

ed to an XO female (Table 2). The C-bands

(data not shown) were pericentromeric and

seen only in some medium and small pairs.

The C-band was always absent in the 1st

pair. The short arm of the X chromosome

and the long arm of the Y chromosome

were heterochromatic. That same karyo-

type, with 2n 62 and the pericentric in-

version systems, was reported in specimensfrom Paraguay by Myers and Carleton

(1981:16, figure 5) and from the states of

Sao Paulo and Rio de Janeiro, Brazil (Al-

meida and Yonenaga-Yassuda 1991); this

karyotype also was described in individuals

referred to as O. delticola from Uruguay

(Brum-Zorrilla et al. 1988) and Argentina

(Espinosa and Reig 1991). O. nigripes Ol-

fers, 1818 (an older name than delticola)

inhabits eastern Paraguay and northern Ar-

gentina, and O. delticola Thomas, 1917presents a complementary distribution to it,

occurring in east-central Argentina, Uru-

guay, and southern Brazil (Musser and

Carleton 1993). These 2 taxa apparently

constitute a unique cytotaxonomic entity

(2n 62, FN 7882), which we found

occupying a broad area of South America,

extending from northern Argentina to Par-

aguay and from Uruguay and southern Bra-

zil to the east on the coast to 15S (Southern

Bahia) and to the west to the Cerrado biome

in central Brazil.

Oligoryzomys eliurus.The individuals

of O. eliurus, another species with 2n 62,

were collected at 4 localities of the Cerrado

biome (Fig. 1; Table 1). Two autosomal

conditions, FN 64 and 66, were ob-

served, both with a large acrocentric pair, a

medium submetacentric pair, a large poly-

morphic X chromosome (acrocentric or

subtelocentric), and a small acrocentric Y

chromosome. The FN 64 karyotype pre-

sented 27 acrocentric pairs of medium to

small sizes, and a medium metacentric pair.

The karyotype with FN

66 has 26 acro-centric pairs of medium to small size and 2

pairs of small metacentrics; the Y chromo-

some was a submetacentric (Fig. 2b; Table

2). The C-bands occurred in the centromer-

ic regions of the majority of the autosomes

but were absent in the 1st pair. In the X

chromosome, the C-band (centromeric) var-

ied in size, and the Y chromosome proved

to be entirely heterochromatic (Fig. 3b).

The NORs were located in the short arms

of 13 pairs (Table 2).

From a chromosomal standpoint, 3 spe-

cies of Oligoryzomys presented 2n 62and a low FN to 2n ratio: in Brazil in the

Cerrado (FN 64, 66this work) and

Caatinga (FN 64, referred to as O. aff.

eliurus by Furtado 1981) biomes and in

Paraguay (FN 64, named O. fornesi by

Myers and Carleton 1981). Musser and


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Carleton (1993) later suggested that the kar-

yotype of O. fornesi from Paraguay would

correspond to that of O. microtis. These 3

taxa share a large acrocentric pair 1, 2627

medium to small acrocentric pairs, and 2 or3 pairs of medium to small biarmed ele-

ments (Table 3). They also have the same

polymorphic X and Y chromomosomes. We

collected our sample in the Brazilian Cer-

rado, a region within the proposed distri-

bution of O. eliurus (central and southeast-

ern BrazilMusser and Carleton 1993).

At 1 of the localities (Ipameri) of the

Cerrado where O. eliurus was trapped, it

was sympatric with O. nigripes. Myers and

Carleton (1981) proposed that these 2 taxa

were subspecifically distinct. Nevertheless,

the karyotype ofO. eliurus is quite different

from that of O. nigripes. Although it bears

the same diploid number (62), O. eliurus

shows autosomal arm numbers of 64 and

66, smaller than the 7882 presented by O.

nigripes. Furthermore, in previous studies,

we verified that heterologous amplifications

of some microsatellite DNA loci of Mus

and Rattus showed bands that were species-

specific in several oryzomyine taxa (Lima-

Rosa et al. 2000). We performed this anal-

ysis with 11 microsatellite DNA heterolo-

gous primers in 10 O. nigripes and 7 O.eliurus specimens. We found that all of the

O. nigripes individuals presented a 192-

base pair band (amplified by primer R65 of

rat, designed by Serikawa et al. 1992). The

occurrence of this band in only this species,

not having been seen in the other species

of the Oligoryzomys genus including O. eli-

urus, reinforces the distinction between

these 2 species.

Oligoryzomys flavescens.In O. flaves-

cens, 2n varied from 64 to 67 and FN from

66 to 72 because of the occurrence of a sys-

tem of 03 accessory chromosomes (B-chromosomes; Table 2). The Giemsa-

stained and C-banded karyotype, the same

as that described by Sbalqueiro et al. (1991:

196, figures 2A and 2C), basically was

composed of a large acrocentric pair, 28

pairs of medium to small acrocentrics, and

2 or 3 small biarmed pairs. The B chro-

mosomes were small submetacentrics to

metacentrics. The X chromosome was a

medium submetacentric (between pairs 1

and 2 in size), and the Y chromosome wasa polymorphic medium subtelocentric to

submetacentric or a small metacentric. That

same basic karyotype also was reported in

Argentina, Uruguay, Bolivia, and in several

localities in southern Brazil (Aniskin and

Volobouev 1999; Brum-Zorrilla et al. 1988;

Sbalqueiro et al. 1991; Vidal-Rioja et al.

1988). That karyotype also was the same as

that of O. fornesi with 2n 6466, which

was collected in Paraguay by Myers and

Carleton (1981). O. fornesi was originally

described by Massoia (1973), who noted its

close relationship to O. flavescens. Later,

based on the chromosomal identity, Sbal-

queiro et al. (1991) considered flavescens

and fornesi to be in the same cytotaxonomic

group. We found O. flavescens with O. ni-

gripes at 4 sites in southern Brazil (locali-

ties 10, 17, 22, and 26; Fig. 1).

Oligoryzomys microtis.The species O.

microtis, trapped in the Amazon biome (lo-

cality 2, Fig. 1), showed 2n 66 and FN

74. This species presented a large acro-

centric pair, 26 medium to small acrocen-

trics, and 5 biarmed pairs. The X chromo-some was a medium acrocentric to subtel-

ocentric (between pairs 2 and 3 in size), and

the Y chromosome was a small acrocentric

(Fig. 2c). The C-bands occurred in the cen-

tromeric regions of the majority of the au-

tosomes but were absent in the 1st pair. In

the X chromosome, the C-band was centro-

meric, and the entire Y chromosome was

heterochromatic (Fig. 3c). The NORs were

located in the short arms of 2 pairs. O. mi-

crotis, whose type locality is the lower So-

limoes River in the Brazilian Amazon, is

distributed throughout central Brazil, southof the SolimoesAmazon rivers, and in the

contiguous lowlands of Peru, Bolivia, Par-

aguay, and Argentina (Musser and Carleton

1993). O. microtis includes 2 chromosome

constitutions, 2n 66, FN 74 in Brazil

(this report) and 2n 64, FN 66 in Peru


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FIG. 4.Oligoryzomys sp., diploid number (2n) 70, fundamental number (FN) 74: a) con-

ventional stained karyotype; b) G-banding; and c) NOR-bearing chromosomes.


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(Aniskin and Volobouev 1999). This last

karyotype also was described in Peru by

Gardner and Patton (1976) as O. longicau-

datus (variant 2). The main difference be-

tween these 2 cytotypes is a biarmed pair1, a feature seen in individuals from Peru

that is absent in the Brazilian specimens.

Oligoryzomys sp.Twenty-one speci-

mens of Oligoryzomys caught exclusively

in the Cerrado biome displayed 2n 70

and FN 74. The karyotype was com-

posed of a pair of large acrocentrics, 30

pairs of medium to small acrocentrics, 2

medium pairs of metacentrics, and a small

pair of metacentrics (Tables 1 and 2). The

X chromosome was a large subtelocentric

(between pairs 1 and 2 in size), and the Ychromosome was a medium-sized subtelo-

centric (Giemsa staining, Fig. 4a; G-bands,

Fig. 4b). The C-bands were at the centro-

meres of the majority of the autosomes, in-

cluding the 1st pair, the 3 biarmed pairs,

and in the X chromosome. The Y chro-

mosome was entirely heterochromatic (Fig.

3d). This karyotype, which belonged to

species group type 1 (1st pair is 1.5 larger

than pair 2) has not been described previ-

ously for the genus Oligoryzomys and may

represent a new form or 1 of the several

forms included as synonyms in the current-

ly recognized species of central and western

Brazil.

The (T2AG3)n telomeric probes hybrid-

ized in situ in both the short and the long

arms of all pairs of chromosomes of the

species O. cf. messorius, O. eliurus, and

Oligoryzomys sp.; the hybridization signal

occurred with the same intensity in both

arms (data not shown). That band locali-

zation also was observed by Silva and

Yonenaga-Yassuda (1997) in the 2 unclas-

sified species of Oligoryzomys that theystudied with this technique.

The 26 taxa of genus Oligoryzomys kar-

yotyped to date are listed in Table 3. The

diploid numbers of the genus vary from 44

to 70, and 17 of the 26 taxa karyotyped

displayed a characteristic 1st pair, about

50% larger than pair 2, with the majority

(12 of 17) being acrocentric.

Several authors found Oligoryzomys to

be a monophyletic genus (Carleton and

Musser 1989; Dickerman and Yates 1995;Myers et al. 1995; Steppan 1995). But the

hierarchical relationships among its species

are controversial because the groups ar-

ranged by morphological characteristics

(Carleton and Musser 1989; Hershkovitz

1966; Myers and Carleton 1981; Olds and

Anderson 1987; Steppan 1995), polymor-

phism of proteins (Dickerman and Yates

1995), and polymorphism of DNA (Lima-

Rosa et al. 2000; Myers et al. 1995) differ.

The karyological data obtained in this work

and in the literature (Table 3) also assemble

the species of the genus in distinct groups.

ACKNOWLEDGMENTS

Conselho Nacional de Desenvolvimento Cien-

tfico e Tecnologico (CNPq), Financiadora de

Estudos e Projetos (FINEP), Fundacao de Am-

paro a Pesquisa do Estado do Rio Grande do Sul

(FAPERGS), and Organization of the American

States (OAS) supported this study. We are grate-

ful to A. R. Langguth, D. A. Sana, J. L. P. Cor-

deiro, and J. R. Marinho for help in the field and

to L. S. Silva, B. A. Carvalho, and F. L. Hadrich

for technical support.

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Submitted 15 May 2000. Accepted 5 February 2001.

Associate Editor was Meredith J. Hamilton.

APPENDIX I

Voucher specimens (Brazil).Oligoryzomys

stramineus: UFPB18241827, 1836, 1837(Mamba, GO, 1429S, 4606W); Oligoryzo-

mys cf. messorius: MN3775037753 (Surumu,

RR, between 03580427N and 6013

6116W); Oligoryzomys nigripes: MN37488

37500 (Ipameri, Caldas Novas, and Corumbaba,

GO, between 17411756S and 4828

4832W), MN37501 (Parque Estadual do Tur-

vo, Derrubadas, RS, between 272720S and

53405410W), MN3750237504 (Aratiba,

RS, 2723S, 5218W), MN3750537511 (Ar-

acuri Ecological Station, Muitos Capoes, RS,

2813S, 5110W), MN37512 (Iva, Tupancire-

ta, RS, 2857S, 5340W), MN37513 (Caxias

do Sul, RS, 2910S, 5110W), MN3751437528 (Tainhas, RS, 2916S, 5018W),

MN3752937531 (Sao Francisco de Paula, RS,

2927S, 5035W), MN3753237538 (Char-

queadas, RS, 2957S, 5137W), MN37539

37574 (Picada Verao and Alto Ferrabraz, Sapi-

ranga, RS, 2938S, 5100W), MN3757537578

(Riozinho, RS, 2938S, 5027W), MN37579

37604 (Faxinal, Torres, RS, 2920

S, 49

45

W),

MN37605 (Rio Jacu, Eldorado do Sul, RS,

3005S, 5136W), MN37606 (Belem Novo,

RS, 3008S, 5110W), MN3760737622 (Oso-

rio, RS, 2953S, 5016W), MN37623 (Traman-

da Lagoon, Tramanda, RS, 2959S, 5008W),

MN37624, MN37625 (Quintao, RS, 3020S,

5016W), MN3762637630 (northern Tapes

Bay, Tapes, RS, 3040S, 5123W), MN37631

37659 (Capao do Leao, Mostardas, RS, 3110S,

5056W), MN3766037682 (Taim Ecological

Station, Rio Grande, RS, 3232S, 5232W),

MN37683 (Costa de Dentro, Santa Catarina Is-

land, SC, 2746S, 4831W), MN3768437690

(Concordia, SC, 2714

S, 52

01

W), MN37691

37693 (Parque Nacional de Iguacu, PR, 2530S,

5430W), UFPB354, UFPB355, UFPB357361

( Mont e Ver de, ES, 1903S, 4157W),

UFPB428, UFPB429 (Rio de Una, 10 km ESE

Sao Jose, BA, 1506S, 3916W); Oligoryzomys

eliurus: MN35992 (55 km N Niquelandia, GO,

1428S, 4827W), MN36746 (40 km SW Min-

acu, GO, 1331S, 4813W), MN36928,

MN37258, MN37347 (40 km NE Uruacu, GO,

1431S, 4908W), MN3769437696 (Ipameri,

Caldas Nova, and Corumbaba, GO, between

17411756S and 48284832W); Oligory-

zomys flavescens: MN37697 (Nonoai, RS,

2725S, 5304W), MN3769837720 (Tainhas,RS, 2916S, 5018W), MN3772137747

(Ch arqu ea da s, R S, 2 957S, 5137W),

MN37748 (Quintao, RS, 3020S, 5016W),

MN37749 (Costa de Dentro, Santa Catarina Is-

land, SC, 2746S, 4831W); Oligoryzomys mi-

crotis: MN3775437759 (Fazenda Sao Bento,

Tartarugalzinho, AP, 0117N, 5048W); Oli-

goryzomys sp.: MN3 59 67, MN3 60 61 ,

MN36082, MN36083, MN36145, MN36147,

MN36161 (55 km N Niquelandia, GO, 1428S,

4827W), MN36220, MN36224, MN36234,

MN36242, MN36345, MN36356, MN36357,

MN36367, MN36426, MN36433, MN36438 (20

km NW Colinas do Sul, GO, 1409S, 4804W),MN36832 (40 km SW Minacu, GO, 1331S,

4813W), MN37282, MN37441 (40 km NE

Uruacu, GO, 1431S, 4908W).

andrades-miranda - chromosomes studies of seven species of oligoryzomys from brazil

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