review of the harvest mice (genus reithrodontomys) in the mexican state of mÉxico

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REVIEW OF THE HARVEST MICE (GENUSREITHRODONTOMYS) IN THE MEXICAN STATE OFMEacuteXICOAuthor(s) Noeacute Gonzaacutelez-Ruiz Joseacute Ramiacuterez-Pulido Hugh H GenowaysSource Western North American Naturalist 67(2)238-250 2007Published By Monte L Bean Life Science Museum Brigham Young UniversityDOI httpdxdoiorg1033981527-0904(2007)67[238ROTHMG]20CO2URL httpwwwbiooneorgdoifull1033981527-090428200729675B2383AROTHMG5D20CO3B2

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Relatively few papers dealing with the genusReithrodontomys from the Mexican state ofMeacutexico have been published The oldest paperknown to us is Merriam (1901) in whichappeared the description of Reithrodontomyschrysopsis based on specimens from VolcanPopocateacutepetl In the same paper Merriam(1901) also described R tolucae from theNevado de Toluca but Howell (1914) in hisrevision of the genus Reithrodontomys recog-nized R tolucae as a subspecies of R chrysop-sis Merriam (1901) also recorded the presenceof R megalotis and R fulvescens in the state ofMeacutexico Hooper (1952) in his review of LatinAmerican harvest mice reported records ofseveral species in the state R tolucae wasretained as a junior synonym of R chrysopsisand R sumichrasti was recorded for the 1sttime from the state of Meacutexico Other papers

contributing to our understanding of the genusin the state are those by Davis (1944) Villa-Ramiacuterez (1953) and Ramiacuterez-Pulido (1969)

Because available information is scarce ourpurpose is to document the data for species ofReithrodontomys in the state of Meacutexico Thesedata have accumulated over the past 35 yearsin the mammal collection of the EscuelaNacional de Ciencias Bioloacutegicas del InstitutoPoliteacutecnico Nacional We present informationfor R chrysopsis R sumichrasti R megalotisR fulvescens (represented by 2 subspeciesmdashR f toltecus and R f mustelinus) and Rei-throdontomys microdon wagneri which is re-corded herein for the 1st time from the stateof Meacutexico With a sample of 590 specimens ofthe 5 species we evaluate compare and con-trast the taxonomy morphometrics repro-duction and habitat characteristics of thesespecies

Western North American Naturalist 67(2) 2007 pp 238ndash250

REVIEW OF THE HARVEST MICE (GENUS REITHRODONTOMYS) IN THE MEXICAN STATE OF MEacuteXICO

Noeacute Gonzaacutelez-Ruiz12 Joseacute Ramiacuterez-Pulido2 and Hugh H Genoways3

ABSTRACTmdashWe examined 590 specimens of Reithrodontomys from 95 localities in the state of Meacutexico Four speciesof the subgenus Reithrodontomys and 1 of the subgenus Aporodon were identified The former subgenus included Rchrysopsis R sumichrasti R megalotis and R fulvescens which has 2 subspeciesmdashR f toltecus and R f mustelinusmdashin the state The representative of the subgenus Aporodon is R microdon wagneri which is recorded for the first time inthe state We give information on taxonomy morphometrics reproduction habitat characteristics and related fauna Adiscriminant analysis correctly classified 100 of specimens from the 6 taxa with 5 canonical variates and accounted for961 of the variance with the first 3 canonical variates

Key words rodents Reithrodontomys distribution taxonomy state of Meacutexico Meacutexico

RESUMENmdashSe examinaron 590 ejemplares de las especies del geacutenero Reithrodontomys procedentes de 95 locali-dades del Estado de Meacutexico Se identificaron cuatro especies del subgeacutenero Reithrodontomys y una del Aporodon delprimero R chrysopsis R sumichrasti R megalotis y R fulvescens de eacutesta uacuteltima se identificaron dos subespecies R ftoltecus y R f mustelinus La especie del subgeacutenero Aporodon es R microdon wagneri que se registra por primera vezpara el Estado De cada taxon se dan medidas comentarios taxonoacutemicos y de la condicioacuten reproductora asiacute como lascaracteriacutesticas del haacutebitat y de la fauna asociada Un anaacutelisis discriminante mostroacute el 100 de certeza en la identifi-cacioacuten de los 6 taxa y los primeros tres vectores canoacutenicos expresaron el 961 de variacioacuten total

Palabras clave roedores Reithrodontomys distribucioacuten taxonomiacutea Estado de Meacutexico Meacutexico

1Universidad Autoacutenoma Metropolitana Unidad Iztapalapa Divisioacuten de CBS Departamento de Biologiacutea Apartado Postal 55-535 Meacutexico 09340 DFMeacutexico

2Laboratorio de Cordados Terrestres Escuela Nacional de Ciencias Bioloacutegicas Instituto Politeacutecnico Nacional Prolongacioacuten de Carpio y Plan de AyalaColonia Santo Tomaacutes Apartado Postal CON 256 11340 DF Meacutexico E-mail jrpxanumuammx

3University of Nebraska State Museum and School of Natural Resources Sciences W436 Nebraska Hall University of NebraskandashLincoln Lincoln NE68588-0514

238

METHODS

We examined and analyzed 590 specimensfrom 95 localities in the state of Meacutexico(Appendix) For each species the specimensexamined are listed by locality In the addi-tional records sections only verified recordswere entered Place of origin reproductivedata and body measurements were obtainedfrom the tag of each specimen Ecological datacapture conditions and information aboutassociated fauna and vegetation were obtainedfrom field notes preserved in the collection

Using a digital caliper precise to 001 mmwe measured skulls according to methods ofHooper (1952) Measurements were taken onlyfrom adult specimens which were identifiedby the degree of wear on the occlusal surfaceof the molars Adults also generally showedworn labial and lingual cuspids and absentreentrant angles of M2 and M3 Measurementswere the following total length (TL) length oftail vertebrae (TV) length of hind foot (HF)length of ear (LE) greatest length of skull (GLS)zygomatic breadth (ZB) cranial breadth (CB)interorbital constriction (IC) breadth of ros-trum (BR) length of nasals (LN) length of max-illary toothrow (LMTR) breadth of zygomaticplate (BZP) and breadth of mesopterygoidfossa (BMF)

Geographic coordinates of the localitiessampled (Appendix) were gathered by 2 meth-ods (1) GPS in situ and (2) a map scaled at1250000 and published by INEGI (InstitutoNacional de Estadiacutestica y Geografiacutea) Thelocalities where specimens were captured and the major types of vegetation are shown in Figure 1

For comparisons between 2 samples weused separate-variance t tests with the correc-tion of Welch to obtain a greater sensitivity todifferences between means than the Tukeytest provides For comparison of ge3 samples weused analysis of variance (ANOVA) coupledwith a Tukey test for multiple comparisons todefine the significance level of differencesamong means minimums and maximums ofthe 6 groups considered The 3 significancelevels that we considered were P-values lt005lt001 and lt0001

Basic statistics (ie mean minimum maxi-mum standard deviation [s] and number ofspecimens measured [n]) were obtained fromGraphPad Prisma (Motulsky 2003) software

In order to compare measurements of speci-mens of all taxa examined we ran a multivari-ate analysis of variance (MANOVA) and todemonstrate the separation among species weused a discriminant analysis with NCSS (Hintze2000) However in both cases we did not in-clude the weight Significance levels in theMANOVA were tested using Wilksrsquo lambdaHotelling-Lawley trace Pillairsquos trace and Royrsquoslargest root

SPECIES ACCOUNTS

Reithrodontomys (Aporodon)microdon wagneri

Hooper 1950

SPECIMENS EXAMINED (3)mdash27 km N 9 kmW Villa del Carboacuten 2670 m (1) Llanos deAculco 3200 m (1) 15 km S 5 km E SanRafael (1)

This is 1 of the rarest species of the genusIts distribution is restricted to the mountain-ous zones of Guatemala the states of Chiapasand Oaxaca and central Meacutexico The sub-species R m wagneri was known previouslyfrom only 4 specimens 2 from a locality inMeacutexico City and 2 from a locality in the stateof Michoacaacuten (Hooper 1952) As a result ourspecimens significantly increased the numberof referred specimens and the number ofknown localities for the taxon These are the1st record for the state of Meacutexico

MORPHOLOGY AND DISTRIBUTIONmdashReithro-dontomys microdon is the only species of thesubgenus Aporodon in central Meacutexico Mem-bers of this subgenus are characterized byhaving the 2nd primary lingual fold of M3very well developed so that it appears as aninternal enamel island Other characters dis-tinguishing this species are as follows zygo-matic plate narrow but broader than the widthof the mesopterygoid fossa zygomatic breadthonly slightly wider than the cranial breathand the rostral region tending to be relativelylong and narrow (Table 1)

Specimens we examined were larger thanthose reported by Hooper (1952) particularlythe body measurements (Table 1) of the 2Michoacaacuten specimens The results of ANOVAshowed that R m wagneri is significantlysmaller than R f toltecus R f mustelinus andR s sumichrasti in BPZ (P lt 005 P lt 0001and P lt 0001 respectively) significantly largerthan R f mustelinus in ZB CB (P lt 0001)

2007] HARVEST MICE IN THE STATE OF MEXICO 239

and IC (P lt 005) and significantly larger thanR s sumichrasti in TV (P lt 0001) and IC (P lt 005)

ECOLOGYmdashThe specimen from Villa delCarbon was a female collected on 11 January1986 There was no evidence of gross repro-ductive activity and the capture location wasin a forest of oak mixed with fir and pine theground was damp with a large amount of mossThe sex of the specimen from Llanos de Aculcois unknown because it was recovered in a re-

gurgitant probably from an owl The specimenfrom San Rafael was a female that showed noactive signs of reproduction This individualwas trapped in a pine-oak forest (Fig 1) withan abundant herbaceous stratum consisting ofclusters of plant shoots Other species taken atthese same sites included Microtus mexicanusNeotomodon alstoni Peromyscus maniculatusP melanotis and P difficilis Apparently thisspecies occupies a specialized habitat nichebecause the few known specimens have been

240 WESTERN NORTH AMERICAN NATURALIST [Volume 67

Fig 1 Index and regional map of the Mexican state of Meacutexico Numbers and dots correspond to the gazetteer in theAppendix

Agricultural Fields

Human Settlement

Coniferous Forest

Shrubs

Grassland

Tropical Deciduous Forest

Halophytic Vegetation

Specimens Examined

2007] HARVEST MICE IN THE STATE OF MEXICO 241T

AB

LE

1 S

elec

ted

exte

rnal

and

cra

nial

mea

sure

men

ts o

f the

gen

us R

eith

rodo

ntom

ysfr

om th

e M

exic

an s

tate

of M

eacutexic

o A

cron

yms

are

expl

aine

d in

the

met

hods

The

bas

ic s

tatis

tics

incl

ude

stan

dard

dev

iatio

n (s

) and

num

ber

of s

peci

men

s ex

amin

ed (n

)

TL

TV

HF

LE

GL

SZ

BC

BIC

BR

LN

LM

TR

BZ

PB

MF

RE

ITH

RO

DO

NT

OM

YSC

HR

YSO

PSIS

CH

RYS

OPS

IS

xndash17

93

100

220

64

180

924

34

131

511

43

337

39

883

365

237

139

Min

imum

157

8119

1523

68

118

111

22

323

361

846

335

204

113

Max

imum

179

311

222

2124

78

131

511

73

353

409

949

392

319

163

s13

53

927

081

187

035

035

018

011

014

027

016

031

012

n10

1011

1111

1111

1111

1111

1111

RE

ITH

RO

DO

NT

OM

YSF

ULV

ESC

EN

SM

UST

ELI

NU

S

xndash17

233

103

1815

521

41

104

210

23

283

697

333

231

871

28M

inim

um15

795

1615

201

910

42

987

321

352

707

312

18

126

Max

imum

189

111

2016

222

810

99

106

83

333

847

853

331

921

31s

160

411

31

20

711

090

560

430

060

160

450

110

060

03n

32

32

33

33

33

33

3R

EIT

HR

OD

ON

TO

MYS

FU

LVE

SCE

NS

TO

LTE

CU

S

xndash18

146

101

520

31

156

223

02

116

210

84

347

426

83

35

218

131

Min

imum

171

8619

1322

44

113

810

31

322

403

796

339

193

117

Max

imum

199

112

2217

241

511

86

113

93

684

268

893

722

461

43s

885

703

075

126

055

016

024

014

022

03

008

016

009

n13

1213

1313

1313

1312

1313

1313

RE

ITH

RO

DO

NT

OM

YSM

EG

ALO

TIS

SAT

UR

ATU

S

xndash15

297

793

618

15

140

922

04

110

910

21

318

376

79

362

235

101

Min

imum

131

6716

1020

81

104

89

622

933

377

313

352

080

69M

axim

um17

089

2018

233

111

810

74

349

428

873

362

235

126

s9

955

021

194

049

029

021

013

376

029

012

015

013

n33

3333

3233

3233

3333

3332

3333

RE

ITH

RO

DO

NT

OM

YSSU

MIC

HR

AST

ISU

MIC

HR

AST

I

xndash16

991

86

188

615

43

227

911

48

107

63

363

848

273

362

211

28M

inim

um16

066

1810

220

210

910

35

298

362

798

308

21

16M

axim

um18

110

221

1923

73

119

811

03

383

402

827

359

248

128

s6

5910

17

095

21

046

032

024

022

012

027

015

015

011

n14

1414

1415

1514

1515

1515

1515

RE

ITH

RO

DO

NT

OM

YSM

ICR

OD

ON

WA

GN

ER

I

xndash18

811

55

1818

221

511

63

111

53

673

847

963

051

451

61M

inim

um18

511

218

1821

77

115

410

89

351

384

765

305

137

158

Max

imum

191

119

1818

225

117

211

48

379

384

819

306

152

163

s4

244

950

00

370

130

30

140

280

011

450

03n

22

22

32

33

13

23

3

found in oak forests with abundant herbaceousvegetation and are generally associated withvery humid zones (Hooper 1952) These con-ditions are similar to those at the localities inthe state of Meacutexico The opinion of Hooper(1952) is that this species is probably arboreal

Reithrodontomys (Reithrodontomys)chrysopsis chrysopsis

Merriam 1900

SPECIMENS EXAMINED (33)mdash4 km S 9 kmE Amecameca 3480 m (5) 11 km N 25 km WCoatepec de Harinas 2820 m (2) Nevado deToluca (1) 11 km S 25 km W San Juan de lasHuertas 3780 m (2) 115 km S 25 km W SanJuan de las Huertas 3780 m (2) 12 km S SanJuan de las Huertas 3850 m (2) 52 km E SanPedro Nexapa 3250 m (1) 4 km S 85 km ESan Pedro Nexapa 3500 m (2) 55 km E SanPedro Nexapa 3250 m (2) 7 km SE San PedroNexapa 3010 m (2) 22 km N 65 km E SanPedro Nexapa 3490 m (3) 16 km N 64 km ESan Pedro Nexapa 3200 m (5) 8 km N 175km E San Simoacuten de Guerrero 2940 m (2)155 km S 7 km W Zinacantepec 3470 m (1)185 km S 9 km W Zinacantepec 3440 m (1)

ADDITIONAL RECORDSmdashVolcaacuten Popocateacute-petl 11500 ft (type locality of R c chrysopsisMerriam 1900 Hooper 1952) Volcaacuten de Toluca11500 ft (type locality of R c tolucae Mer-riam 1901 Hooper 1952 Howell 1914) MonteRiacuteo Friacuteo 55 km ESE Meacutexico City 10500 ftLadera N del Mt Popocateacutepetl 13500 ft (Davis1944) 8 mi ESE Amecameca 11500 ft 28 miESE Meacutexico City Monte Riacuteo Friacuteo 10500 ft19 mi E Amecameca Paso Cortes 11300 ftSalazar 9000 ft Volcaacuten Ixtacciacutehuatl 13500 ftslopes W and SW del Volcaacuten de Toluca [=Nevado de Toluca] 11400ndash12200 ft (Hooper1952) Paso de Cortes (Villa-Ramiacuterez 1953)Zoquiapan (Mass et al 1981) 6 km E Ame-cameca Vivero Izta-Popo (Leoacuten and Romo1991)

MORPHOLOGY DISTRIBUTION AND TAXO-NOMIC COMMENTSmdashReithrodontomys chrys-opsis is an endemic Mexican species distrib-uted throughout the Transvolcanic Belt fromVeracruz to Colima In the state of Meacutexico itoccurs on the volcanoes of Nevado de TolucaPopocateacutepetl and Ixtacciacutehualt Hooper (1952)concluded that the taxon described by Mer-riam (1901) as R tolucae is a junior synonymof R c chrysopsis and we agree with thisassessment Examination of a large sample of

specimens from the area surrounding VolcanNevado de Toluca and comparison of these spe-cimens with a series from the Sierra Nevadarevealed no morphological differences to jus-tify a different conclusion

Reithrodontomys chrysopsis chrysopsis wassignificantly larger than R megalotis saturatusin TL TV HF LE and IC (P lt 0001) R ssumichrasti in HF LE LN BZP (P lt 0001)TL and TV (P lt 005) R fulvescens musteli-nus in HF LN and BZP (P lt 0001) R ftoltecus in LN (P lt 0001) LE (P lt 001) andBZP (P lt 005) and R microdon wagneri inLN (P lt 0001) and HF (P lt 001 Table 1)

ECOLOGYmdashTwo lactating females weretaken in May and July and a young individualwas obtained in November Average testislengths of males by month of collection wereas follows March 75 mm (n = 4 range 70ndash80 mm) April 80 mm (n = 1) May 50 mmand 40 mm (n = 2) July 67 mm (n = 3range 30ndash100 mm) October 46 mm (n = 8range 30ndash70 mm) November 40 mm (n = 1)December 40 mm (n = 1) Although thesedata are sketchy it appears that individuals ofR chrysopsis are capable of reproduction fromat least March through July and possibly aslate as September

This species was collected primarily in pineforests although in San Simon de Guerrero itwas collected in an Abies forest with tall grass(Fig 1) Generally this species is captured inrugged terrain where the climate is humidIndividuals of this species were obtainedtogether with Reithrodontomys megalotis Rsumichrasti Peromyscus melanotis Neotomodonalstoni and Microtus mexicanus

Reithrodontomys (Reithrodontomys)fulvescens JA Allen 1894

Two subspecies of Reithrodontomys fulves-cens have been recorded for the state of Meacutex-ico (Ramiacuterez-Pulido et al 1995 1997 Chaacutevezand Ceballos 1998) The primary external dif-ference between R f toltecus and R f muste-linus was in dorsal colorationmdashthe former isgrayish orange whereas the latter is brightlight orange

Reithrodontomys fulvescens toltecus Merriam 1901

SPECIMENS EXAMINED (126)mdashEl Angosta-dero 23 km NW Acambay 2700 m (4) 85 km


N 15 km E Acambay 2740 m (1) 125 km N45 km W Acambay 2450 m (9) 35 km N 19km Aculco 2650 m (1) 5 km W Atenco (2) 11km N Barrientos 2300 m (3) 45 km E Chico-nautla 2350 m (2) 3 km E 2 km S Chilpa2270 m (18) 2 km S Coacalco 2440 m (7) 5km E Jilotepec 2460 m (14) 85 km N 25 kmE Jilotepec 2320 m (15) 1 km S 2 km ENicolaacutes Romero 2350 m (2) 25 km W SanFrancisco Tepojaco 2350 m (20) 25 km N SanJuan Zitlaltepec 2250 m (1) Santa Mariacutea delas Cuevas 2480 m (15) 2 km NW SantiagoTolman 2600 m (3) 65 km S 6 km W Temas-calcingo 2560 m (3) 2 km NW Tlanepantla2300 m (4) 95 km N Zumpango 2270 m (2)

ADDITIONAL RECORDSmdash17ndash22 km E Meacutex-ico City 7500 ft (Davis 1944) Hacienda Coacuter-doba 8300 ft 10ndash13 mi SE Meacutexico City 7500ft (Hooper 1952) Rancho Coacuterdoba 2650 m(Hooper 1947) 4 km ENE Tlalmanalco 2290m (Hall 1981)

MORPHOLOGYmdashReithrodontomys f tolte-cus differs significantly from R f mustelinus inGLS ZB CB LN (ANOVA P lt 0001) andHF (P lt 001) whereas t tests showed differ-ences in IC (P lt 005) RB (P lt 001) andBZP (P lt 0001) Reithrodontomys f toltecusaveraged larger than R f mustelinus in allmeasurements

ECOLOGYmdashA female taken in February waspregnant with 4 embryos (crown-rump length3 mm) and a female taken in March was lac-tating The remainder of the females collectedin January February March April Novemberand December showed no gross signs of re-productive activity although 2 young specimenswere taken in February and 1 in NovemberMales taken in September and November hadthe largest testes which may indicate thatbreeding took place during these monthsAverage testis lengths of males by month ofcollection were as follows January 32 mm (n= 22 range 20ndash60 mm) February 42 mm (n= 20 range 20ndash90 mm) March 52 mm (n =11 range 25ndash100 mm) April 30 mm (n = 1)September 83 mm (n = 4 range 30ndash110mm) November 70 mm (n = 3 range 30ndash90mm) December 49 mm (n = 9 range 20ndash110 mm)

The primary difference in habitat betweenthe subspecies was that R f toltecus was col-lected in cold areas of the Mexican Transvol-canic Belt at heights ranging from 2250 m to2740 m in coniferous forests and xerophytic

shrubland whereas R f mustelinus was foundin the warm areas of the Balsas Basin in thesouthwestern part of the state between 1180m and 1900 m in the low tropical deciduousforest In fact Reithrodontomys f mustelinusis the only species of this genus located in thisprovince in the Balsas Basin (Saacutenchez 1993)In most instances this subspecies was found inxerophytic shrub and occasionally in oak forestand around Acambay in very altered forestsand farmlands (Fig 1) Species that were caughtge5 times together with R fulvescens toltecuswere Peromyscus gratus P maniculatus andBaiomys taylori Other associated species takenless frequently were Peromyscus melanophrysP difficilis P levipes Perognathus flavus Sigmo-don hispidus Liomys irroratus and R mega-lotis

Reithrodontomys fulvescens mustelinus Howell 1914

SPECIMENS EXAMINED (13)mdash1 km SWZacazonapan 1320 m (1) 3 km S 8 km W Sul-tepec 1300 m (2) 5 km S 5 km W PalmarChico 1180 m (1) 2 km E Tonatico (1) 5 kmS 55 km E Tonatico 1590 m (3) Cantildeada deNanchititla 1900 m (1) Jalmolonga 1600 m(1) Zumpahuacaacuten 1630 m (3)

ADDITIONAL RECORDmdash4 km SSE Tonatico1670 m (Saacutenchez 1993)

ECOLOGYmdashA female obtained in Februarydid not show signs of reproductive activityThree males collected in February had testesthat were 5 2 and 8 mm long respectivelyOne male in October had testes 9 mm longand 6 males in November had testes averaging30 mm long (range 20ndash50 mm)

This subspecies was found in ravines in thesouthwestern part of the state and was alwaysrelated to the low tropical deciduous forest(Fig 1) Species collected together with thissubspecies were Liomys pictus Oryzomyscouesi Peromyscus melanophrys and Baiomysmusculus

Reithrodontomys (Reithrodontomys)megalotis saturatus

Allen and Chapman 1897

SPECIMENS EXAMINED (387)mdashEl Angosta-dero 23 km NW Acambay 2700 m (1) 125 kmN 45 km W Acambay 2450 m (39) 9 km N75 km E Acambay 2740 m (5) 8 km N 3 kmW Aculco 2250 m (1) 35 km N 19 km Aculco


2650 m (24) 6 km S Amecameca 2450 m (1) 5 km W Atenco m (6) 1 km E Capulhuac 2600m (18) 1 km S 2 km W Chimalpa 2900 m (1)11 km S 25 km W Coatepec de Harinas 2870m (4) 15 km N 55 km E Ecatzingo 2970 m(2) 25 km N 15 km W Huixquilucan 2800 m(7) 3 km S Huixquilucan 3000 m (1) 5 km EJilotepec 2460 m (2) Cerro Ayaqueme 6 kmN 65 km W Juchitepec 2910 m (6) 4 km WLa Providencia 3050 m (3) Cuesta del Car-men 3 km N 1 km W Lengua de Vaca 2760 m(5) 3 km E Popo Park 2480 m (17) Nevado deToluca 4 km S 2 km W Raiacuteces 3350 m (1) 3km W Riacuteo Friacuteo 3000 m (4) 9 km W Riacuteo Friacuteo3050 m (1) 2 km S 5 km W Riacuteo Friacuteo 3100 m(1) 3 km S 4 km W Riacuteo Friacuteo 3290 m (1)Salazar 3050 m (1) San Felipe del Progreso2620 m (4) 1 km S 3 km W San Joseacute delRincoacuten 2960 m (15) 12 km S San Juan de lasHuertas 3850 m (2) 115 km S 25 km W SanJuan de las Huertas 3720 m (5) 25 km N SanJuan Zitlaltepec 2250 m (1) 7 km SE SanNicolas Coatepec 2740 m (3) 025 km S 225km E San Pedro Atlapulco 3100 m (4) 15 kmE San Pedro Nexapa 2700 m (6) 15 km N 4km E San Pedro Nexapa 2940 m (8) 6 km E4 km S San Pedro Nexapa 3050 m (1) SanSebastian Shala FES-Cuatitlaacuten (2) 8 km N175 km E San Simoacuten de Guerrero 2940 m (1)Santa Mariacutea de las Cuevas 2480 m (2) 5 km SSantiago Tianguistenco 2620 m (24) 4 km N15 km E Santiago Tilapa 2750 m (5) 4 km N1 km W Temascalcingo 2360 m (11) 65 km S6 km W Temascalcingo 2560 m (1) 4 km NTemoaya 2870 m (25) 75 km E Tenango deArista 2600 m (11) 6 km S 85 km W Tenangode Arista 2750 m (35) Texcalyacac 2570 m(1) 16 km SW Texcoco 2220 m (1) 600 m SWTlamacas 4000 m (1) 3 km S Tlapacoya 2250m (12) 9 km N Villa del Carboacuten 2400 m (2) 6km NW Villa Victoria 2400 m (11) 45 km S10 km W Villa Victoria 2560 m (14) 6 km N25 km W Villa Victoria 2710 m (19) CriaderoSan Cayetano 8 km NW Villa Victoria m (1)15 km S 7 km W Zinacantepec 3470 m (6)95 km N Zumpango 2270 m (1)

ADDITIONAL RECORDSmdash5 km NW Texcoco7600 ft Monte Riacuteo Frio 45ndash55 km ESE Meacutex-ico City 10500 ft Lago Zempoala 45 km SWMeacutexico City 9400 ft (Davis 1944) Rancho Coacuter-doba 2560ndash2600 m Riacuteo Friacuteo 2900ndash3000 m(Hooper 1947) 5 mi E Amecameca 9600 ftAtlacomulco 8200 ft Lerma 8500 ft Lago de

Zempoala 9400 ft Rancho Coacuterdoba 8200ndash8700ft vicinity of Riacuteo Friacuteo 9500ndash10500 ft Salazar10000 ft 3 mi NW Texcoco 7600 ft TolucaValley 9000 ft 15 mi SW Toluca 10000 ft Nand NW slope Volcaacuten Popocateacutepetl 10800ndash13000 ft N slope Volcaacuten de Toluca 11400 ft(Hooper 1952) Paso de Cortes 35 km E Ame-cameca 3450 m Cerro la Caldera 17 km ESECiudad de Meacutexico 2350 m N slope Popoca-teacutepetl 2 km E Villa de Guadalupe 2350 m(Villa-Ramiacuterez 1953) Lengua de Vaca San Caye-tano (Hooper 1957) Laguna Prieta 2900 m05 km SW Laguna Prieta 2900 m CerroOcuilapan 3200 m (Ramiacuterez-Pulido 1969)Cerro Gordo 8 km N San Juan Teotihuacaacuten2800 m 1 km E San Juan Teotihuacaacuten 2280m 5 km NE Zumpango 2260 m (Ceballos andGalindo 1984) 8 km E Zacualpan El Volcaacutende Toluca 11500 ft Laguna Quila Lagunasde Zempoala Puentecillas (Leoacuten and Romo1991) Unidad de Evaluacioacuten y Monitoreo dela Biodiversidad San Cayetano (Cervantes etal 1995) 6 km N 25 km W Villa Victoria2810 m (Gaona et al 2000)

MORPHOLOGYmdashReithrodontomys m satu-ratus is significantly smaller than R f toltecus inTL TV GLS CB ZB IC and LN (ANOVA Plt0001) R f mustelinus in TV (P lt 0001)ZB (P lt 001) TL and LN (P lt 005) Rmicrodon wagneri in TL TV CB and IC (P lt0001) and R s sumichrasti in TL TV GLSZB CB IC (P lt 0001) and LN (P lt 001)Reithrodontomys m saturatus is significantlylarger than R f mustelinus and R microdonwagneri in HF (P lt 0001 and P lt 005 re-spectively)

ECOLOGYmdashReithrodontomys megalotis isthe most abundant mammalian species in thestate of Meacutexico and is probably also the mostabundant rodent in the Basin of Meacutexico (Villa-Ramiacuterez 1953 Saacutenchez 1993) It has great en-vironmental plasticity and has been collectedin all types of vegetation in the state exceptfor the low tropical forest of the southwesternportion of the state (Fig 1) In many instancesthis species was taken in temperate forestsprimarily pine-oak oak and Abies forests andalso often in xerophytic shrubland In fact itwas most frequently captured in altered envi-ronments most commonly in agricultural fields(Fig 1) Reithrodontomys megalotis megalotisoccupies most areas of the Mexican high pla-teau and its southern distribution reaches the


boundaries of the state of Meacutexico Thereforethe examined specimens of R m saturatus fromAculco confirm the distribution suggested byHooper (1952)

Six pregnant females were taken 1 in Feb-ruary with 4 embryos (crown-rump length 15mm) 1 in March with 3 embryos (crown-rumplength 12 mm) 3 in November with 1 5 6embryos respectively (crown-rump lengths 14mm 19 mm and 2 mm respectively) and 1 inDecember with 3 embryos (crown-rump length22 mm) Moreover 4 lactating females werecollected in January March September andNovember Apparently this species has a sin-gle breeding season lasting from September toMarch however because data are limited itwill be necessary to verify these observationsduring field research The mean testis lengthsof males by month of collection were as fol-lows January 40 mm (n = 39 range 21ndash101mm) February 50 mm (n = 7 range 20ndash80mm) March 42 mm (n = 46 range 20ndash90mm) April 44 mm (n = 25 range 20ndash80mm) May 67 mm (n = 7 range 50ndash 100mm) June 59 mm (n = 10 range 40ndash80mm) September 110 mm (n = 1) November39 mm (n = 29 range 10ndash110 mm) Decem-ber 34 mm (n = 14 range 10ndash110 mm)

Species frequently captured along with Rmegalotis include Liomys irroratus Peromys-cus difficilis P levipes P maniculatus P melan-otis Neotomodon alstoni Baiomys taylori andMicrotus mexicanus Other species less fre-quently captured with R megalotis were Per-ognathus flavus Reithrodontomys fulvescensPeromyscus difficilis P maniculatus P melano-phrys P gratus and Sigmodon hispidus

Reithrodontomys (Reithrodontomys)sumichrasti sumichrasti

(Saussure 1861)

SPECIMENS EXAMINED (28)mdash2 km NE Chi-malpa 2700 m (1) 25 km NE Ecatzingo 2600m (3) Cuesta del Carmen 3 km N 1 km WLengua de Vaca 2760 m (6) 25 km E PopoPark 2470 m (1) 3 km E Popo Park 2480 m(2) 5 km N 2 km E Real de Arriba 1980 m (2)115 km S 25 km W San Juan de las Huertas3780 m (1) 1 km N 06 km E San Juan Tehu-ixtlaacuten 2480 m (2) Cerro Ayaqueme 6 km N65 km W Juchitepec 2910 m (1) 10 km N124 km E Temascaltepec 2450 m (2) 3 km STemascaltepec 1900 m (2) 6 km S 85 km W

Tenango de Arista 2750 m (2) 75 km E Ten-ango de Arista 2600 m (1) 3 km SW Valle deBravo 2060 m (1) Criadero San Cayetano 8km NW Villa Victoria (1)

ADDITIONAL RECORDSmdashHacienda Coacuterdoba8300 ft (Hooper 1952) Zoquiapan (Mass et al1981) ladera W Popocateacutepetl km 15 carreteraAmecameca-Tlamacas 3220 m (Ceballos andGalindo 1984)

MORPHOLOGY DISTRIBUTION AND TAXONOM-IC COMMENTSmdashHooper (1952) in his revisionof the Latin American Reithrodontomys sug-gested that it is R sumichrasti nerterus thatoccurs in the state of Meacutexico because this sub-species is distributed in eastern MichoacaacutenHowever the only feature that supports therecognition of R s nerterus as a subspecies isa lighter ventral coloration than found in R ssumichrasti (Hooper 1952) The specimensfrom the state of Meacutexico were compared withspecimens of R s nerterus and R s sumi-chrasti but no important differences werefound between them in fact ventral colorationwas highly variable even in populations close tothe type localities For example 3 of 10 speci-mens from Pinal de Amoles Quereacutetaro had adark ventral coloration and 5 of 15 specimensfrom Colima and Jalisco showed a pale venterFurthermore the specimens from westernPuebla Meacutexico and Meacutexico City showed anintermediate color between R s sumichrastiand R s nerterus (Hooper 1952) Owing to thehighly variable ventral coloration in these pop-ulations the differences between R s ner-terus and R s sumichrasti may be clinal varia-tion or simply individual variation This raisesquestions about the validity of R s nerterushowever it will be necessary to carry out a morecareful review to solve this problem

In the state of Meacutexico 14 adult and sub-adult specimens showed a pale ventral col-oration and 10 showed a dark coloration butthis variation did not appear to have any rela-tion to the date of capture nor to the geo-graphic areas of the captures We tentativelyassign our specimens to R s sumichrasti as themost conservative course of action to follow

Reithrodontomys s sumichrasti is signifi-cantly smaller than R f toltecus in TL TV(ANOVA P lt 005) and HF (P lt 001) andsignificantly larger than R f mustelinus inGLS ZB LN (P lt 0001) and CB (P lt 001)

ECOLOGYmdashA female taken in May carried3 embryos (crown-rump length 24 mm) and in


August a lactating female and a young individ-ual were captured Five females captured inFebruary March and November showed nogross reproductive activity Mean testis lengthsfor males by month of collection were as fol-lows February 70 mm (n = 1) March 45mm (n = 5 range 30ndash50 mm) May 45 mmand 50 mm (n = 2) July 73 mm (n = 3range 60ndash90 mm) August 90 mm and 90mm (n = 2) October 100 mm (n = 1) andNovember 20 mm (n = 1)

In general this species was found in pinepine-oak oak and Abies forests but they alsowere collected in grass and farming areas (Fig1) In Tenango de Arista a specimen was cap-tured in its nest under a log This species wastaken together with Liomys irroratus Reithro-dontomys chrysopsis R megalotis Peromyscusaztecus P difficilis P levipes P maniculatus Pmelanotis Neotomodon alstoni and Microtusmexicanus

DISCUSSION

The state of Meacutexico can be divided into 2main regions based on its physiographic andclimatic characteristics and types of vegeta-tion The largest region is the temperate zonecovering the northern and central parts of thestate and belonging to the Mexican Transvol-canic Belt Its elevations range from 2200 m to5400 m at the summit of the Popocateacutepetl Vol-cano and climate of the region is both tem-perate and cold In this province pine-oakforests prevail although xerophytic shrub alsois abundant especially in the northern part ofthe state along the boundary with the state ofHidalgo In this zone the vegetation has suf-fered intensive alteration and destruction dueto human activities uncontrolled felling oftrees creation of new farm and pasture landsand fundamentally the resulting soil erosionThe hot region is located in the southwesternpart of the state and is directly related to theprovince of the Balsas Basin Vegetation isalmost exclusively thorn deciduous forest theclimate is hot and the elevation is lt2200 mwith the lowest areas being at 450 m

Even though species of Reithrodontomysare located in both regions the records of Rchrysopsis R megalotis R microdon and Rsumichrasti confirm that they belong to thetemperate zone On the other hand R fulves-cens shows a clear ecological separation in the

distribution of its subspecies Reithrodonto-mys f toltecus has a wide distribution but onlyin the temperate zone and R f mustelinus be-longs exclusively to the Balsas Basin (Fig 1)The existence of R f mustelinus as the onlytaxon in the Balsas Basin probably is the resultof its greater tolerance of environmental con-ditions that restrict the other species in thesame genus (Saacutenchez 1993)

The co-occurrence of several Reithrodonto-mys species in the Mexican Transvolcanic Beltportion of the state of Meacutexico allows evalua-tion of the geographic distribution of eachspecies in relation to a definite habitat typeFor example R chrysopsis is found only onthe sides of the large volcanoes Nevado deToluca Popocateacutepetl and Iztacciacutehuatl above2800 m ( = 3375 m) with pine forest associa-tions but primarily with fir forest (Abies reli-giosa Fig 1) Reithrodontomys megalotis andR sumichrasti are found in all types of vegeta-tion of the temperate region of the state oakforest pine forest and fir forest with theirrespective ecotones and understory typesBoth species are distributed over a large alti-tudinal range which varies from 2000 m to3330 m for R megalotis and 1900 m to 3780 mfor R sumichrasti Reithrodontomys fulvescenstoltecus is restricted to xerophytic shrublandat locations where elevations range from 2250m to 2740 m R microdon is a species forwhich few specimens are known it has habitsof a tree-dwelling species making its capturevery difficult (Hooper 1952) The rare locali-ties of this taxon are found in very humidplaces in oak forests (Fig 1) with 15-m-highunderstory

Notwithstanding the types of vegetationshared by the different species the selectionof their habitats is distinctive In fact only in18 out of 95 localities (19) were 2 or morespecies collected together and at all of theselocalities the most common species was Rmegalotis It was sympatric with R chrysopsisat 3 localities with R sumichrasti at 7 locali-ties and with R fulvescens toltecus at 8 Onlyat 1 locality (115 km S 25 km W San Juan delas Huertas) were individuals from 3 speciescollected in the same trap line (R chrysopsisR sumichrasti and R megalotis) R megalotisis a species with great environmental plastic-ity It is the most abundant species and theonly one found in altered zones such as agri-cultural fields or pasture (Hooper 1952)


According to results from MANOVA (4commonly used test statistics Wilksrsquo lambdaHotelling-Lawley trace Pillairsquos trace and Royrsquoslargest root) these species have significant dif-ferences (P lt 0001) among the mean vectorsfrom the 6 taxa To understand the relation-ship of the significant differences among the 6taxa of Reithrodontomys we performed a dis-criminant analysis (Fig2) The discriminantanalysis correctly classified 100 of the speci-mens from the 6 taxa with 5 canonical variatesThree variates accounted for 961 of the vari-ance and the canonical correlations were 095086 and 077 respectively (Table 2) The highlynegative values in canonical variate 1 for allmeasurements except breadth of zygomaticplate suggest that size plays an important rolein identification of Reithrodontomys speciesCranial breadth greatest length of skull zygo-matic breadth total length and length of tailvertebrae are particularly important Theanalysis indicates that as other measurementsbecome larger the breadth across the zygo-matic plate varies in the opposite directionand becomes narrower Reithrodantomys chrys-opsis R sumichrasti and R megalotis are fairlywell separated along the 1st canonical variate

with only some overlap between the first 2species The taxa R f toltecus R f mustelinusand R microdon broadly overlap the other taxaalong the 1st variate

The highest positive weightings along canon-ical variate 2 are for length of tail vertebraeand total length whereas the highest negativevalues were for breadth across the zygomaticplate and zygomatic breadth The 2 taxa clearlyseparated from the others along canonicalvariate 2 were R microdon and R f mustelinusThe 2 subspecies of R fulvescens were clearlyseparated along canonical variate 2 (Fig 2)Using only the first 2 canonical variates(accounting for 851 of total variance) onlythe taxa R f toltecus and R sumichrasti werenot separated from each other This seems toindicate that these 2 taxa most closely resem-ble each other in external and cranial morpho-metrics among the 6 taxa occurring in thestate of Meacutexico Ultimately however all taxacan be differentiated based on the measure-ments given herein

There is good evidence of a relationshipbetween the morphometric separation and habi-tat preferences among species of the genusReithrodontomys in the state of Meacutexico In fact


Fig 2 First and 2nd canonical variates resulting from disciminant analysis showing relationships among 65 specimensof Reithrodontomys R chrysopsis (n = 9) R fulvescens mustelinus (n = 1) R f toltecus (n = 11) R megalotis (n = 30)R microdon (n = 1) R sumichrasti (n = 13) The canonical correlation values from the 1st and 2nd canonical correla-tion are in Table 2

CV

2 (

212

)

CV 1 (639)

the species that share habitats exhibit notice-able morphometric differences For example R fulvenscens toltecus and R sumichrasti withclose morphometric similarities show consider-able differences in habitat utilization in con-trast R fulvenscens toltecus and R megalotiscan be found in the same locality but the mor-phometric differences are very strong (Fig 2)The sequential expansion-retraction eventspostulated by Hooper (1952) would seem toexplain the evolutionary processes that deter-mined the taxonomic patterns and distributionof Reithrodontomys species in the Transvol-canic Belt These events probably were re-sponsible for habitat selection in the differentspecies of the genus as well as for size diversi-fication within the genus

ACKNOWLEDGMENTS

Our grateful thanks go to M en C TiculAacutelvarez (deceased) former Curator and BioacutelJuan Carlos Loacutepez Vidal Curator of the Colec-cioacuten Mastozooloacutegica de Escuela Nacional deCiencias Bioloacutegicas IPN for permission toexamine the specimens under their care Wegive special thanks to 2 anonymous reviewerswho enhanced our work through their valu-able comments This project was partiallyfunded by grants from Consejo Nacional deCiencia y Tecnologiacutea (CONACyT No 39619-Q-A to JRP Our special thanks go to MiguelBravo Rivera and Diana Iraacuten Loacutepez for pre-paring Fig 1

LITERATURE CITED

CEBALLOS GG AND CL GALINDO 1984 Mamiacuteferos sil-vestres de la Cuenca de Meacutexico Edit Limusa Meacutex-ico

CERVANTES FA G MATAMOROS AND I MARTIacuteNEZ MATEOS1995 Mamiacuteferos silvestres de la Unidad de Evalu-acioacuten y Monitoreo de la Biodiversidad ldquoIng LuisMaciacuteas Arellanordquo San Cayetano Estado de MeacutexicoAnales del Instituto de Biologiacutea Universidad Nacio-nal Autoacutenoma de Meacutexico Serie Zoologiacutea 66233ndash239

CHAacuteVEZ C AND G CEBALLOS 1998 Diversidad y estadode conservacioacuten de los mamiacuteferos del Estado deMeacutexico Revista Mexicana de Mastozoologiacutea 3113ndash134

DAVIS WB 1944 Notes on Mexican mammals Journal ofMammalogy 25370ndash403

GAONA S C GALINDO-GALINDO AND M GONZAacuteLEZ-ESCAMILLA 2000 Ampliacioacuten y confirmacioacuten de ladistribucioacuten de la ardilla voladora Glaucomys volansgoldmani (Nelson 1904) Vertebrata Mexicana 89ndash15

HALL ER 1981 The mammals of North America 2nd edi-tion Volume 1 John Wiley amp Sons Inc New York

HINTZE JL 2000 NCSS 2000 statistical system for Win-dows Userrsquos guide NCSS Kaysville UT Availablefrom httpwwwncsscom

HOOPER ET 1947 Notes on Mexican mammals Journalof Mammalogy 2840ndash57

______ 1952 A systematic review of the harvest mice(genus Reithrodontomys) of Latin America Miscella-neous Publications Museum of Zoology Universityof Michigan 771ndash225

______ 1957 Records of Mexican mammals OccasionalPapers of the Museum of Zoology University ofMichigan 5861ndash9

HOWELL AH 1914 Revision of the American harvestmice (genus Reithrodontomys) North American Fauna361ndash97

LEOacuteN P L AND E ROMO V 1991 Cataacutelogo de mamiacuteferos(Vertebrata Mammalia) Serie Cataacutelogos del Museo


TABLE 2 Results of a discriminant analysis of 65 specimens of Reithrodontomys from the state of Meacutexico with 100correct classification Acronyms are explained in the methods

Variable CV1a CV2 CV3

TL ndash0432311 0289363 ndash0147233TV ndash044583 0316854 ndash0056913HF ndash0344824 0024895 ndash0385178LE ndash0266006 ndash0085493 0105441GLS ndash0538076 ndash0286695 ndash0124636ZB ndash0494822 ndash0395939 ndash0086483CB ndash0668076 ndash0269575 ndash0006933IC ndash02413 022392 ndash0067133BR ndash0192346 0254884 ndash0568324LN ndash0399501 ndash0185197 ndash0071697MTR ndash0212583 ndash0291371 ndash0439342BZP 0068433 ndash0405147 ndash0245465BMF ndash051557 0161743 0125782Canonical correlation 095 086 077Eigenvalues 835 278 144 variance explained 639 212 11aCanonical variate


de Zoologiacutea ldquoAlfonso L Herrerardquo Facultad de Cien-cias Universidad Nacional Autoacutenoma de MeacutexicoCataacutelogo 21ndash68

MASS J R PATROacuteN A SUAacuteREZ S BLANCO G CEBALLOSC GALINDO AND A PESCADOR 1981 Ecologiacutea de laEstacioacuten Experimental Zoquiapan (Descripcioacuten gen-eral vegetacioacuten y fauna) Universidad AutoacutenomaChapingo Direccioacuten de Difusioacuten Cultural Departa-mento de Bosques Coleccioacuten Cuadernos Universi-tarios 21ndash114

MERRIAM CH 1900 Description of a new harvest mouse(Reithrodontomys) from Meacutexico Proceedings of theBiological Society of Washington 13152

______ 1901 Descriptions of 23 new harvest mice (genusReithrodontomys) Proceedings of Washington Acad-emy of Sciences 3547ndash558

MOTULSKY HJ 2003 Prism 4 statistics guidemdashstatisticalanalyses for laboratory and clinical researchersGraphPad Prisma Software Inc San Diego CA

RAMIacuteREZ-PULIDO J 1969 Contribucioacuten al estudio de losmamiacuteferos del Parque Nacional ldquoLagunas de Zem-poalardquo Morelos Meacutexico Anales del Instituto de

Biologiacutea Universidad Nacional Autoacutenoma de Meacutex-ico 40253ndash290

______ 1995 Sinopsis de los mamiacuteferos del Estado deMeacutexico Revista de la Sociedad Mexicana de Histo-ria Natural 46205ndash246

RAMIacuteREZ-PULIDO J A CASTRO-CAMPILLO AND U AGUIL-ERA 1997 Capiacutetulo III Mamiacuteferos Pages 159ndash201in Lista taxonoacutemica de los vertebrados terrestres delEstado de Meacutexico Universidad Autoacutenoma del Estadode Meacutexico Coleccioacuten Ciencias y Teacutecnicas 321ndash201

SAacuteNCHEZ O 1993 Anaacutelisis de algunas tendencias eco-geograacuteficas del geacutenero Reithrodontomys (RodentiaMuridae) Pages 25ndash44 in RA Medelliacuten and G Cebal-los editors Avances en el estudio de los mamiacuteferosde Meacutexico Asociacioacuten Mexicana de MastozoologiacuteaPublicaciones Especiales 11ndash464

VILLA-RAMIacuteREZ B 1953 Mamiacuteferos silvestres del Valle deMeacutexico Anales del Instituto de Biologiacutea UniversidadNacional Autoacutenoma de Meacutexico 23269ndash492

Received 15 March 2006Accepted 31 October 2006

APPENDIX Localities of specimens of the genus Reithrodontomys housed in the Mammals Collection of the NationalSchool of Biological Sciences (Coleccioacuten de Mamiacuteferos de la Escuela Nacional de Ciencias Bioloacutegicas del InstitutoPoliteacutecnico Nacional) The elevation is in meters (m) and the geographic coordinates are degrees minutes and secondsLocalities are ordered according to latitude

No Locality Latitude Longitude

1 8 km N 3 km W Aculco 2250 m 20deg10prime13Prime 99deg50prime56Prime2 35 km N 19 km E Aculco 2650 m 20deg07prime51Prime 99deg39prime21Prime3 El Angostadero 23 km NW Acambay 2700 m 20deg06prime06Prime 99deg41prime45Prime4 Llanos de Aculco 3200 m 20deg05prime54Prime 99deg49prime37Prime5 125 km N 45 km W Acambay 2450 m 20deg04prime05Prime 99deg53prime02Prime6 9 km N 75 km E Acambay 2740 m 20deg02prime09Prime 99deg46prime33Prime7 85 km N 15 km E Acambay 2740 m 20deg01prime55Prime 99deg49prime44Prime8 85 km N 25 km E Jilotepec 2320 m 20deg01prime46Prime 99deg30prime43Prime9 5 km E Jilotepec 2460 m 19deg57prime20Prime 99deg30prime03Prime

10 4 km N 1 km W Temascalcingo 2360 m 19deg57prime02Prime 100deg00prime36Prime11 95 km N Zumpango 2270 m 19deg52prime56Prime 99deg05prime52Prime12 Santa Mariacutea Cuevas 2480 m 19deg52prime00Prime 99deg05prime47Prime13 65 km S 6 km W Temascalcingo 2560 m 19deg51prime23Prime 100deg03prime00Prime14 25 km N San Juan Zitlaltepec 2250 m 19deg49prime55Prime 99deg08prime52Prime15 9 km N Villa del Carboacuten 2400 m 19deg48prime30Prime 99deg27prime43Prime16 025 km S 225 km E San Pedro Atlapulco 3100 m 19deg47prime42Prime 99deg51prime14Prime17 27 km N 9 km W Villa del Carboacuten 2670 m 19deg45prime08Prime 99deg22prime55Prime18 2 km NW Santiago Tolman 2600 m 19deg44prime25Prime 98deg48prime03Prime19 San Felipe del Progreso 2620 m 19deg42prime45Prime 99deg57prime04Prime20 2 km S 3 km E Chilpa 2270 m 19deg42prime05Prime 100deg00prime36Prime21 4 km W La Providencia 3050 m 19deg41prime52Prime 99deg47prime24Prime22 25 km W San Francisco Tepojaco 2350 m 19deg39prime30Prime 97deg13prime08Prime23 1 km S 3 km W San Joseacute del Rincoacuten 2960 m 19deg39prime05Prime 100deg10prime11Prime24 San Sebastiaen Shala FES Cuatitlaacuten 19deg38prime46Prime 99deg12prime41Prime25 3 km E Popo Park 2480 m 19deg38prime45Prime 98deg45prime07Prime26 1 km S 2 km E Nicolaacutes Romero 2350 m 19deg36prime56Prime 99deg17prime41Prime27 2 km S Coacalco 2440 m 19deg36prime55Prime 99deg05prime34Prime28 11 km N Barrientos 2300 m 19deg35prime38Prime 99deg13prime22Prime29 45 km E Chiconautla 2350 m 19deg35prime14Prime 99deg02prime20Prime30 2 km NE Chimalpa 2700 m 19deg34prime55Prime 98deg52prime44Prime31 1 km S 2 km W Chimalpa 2900 m 19deg33prime36Prime 98deg54prime36Prime32 2 km NW Tlanepantla 2300 m 19deg32prime58Prime 99deg12prime25Prime33 4 km N Temoaya 2870 m 19deg30prime16Prime 99deg35prime34Prime34 6 km N 25 km W Villa Victoria 2710 m 19deg29prime32Prime 99deg58prime37Prime35 Criadero San Cayetano 8 km NW Villa Victoria 19deg28prime59Prime 100deg03prime00Prime


APPENDIX Continued


36 Cuesta del Carmen 3 km N 1 km W Lengua de Vaca 2760 m 19deg28prime39Prime 100deg11prime24Prime37 6 km NW Villa Victoria 2400 m 19deg28prime32Prime 99deg57prime25Prime38 16 km SW Texcoco 2220 m 19deg24prime13Prime 98deg46prime44Prime39 45 km S 10 km W Villa Victoria 2560 m 19deg23prime48Prime 99deg54prime17Prime40 25 km N 15 km W Huixquilucan 2800 m 19deg23prime01Prime 99deg21prime50Prime41 9 km W Riacuteo Friacuteo 3050 m 19deg21prime09Prime 98deg45prime03Prime42 2 km S 5 km W Riacuteo Friacuteo 3100 m 19deg20prime04Prime 98deg42prime53Prime43 3 km S Huixquilucan 3000 m 19deg20prime03Prime 99deg20prime59Prime44 3 km W Riacuteo Friacuteo 3000 m 19deg19prime57Prime 98deg41prime49Prime45 3 km S 4 km W Riacuteo Friacuteo 3290 m 19deg19prime31Prime 98deg42prime21Prime46 Salazar 3050 m 19deg18prime25Prime 99deg23prime17Prime47 3 km S Tlapacoya 2250 m 19deg18prime08Prime 98deg52prime26Prime48 5 km W Atenco 19deg16prime13Prime 99deg34prime37Prime49 4 km N 15 km E Santiago Tilapa 2750 m 19deg13prime17Prime 99deg24prime21Prime50 15 km S 5 km E San Rafael 19deg11prime43Prime 98deg42prime43Prime51 1 km E Capulhuac 2600 m 19deg11prime29Prime 99deg27prime18Prime52 3 km SW Valle de Bravo 2060 m 19deg10prime22Prime 100deg06prime36Prime53 Cerro Ayaqueme 6 km N 65 km W Juchitepec 2910 m 19deg09prime14Prime 98deg56prime20Prime54 15 km S 7 km W Zinacantepec 3470 m 19deg08prime57Prime 99deg40prime11Prime55 11 km S 25 km W San Juan de las Huertas 3780 m 19deg08prime45Prime 99deg46prime40Prime56 115 km S 25 km W San Juan de las Huetas 3780 m 19deg08prime45Prime 99deg46prime58Prime57 155 km S 7 km W Zinacatepec 3470 m 19deg08prime43Prime 99deg40prime11Prime58 12 km S San Juan de las Huertas 3850 m 19deg08prime15Prime 99deg45prime21Prime59 5 km S Santiago Tianguistenco 2620 m 19deg08prime07Prime 99deg28prime04Prime60 10 km N 124 km E Temascaltepec 2450 m 19deg08prime02Prime 99deg55prime40Prime61 Texcalyacac 2570 m 19deg07prime52Prime 99deg30prime05Prime62 185 km S 9 km W Zinacatepec 3440 m 19deg07prime47Prime 99deg39prime10Prime63 Nevado de Toluca 4 km S 2 km W Raiacuteces 3350 m 19deg07prime26Prime 99deg49prime19Prime64 Nevado de Toluca 19deg07prime25Prime 99deg49prime15Prime65 600 m SW Tlamacas 4000 m 19deg06prime15Prime 98deg46prime15Prime66 22 km N 65 km E San Pedro Nexapa 3490 m 19deg06prime10Prime 98deg40prime33Prime67 75 km E Tenango de Arista 2600 m 19deg06prime09Prime 99deg39prime21Prime68 16 km N 64 km E San Pedro Nexapa 3200 m 19deg05prime51Prime 98deg40prime37Prime69 7 km SE San Nicolas Coatepec 2740 m 19deg05prime49Prime 99deg22prime44Prime70 15 km N 4 km E San Pedro Nexapa 2940 m 19deg05prime47Prime 98deg41prime56Prime71 4 km S 9 km E Amecameca 3480 m 19deg05prime33Prime 98deg40prime48Prime72 8 km N 175 km E San Simoacuten de Guerrero 2940 m 19deg05prime30Prime 99deg52prime04Prime73 5 km N 2 km E Real de Arriba 1980 m 19deg05prime04Prime 99deg59prime06Prime74 15 km E San Pedro Nexapa 2700 m 19deg04prime59Prime 98deg43prime15Prime75 52 km E San Pedro Nexapa 3250 m 19deg04prime59Prime 98deg41prime16Prime76 55 km E San Pedro Nexapa 3250 m 19deg04prime59Prime 98deg41prime05Prime77 6 km S Amecameca 2450 m 19deg04prime24Prime 98deg45prime43Prime78 1 km SW Zacazonapan 1320 m 19deg03prime59Prime 100deg14prime23Prime79 25 km E Popo Park 2470 m 19deg03prime51Prime 98deg45prime25Prime80 1 km N 06 km E San Juan Tehuixtlaacuten 2480 m 19deg03prime41Prime 98deg46prime19Prime81 6 km S 85 km W Tenango de Arista 2750 m 19deg02prime52Prime 99deg30prime39Prime82 6 km E 4 km S San Pedro Nexapa 3050 m 19deg02prime50Prime 98deg40prime48Prime83 4 km S 85 km E San Pedro Nexapa 3500 m 19deg02prime49Prime 98deg39prime28Prime84 7 km SE San Pedro Nexapa 3010 m 19deg02prime45Prime 98deg40prime58Prime85 3 km S Temascaltepec 1900 m 19deg00prime56Prime 100deg02prime24Prime86 25 km NE Ecatzingo 2600 m 18deg58prime19Prime 98deg44prime13Prime87 15 km N 55 km E Ecatzingo 2970 m 18deg58prime12Prime 98deg42prime10Prime88 Jalmolonga 1600 m 18deg55prime06Prime 99deg29prime37Prime89 Cantildeada de Nanchitita 1900 m 18deg51prime47Prime 100deg27prime00Prime90 Zumpahuacan 1630 m 18deg50prime11Prime 99deg34prime43Prime91 3 km S 8 km W Sultepec 1300 m 18deg49prime53Prime 100deg01prime47Prime92 11 km S 25 km W Coatepec de Harinas 2870 m 18deg49prime30Prime 99deg47prime27Prime93 2 km E Tonatico 18deg48prime02Prime 99deg39prime03Prime94 5 km S 55 km E Tonatico 1590 m 18deg46prime38Prime 99deg37prime01Prime95 5 km S 5 km W Palmar Chico 1180 m 18deg38prime57Prime 100deg24prime36Prime

Relatively few papers dealing with the genusReithrodontomys from the Mexican state ofMeacutexico have been published The oldest paperknown to us is Merriam (1901) in whichappeared the description of Reithrodontomyschrysopsis based on specimens from VolcanPopocateacutepetl In the same paper Merriam(1901) also described R tolucae from theNevado de Toluca but Howell (1914) in hisrevision of the genus Reithrodontomys recog-nized R tolucae as a subspecies of R chrysop-sis Merriam (1901) also recorded the presenceof R megalotis and R fulvescens in the state ofMeacutexico Hooper (1952) in his review of LatinAmerican harvest mice reported records ofseveral species in the state R tolucae wasretained as a junior synonym of R chrysopsisand R sumichrasti was recorded for the 1sttime from the state of Meacutexico Other papers

contributing to our understanding of the genusin the state are those by Davis (1944) Villa-Ramiacuterez (1953) and Ramiacuterez-Pulido (1969)

Because available information is scarce ourpurpose is to document the data for species ofReithrodontomys in the state of Meacutexico Thesedata have accumulated over the past 35 yearsin the mammal collection of the EscuelaNacional de Ciencias Bioloacutegicas del InstitutoPoliteacutecnico Nacional We present informationfor R chrysopsis R sumichrasti R megalotisR fulvescens (represented by 2 subspeciesmdashR f toltecus and R f mustelinus) and Rei-throdontomys microdon wagneri which is re-corded herein for the 1st time from the stateof Meacutexico With a sample of 590 specimens ofthe 5 species we evaluate compare and con-trast the taxonomy morphometrics repro-duction and habitat characteristics of thesespecies

Western North American Naturalist 67(2) 2007 pp 238ndash250

REVIEW OF THE HARVEST MICE (GENUS REITHRODONTOMYS) IN THE MEXICAN STATE OF MEacuteXICO

Noeacute Gonzaacutelez-Ruiz12 Joseacute Ramiacuterez-Pulido2 and Hugh H Genoways3

ABSTRACTmdashWe examined 590 specimens of Reithrodontomys from 95 localities in the state of Meacutexico Four speciesof the subgenus Reithrodontomys and 1 of the subgenus Aporodon were identified The former subgenus included Rchrysopsis R sumichrasti R megalotis and R fulvescens which has 2 subspeciesmdashR f toltecus and R f mustelinusmdashin the state The representative of the subgenus Aporodon is R microdon wagneri which is recorded for the first time inthe state We give information on taxonomy morphometrics reproduction habitat characteristics and related fauna Adiscriminant analysis correctly classified 100 of specimens from the 6 taxa with 5 canonical variates and accounted for961 of the variance with the first 3 canonical variates

Key words rodents Reithrodontomys distribution taxonomy state of Meacutexico Meacutexico

RESUMENmdashSe examinaron 590 ejemplares de las especies del geacutenero Reithrodontomys procedentes de 95 locali-dades del Estado de Meacutexico Se identificaron cuatro especies del subgeacutenero Reithrodontomys y una del Aporodon delprimero R chrysopsis R sumichrasti R megalotis y R fulvescens de eacutesta uacuteltima se identificaron dos subespecies R ftoltecus y R f mustelinus La especie del subgeacutenero Aporodon es R microdon wagneri que se registra por primera vezpara el Estado De cada taxon se dan medidas comentarios taxonoacutemicos y de la condicioacuten reproductora asiacute como lascaracteriacutesticas del haacutebitat y de la fauna asociada Un anaacutelisis discriminante mostroacute el 100 de certeza en la identifi-cacioacuten de los 6 taxa y los primeros tres vectores canoacutenicos expresaron el 961 de variacioacuten total

Palabras clave roedores Reithrodontomys distribucioacuten taxonomiacutea Estado de Meacutexico Meacutexico

1Universidad Autoacutenoma Metropolitana Unidad Iztapalapa Divisioacuten de CBS Departamento de Biologiacutea Apartado Postal 55-535 Meacutexico 09340 DFMeacutexico

2Laboratorio de Cordados Terrestres Escuela Nacional de Ciencias Bioloacutegicas Instituto Politeacutecnico Nacional Prolongacioacuten de Carpio y Plan de AyalaColonia Santo Tomaacutes Apartado Postal CON 256 11340 DF Meacutexico E-mail jrpxanumuammx

3University of Nebraska State Museum and School of Natural Resources Sciences W436 Nebraska Hall University of NebraskandashLincoln Lincoln NE68588-0514

238

METHODS







SPECIES ACCOUNTS


Hooper 1950











Agricultural Fields

Human Settlement

Coniferous Forest

Shrubs

Grassland



Specimens Examined


AB

LE

1 S

elec

ted

exte

rnal

and

cra

nial

mea

sure

men

ts o

f the

gen

us R

eith

rodo

ntom

ysfr

om th

e M

exic

an s

tate

of M

eacutexic

o A

cron

yms

are

expl

aine

d in

the

met

hods

The

bas

ic s

tatis

tics

incl

ude

stan

dard

dev

iatio

n (s

) and

num

ber

of s

peci

men

s ex

amin

ed (n

)

TL

TV

HF

LE

GL

SZ

BC

BIC

BR

LN

LM

TR

BZ

PB

MF

RE

ITH

RO

DO

NT

OM

YSC

HR

YSO

PSIS

CH

RYS

OPS

IS

xndash17

93

100

220

64

180

924

34

131

511

43

337

39

883

365

237

139

Min

imum

157

8119

1523

68

118

111

22

323

361

846

335

204

113

Max

imum

179

311

222

2124

78

131

511

73

353

409

949

392

319

163

s13

53

927

081

187

035

035

018

011

014

027

016

031

012

n10

1011

1111

1111

1111

1111

1111

RE

ITH

RO

DO

NT

OM

YSF

ULV

ESC

EN

SM

UST

ELI

NU

S

xndash17

233

103

1815

521

41

104

210

23

283

697

333

231

871

28M

inim

um15

795

1615

201

910

42

987

321

352

707

312

18

126

Max

imum

189

111

2016

222

810

99

106

83

333

847

853

331

921

31s

160

411

31

20

711

090

560

430

060

160

450

110

060

03n

32

32

33

33

33

33

3R

EIT

HR

OD

ON

TO

MYS

FU

LVE

SCE

NS

TO

LTE

CU

S

xndash18

146

101

520

31

156

223

02

116

210

84

347

426

83

35

218

131

Min

imum

171

8619

1322

44

113

810

31

322

403

796

339

193

117

Max

imum

199

112

2217

241

511

86

113

93

684

268

893

722

461

43s

885

703

075

126

055

016

024

014

022

03

008

016

009

n13

1213

1313

1313

1312

1313

1313

RE

ITH

RO

DO

NT

OM

YSM

EG

ALO

TIS

SAT

UR

ATU

S

xndash15

297

793

618

15

140

922

04

110

910

21

318

376

79

362

235

101

Min

imum

131

6716

1020

81

104

89

622

933

377

313

352

080

69M

axim

um17

089

2018

233

111

810

74

349

428

873

362

235

126

s9

955

021

194

049

029

021

013

376

029

012

015

013

n33

3333

3233

3233

3333

3332

3333

RE

ITH

RO

DO

NT

OM

YSSU

MIC

HR

AST

ISU

MIC

HR

AST

I

xndash16

991

86

188

615

43

227

911

48

107

63

363

848

273

362

211

28M

inim

um16

066

1810

220

210

910

35

298

362

798

308

21

16M

axim

um18

110

221

1923

73

119

811

03

383

402

827

359

248

128

s6

5910

17

095

21

046

032

024

022

012

027

015

015

011

n14

1414

1415

1514

1515

1515

1515

RE

ITH

RO

DO

NT

OM

YSM

ICR

OD

ON

WA

GN

ER

I

xndash18

811

55

1818

221

511

63

111

53

673

847

963

051

451

61M

inim

um18

511

218

1821

77

115

410

89

351

384

765

305

137

158

Max

imum

191

119

1818

225

117

211

48

379

384

819

306

152

163

s4

244

950

00

370

130

30

140

280

011

450

03n

22

22

32

33

13

23

3



Merriam 1900






































(Saussure 1861)











DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued



METHODS







SPECIES ACCOUNTS


Hooper 1950











Agricultural Fields

Human Settlement

Coniferous Forest

Shrubs

Grassland



Specimens Examined


AB

LE

1 S

elec

ted

exte

rnal

and

cra

nial

mea

sure

men

ts o

f the

gen

us R

eith

rodo

ntom

ysfr

om th

e M

exic

an s

tate

of M

eacutexic

o A

cron

yms

are

expl

aine

d in

the

met

hods

The

bas

ic s

tatis

tics

incl

ude

stan

dard

dev

iatio

n (s

) and

num

ber

of s

peci

men

s ex

amin

ed (n

)

TL

TV

HF

LE

GL

SZ

BC

BIC

BR

LN

LM

TR

BZ

PB

MF

RE

ITH

RO

DO

NT

OM

YSC

HR

YSO

PSIS

CH

RYS

OPS

IS

xndash17

93

100

220

64

180

924

34

131

511

43

337

39

883

365

237

139

Min

imum

157

8119

1523

68

118

111

22

323

361

846

335

204

113

Max

imum

179

311

222

2124

78

131

511

73

353

409

949

392

319

163

s13

53

927

081

187

035

035

018

011

014

027

016

031

012

n10

1011

1111

1111

1111

1111

1111

RE

ITH

RO

DO

NT

OM

YSF

ULV

ESC

EN

SM

UST

ELI

NU

S

xndash17

233

103

1815

521

41

104

210

23

283

697

333

231

871

28M

inim

um15

795

1615

201

910

42

987

321

352

707

312

18

126

Max

imum

189

111

2016

222

810

99

106

83

333

847

853

331

921

31s

160

411

31

20

711

090

560

430

060

160

450

110

060

03n

32

32

33

33

33

33

3R

EIT

HR

OD

ON

TO

MYS

FU

LVE

SCE

NS

TO

LTE

CU

S

xndash18

146

101

520

31

156

223

02

116

210

84

347

426

83

35

218

131

Min

imum

171

8619

1322

44

113

810

31

322

403

796

339

193

117

Max

imum

199

112

2217

241

511

86

113

93

684

268

893

722

461

43s

885

703

075

126

055

016

024

014

022

03

008

016

009

n13

1213

1313

1313

1312

1313

1313

RE

ITH

RO

DO

NT

OM

YSM

EG

ALO

TIS

SAT

UR

ATU

S

xndash15

297

793

618

15

140

922

04

110

910

21

318

376

79

362

235

101

Min

imum

131

6716

1020

81

104

89

622

933

377

313

352

080

69M

axim

um17

089

2018

233

111

810

74

349

428

873

362

235

126

s9

955

021

194

049

029

021

013

376

029

012

015

013

n33

3333

3233

3233

3333

3332

3333

RE

ITH

RO

DO

NT

OM

YSSU

MIC

HR

AST

ISU

MIC

HR

AST

I

xndash16

991

86

188

615

43

227

911

48

107

63

363

848

273

362

211

28M

inim

um16

066

1810

220

210

910

35

298

362

798

308

21

16M

axim

um18

110

221

1923

73

119

811

03

383

402

827

359

248

128

s6

5910

17

095

21

046

032

024

022

012

027

015

015

011

n14

1414

1415

1514

1515

1515

1515

RE

ITH

RO

DO

NT

OM

YSM

ICR

OD

ON

WA

GN

ER

I

xndash18

811

55

1818

221

511

63

111

53

673

847

963

051

451

61M

inim

um18

511

218

1821

77

115

410

89

351

384

765

305

137

158

Max

imum

191

119

1818

225

117

211

48

379

384

819

306

152

163

s4

244

950

00

370

130

30

140

280

011

450

03n

22

22

32

33

13

23

3



Merriam 1900






































(Saussure 1861)











DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued








Agricultural Fields

Human Settlement

Coniferous Forest

Shrubs

Grassland



Specimens Examined


AB

LE

1 S

elec

ted

exte

rnal

and

cra

nial

mea

sure

men

ts o

f the

gen

us R

eith

rodo

ntom

ysfr

om th

e M

exic

an s

tate

of M

eacutexic

o A

cron

yms

are

expl

aine

d in

the

met

hods

The

bas

ic s

tatis

tics

incl

ude

stan

dard

dev

iatio

n (s

) and

num

ber

of s

peci

men

s ex

amin

ed (n

)

TL

TV

HF

LE

GL

SZ

BC

BIC

BR

LN

LM

TR

BZ

PB

MF

RE

ITH

RO

DO

NT

OM

YSC

HR

YSO

PSIS

CH

RYS

OPS

IS

xndash17

93

100

220

64

180

924

34

131

511

43

337

39

883

365

237

139

Min

imum

157

8119

1523

68

118

111

22

323

361

846

335

204

113

Max

imum

179

311

222

2124

78

131

511

73

353

409

949

392

319

163

s13

53

927

081

187

035

035

018

011

014

027

016

031

012

n10

1011

1111

1111

1111

1111

1111

RE

ITH

RO

DO

NT

OM

YSF

ULV

ESC

EN

SM

UST

ELI

NU

S

xndash17

233

103

1815

521

41

104

210

23

283

697

333

231

871

28M

inim

um15

795

1615

201

910

42

987

321

352

707

312

18

126

Max

imum

189

111

2016

222

810

99

106

83

333

847

853

331

921

31s

160

411

31

20

711

090

560

430

060

160

450

110

060

03n

32

32

33

33

33

33

3R

EIT

HR

OD

ON

TO

MYS

FU

LVE

SCE

NS

TO

LTE

CU

S

xndash18

146

101

520

31

156

223

02

116

210

84

347

426

83

35

218

131

Min

imum

171

8619

1322

44

113

810

31

322

403

796

339

193

117

Max

imum

199

112

2217

241

511

86

113

93

684

268

893

722

461

43s

885

703

075

126

055

016

024

014

022

03

008

016

009

n13

1213

1313

1313

1312

1313

1313

RE

ITH

RO

DO

NT

OM

YSM

EG

ALO

TIS

SAT

UR

ATU

S

xndash15

297

793

618

15

140

922

04

110

910

21

318

376

79

362

235

101

Min

imum

131

6716

1020

81

104

89

622

933

377

313

352

080

69M

axim

um17

089

2018

233

111

810

74

349

428

873

362

235

126

s9

955

021

194

049

029

021

013

376

029

012

015

013

n33

3333

3233

3233

3333

3332

3333

RE

ITH

RO

DO

NT

OM

YSSU

MIC

HR

AST

ISU

MIC

HR

AST

I

xndash16

991

86

188

615

43

227

911

48

107

63

363

848

273

362

211

28M

inim

um16

066

1810

220

210

910

35

298

362

798

308

21

16M

axim

um18

110

221

1923

73

119

811

03

383

402

827

359

248

128

s6

5910

17

095

21

046

032

024

022

012

027

015

015

011

n14

1414

1415

1514

1515

1515

1515

RE

ITH

RO

DO

NT

OM

YSM

ICR

OD

ON

WA

GN

ER

I

xndash18

811

55

1818

221

511

63

111

53

673

847

963

051

451

61M

inim

um18

511

218

1821

77

115

410

89

351

384

765

305

137

158

Max

imum

191

119

1818

225

117

211

48

379

384

819

306

152

163

s4

244

950

00

370

130

30

140

280

011

450

03n

22

22

32

33

13

23

3



Merriam 1900






































(Saussure 1861)











DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued




AB

LE

1 S

elec

ted

exte

rnal

and

cra

nial

mea

sure

men

ts o

f the

gen

us R

eith

rodo

ntom

ysfr

om th

e M

exic

an s

tate

of M

eacutexic

o A

cron

yms

are

expl

aine

d in

the

met

hods

The

bas

ic s

tatis

tics

incl

ude

stan

dard

dev

iatio

n (s

) and

num

ber

of s

peci

men

s ex

amin

ed (n

)

TL

TV

HF

LE

GL

SZ

BC

BIC

BR

LN

LM

TR

BZ

PB

MF

RE

ITH

RO

DO

NT

OM

YSC

HR

YSO

PSIS

CH

RYS

OPS

IS

xndash17

93

100

220

64

180

924

34

131

511

43

337

39

883

365

237

139

Min

imum

157

8119

1523

68

118

111

22

323

361

846

335

204

113

Max

imum

179

311

222

2124

78

131

511

73

353

409

949

392

319

163

s13

53

927

081

187

035

035

018

011

014

027

016

031

012

n10

1011

1111

1111

1111

1111

1111

RE

ITH

RO

DO

NT

OM

YSF

ULV

ESC

EN

SM

UST

ELI

NU

S

xndash17

233

103

1815

521

41

104

210

23

283

697

333

231

871

28M

inim

um15

795

1615

201

910

42

987

321

352

707

312

18

126

Max

imum

189

111

2016

222

810

99

106

83

333

847

853

331

921

31s

160

411

31

20

711

090

560

430

060

160

450

110

060

03n

32

32

33

33

33

33

3R

EIT

HR

OD

ON

TO

MYS

FU

LVE

SCE

NS

TO

LTE

CU

S

xndash18

146

101

520

31

156

223

02

116

210

84

347

426

83

35

218

131

Min

imum

171

8619

1322

44

113

810

31

322

403

796

339

193

117

Max

imum

199

112

2217

241

511

86

113

93

684

268

893

722

461

43s

885

703

075

126

055

016

024

014

022

03

008

016

009

n13

1213

1313

1313

1312

1313

1313

RE

ITH

RO

DO

NT

OM

YSM

EG

ALO

TIS

SAT

UR

ATU

S

xndash15

297

793

618

15

140

922

04

110

910

21

318

376

79

362

235

101

Min

imum

131

6716

1020

81

104

89

622

933

377

313

352

080

69M

axim

um17

089

2018

233

111

810

74

349

428

873

362

235

126

s9

955

021

194

049

029

021

013

376

029

012

015

013

n33

3333

3233

3233

3333

3332

3333

RE

ITH

RO

DO

NT

OM

YSSU

MIC

HR

AST

ISU

MIC

HR

AST

I

xndash16

991

86

188

615

43

227

911

48

107

63

363

848

273

362

211

28M

inim

um16

066

1810

220

210

910

35

298

362

798

308

21

16M

axim

um18

110

221

1923

73

119

811

03

383

402

827

359

248

128

s6

5910

17

095

21

046

032

024

022

012

027

015

015

011

n14

1414

1415

1514

1515

1515

1515

RE

ITH

RO

DO

NT

OM

YSM

ICR

OD

ON

WA

GN

ER

I

xndash18

811

55

1818

221

511

63

111

53

673

847

963

051

451

61M

inim

um18

511

218

1821

77

115

410

89

351

384

765

305

137

158

Max

imum

191

119

1818

225

117

211

48

379

384

819

306

152

163

s4

244

950

00

370

130

30

140

280

011

450

03n

22

22

32

33

13

23

3



Merriam 1900






































(Saussure 1861)











DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued





Merriam 1900






































(Saussure 1861)











DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued




























(Saussure 1861)











DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued





DISCUSSION













CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued









CV

2 (

212

)

CV 1 (639)


ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued




ACKNOWLEDGMENTS


LITERATURE CITED



































APPENDIX Continued





















APPENDIX Continued



review of the harvest mice (genus reithrodontomys) in the mexican state of mÉxico

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