Journal of Mammalogy, 87(4):683–689, 2006

INTRASPECIFIC VARIATION IN PEROMYSCUS ZARHYNCHUS

(RODENTIA: MURIDAE) FROM CHIAPAS, MEXICO
CONSUELO LORENZO,* LUCERO CUAUTLE, EDUARDO ESPINOZA, AND MARICELA GARCÍA
El Colegio de la Frontera Sur, Departamento de Ecologı́a y Sistemática Terrestre, Carretera
Panamericana y Periférico Sur s/n. C. P. 29290, San Cristóbal de Las Casas, Chiapas, México

In this study we provide data on morphological and morphometric variation in specimens of Peromyscus zarhynchus
from 6 different geographically isolated localities in Chiapas, Mexico. We found significant differences in the mean
greatest skull length in all specimens examined of P. zarhynchus, but we did not find differences in other cranial and
external variables related to age, sex, and individual variation. The individuals examined from different populations
of P. zarhynchus showed distinctive morphometric characters correlated with their distribution, and probably to
feeding changes related to different types of vegetation and weather in Chiapas.

Key words: Chiapas, Mexico, morphological variation, morphometric variation, Muridae, Peromyscus zarhynchus,
Rodentia

The Chiapan deer mouse (Peromyscus zarhynchus) is an
endemic and monotypic species from Mexico, and it is a
species of special protection according to the current Mexican
Official Norm (NOM-059-ECOL-2001, Secretarı́a de Medio
Ambiente y Recursos Naturales 2001, available at http://www.
ine.gob.mx/ueajei/norma59a.html). The distribution range of
the species is located between 1,500 and 2,900 m in elevation
and is restricted to the northern mountains of Chiapas and
Lagunas de Montebello National Park in the eastern part of the
state of Chiapas, Mexico (Horváth and Navarrete 1997; Retana
and Lorenzo 2002). This mouse belongs with 13 other species
in the mexicanus species group in the family Muridae (Rogers
and Engstrom 1992). P. zarhynchus is associated with cloud
forest; pine forest with Pinus, Quercus, and Liquidambar;
secondary forest of these 3 types; and deforested areas (Hall
1981; Horváth and Navarrete 1997; Huckaby 1980).
To know the limits between species in the mexicanus species
group of genus Peromyscus, multivariate techniques were
applied and showed that there are remarkable differences in
greatest skull lengths of P. zarhynchus, P. guatemalensis, and P.
mexicanus, which allows us to tell them apart (Huckaby 1980).
Based on allozymic variation in 28 loci, genetic distances
between P. zarhynchus and other species of the mexicanus
group are low (Rogers and Engstrom 1992). Distance values
(Roger’s distance) range from 0.10 between P. mexicanus
and P. zarhynchus to 0.31 between P. melanocarpus and
* Correspondent: clorenzo@sclc.ecosur.mx
Ó 2006 American Society of Mammalogists
www.mammalogy.org
683
P. zarhynchus (Rogers and Engstrom 1992). Similarly, other
studies of allozymic variation in 18 loci show that distance
values range from 0.07 between P. zarhynchus from Huitepec
and P. mexicanus to 0.33 between P. zarhynchus from Cerro
Tzontehuitz and P. guatemalensis (Garcı́a 2001). On the other
hand, genetic differentiation was observed in 22 loci in separate
populations of P. zarhynchus from Huitepec Ecological
Reserve, Cerro Tzontehuitz, Coapilla, and Lagunas de Mon-
tebello National Park, and particularly in the last 2 localities,
which constitute the extremes of the species’ known dis-
tribution range (Robles 2003). The most genetically similar
populations of P. zarhynchus were those from the closest
localities of Huitepec Ecological Reserve and Cerro Tzonte-
huitz (12.5 km in straight line), with a Roger’s distance value
of 0.06. In contrast, the most genetically dissimilar populations
were those from Coapilla and Lagunas de Montebello National
Park, which are more geographically separated (188 km
in straight line), with a Roger’s distance of 0.09 (Robles 2003).
There are no fossil records of P. zarhynchus that would
allow us to establish its origin in geologic time with accuracy.
No one has reported morphological variation in P. zarhynchus,
but specimens from the type locality (Tumbala, Chiapas)
appear slightly darker in color than specimens from other
localities, perhaps because of moister habitats in the former
(McClellan and Rogers 1997). In addition, adult specimens
from the vicinity of Rayon, Chiapas, have a pronounced
ochraceous-tawny ventral color. No sexual dimorphism in
either external or cranial measurements has been reported in
P. zarhynchus (McClellan and Rogers 1997).
We analyzed the morphological and morphometric variation
of P. zarhynchus from several geographically separated locali-
ties in Chiapas. Our objectives were to know the variation
684 JOURNAL OF MAMMALOGY
FIG. 1.—Populations of Peromyscus zarhynchus in Chiapas,
México: 1, Rayon; 2, Coapilla; 3, Tzontehuitz; 4, Oxchuc; 5, Huitepec
Ecological Reserve; 6, Lagunas de Montebello National Park. Letters
indicate regions: A ¼ mountains of northern Chiapas; B ¼ Central
Plateau of Chiapas.
patterns in external traits of the specimens examined and to
assess the variation patterns in cranial size and shape.
MATERIALS AND METHODS
Museum specimens (n ¼ 190; 66 males and 124 females) of
P. zarhynchus were examined from 6 geographic localities (Fig. 1),
in 4 mammal collections in Mexico (Appendix I).
The age of specimens was estimated on the basis of tooth wear and
eruption patterns as defined by Hoffmeister (1951). Accordingly, each
specimen was assigned to 1 of the following age classes: juveniles
(age class 1), subadults (age class 2), and adults (age class 3). Four
conventional external measurements were obtained from the labels of
190 specimens: total length, length of tail, length of hind foot, and
length of ear.
Morphological variables.—Eleven qualitative morphological vari-
ables were recorded for 46 individuals (32 males and 14 females) from
5 localities in Chiapas (Fig. 1; Appendix I). We measured the
following variables in accordance with our observations of specimens
of P. zarhynchus: 1, length of dorsal fur (short or long); 2, scales of the
tail (marked or not marked); 3, color of hind feet (white or dark);
4, reddish spot on the chest (absent or present); 5, color of tail (spotted
or unspotted); 6, ventral scales of the tail (marked or not marked);
7, frontal bone (flat or not flat); 8, frontal process (continuous or
discontinuous); 9, palatine bone (long or short); 10, posterior border of
the palatine (extending in cusps or rounded); 11, palatine foramen
(extending through maxillary toothrows or not). Variables 6–11 were
observed with a Zeiss stereoscopic microscope (Stemi DV4, Thron-
wood, New York). All qualitative morphological variables were
recorded binomially (0 or 1) and considered as categorical explanatory
variables for the construction of a classification tree using the software
(S_Plus; Mathsoft, Inc. 2000, available at http://www.mirror.ac.uk/
sites/lib.stat.cmu.edu).
Vol. 87, No. 4
Morphometric variables.—Localities are shown in Fig. 1, and
number of specimens measured are listed in Appendix I. Twenty-
seven skull measurements (Robinson and Dippenaar 1987; Williams
and Ramı́rez-Pulido 1984) were taken to the nearest 0.1 mm with
a dial caliper: 1, greatest skull length (GLS); 2, condylobasal length
(CBL); 3, skull height (SKH); 4, bullar length (BUL); 5, shield-bullae
depth (SBD); 6, diastema length (DIL); 7, rostral height (ROH); 8,
palatine bridge length (PBL); 9, postpalatal length (POL); 10, occipital
length (OCL); 11, rostral breadth (BRR); 12, maxillary toothrow
length (MTL); 13, maxillary toothrow breadth (MTB); 14, superior
molar breadth (SMB); 15, postdental breadth (PDB); 16, zygomatic
breadth (ZYB); 17, braincase breadth (BAB); 18, nasal length (NAL);
19, anterior frontal breadth (ANB); 20, eye socket length (ESL);
21, eye socket breadth (ESB); 22, nasal breadth (NAB); 23, mandible
height (MAH); 24, dentary height (DEH); 25, mandibular ramus
height (MRH); 26, mandible length (MAL), and 27, mandibular
toothrow length (MDL). Statistical analyses were performed using
Statistical Analysis System (SAS version 8.2—SAS Institute Inc.
1999–2001), SPSS (SPSS Inc. 1998), and JMP_IN (SAS Institute Inc.
1997) software.
Variation in cranial size was estimated 1st by comparing means of
the greatest skull length (GLS) of 181 specimens from all localities
with a single-classification analysis of variance (ANOVA).
Nongeographic variation.—One hundred ninety individuals from
Chiapas, Mexico (Fig. 1) were analyzed to determine variation be-
tween age classes, between sexes, and between individuals. Age
variation was evaluated comparing the 3 age classes for each external
and skull measurement with a single-classification analysis of variance
(F; ANOVA), considering males and females separately. When the
difference between at least 2 age classes was significant (P
0.05),
Duncan’s multiple range test was used to determine maximally non-
significant subsets of age classes (Engstrom et al. 1987).
Sex variation was evaluated comparing all variables for males and
females of each age classes using the nonparametric Wilcoxon test (Z;
P
0.05). Individual variation was evaluated comparing the coef-
ficient of variation (CV) of each variable between age classes and
between sexes. This statistic has been used to evaluate phenotypic
variation of populations and assumes that the measurement error is
constant for all variables comparative.
Geographic variation.—For each morphometric variable in each
grouped locality, standard statistics were calculated. External measure-
ments were excluded from further geographic analysis because an
unknown proportion of variance of these characters was associated
with differences in measuring techniques among preparators. A prin-
cipal component analysis was used to obtain principal axes that
summarize the directions of greatest variation among all samples.
Canonical variates analysis was used to order the populations of P.
zarhynchus along the axes of maximum differentiation by maximizing
between-population variation with respect to within-population varia-
tion. A discriminant function analysis was performed to allocate each
individual to the closest locality, regardless of its original locality.
RESULTS
Morphological analyses.— The 46 individuals of P. zarhyn-
chus examined showed variation in 5 qualitative morphological
variables (color of tail, length of dorsal fur, color of hind feet,
scales of the tail, and posterior palatine border), observed in
the classification tree (Fig. 2). The misclassification error rate
was 0 for all individuals examined (n ¼ 46). The 16 specimens
from Huitepec Ecological Reserve stand out and were con-
August 2006 LORENZO ET AL.—VARIATION IN PEROMYSCUS ZARHYNCHUS 685

TABLE 1.—Sex variation for the cranial variables of Peromyscus

zarhynchus that showed significant differences between females and
males in the only 2 localities in Chiapas that showed sex variation,
using the nonparametric Wilcoxon test (Z; P
0.05).

Locality Variable Z d.f. P

Coapilla GLS 2.77 28 0.011
SKH 2.42 28 0.023
SBD 2.23 28 0.035
BAB 2.22 25 0.035
NAL 2.57 27 0.016
MAL 2.77 28 0.010
Tzontehuitz BUL 2.47 95 0.015
PDB 2.09 95 0.039
BAB 2.02 95 0.046
MRH 2.21 93 0.029

ANOVA were 3 variables in Rayon, 13 variables in Coapilla,

FIG. 2.—Phenogram illustrating the phenetic relationships among
5 populations of Peromyscus zarhynchus in Chiapas, México, using
morphological variables considered as categorical explanatory varia-
bles (using the software S_Plus). Individuals from Lagunas de
Montebello National Park and Coapilla showed great morphologic
variation and were not clumped together.
sidered separate by the variable color of the tail (Fig. 2). For the
rest of the variables, length of dorsal fur is the most important
variable explaining variation in morphological levels, followed
by color of hind feet, scales of the tail, and posterior palatine
border. Specimens from Huitepec Ecological Reserve were the
most different of the groups from all localities, and the
individuals from Oxchuc and Cerro Tzontehuitz were clumped
together. Specimens from Lagunas de Montebello National
Park and Coapilla had great morphologic variation and were
not clumped together (Fig. 2).
Morphometric analyses.— The size pattern in greatest length
of skull (GLS) from smallest to largest (in mm; mean 6 SD) in
181 individuals considered as a single sample was Coapilla
(35.3 6 1.70), Rayon (35.4 6 1.72), Cerro Tzontehuitz
(35.8 6 0.99), Huitepec Ecological Reserve (36.2 6 1.04),
Oxchuc (36.3 6 1.13), and Lagunas de Montebello National
Park (37.6 6 1.71). Specimens from Lagunas de Montebello
National Park were on average larger in GLS than specimens
from Coapilla (by 2.33 mm, or 3.19%). Differences between
localities were statistically significant with ANOVA (F ¼ 9.87,
d.f. ¼ 180, P , 0.001).
Nongeographic variation.— One hundred ninety individuals
were used to estimate within-sample variation of the 27 mea-
sured characters. In analysis of age variation, age class 1 was
not included because only 1 individual was reported from each
of the following localities: Cerro Tzontehuitz, Oxchuc, and
Huitepec Ecological Reserve.
Statistically significant differences in cranial variables between
age classes 2 and 3 by locality according to a single-classification
15 variables in Cerro Tzontehuitz, 5 variables in Oxchuc, and
8 variables in Huitepec Ecological Reserve (Appendix II).
Sex variation.— No significant differences were found
between females and males in the majority of cranial variables.
Exceptions were 6 variables for Coapilla and 4 variables for
Cerro Tzontehuitz (Table 1).
Individual variation.— For individual variation, the coeffi-
cient of variation was less than 10% in almost all variables for
all localities. The exceptions were BUL (25.99%) and NAB
(10.60%) for Rayon; length of tail (11.73%) and length of ear
(16.58%) for Coapilla; BRR (13.73%) and NAB (10.17%) for
Cerro Tzontehuitz; MTL (10.35%), SMB (10.35%), and NAB
(12.52%) for Huitepec Ecological Reserve; and length of ear
(10.80%), ESB (11.51%), and NAB (14.69%) for Lagunas de
Montebello National Park.
Geographic variation.— Because of the low levels of
variation recorded, skull measurements of all male and female
specimens of age classes 2 and 3 of each locality were pooled
for multivariate analyses. Using principal component analysis,
the first 3 factors explained 53.9% of the total variation be-
tween the 27 cranial variables for the specimens of the 6 local-
ities. Component 1 explained 37.2% of the total intraspecific
variation, component 2 explained 10.3% of the total variation,
and component 3 explained 6.4%, with a cumulative proportion
of 53.9% in these first 3 components that explained the
difference between localities. All factor scores in component
1 showed similar and positive values (except NAB with nega-
tive value) that can be assigned to a size factor. Variables that
contributed greatly to this separation were GLS, DIL, PBL, and
ZYB. Components 2 and 3 showed characters with positive and
negative values. This condition may be attributed to a shape
factor (Table 2). Distances from individuals of each locality
to its centroid were calculated to allocate each individual to
the closest locality, regardless of its original locality. The
probability for group classification showed 100% correct
classification for individuals of each locality.
The canonical variates analysis ordered the populations of
P. zarhynchus along the axes of maximum differentiation by
maximizing between-population variation with respect to
within-populations variation. Individuals of different popula-
686 JOURNAL OF MAMMALOGY Vol. 87, No. 4

TABLE 2.—First 3 principal components of Peromyscus zarhynchus

in the ages classes 2 and 3, including males and females, and based on
27 cranial variables. Asterisks (*) indicate variables that explained the
greatest percentage of variation.

Variable Component 1 Component 2 Component 3

Greatest skull length (GLS) 0.289* 0.002 0.128
Condylobasal length (CBL) 0.166 0.033 0.027
Skull height (SKH) 0.112 0.478* 0.018
Bullar length (BUL) 0.083 0.196 0.252
Shieldbullae depth (SBD) 0.190 0.190 0.145
Diastema length (DIL) 0.270* 0.014 0.154
Rostral height (ROH) 0.219 0.097 0.081
Palatine bridge length (PBL) 0.267* 0.046 0.050
Postpalatal length (POL) 0.257 0.176 0.151
Occipital length (OCL) 0.227 0.071 0.042
Rostral breadth (BRR) 0.189 0.184 0.072
Maxillary toothrow length (MTL) 0.105 0.049 0.399*
Maxillary toothrow breadth
(MTB) 0.215 0.047 0.194
Superior molar breadth (SMB) 0.125 0.202 0.243
Postdental breadth (PDB) 0.026 0.114 0.436*
Zygomatic breadth (ZYB) 0.267 0.020 0.034
Braincase breadth (BAB) 0.192 0.022 0.104
Nasal length (NAL) 0.115 0.150 0.011 FIG. 3.—Canonical analysis for males and females of Peromyscus
Anterior frontal breadth (ANB) 0.153 0.088 0.125 zarhynchus of age classes 2 and 3, for 6 localities in Chiapas, México:
Eye socket length (ESL) 0.211 0.119 0.139 1, Rayon; 2, Coapilla; 3, Cerro Tzontehuitz; 4, Oxchuc; 5, Huitepec
Eye socket breadth (ESB) 0.109 0.400* 0.019 Ecological Reserve; 6, Lagunas de Montebello National Park.
Nasal breadth (NAB) 0.007 0.440* 0.122
Mandible height (MAH) 0.187 0.182 0.028 Nongeographic variation.— Variability due to age and sex in
Dentary height (DEH) 0.243 0.002 0.169
our data was small, although it was greatest in specimens of
Mandibular ramus height (MRH) 0.164 0.210 0.277
Mandible length (MAL) 0.204 0.040 0.186 Coapilla and Cerro Tzontehuitz. No significant variation was
Mandibular toothrow length found among individuals; therefore, no overall sexual dimor-
(MDL) 0.100 0.028 0.391* phism or individual variation was considered. However, sig-
nificant differences were observed in the greatest skull length
of individuals from all localities.
tions were discriminated by the 1st factor, where SKH (0.7304),
Geographic variation.— Multivariate analyses indicated that
NAB (0.6470), and ESB (0.5428) were important characters,
cranial size is the most important factor (37% of total variation)
which contributed to separating the pair Rayon and Cerro
contributing to morphometric differentiation among individuals
Tzontehuitz from the other localities (Fig. 3). The 2nd vector
from different populations of P. zarhynchus. Specimens from
was strongly influenced by SBD (0.6322), SMB (0.3251),
Lagunas de Montebello National Park were the largest indi-
and NAL (0.2906).
viduals, whereas individuals from Coapilla were the smallest.
Quadratic generalized distances between the individuals’
The variables explaining most of this variation were greatest
centroids were computed as part of the analysis to assess their
skull length (GLS), diastema length (DIL), and palatine bridge
affinities. The greatest distance (20.23) was recorded between
length (PBL). Diastema length and palatine bridge length are
individuals of Rayon and Lagunas de Montebello National
directly related to feeding habits of rodents because these
Park, whereas the shortest distance (0.64) was found between
variables are based on osseous structures important for mas-
individuals of Rayon and Cerro Tzontehuitz (Table 3).
tication. Changes in length of these variables could be related
The phenogram based on average taxonomic distances
to changes in the greatest skull length. The shape factor also
computed from the morphometric characters includes 2 well-
contributed in smaller proportion to intraspecific morphometric
delineated clusters (Fig. 4). The specimens from Rayon, Cerro
differences among individuals with no more than 17% of total
Tzontehuitz, and Coapilla form a cluster at 4.52, with Rayon and
variation.
Cerro Tzontehuitz the most similar localities (1.75). Oxchuc,
Maximization of multidimensional distances among individ-
Huitepec Ecological Reserve, and Lagunas de Montebello
uals produced by the discriminant analysis clearly separated
National Park were grouped together in a cluster at 4.10.
the populations of Rayon and Lagunas de Montebello National
Park from one another. The shortest distances were between
DISCUSSION Rayon and Cerro Tzontehuitz, and individuals from both lo-
Morphological analyses.— Variation in morphological var- calities showed a distinctive spatial position from other locali-
iables among populations of P. zarhynchus was evident, and ties in the plot of canonical variables.
variables that explained this variation were tail color and length The discriminatory power of the analysis was excellent
of dorsal fur. because 100% of the specimens were correctly classified to
August 2006 LORENZO ET AL.—VARIATION IN PEROMYSCUS ZARHYNCHUS
TABLE 3.—Quadratic generalized distances between individuals of
age classes 2 and 3, including males and females, of Peromyscus
zarhynchus from 6 localities from Chiapas using 27 cranial variables.
Locality
Locality Rayon Coapilla Tzontehuitz Oxchuc Huitepec Montebello
Rayon 0
Coapilla 12.22 0
Tzontehuitz 0.64 11.77 0
Oxchuc 11.97 2.96 12.49 0
Huitepec 15.39 3.69 15.37 1.19 0
Montebello 20.23 5.69 18.62 4.73 4.08 0 9 cranial variables that explain the greatest percentage of variation in
localities. The resulting cluster analysis describes the morpho-
metric relationships among the 6 populations of P. zarhynchus.
The individuals of Rayon and Cerro Tzontehuitz are morpho-
metrically more similar to each other than to any other popu-
lation, with a slight separation between these populations and
Coapilla (population with the smallest individuals). The popu-
lations of Oxchuc and Huitepec Ecological Reserve are
morphometrically closely similar and clumped together with
Lagunas de Montebello National Park (population with the
largest individuals; Fig. 4).
Individual variation in cranial size and slight differences
in cranial shape produce the morphometric distances among
populations of P. zarhynchus, showing a trend in the pattern
of cranial size where the individuals from eastern Chiapas are
larger than individuals of northwestern Chiapas. It is probable
that these cranial differences are influenced by variation in
vegetation types and weather in Chiapas, representing different
environmental conditions in the 2 areas where P. zarhynchus
occurred. Populations of southeastern Chiapas, located in the
Central Plateau region, are associated with cloud forest; sec-
ondary forest; and pine forest with Pinus, Quercus, and
Liquidambar; and with semiwarm, humid weather with rains
in summer, and cooler (mean annual temperature of 188C) areas
than the north mountains region, where individuals are
associated with wet, highland forests with subtropical, very
humid weather and a mean annual temperature of 228C (Instituto
Nacional de Estadı́stica, Geografı́a e Informática 1982).
In summary, examination of our data suggests that within
populations, individual variation of P. zarhynchus makes it dif-
ficult to separate individuals by locality; that cranial morpho-
metric differentiation among populations of P. zarhynchus is
mainly due to variation in cranial size and shape; and that the
morphological discontinuity among geographic populations of
P. zarhynchus is associated with changes in skull size and shape.
Our results imply that morphological variation and skull size
and shape differences among populations of P. zarhynchus
could be the result of local adaptation and geographic isolation
due to habitat fragmentation.
Future efforts must include more regular and uniform
geographic sampling of populations of P. zarhynchus, par-
ticularly in the area between the northern and southern popu-
lations in the regions of Coapilla and Lagunas de Montebello
National Park, where individuals displayed remarkable varia-
tion. This information coupled with detailed descriptions of
687
FIG. 4.—Phenogram illustrating phenetic relationships among 6
populations of Peromyscus zarhynchus in Chiapas, México, based on
the principal component analysis. The phenogram was computed with
the unweighted pair-group method with arithmetic averages (UPGMA).
morphological, chromosomal, and molecular variation at local
and geographic scales should shed light on the patterns and
processes of differentiation in this species.
RESUMEN
En este estudio proporcionamos datos sobre la variación
morfológica y morfométrica en Peromyscus zarhynchus de 6
diferentes localidades geográficamente aisladas en Chiapas.
Existen diferencias estadı́sticamente significativas en el prom-
edio de la longitud mayor del cráneo en todos los ejemplares
examinados de P. zarhynchus, pero no existen diferencias en el
resto de las variables craneales y externas, con respecto a la
edad, sexo y variación individual. Los ejemplares examinados
de las diferentes poblaciones de P. zarhynchus muestran carac-
teres morfométricos distintivos relacionados con su distrib-
ución, y probablemente con cambios en hábitos alimentarios de
acuerdo a los diferentes tipos de vegetación y clima presentes
en Chiapas.
ACKNOWLEDGMENTS
F. A. Cervantes from Colección Nacional de Mamı́feros, Instituto
de Biologı́a, Universidad Nacional Autónoma de México; G. Rivera
from the Museo de Zoologı́a of the Universidad de Ciencias y Artes de
Chiapas; and E. Morales and A. Riechers from the Museo de Zoologı́a
of the Instituto de Historia Natural y Ecologı́a were kind enough to
permit us to examine the specimens of P. zarhynchus. A. Morón, E.
Naranjo, and F. A. Cervantes provided valuable comments on several
drafts of the manuscript. A. Flamenco prepared Fig. 1, and D. Dı́az
improved the quality of all figures. R. Palacios gave assistance with
the computer analysis of morphological variables. F. Barragán, J.
Bolaños, and P. Robles assisted in the laboratory work.
LITERATURE CITED
ENGSTROM, M. D., H. H. GENOWAYS, AND P. K. TUCKER. 1987.
Morphological variation, karyology, and systematic relationships of
Heteromys gaumeri (Rodenita: Heteromyidae). Pp. 289–303 in
Studies in neotropical mammalogy: essays in honor of Philip
Hershkovitz (B. D. Patterson and R. M. Timm, eds.). Fieldiana:
Zoology 39:i–vii þ 1–506.
GARCÍA, B. M. 2001. Diferenciación genética en tres especies de
Peromyscus del grupo mexicanus (Rodentia:Muridae) en Chiapas.
B.S. thesis, Instituto Tecnológico Agropecuario de Hidalgo,
Hidalgo, México.
688 JOURNAL OF MAMMALOGY
HALL, E. R. 1981. The mammals of North America. Vol. 2. 2nd ed.
John Wiley & Sons, Inc., New York.
HOFFMEISTER, D. F. 1951. A taxonomic and evolutionary study of the
pinion mouse, Peromyscus truei. Illinois Biological Monograph
21:1–104.
HORVÁTH, A., AND D. A. NAVARRETE. 1997. Ampliación del área de
distribución de Peromyscus zarhynchus Merriam, 1898 (Rodentia:
Muridae). Revista Mexicana de Mastozoologı́a 2:122–125.
HUCKABY, D. G. 1980. Species limits in the Peromyscus mexicanus
group (Mammalia: Rodentia: Muroidea). Contributions in Science,
Natural History Museum of Los Angeles County 326:1–24.
INSTITUTO NACIONAL DE ESTADÍSTICA, GEOGRAFÍA E INFORMÁTICA. 1982.
Cartas de climas, escala 1:1,000,000. Cartas temáticas de efectos
climáticos, escala 1:250,000. Procesado por el Laboratorio de
Información Geográfica de El Colegio de la Frontera Sur, Distrito
Federal, México.
MATHSOFT, INC. 2000. S_Plus. Mathsoft, Inc., Chicago, Illinois.
MCCLELLAN, D. A., AND D. S. ROGERS. 1997. Peromyscus zarhynchus.
Mammalian Species 562:1–3.
RETANA, O., AND C. LORENZO. 2002. Lista de los mamı́feros terrestres
de Chiapas: endemismos y estado de conservación. Acta Zoológica
Mexicana 85:25–49.
ROBINSON, T. J., AND N. J. DIPPENAAR. 1987. Morphometrics of the
South African Leporidae. II: Lepus Linnaeus, 1758, and Bunolagus
Thomas, 1929. Annals Transvaal Museum 34:379–404.
ROBLES, Z. P. 2003. Variación genética en poblaciones aisladas de
Peromyscus zarhynchus (Rodentia: Muridae) en Chiapas, México.
B.S. thesis, Universidad de Ciencias y Artes de Chiapas, México.
ROGERS, D. S., AND M. D. ENGSTROM. 1992. Evolutionary implications
of allozymic variation in tropical Peromyscus of the mexicanus
species group. Journal of Mammalogy 73:55–69.
SAS INSTITUTE INC. 1997. JMP start statistics, a guide to statistics and
data analysis using JMP and JMP IN software. Version 3.2.2. SAS
Institute Inc., Cary, North Carolina.
SAS INSTITUTE INC. 1999–2001. SAS version 8.2. SAS Institute Inc.,
Cary, North Carolina.
SECRETARÍA DE MEDIO AMBIENTE Y RECURSOS NATURALES. 2001. Norma
Oficial Mexicana: NOM-059-ECOL-2001. Protección ambiental—
especies nativas de México de flora y fauna silvestres—categorı́as
de riesgo y especificaciones para su inclusión, exclusión o
cambio—lista de especies en riesgo. Diario Oficial de la Federación,
Distrito Federale, Mexico.
SPSS INC. 1998. SPSS. Version 8.0 for Windows. Prentice-Hall, Inc.,
Englewood Cliffs, New Jersey.
WILLIAMS, S. L., AND J. RAMÍREZ-PULIDO 1984. Morphometric
variation in the volcano mouse, Peromyscus (Neotomodon) alstoni
(Mammalia, Cricetidae). Annals of Carnegie Museum 53:163–183.
Submitted 11 July 2005. Accepted 19 December 2005.
Associate Editor was Eric A. Rickart.
APPENDIX I
Morphometric analyses.—The 190 specimens examined are listed
below by locality and museum acronym. ECO-SC-M ¼ Colección
Mastozoológica de El Colegio de La Frontera Sur: México, Chiapas.
Cerro La Ventana, km 16 road Coapilla–Tapalapa, Municipio de
Coapilla, 2,200 m (ECO-SC-M 159, 1246, 1247, 1248, 1249, 1250,
Vol. 87, No. 4
1252, 1253, 1254, 1444, 1445, 1446, 1447, 1448); México, Chiapas.
Santo Tomas Oxchuc, Mercado Indı́gena Municipio Oxchuc, 2,000 m
(ECO-SC-M 811, 812, 813, 814, 1026, 1028, 1037); México, Chiapas.
Huitepec Ecological Reserve, 2 km (by road) NE San Juan Chamula,
Municipio San Cristóbal de Las Casas, 2,340 m (ECO-SC-M 529,
530, 531, 532, 581, 582, 583, 586, 591, 592, 595, 596, 599, 600, 601,
602, 603, 606, 607, 608, 609); México, Chiapas. Las Grutas, Lagunas
de Montebello Nacional Park, 3.5 km N El Vivero, Municipio La
Independencia 1,513 m (ECO-SC-M 349, 350, 351, 352, 353, 465,
466, 501, 1215, 1216, 1217, 1218, 1219, 1221, 1227, 1424, 1425,
1426, 1427, 1428); México, Chiapas. Cerro Tzontehuitz, 12.5 km SW
de San Cristóbal de Las Casas, Municipio San Juan Chamula (ECO-
SC-M 695, 696, 1438, 1439). CZRMA ¼ Colección Zoológica
Regional del Instituto de Historia Natural y Ecologı́a: México,
Chiapas. Huitepec Ecological Reserve, 2 km (by road) NE San Juan
Chamula, Municipio San Cristóbal de Las Casas, 2,340 m (CZRMA
1850). MZUNI ¼ Colección de Mamı́feros de la Escuela de Biologı́a
de la Universidad de Ciencias y Artes: México, Chiapas, Cerro La
Ventana, km 16 road Coapilla–Tapalapa, Municipio de Coapilla
(MZUNI 742, 743, 745, 746, 747, 748, 749, 750, 1011, 1014, 1015,
1016, 1018, 1019, 1188); México, Chiapas. Huitepec Ecological
Reserve, 2 km (by road) NE San Juan Chamula, Municipio San
Cristóbal de Las Casas, 2,340 m (MZUNI 276, 277, 284). CNMA ¼
Colección Nacional de Mamı́feros del Instituto de Biologı́a de la
UNAM: México, Chiapas. 5 km al E de Rayon, Municipio de Rayon
(CNMA 31592, 31593, 31594, 31597, 31598, 31629, 33040, 33041);
México, Chiapas. Cerro Tzontehuitz 13 km al NE de San Cristóbal de
Las Casas, Municipio San Juan Chamula (CNMA 35372, 35373,
35374, 35375, 35376, 35377, 35378, 35379, 35380, 35381, 35382,
35383, 35384, 35385, 35386, 35387, 35388, 35389, 35390, 35391,
35392, 35393, 35394, 35395, 35396, 35397, 35398, 35399, 35400,
35401, 35402, 35403, 35404, 35405, 35406, 35407, 35408, 35409,
35410, 35411, 35412, 35413, 35414, 35415, 35416, 35417, 35418,
35419, 35420, 35421, 35422, 35423, 35424, 35425, 35426, 35427,
35428, 35429, 35430, 35431, 35432, 35433, 35434, 35435, 35436,
35437, 35438, 35439, 35440, 35441, 35442, 35443, 35444, 35445,
35446, 35447, 35448, 35449, 35450, 35451, 35452, 35453, 35454,
35455, 35456, 35457, 35458, 35459, 35460, 35461, 35462, 35463,
35547, 35548, 35549); México, Chiapas. Huitepec Ecological
Reserve, 5 km AL W de San Cristóbal de Las Casas (CNMA 30699).
Morphological analyses.—The 46 specimens examined are listed
below by locality and museum acronym. ECO-SC-M ¼ Colección
Mastozoológica de El Colegio de La Frontera Sur: México, Chiapas,
Cerro La Ventana, km 16 road Coapilla–Tapalapa, Municipio de
Coapilla, 2,200 m (ECO-SC-M 159, 1444, 1446, 1447, 1448);
México, Chiapas. Santo Tomas Oxchuc, Mercado Indı́gena Municipio
Oxchuc, 2,000 m (ECO-SC-M 806, 811, 813, 814, 1028); México,
Chiapas. Huitepec Ecological Reserve, 2 km (by road) NE San Juan
Chamula, Municipio San Cristóbal de Las Casas, 2,340 m (ECO-SC-
M 582, 583, 586, 591, 592, 595, 596, 600, 601, 602, 603, 606, 607,
608, 609); México, Chiapas. Las Grutas, Lagunas de Montebello
Nacional Park, 3.5 km N El Vivero, Municipio La Independencia
1,513 m (ECO-SC-M 349, 350, 351, 352, 353, 501, 1215, 1216, 1217,
1218, 1219, 1221, 1227, 1424, 1426, 1427, 1428); México, Chiapas.
Cerro Tzontehuitz, 12.5 km SW de San Cristóbal de Las Casas,
Municipio San Juan Chamula (ECO-SC-M 695, 696, 1438, 1439).
August 2006 LORENZO ET AL.—VARIATION IN PEROMYSCUS ZARHYNCHUS 689

APPENDIX II
Statistically significant differences in cranial variables between age classes 2 and 3 by locality in Peromyscus zarhynchus in Chiapas, according
to a single-classification analysis of variance (F-statistics, P ¼ levels of significance, d.f. ¼ degrees of freedom).

Huitepec Ecological
Rayon Coapilla Cerro Tzontehuitz Oxchuc Reserve
Variable F d.f. P F d.f. P F d.f. P F d.f. P F d.f. P
BAB 8.31 96 0.0040
BRR 7.03 8 0.0320 11.96 7 0.0130
CBL 11.38 24 0.0020 29.68 96 ,0.0001
DEH 13.73 28 0.0010 19.13 93 ,0.0001 6.70 7 0.04100
DIL 13.21 27 0.0010 29.59 95 ,0.0001 8.89 24 0.0060
ESB 8.63 28 0.0060 4.32 95 ,0.0400
ESL 8.15 8 0.0240 8.42 28 0.0070 15.85 95 ,0.0001
GLS 8.71 24 0.0070 22.54 94 ,0.0001 14.65 24 0.0009
MAH 17.6 28 0.0030
MAL 0.434 93 0.0220
MTB 20.69 28 0.0001 26.33 95 ,0.0001 7.68 7 0.0320
NAL 14.77 27 0.0007 7.44 95 0.0070 13.03 7 0.0110 7.52 24 0.0110
OCL 10.84 25 0.0030
PBL 21.00 95 ,0.0001 11.47 24 0.0020
POL 18.11 94 ,0.0001 4.74 24 0.0400
ROH 13.75 27 0.0010 18.87 95 ,0.0001 5.20 24 0.0320
SBD 4.40 95 0.0380
SKH 9.56 24 0.0050
SMB 6.94 24 0.0140
ZYB 7.10 8 0.0320 8.89 25 0.0060 16.34 96 0.0001 11.36 7 0.0150 5.65 24 0.0260