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FIELDIANA
Zoology
NEW SERIES, NO. 74
Intraspecific and Interspecific Variation in the
Cryptotis nigrescens Species Complex of
Small-Eared Shrews (Insectivora: Soricidae), with the
Description of a New Species from Colombia
Neal Woodman
Robert M. Timm
Museum of Natural History and
Department of Systematics and Ecology
University of Kansas
Lawrence, Kansas 66045-2454
Accepted April 16, 1993
Published September 30, 1993
Publication 1452
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 1993 Field Museum of Natural History
ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
Abstract 1
Resumen 1
Introduction 1
Methods 2
Sexual Variation 3
Systematic Descriptions 5
Cryptotis mayensis (Merriam, 1901) 7
Cryptotis merriami Choate, 1970 14
Cryptotis nigrescens (J. A. Allen, 1895) ... 17
Cryptotis mera Goldman, 1912 22
Cryptotis colombiana, new species 24
Acknowledgments 27
Literature Cited 28
Specimens Examined 29
1 0. Skull and mandible of C. merriami .... 16
1 1 . Map showing the distribution of C.
merriami 17
12. Plots of PC A factor scores of C. merria-
mi and C. nigrescens 18
13. Skull and mandible of C. nigrescens .... 19
14. Map showing the distribution of C. ni-
grescens 20
1 5. Skull and mandible of C. mera 22
16. Skull and mandible of C colombiana . . 24
1 7. Map showing the distribution of C.
mera and the type locality of C. colom-
biana 25
18. Tympanic region of the skulls of C.
merriami, C nigrescens, and C. colom-
biana 26
19. Plot of PCA factor scores of C. colom-
biana, C mera, and C. nigrescens 26
List of Illustrations
1. Cranial measurements used in this
study 4
2. Plot of PCA factor scores for male and
female C. nigrescens from Monteverde,
Costa Rica 8
3. Plot of PCA factor scores for male and
female C. nigrescens from San Felix,
Panama 8
4. Skull and mandible of C. mayensis .... 9
5. Map showing the distribution of C.
mayensis 10
6. Plot of C mayensis and C. merriami on
factor axes 1 and 3 from PCA of cranial
variables 11
7. Plot of C. mayensis and C merriami on
factor axes 1 and 2 from PCA of man-
dibular variables 14
8. Plot of C mayensis and C. merriami on
canonical axes 1 and 2 from discrimi-
nant function analysis of cranial vari-
ables 14
9. Plot of C. mayensis and C. merriami on
canonical axes 1 and 2 from discrimi-
nant function analysis of mandibular
variables 15
List of Tables
1 . Selected measurements of male and fe-
male C. nigrescens 6
2. Factor loadings from PCA of male and
female C. nigrescens from Monteverde,
Costa Rica 7
3. Factor loadings from PCA of male and
female C. nigrescens from San Felix,
Panama 7
4. Selected measurements of members of
the C. nigrescens group 12
5. Factor loadings from PCA of cranial
variables of C. mayensis and C. merriami 1 3
6. Factor loadings from PCA of mandibular
variables of C. mayensis and C merriami 14
7. Factor loadings from PCA of cranial
variables of C. merriami and C. nigres-
cens 17
8. Factor loadings from PCA of cranial
variables of C. colombiana, C mera, and
C. nigrescens 27
in
Back cover: Cryptotis colombiana
Intraspecific and Interspecific Variation in the
Cryptotis nigrescens Species Complex of
Small-Eared Shrews (Insectivora: Soricidae), with the
Description of a New Species from Colombia
Neal Woodman and Robert M. Timm
Abstract
The Cryptotis nigrescens species complex of small-eared shrews previously was considered
to consist of a single, wide-ranging species with three subspecies distributed from southern
Mexico to Panama. Our study of the patterns of morphological variation within this complex
indicates that it is much more diverse and speciose than earlier believed. The group includes
populations that occur only at high elevations (> 500 m) as well as the only taxon in the genus
restricted to elevations below 100 m. Based on our investigations, we recognize five species:
C. mayensis occurs on the Yucatan Peninsula and is known from an isolated locality in Guerrero;
C. merriami has a patchy distribution from Chiapas, Mexico, to northern Costa Rica; C.
nigrescens inhabits conterminous highlands in Costa Rica and Panama; C. mera is known from
two isolated mountaintops along the Panama/Colombia border; and we describe a new species
from the Central Cordillera of Colombia, which extends the known distribution of the complex
to the southeast. In addition, recent studies indicate that C. hondurensis and C. magna may
be imbedded phylogenetically within the C. nigrescens complex.
Resumen
Cryptotis nigrescens es un complejo de especies que solia considerarse como una sola especie
con tres subespecies distribuidas a lo largo de America Latina desde el sur de Mexico hasta
Panama. Nuestro estudio de patrones de variacion morfologica en este complejo indica que es
diverso y con mas especies de las que se habia pensado antes. El grupo incluye poblaciones que
habitan solamente en elevaciones por encima de los 500 m, y tambien el unico taxon en el
genero que esta restringido a elevaciones bajo 100 m. Basados en nuestras investigaciones,
reconocemos cinco especies. La primera, C. mayensis, se encuentra en la Peninsula de Yucatan
y en una localidad aislada en Guerrero, Mexico. La segunda, C. merriami, tiene una distribucion
discontinua desde Chiapas en Mexico hasta el norte de Costa Rica. La especie C. nigrescens
habita en las cordilleras de Costa Rica y Panama. Se sabe que C. mera vive en las cimas de
dos montanas aisladas en la frontera entre Panama y Colombia. La ultima es una especie nueva
que describimos en esta publication y que mora en la Cordillera Central de Colombia y que
extiende la distribucion del complejo hacia el sureste. Ademas, estudios recientes indican que
las especies C. hondurensis y C. magna pueden estar relacionadas filogeneticamente con el
complejo C. nigrescens.
Introduction America. The genus reaches its greatest diversity
in Mexico and Central America. Hall and Kelson
Small-eared shrews of the genus Cryptotis have (1959) recorded 25 species of Cryptotis in this re-
a wide distribution, occurring in parts of North gion. In his revision of the Middle American Cryp-
America, Central America, and northwestern South totis, Choate (1970) recognized eight species of
FIELDIANA: ZOOLOGY, N.S., NO. 74, SEPTEMBER 30, 1993, PP. 1-30 1
small-eared shrews in Mexico and Central Amer-
ica, and his taxonomy has been followed by most
subsequent authors, including Hall (1981), Hona-
cki et al. (1982), Corbet and Hill (1991), and Wil-
son and Reeder (1993). We (Woodman & Timm,
1 992, submitted) recognized four additional species
from Guatemala, Honduras, and Mexico, bringing
the total to 1 2 species.
One poorly understood Central American group
of Cryptotis is the C. nigrescens complex, which
was known to occur from Guerrero, Mexico,
through Central America to the Panama/Colom-
bia border. Most species of small-eared shrews
that occur from central Mexico through northern
South America are high-elevation forms, inhab-
iting regions over 500 m. The C. nigrescens com-
plex, which includes both high-elevation and low-
elevation taxa, is an exception. Choate (1970)
provided the only recent comprehensive treat-
ment of the taxonomy and distribution of C. ni-
grescens. He recognized one widely distributed
species, C. nigrescens, which included three sub-
species: C. n. mayensis, occurring on the Yucatan
Peninsula and known from one locality in Guer-
rero, Mexico; C. n. merriami, which Choate de-
scribed and is distributed from Chiapas, Mexico,
to Honduras and El Salvador; and C. n. nigrescens,
inhabiting Costa Rica and Panama to the Panama/
Colombia border. Of the previously recognized
species, Choate (1970) synonymized C. mera, C.
micrura, C. tersus, and C zeteki with C. n. ni-
grescens. Cryptotis mayensis was reduced to a sub-
species of C. nigrescens.
In his review of the Cryptotis of Middle Amer-
ica, Choate (1970) had few complete specimens of
C. nigrescens. Of the 237 C n. mayensis available
at that time, all but 10 were mandibles and partial
crania from owl pellets, cave deposits, or Mayan
ruins. Similarly, 15 of the 25 specimens available
for C. n. merriami were from owl pellets, and only
24 skins with accompanying skulls of C. n. ni-
grescens were available for study. Recent collect-
ing in Mexico has added important new specimens
of C n. mayensis and C. n. merriami, and work
in Panama and Costa Rica has tremendously in-
creased the number of C. n. nigrescens in museum
collections. This wealth of new material led us to
review this complex more thoroughly than was
possible previously.
Cryptotis nigrescens was considered to be dis-
tinguished by its short rostrum; bulbous dentition;
equal development of anterior and posterior por-
tions of M 1 ; unrecessed posterior margins of P4,
M 1 , and M2; simple M3; and lack of an entoconid
in m3 (Choate, 1970). However, there is much
variation in these characters both within this taxon
and within the genus, and their polarizations are
difficult to define.
Systematic relationships within the genus Cryp-
totis remain mostly unresolved. Woodman (1992)
showed that the C. nigrescens group may be para-
phyletic with respect to C. hondurensis and C.
magna. Cryptotis hondurensis recently was de-
scribed from montane pine forests in Honduras
(Woodman & Timm, 1992). Cryptotis magna was
described by Merriam (1895) from mountains of
southern Oaxaca, Mexico. A more comprehensive
redescription of this species was provided by
Choate ( 1 970), and it was illustrated by Robertson
and Rickart (1975), who summarized its biology.
In this paper, we use the "C nigrescens group" to
refer informally to only those shrews previously
called C. nigrescens.
The purposes of this paper are to ( 1 ) revise the
taxonomy and delineate the distributions of taxa
previously included in C. nigrescens; (2) describe
and illustrate a species new to science; and (3)
review the available information on reproduction,
elevational distribution, and habitat for the mem-
bers of this complex.
Methods
Cranial and mandibular measurements were re-
corded to the nearest 0. 1 mm using either an ocular
micrometer in a binocular microscope or a hand-
held dial caliper. Skin measurements are those re-
corded by the collector, except for head and body
length (HB), which we calculated by subtracting
the recorded tail length (TL) from the total length.
All measurements are in millimeters. Dental ter-
minology follows Choate (1970). The following
craniomandibular measurements (fig. 1 ) were used
in our analyses: condylobasal length, not including
the upper incisors (CBL); cranial breadth (CB);
breadth of zygomatic plate (ZP); interorbital
breadth (IO); breadth of palate across first uni-
cuspids (U1B); breadth of palate across third uni-
cuspids (U3B); breadth of palate across second
molars (M2B); palatal length (PL); upper tooth row
length, Ul to M3, parallel to the long axis of the
skull (TR); unicuspid tooth row length, parallel to
the unicuspid tooth row (UTR); molariform tooth
row length, P4 to M3, parallel to the long axis of
the skull (MTR); posterior width of M 1 , across
hypocone and metastyle (WM1); mandibular
FIELDIANA: ZOOLOGY
length, from inferior sigmoid notch to posterior
edge of mental foramen (ML); height of coronoid
process (HCP); height of coronoid valley (HCV);
height of articular condyle (HAC); breadth of ar-
ticular condyle (BAC); articular condyle to pos-
terior edge of m3 (AC3); lower tooth row length,
p3 to m3 (TRD); and length of lower first molar
(mlL). Unless otherwise stated, univariate statis-
tics are mean ± standard deviation. All capitalized
color names follow Ridgway (1912). Localities and
elevations were taken directly from specimen tags
and represent the descriptions of the original col-
lectors. Corrections and additions are provided in
brackets. We did not convert distances measured
in miles or elevations measured in feet to the met-
ric system to avoid inferring a level of accuracy
greater than that originally recorded by the col-
lector. Specimens from archeological or paleon-
tological contexts were not used in statistical anal-
yses or plotted on maps because of the possibility
of temporal variation in size, shape, or geographic
distribution. Number of specimens from archeo-
logical or paleontological sites is calculated as the
minimum number of individuals.
Multivariate analyses were used to look at pat-
terns of variation in overall similarity among spe-
cific members of the C. nigrescens complex. Al-
though these analyses often are useful for
distinguishing among known groups, and occa-
sionally point out variables useful for distinguish-
ing taxa, they were not relied upon to determine
species or provide taxonomic diagnoses. In gen-
eral, members of the C. nigrescens complex were
found to be very conservative in overall form,
particularly as determined by principal compo-
nents analyses (PCA), which is a descriptive rather
than a discriminatory tool. Most variation de-
tected by PCA was in size rather than shape. De-
spite this similarity in shape, species differ dis-
tinctly in their possession of specific characters,
and we used these characters to separate them.
Analyses of variance (ANOVA), correlation ma-
trices, and multivariate analyses were carried out
using BMDP on the University of Kansas Aca-
demic Computing Services' IBM VM/CMS com-
puter system. All data were log-transformed prior
to carrying out these analyses.
Elevational and geographic distributions and the
correlation of character states to these distribu-
tions were examined for all members of the C.
nigrescens group.
Specimens used in our analyses are listed in the
Specimens Examined section. These specimens are
deposited in the following institutions: American
Museum of Natural History, New York (amnh);
Angelo State Natural History Collections, San An-
gelo, Texas (asnhc); Escuela Nacional de Ciencias
Biologicas, Mexico (encb); Field Museum of Nat-
ural History, Chicago (fmnh); Instituto de Biolo-
gia, Universidad Nacional Autonoma de Mexico,
Mexico City (ibunam); Instituto Nacional de Bio-
diversidad, Santo Domingo de Heredia, Costa Rica
(iNBio); University of Kansas Museum of Natural
History, Lawrence (ku); Los Angeles County Mu-
seum, Los Angeles (lacm); Museum of Compar-
ative Zoology, Harvard University, Cambridge
(mcz); Museum of the High Plains, Fort Hays State
University, Hays, Kansas (mhp); James Ford Bell
Museum of Natural History, University of Min-
nesota, Minneapolis (mmnh); Museo Nacional de
Costa Rica, San Jose (mncr); Museum of Verte-
brate Zoology, University of California, Berkeley
(mvz); Royal Ontario Museum, Toronto (rom);
University of Iowa Museum of Natural History,
Iowa City (sui); Departamento de Biologia, Uni-
versidad Autonoma Metropolitana Iztapalapa,
Mexico (uami); University of Michigan Museum
of Zoology, Ann Arbor (ummz); Universidad Na-
cional Autonoma de Honduras, Tegucigalpa
(unah); and U.S. National Museum, Washington,
D.C. (usnm).
Sexual Variation
Male and female Cryptotis are difficult to tell
apart, and few tests for sexual dimorphism within
the genus have been carried out, at least in part
because there are few good series of individuals of
known sex. The two studies that have investigated
sexual dimorphism in Cryptotis indicate that sex-
ual differences in mensural variables are minor at
most. Choate ( 1 970) found only one of seven vari-
ables (length of maxillary tooth row) that he ex-
amined to be significantly different between the
sexes in his morphometric analysis of C. mexicana
from near Jalapa, Veracruz, Mexico. Only 1 of 19
variables differed significantly between males and
females of C. gracilis in Costa Rica and Panama
(Woodman, 1 992); zygomatic plate length was sig-
nificantly longer in females.
We tested for secondary sexual variation in C.
nigrescens using our two largest series of sexed
individuals. One included 1 7 males ( 1 4 crania, 1 6
skins) and 16 females (12 crania, 16 skins) from
Monteverde, Costa Rica, and the other consisted
of 13 males (13 crania, 9 skins) and 10 females
WOODMAN & TIMM: SMALL-EARED SHREWS
CBL
Fig. 1. Cranial and mandibular measurements used in this study. Abbreviations of variables are explained in the
Methods section of text.
( 1 0 crania, 8 skins) from near San Felix (including
Cerro Bollo), Panama. A correlation matrix was
calculated for 1 9 craniomandibular variables (CBL,
CB, ZP, IO, U1B, M2B, PL, TR, UTR, MTR,
WM1, ML, HCP, HAC, BAC, TRD, mlL, AC3,
HAV). Because sample sizes were different for skin
variables and for craniomandibular variables, skin
measurements (HB, TL) were not included in the
correlation matrix. Variables then were culled so
that no two variables in the remaining data set had
a correlation coefficient of 0.75 or greater. This
yielded eight variables (CBL, ZP, UTR, MTR,
WM1, HAC, BAC, HCV); all other variables cor-
related strongly with CBL, except HCV, which was
strongly correlated with HAC. A two-way ANO-
VA was calculated for each of these eight cranio-
mandibular variables and the two skin variables.
Both sex and locality (Monteverde vs. San Felix)
were tested as sources of variance to separate ef-
fects of sexual dimorphism and geographic vari-
ation. Statistical significance was determined using
a Brown-Forsythe test for equality of means, which
does not assume equality of the variances. Pro-
tected alpha values were not used despite a lack
of independence among the variables tested, be-
cause protected values would have increased the
likelihood of making Type II errors (i.e., incor-
rectly accepting no difference between males and
females or between localities).
Only one variable, UTR (P - 0.0042), exhibited
a statistically significant difference between sexes,
with females larger than males. In addition, CBL
(P = 0.0566) was close to being significantly dif-
ferent; again, females were larger than males. Five
of the 10 variables (ZP, MTR, WM 1 , BAC, HCV)
had P > 0.30, indicating no distinction between
the sexes. The remaining three variables (HB, TL,
HAC) had probabilities falling between 0.10 and
0.30, providing no clear evidence of whether a
difference existed between males and females. Fe-
males averaged slightly larger than males for five
measurements in the groups from both Monte-
verde (TL, CBL, ZP, UTR, HAC) and San Felix
(CBL, UTR, MTR, HAC, mlL), although not for
all the same variables. Males from Monteverde
were larger than females for HB, and males from
FIELDIANA: ZOOLOGY
San Felix were larger for HB and TL. However,
actual differences between male and female means
in both groups were small (table 1). If we use the
differences, taken to four decimal places, between
the male and female means for the eight cranio-
mandibular variables, we find that they average
0.0470 (range = 0.0 1 69-0.0892) for the specimens
from San Felix and 0.0926 (range - 0.0 1 07-0.4630)
for specimens from Monteverde. Most are below
our ability to measure accurately.
In contrast to the sex term, 9 of the 10 variables
tested were significantly different between locali-
ties. Seven of these variables (HB, TL, CBL, UTR,
WM 1 , H AC, BAC) had P < 0.00 1 , and two (MTR,
m 1 L) had P < 0.05. Only ZP showed no difference
between the populations at Monteverde and San
Felix (P = 0.3531). For all variables, specimens
from San Felix averaged larger than those from
Monteverde. In fact, males from San Felix aver-
aged larger than females from Monteverde for all
variables except ZP, UTR, and m 1 L.
The interaction between sex and locality was not
significant for any variables, and P < 0.30 for only
TL (P = 0.0734), CBL (P = 0.0947), and ZP (P =
0.2593).
Body mass has been shown to be a good cor-
relate of other estimators of size in some other
small mammals (Iskjaer et al., 1989). Average
weights of males and females from Monteverde
were the same (table 1), giving no indication of
sexual dimorphism. However, our use of body mass
was hampered by the imprecision with which very
small animals can be weighed in the field and small
sample sizes available. Geographic variation could
not be evaluated, because weights were not avail-
able for specimens from San Felix.
To test whether all craniomandibular variables
together would show size differentiation between
males and females, we ran PCAs separately for
specimens from Monteverde and from San Felix
using the original 19 craniomandibular variables
and the same individuals as for the ANOVAs.
Because all variables generally load heavily on fac-
tor axis 1 in a PCA, this axis provides a good
estimator of overall size. Therefore, factor 1 scores
should prove to be useful in determining any ob-
vious patterns of size differentiation between the
sexes. In the PCA of specimens from Monteverde,
factor 1 estimates size and factor 2 is a shape score
representing contrast between a combination of
m 1 L and WM 1 vs. the negatively weighted AC3
(table 2). A plot of factor 1 and factor 2 scores for
these specimens (fig. 2) indicates nearly complete
overlap of males and females on both the size and
shape axes. In the PCA of specimens from San
Felix, factor 1 represents size and factor 2 is a
contrast between shape of the articular condyle of
the mandible (HCV and HAC) vs. a negatively
weighted mlL (table 3). In the plot of factor 1 and
factor 2 scores for these specimens (fig. 3), males
and females exhibit complete overlap along the
shape axis. Along the size axis, however, there is
separation of the two sexes. Males range along the
entire length of factor axis 1 but are more con-
centrated at the lower end of the scale. Females
all cluster toward the upper end of the scale.
Our analysis of sexual variation in C. nigrescens
shows no clear pattern of sexual dimorphism in
size within this species. Although females gener-
ally averaged larger in most craniomandibular
measurements in which there were differences be-
tween the sexes, males averaged larger in HB, es-
pecially among specimens from San Felix. The
overlap between males and females for all mea-
surements was great. Only two variables were sig-
nificant or close to being significant when tested
for differences between the sexes, and the statis-
tical significance of those variables may have been
a result of multiple comparison of dependent
means. No other variables tested were clearly dif-
ferent. PCA indicated a tendency for male and
female C. nigrescens from San Felix to separate
out on size, but this was not the case for those
from Monteverde, and the size range for males
from San Felix overlapped completely that of fe-
males.
These results indicate that any sexual variation
that exists is beyond our ability to measure ac-
curately and precisely in these diminutive mam-
mals. Differences between sexes were considerably
less than differences between geographic localities,
and they did not influence statistical tests among
populations or contribute significantly to overall
patterns of geographic variation in which we were
interested. Possessing few sexed individuals from
each locality, we could not partition our sample
into tooth-wear classes (Rudd, 1 955) and thus pro-
portion out possible effects of age-related differ-
ences. However, because sexual variation did not
inhibit our ability to detect geographic variation
within C. nigrescens, we used males, females, and
unsexed individuals in all subsequent analyses.
Systematic Descriptions
In the following descriptions, species are ar-
ranged geographically, roughly from northwest to
WOODMAN & TIMM: SMALL-EARED SHREWS
Table 1 . Selected measurements of male and female C. nigrescens from Monteverde, Costa Rica, and San Felix,
Panama. Abbreviations of measurements are explained in the Methods section of text. The statistics presented are
mean ± standard deviation of the mean, and observed extremes. Number of individuals for each species is in
parentheses.
HB
TL
CBL
CB
ZP
IO
U1B
M2B
PL
TR
UTR
MTR
WM1
ML
HCP
HCV
HAC
Mont
everde
San Feli>
Males
Females
Males
Females
68 ± 5
67 ± 7
77 ± 3
74 ± 4
60-76
56-79
72-83
68-79
(16)
(16)
(9)
(8)
28 ± 4
30 ± 2
34 ± 2
33 ± 2
20-34
27-33
29-37
30-36
(16)
(16)
(9)
(8)
17.9 ± 0.5
18.4 ±0.5
19.2 ± 0.4
19.3 ± 0.4
16.9-18.7
17.7-19.3
18.7-20.2
18.3-19.8
(14)
(12)
(13)
(10)
9.0 ± 0.2
9.0 ± 0.3
9.7 ± 0.2
9.7 ± 0.2
8.7-9.4
9.4-10.0
9.4-10.3
9.2-10.0
(14)
(12)
(13)
(10)
1.9 ±0.2
2.0 ± 0.1
2.0 ± 0.2
2.0 ± 0.2
1.6-2.2
1.8-2.2
1.5-2.5
1.6-2.2
(14)
(12)
(13)
(10)
4.2 ± 0.1
4.2 ± 0.2
4.6 ± 0.1
4.6 ± 0.2
4.0-4.3
4.0-4.7
4.4-4.9
4.3^.8
(14)
(12)
(13)
(10)
2.3 ± 0.1
2.3 ± 0.1
2.6 ±0.1
2.6 ± 0.1
2.2-2.5
2.2-2.5
2.5-2.8
2.5-2.7
(14)
(12)
(13)
(10)
5.3 ± 0.1
5.3 ± 0.2
5.7 ±0.1
5.8 ± 0.1
5.1-5.6
4.9-5.6
5.5-5.9
5.5-6.0
(14)
(12)
(13)
(10)
7.6 ± 0.2
7.9 ± 0.3
8.1 ± 0.2
8.3 ± 0.3
7.2-7.9
7.4-8.6
7.8-8.3
7.8-8.8
(14)
(12)
(13)
(10)
6.8 ± 0.2
7.1 ±0.2
7.4 ± 0.2
7.5 ± 0.3
6.4-7.2
6.7-7.3
7.2-7.6
6.9-7.8
(14)
(12)
(13)
(10)
2.3 ± 0.2
2.4 ± 0.1
2.4 ±0.1
2.5 ± 0.1
2.0-2.5
2.3-2.5
2.2-2.6
2.4-2.6
(14)
(12)
(13)
(10)
5.2 ± 0.2
5.2 ± 0.2
5.3 ±0.1
5.4 ± 0.2
4.8-5.5
4.8-5.6
5.2-5.5
4.9-5.7
(14)
(12)
(13)
(10)
1.7 ± 0.1
1.7 ± 0.1
1.8 ±0.05
1.8 ± 0.1
1.5-1.8
1.6-1.8
1.7-1.9
1.7-1.9
(14)
(12)
(13)
(10)
6.1 ± 0.2
6.3 ± 0.2
6.4 ± 0.3
6.5 ± 0.2
5.7-6.5
5.9-6.6
6.0-6.9
6.1-6.9
(14)
(12)
(13)
(10)
4.3 ± 0.2
4.4 ± 0.2
4.7 ± 0.1
4.8 ± 0.1
4.0-4.6
4.1^.7
4.6-5.0
4.6-5.0
(14)
(12)
(13)
(10)
2.6 ± 0.1
2.7 ±0.1
2.8 ±0.1
2.9 ± 0.1
2.4-2.8
2.5-2.9
2.6-3.0
2.8-3.0
(14)
(12)
(13)
(10)
3.6 ± 0.2
3.7 ± 0.2
3.8 ± 0.2
3.9 ±0.1
3.3-3.9
3.4-4.0
3.5^1.1
3.7-4.0
(14)
(12)
(13)
(10)
FIELDIANA: ZOOLOGY
Table 1. Continued.
BAC
AC3
TRD
mlL
Weight (g)
Monteverde
San Felix
Males
Females
Males
Females
2.9 ± 0.1
2.9 ± 0.1
3.0 ± 0.1
3.0 ± 0.1
2.7-3.1
2.7-3.1
2.9-3.2
2.9-3.2
(14)
(12)
(13)
(10)
4.4 ± 0.2
4.5 ± 0.2
4.6 ± 0.2
4.7 ± 0.05
4.1-4.7
4.2-4.8
4.4-5.1
4.7^.8
(14)
(12)
(13)
(10)
5.4 ± 0.2
5.5 ± 0.2
5.8 ±0.1
5.9 ± 0.2
5.1-5.6
5.2-5.8
5.7-6.2
5.3-6.2
(14)
(12)
(13)
(10)
1.7 ± 0.1
1.7 ± 0.1
1.7 ± 0.1
1.8 ± 0.1
1.6-1.8
1.6-1.8
1.6-1.9
1.6-1.9
(14)
(12)
(13)
(10)
6 ± 1
6 ± 1
4-8
5-7
—
—
(13)
(9)
southeast. Accounts are not provided for C. hon-
durensis or C. magna, although there is evidence
that these two species may be included within the
C. nigrescens group (Woodman, 1992). However,
comparisons of these two species are made with
the species described below.
Cryptotis mayensis (Merriam, 1901)
(fig. 4)
Blarina mayensis C. H. Merriam, 29 November 1901,
Proc. Washington Acad. Sci. 3:559.
Crvptotis mavensis: G. S. Miller, Jr., 31 December
1912, Bull. U.S. Natl. Mus. 79:26; R. T. Hatt, 18
August 1938, J. Mamm. 19:334; P. Hershkovitz,
1 0 July 1951, Fieldiana Zool. 3 1 :522; R. T. Hatt et
al., March 1953, Cranbrook Inst. Sci. Bull. 33:59;
E. R. Hall and K. R. Kelson. 31 March 1959, The
Mammals of North America 1:61; T. Alvarez and
A. Martinez G., 4 August 1 967, Southwest. Nat. 1 2:
205; R. L. Peterson, 26 November 1968, J. Mamm.
49:796.
Blarina mexicana: G. F. Gaumer, 1917, Monografia
de los mamiferos de Yucatan, p. 249 (part).
Cryptotis micrura: A. Murie, 15 July 1935. Misc. Publ.
Mus. Zool., Univ. Michigan 26: 1 7 (part); E. R. Hall
Table 2. Factor loadings from PCA of male and fe-
male C. nigrescens from Monteverde, Costa Rica. Ab-
breviations of measurements are explained in the Meth-
ods section of the text.
Table 3. Factor loadings from PCA of male and fe-
male C. nigrescens from the vicinity of San Felix, Pan-
ama.
Variable
Factor 1
Factor 2
Variable
Factor 1
Factor 2
CBL
0.875
-0.179
TRD
0.901
-0.291
TR
0.860
0.221
MTR
0.879
-0.167
PL
0.857
-0.090
CBL
0.872
-0.278
HCP
0.824
-0.260
TR
0.836
-0.051
ML
0.786
-0.175
PL
0.817
0.117
TRD
0.755
0.485
HCP
0.738
0.474
U1B
0.728
0.007
IO
0.716
0.088
HAC
0.708
-0.284
BAC
0.713
0.379
IO
0.692
0.102
CB
0.708
-0.248
BAC
0.667
-0.243
ML
0.700
-0.417
HCV
0.663
-0.390
M2B
0.682
0.176
M2B
0.658
0.440
AC3
0.570
-0.032
UTR
0.652
0.160
HCV
0.566
0.634
MTR
0.642
0.392
U1B
0.527
0.324
AC3
0.606
-0.589
ZP
0.496
-0.219
CB
0.596
-0.218
UTR
0.459
-0.123
ZP
0.539
-0.283
HAC
0.449
0.631
mlL
0.535
0.584
WM1
0.443
-0.290
WM1
0.445
0.582
mlL
0.340
-0.703
WOODMAN & TIMM: SMALL-EARED SHREWS
2 -
O
•
1 -
o
o
o
•
o
* °
oo
0 -
o
•
• •
•
-1 -
•
o
o
*
-2 -
O
•
males
females
o
■ i
-3-2-10 1 2 3
factor 1
Fig. 2. Plot of PCA factor 1 and factor 2 scores for
males and females of C. nigrescens from Monteverde,
Costa Rica.
o
u
o
2 -
O
*
o
1 -
o
o
•
•
o
0 -
o
o
o
O
o
o
•
•
-1 -
O ma
es
•o
•
•
• females
•
i
1
-2
-1
1
factor 1
Fig. 3. Plot of PCA factor 1 and factor 2 scores for
males and females of C. nigrescens from the vicinity of
San Felix, Panama.
and K. R. Kelson, 31 March 1959, The Mammals
of North America 1 :62 (part).
Cryptotis nigrescens mayensis: J. R. Choate, 30 De-
cember 1970, Univ. Kansas Publ. Mus. Nat. Hist.
19:275; E. R. Hall, 3 April 1981, The Mammals of
North America 1:63; R. C. Dowler and M. D. Eng-
strom, 15 December 1988, Ann. Carnegie Mus. 57:
160.
Holotype— Skin and skull of subadult female,
U.S. National Museum of Natural History no.
1 08087, obtained by E. W. Nelson and E. A. Gold-
man (collector number 14495) 5 February 1901
from Mexico, Yucatan, Chichen Itza.
Distribution— Known primarily from below
1 00 m on the Yucatan Peninsula of Mexico and
adjacent Belize and Guatemala (fig. 5). Also known
from remains isolated from owl pellets collected
in a cave at 650 m, in the Canon del Zopilote,
Guerrero, Mexico.
Description— A medium-sized Cryptotis, HB
averaging 69 (table 4); tail short, averaging 41%
(± 5%, n = 1 1) of HB; dorsal hairs about 3 mm
long, ranging from about 2 to 4 mm, three-banded;
dorsum often has mottled, salt and pepper ap-
pearance; dorsal pelage variable: Pale Ecru-Drab
and Drab, Light Drab, Light Mouse Gray to Deep
Mouse Gray, Pale Mouse Gray or Mouse Gray
with touches of Olive Brown; lateral pelage from
Smoke Gray to Mouse Gray; venter slightly paler
than dorsum, may include one or more of the
following: Pale Olive Gray, Light Olive Gray, Light
Grayish Olive, Pale Mouse Gray to Light Mouse
Gray.
Rostrum narrow and of normal length (PL/CBL
= 43.4% ± 1.2, n = 10); interorbital area mod-
erately wide; usually only one dorsal foramen (8 1 %,
n = 65) located on either the right or left frontal,
small to medium in size; a well-developed fora-
men leading to a ventral extension of the sinus
canal typically present posterior to the dorsal ar-
ticular facet on one (8%, n = 62) or both (86%)
sides of the skull; normally no foramen dorsal to
dorsal articular facet (93%, n = 61); anterior pro-
cess of the petromastoid low and very narrow (fig.
18 A); paroccipital process prominent; zygomatic
plate broad in proportion to CBL (1 1.8% ± 0.7,
n = 10) and PL (27.8% ± 1.6, n = 72); anterior
border of zygomatic plate from parastyle/meso-
style valley to mesostyle of M 1 , posterior border
from metastyle of M2 to middle of M3, and from
posterior one-third of base of maxillary process to
posterior to maxillary process; palate long and nar-
row; upper tooth row crowded; unicuspids mas-
sive; U4 normally displaced medially, so that U3
and P4 in contact or nearly so; U4 not visible in
lateral view of skull; lateral view of U3 also some-
times partially obstructed by P4; posterior borders
of P4, M 1 , and M2 unrecessed or only very slightly
recessed; M3 with well-developed paracrista and
paracone, reduced precentrocrista, poorly devel-
oped and normally uncolored mesostyle and pro-
tocone, and hypocone absent or vestigial and in-
corporated into the posterior cingulum; dentition
bulbous.
Mandible large; horizontal ramus deep; coro-
noid process high (HCP/ML = 84.3% ± 3.3, n =
FIELDIANA: ZOOLOGY
Fig. 4. Dorsal, ventral, and lateral views of the skull and lateral view of the skull and mandible of C. tnayensis
(asnhc 6071). Scale bar = 5 mm.
21) and broad, joins mandible at steep angle;
viewed posteriorly, horizontal and vertical
branches of articular condyle short and broad;
lower sigmoid notch very shallow, not extending
beneath ventral border of articular process; pos-
terior border of lower incisor extends nearly to
posterior border of cingulum of p4; lower denti-
tion wide; only hypoconid in talonid of m3.
Comparisons— Cryptotis mayensis has the pal-
est pelage of any member of the C. nigrescens group,
appearing medium gray (often speckled with
patches of white) rather than dark brown to black.
Cryptotis magna — C. mayensis is much smaller
(HB averaging 69 vs. 86; table 4) and has much
paler pelage; much shorter tail (averaging 41% of
HB vs. 53%); skull much smaller in all dimensions,
but zygomatic plate relatively broader (averaging
1 1.8% of CBL vs. 10.1% ± 0.6); usually only one
dorsal foramen; well -developed foramen posterior
to dorsal articular facet (foramen present but al-
ways vestigial in C. magna); rarely possesses fo-
ramina dorsal to dorsal articular facet; no ento-
conid in talonid of m3 (vestigial entoconid present
in C. magna).
Cryptotis hondurensis—C. mayensis is much
larger (HB averaging 69 vs. 61 ± 5; table 4) and
has much paler pelage; shorter tail (averaging 41%
vs. 46%); M3 less complex, lacking metacone; den-
tition bulbous; coronoid process of mandible much
higher and broader.
Remarks— The pelage of Cryptotis mayensis is
paler than in any other member of the genus, in-
cluding C. parva. from which the coloration differs
in being more gray and less olive. The overall
appearance of C mayensis is a medium brownish
gray. Close inspection of the dorsal pelage of C
mayensis reveals that the individual hairs are three-
banded. The proximal one-half to three-quarters
of the hairs is a medium gray, followed by a narrow
band of light gray. The distal tip (about one-eighth
of the total length) of the hair is brownish gray.
The transition between bands is not sharp, with
the colors instead tending to grade into each other.
The great extent of lighter, basal coloration in the
hairs of this taxon probably accounts for its overall
paler appearance and greater reflectance.
Goldman (1951) described the northern Yuca-
tan habitat around Chichen Itza, from which the
WOODMAN & TIMM: SMALL-EARED SHREWS
Fig. 5. Map of southern Mexico, Belize, and Guatemala showing the distribution of C. mayensis.
holotype of C. mayensis was collected, as stunted
tropical forest about 25-40 ft high with dense,
brushy undergrowth. He noted that the weather is
highly seasonal, with a long, hot, dry season from
about December to June and a rainy season marked
by torrential afternoon showers. Dowler and Eng-
strom (1988) reported a specimen from 7.5 km W
Escarcega, Campeche, as having been trapped in
mature, transitional deciduous-evergreen forest.
In the same area they also collected Hetewmys
gaumeri, Ototylomys phyllotis, Peromyscus yuca-
tanicus, and Oryzomys melanotis. Two C. may-
ensis from 60 km SE of Dzibalchen, Campeche,
were taken in pitfalls set in transitional deciduous-
evergreen forest dominated by escobo palms (Cry-
sophila) and Sabal palms (M. D. Engstrom, in
litt.). Alvarez and Martinez (1967) captured a C.
mayensis along a road in tropical rain forest op-
posite a cornfield, 2 km SE Laguna Chickanka-
naab, Quintana Roo.
Few reproductive data are available for C. may-
ensis, and no pregnant or lactating females have
been recorded. Males captured on 4 June, 24 June,
and 1 9 August showed no indications of having
lateral glands. When present, these paired glan-
dular areas lack long guard hairs and underfur but
have a sparse covering of short, fine hairs (Wood-
man & Timm, submitted). Both males and females
possess lateral glands, but in females they are much
smaller and more difficult to see (Murariu, 1976;
Bee et al., 1980). It is likely that these glands serve
a function in sexual communication. Eadie (1938)
found that the lateral glands of male Blarina showed
increased activity with enlargement of the testes.
Cryptotis mayensis is rarely collected, and there
are few complete specimens in museum collec-
tions. However, it is unlikely that this shrew is
truly rare, because remains from owl pellets are
abundant. Of 122 specimens examined in our
study, 76 came from owl pellets, 33 are from ar-
cheological contexts, and 1 3 are standard museum
specimens.
Cryptotis mayensis is distinctive in being the
only member of the genus restricted to lowland
areas. On the Yucatan Peninsula the species is not
known to occur above 100 m. The only specimens
of C. mayensis from outside of the Yucatan Pen-
insula are crania and unassociated mandibles col-
lected by William Lopez-Forment C. in 1969 from
below a barn owl (Tyto alba) roost in Macuiltzingo
10
FIELDIANA: ZOOLOGY
Cave [Cueva del Canon de Zopilote], located at
650 m in the Canon de Zopilote, Guerrero. Mexico
(Choate, 1970). The precise locality and habitat
where the shrews were captured by owls is un-
known, and subsequent work in Macuiltzingo Cave
has not produced additional specimens (Lopez-
Forment & Urbano, 1977; W. Lopez-Forment C,
pers. comm.), suggesting a change in feeding pat-
terns by the owls or local extinction of the pop-
ulation of shrews upon which they were feeding.
When one of us (Woodman) visited the cave with
W. Lopez-Forment C. in August 1991. no owls
were occupying the cave, and there was no evi-
dence of any recent use of the cave by owls. Despite
repeated collecting in the region surrounding the
cave (Choate, 1970; W. Lopez-Forment C, pers.
comm.; fieldwork by Woodman in 1991), no ad-
ditional specimens of these shrews have been ob-
tained.
The presence of shrews identifiable as C. may-
ensis in Guerrero presents an interesting biogeo-
graphical problem. These shrews are from an area
that clearly is isolated geographically from the Yu-
catan Peninsula and is climatically and vegeta-
tionally distinct. Choate (1970) noted the great
distance (> 950 km) and extensive biogeograph-
ical barriers between Zopilote Canyon and the
nearest known locality of C. mayensis on the Yu-
catan Peninsula and suggested that the specimens
from Zopilote Canyon eventually might prove to
be subspecifically distinct. All known specimens
from Guerrero are mandibles and incomplete
skulls, making comparisons with other taxa, in-
cluding C. mayensis, difficult.
Because the specimens from Guerrero were from
far outside the range of all other populations on
the Yucatan Peninsula, and because of the geo-
graphical barriers and habitat differences between
the two areas, we considered that specimens from
Guerrero might prove to be either misidentified
or an undescribed species. The fragmentary nature
of the specimens from Guerrero limited the mea-
surements that could be taken from them and did
not yield characters useful for separating them from
C. mayensis on the Yucatan Peninsula. We carried
out PCAs and discriminant function analyses in
order to test the overall similarity of the two groups
of specimens. Crania and mandibles were tested
separately, because these elements are disasso-
ciated in all known specimens from Guerrero.
Analyses of crania included six variables (ZP, IO,
M2B, PL, MTR, WM1) measured on 20 C. may-
ensis from the Yucatan Peninsula, 1 6 C. merriami,
and 20 specimens from Guerrero. Analyses carried
rO
o
2 -
1 -
0 -
o
• o
°~ °
° o
o •
•v
o ©
•
o.<§>*8
V v
V
V
V
-1 -
•
•
V
• o
w
V
-2 -
•
V
V
V
-3 -
O
Guerrero
•
C.
mayensis
-A .
V
C.
merriami
-2-10 1 2
factor 1
Fig. 6. Plot of specimens of C. mayensis from Guer-
rero and the Yucatan Peninsula and C. merriami on
factor axes 1 and 3 from PCA of cranial variables.
out on mandibles utilized eight variables (ML,
HCP, HCV, HAC, BAC, TRD, mlL, AC3) from
20 C. mayensis, 24 C. merriami, and six mandi-
bles from Guerrero. Specimens of C. merriami
were included to determine whether or not the
variables used were useful for distinguishing groups.
PCAs indicated wide overlap of C. mayensis
from the Yucatan with specimens from Guerrero.
In the PCA of crania, all variables except ZP load-
ed out heavily on factor 1 , providing a measure
of overall size of the specimens (table 5). Factor
3 was a shape axis, providing a contrast between
PL and M2B. Factor 2 from this analysis repre-
sented a contrast between a very heavily weighted
ZP and a slightly negatively weighted WM 1 ; it did
not aid in separating any groups. In a plot of factor
scores (fig. 6), C. mayensis and specimens from
Guerrero are essentially indistinguishable. Cryp-
totis mayensis and C. merriami are separated by
a combination of size and shape. Cryptotis mer-
riami tends to be larger cranially than C mayensis,
and, at any given size, C. mayensis has a longer
but narrower palate than C. merriami.
In the PCA carried out using mandibles, factor
1 is a size axis and factor 2 is a contrast between
a combination of variables representing length of
the mandible (mlL, TRD, ML) and HCP (table
6). A plot of specimens on factor 1 and factor 2
(fig. 7) shows C. mayensis from the Yucatan Pen-
insula and specimens from Guerrero again inter-
mixed. In contrast to the PCA using cranial vari-
ables, C. merriami and C mayensis do not separate
out on the size axis but can be discriminated by
WOODMAN & TIMM: SMALL-EARED SHREWS
11
Table 4. Selected measurements of members of the C. nigrescens group. Abbreviations of measurements are
explained in the Methods section of text. The statistics presented are mean ± standard deviation of the mean, and
observed extremes. Number of individuals for each species is in parentheses.
HB
TL
TLas°/oofHB
CBL
CB
ZP
IO
U1B
U3B
M2B
PL
TR
UTR
MTR
WM1
ML
C.
hondurensis
C.
(holotype and
C.
colombiana
C. mayensis
C. magna
C. merriami
2 paratypes)
nigrescens
C. mera
(holotype)
69 ± 8
86 ± 6
69 ± 4
55, 63, 64
68 ± 7
69 ± 2
76
61-90
70-100
60-77
55-83
67-73
(11)
(47)
(23)
(70)
(6)
28 ± 3
46 ± 4
29 ±4
29, 27, 27
29 ± 4
27 ± 2
27
24-33
34-53
22-38
19-37
24-31
(11)
(47)
(23)
(70)
(6)
41 ± 5
54 ± 6
43 ± 6
53, 43, 42
43 ± 5
39 ±4
36
31-49
34-64
32-55
28-53
34-46
(ID
(47)
(23)
(70)
(6)
19.0 ± 0.4
22.8 ± 0.4
19.4 ±0.4
19.2, -, -
18.6 ± 0.7
18.3 ± 0.4
19.9
18.2-19.6
21.9-23.4
18.7-20.0
16.9-20.2
17.7-18.7
(10)
(17)
(16)
(70)
(6)
9.2 ± 0.3
11.6 ± 0.2
9.6 ± 0.2
9.2, -, -
9.2 ± 0.4
9.4 ± 0.3
10.0
8.8-9.8
11.2-12.3
9.2-10.0
8.0-10.3
9.1-9.8
(10)
(17)
(18)
(70)
(6)
2.3 ± 0.1
2.3 ±0.1
2.3 ± 0.2
2.2,2.0, 2.1
2.0 ± 0.2
2.0 ±0.1
2.0
2.0-2.5
2.1-2.6
2.0-2.6
1.5-2.5
1.9-2.2
(21)
(17)
(21)
(70)
(6)
4.4 ± 0.2
5.6 ±0.1
4.7 ± 0.1
4.3,4.5, -
4.4 ± 0.2
4.5 ± 0.2
4.9
4.1-4.8
5.4-5.8
4.4^.9
3.9-4.9
4.3-4.7
(21)
(17)
(21)
(70)
(6)
2.6 ± 0.1
3.2 ±0.1
2.6 ± 0.2
2.4, 2.4, -
2.4 ± 0.2
2.5 ± 0.1
2.6
2.2-2.7
3.0-3.4
2.1-2.8
2.1-2.8
2.4-2.6
(21)
(17)
(21)
(70)
(6)
3.0 ±0.1
3.7 ± 0.2
3.0 ± 0.2
2.9, 3.0, -
2.8 ± 0.2
2.9 ±0.1
3.2
2.8-3.3
3.5-3.9
2.7-3.1
2.4-3.2
2.7-3.0
(21)
(17)
(6)
(70)
(6)
5.5 ±0.1
6.8 ± 0.1
5.8 ± 0.2
5.4, 5.4, -
5.5 ± 0.3
5.6 ±0.1
6.3
5.2-5.7
6.5-7.0
5.2-6.2
4.9-6.0
5.5-5.8
(21)
(17)
(21)
(70)
(6)
8.3 ± 0.3
10.1 ±0.3
8.4 ± 0.3
8.4, 7.8, -
7.9 ± 0.3
7.8 ± 0.1
8.8
7.8-9.0
9.6-10.8
7.9-8.9
7.2-8.8
7.6-7.8
(21)
(17)
(21)
(70)
(6)
7.3 ± 0.2
9.1 ±0.2
7.4 ± 0.2
7.1, 7.1, 7.1
7.1 ± 0.3
7.1 ± 0.1
7.7
6.9-7.8
8.8-9.3
6.8-7.7
6.4-7.8
6.9-7.2
(21)
(17)
(21)
(70)
(6)
2.7 ± 0.2
3.2 ± 0.1
2.5 ±0.1
2.6, 2.4, 2.4
2.4 ± 0.1
2.4 ± 0.1
2.5
2.1-3.1
3.1-3.4
2.2-2.7
2.0-2.6
2.3-2.5
(21)
(17)
(21)
(70)
(6)
5.0 ±0.1
5.1 ±0.6
5.2 ± 0.2
5.1, 5.1, 5.2
5.2 ± 0.2
5.1 ±0.05
5.8
4.8-5.3
5.1-5.2
5.0-5.5
4.7-5.7
5.1-5.2
(21)
(17)
(17)
(70)
(6)
1.7 ± 0.0
1.8 ± 0.1
1.9 ±0.1
1.7, 1.7, 1.9
1.8 ± 0.1
1.8 ± 0.05
1.8
1.7-1.8
1.7-1.9
1.7-2.0
1.5-1.9
1.8-1.9
(21)
(17)
(21)
(70)
(6)
6.2 ± 0.2
7.3 ± 0.2
6.5 ± 0.3
6.1, 5.8, 6.3
6.2 ± 0.3
5.9 ± 0.4
6.9
5.8-6.8
6.8-7.6
6.0-7.2
5.5-6.9
5.2-6.2
(24)
(17)
(21)
(70)
(6)
12
FIELDIANA: ZOOLOGY
Table 4. Continued.
HCP
HCV
HAC
BAC
AC3
TRD
mlL
C.
hondurensis
C.
(holotype and
C.
colombiana
C. mayensis
C. magna
C. merriami
2 paratypes)
nigrescens
C. mera
(holotype)
5.3 ± 0.3
6.1 ±0.2
5.0 ± 0.2
4.8, 4.4, 4.6
4.5 ± 0.3
4.5 ±0.1
4.8
4.8-5.7
5.7-6.3
4.6-5.3
4.0-5.0
4.3^t.6
(24)
(17)
(21)
(70)
(6)
2.9 ±0.1
3.4 ± 0.1
2.9 ± 0.2
2.7, 2.7, 2.7
2.7 ± 0.2
2.7 ± 0.1
2.9
2.7-3.2
3.2-3.7
2.6-3.3
2.3-3.0
2.6-2.8
(24)
(17)
(21)
(70)
(6)
3.9 ± 0.2
4.6 ± 0.1
3.8 ± 0.2
3.6, 3.6, 3.8
3.7 ± 0.2
3.7 ±0.1
4.1
3.4-4.4
4.4-4.8
3.5^.3
3.3-4.1
3.6-3.8
(24)
(17)
(21)
(70)
(6)
3.1 ± 0.1
3.6 ±0.1
3.0 ±0.1
3.0, 2.8. 2.8
2.9 ± 0.1
3.0 ±0.1
3.1
2.8-3.3
3.4-3.8
2.6-3.2
2.6-3.2
2.9-3.1
(24)
(17)
(21)
(70)
(6)
4.8 ± 0.2
5.8 ± 0.2
4.6 ± 0.2
4.3, 4.3, 4.4
4.5 ± 0.2
4.4 ± 0.1
4.9
4.4-5.3
5.3-6.3
4.3-4.9
3.8-5.1
4.3-4.6
(24)
(17)
(21)
(70)
(6)
5.5 ± 0.2
6.9 ± 0.1
5.7 ± 0.2
5.5, 5.3, 5.7
5.6 ± 0.3
5.5 ±0.1
6.0
5.3-5.8
6.6-7.1
5.3-5.9
5.1-6.2
5.4-5.7
(24)
(17)
(21)
(70)
(6)
1.7 ± 0.1
2.2 ± 0.1
1.8 ± 0.1
1.7, 1.7, 1.7
1.7 ± 0.1
1.8 ±0.1
1.9
1.6-1.9
2.1-2.4
1.7-1.9
1.6-1.9
1.7-1.8
(24)
(17)
(21)
(70)
(6)
factor 2 scores. Cryptotis merriami has a relatively
longer mandible and tooth row and a shorter cor-
onoid process than C. mayensis and the specimens
from Guerrero.
Discriminant function analysis of crania failed
to distinguish effectively among the three groups
of shrews. This analysis utilized a three- variable
model (WM1, M2B, PL), which correctly identi-
fied fewer than 70% of specimens using a jack-
knifed classification. A plot of specimens on ca-
nonical axes 1 and 2 (fig. 8) shows some distinction
between C. merriami and the other two groups,
but there is broad overlap between C. mayensis
and the specimens from Guerrero. The pattern of
linear groupings in this plot is due to one variable,
WM 1 , which loaded out equally on each axis and
for which there were only five states. These qual-
ities made it act as a discrete variable rather than
as a continuous variable. However, this does not
affect the validity of the analysis nor the conclu-
sion that C. mayensis from the Yucatan Peninsula
and from Guerrero are difficult to distinguish.
The discriminant function analysis of mandi-
bles utilized four variables (ML, HCP, HAC, BAC),
which correctly classified 96% of C. merriami, 80%
of C. mayensis from the Yucatan, and all of the
specimens from Guerrero. Intermixing between
the two described taxa was minimal; only one C.
merriami and one C. mayensis were incorrectly
identified as the other taxon. However, three C.
mayensis were misclassified as belonging to the
group of specimens from Guerrero. A plot of spec-
imens on canonical axes 1 and 2 shows clustering
of individuals belonging to each of the three groups
but not complete separation among groups (fig. 9).
Cryptotis merriami can be distinguished from the
other two groups along axis 1 , whereas C. mayensis
and the specimens from Guerrero cluster sepa-
rately along axis 2.
Although there is some indication that C. may-
Table 5. Factor loadings from PCA of cranial vari-
ables of C. mayensis from the Yucatan Peninsula and
Guerrero and C merriami.
Variable
Factor 1
Factor 2
Factor 3
MTR
0.892
-0.099
0.096
M2B
0.869
0.047
-0.311
WM1
0.839
-0.233
-0.162
IO
0.828
0.003
-0.180
PL
0.681
0.071
0.708
ZP
0.198
0.970
-0.065
WOODMAN & TIMM: SMALL-EARED SHREWS
13
2 -
1 -
V
V
V
V
V
0 -
V
V
O y#
•
V
-1 -
•
°*o°
• •
o
• •
•
-2 -
O Guerrero
•
•
• C. mayensis
v C. merriami
3 -\
i i
1 1
1 1
factor 1
Fig. 7. Plot of specimens of C. mayensis from Guer-
rero and the Yucatan Peninsula and C. merriami on
factor axes 1 and 2 from PCA of mandibular variables.
ensis and the shrews from Guerrero exhibit some
differentiation, they are very similar, and at this
time it seems best to refer the specimens from
Guerrero to C. mayensis. We do this tentatively,
however, and we believe that once more complete
specimens are available the shrews from Guerrero
will prove to be specifically distinct from C. may-
ensis. Alternatively, the shrews from Guerrero may
represent a disjunct population of C. mayensis.
This is supported by another species found with
the shrews in the owl pellets from Macuiltzingo
Cave, Ototylomys phyllotis, which otherwise is un-
known north of the Isthmus of Tehuantepec. The
specimens from Guerrero are indistinguishable
from O. p. phyllotis from the Yucatan Peninsula,
which in turn are easy to separate from O. p. con-
nectens in Chiapas (M. D. Engstrom, in litt.).
Because of possible geographic variation, and
because of their fragmentary nature, specimens
Table 6. Factor loadings from PCA of mandibular
variables of C. mayensis from the Yucatan Peninsula
and Guerrero and C. merriami.
Variable
Factor 1
Factor 2
HAC
0.873
-0.265
HCV
0.817
-0.005
BAC
0.814
-0.144
AC3
0.723
-0.290
HCP
0.701
-0.529
ML
0.672
0.560
TRD
0.383
0.813
mlL
0.285
0.858
/
(P
O Guerrero O
• C. mayensis
v C. merriami
canonical axis 1
Fig. 8. Plot of specimens of C. mayensis from Guer-
rero and the Yucatan Peninsula and C. merriami on
canonical axes 1 and 2 from discriminant function anal-
ysis of cranial variables.
from Guerrero were not included with other C.
mayensis in calculations of univariate statistics or
in other tests involving this taxon.
One specimen from Guerrero (ibunam 11039)
has an auxiliary unicuspid (RU5 or a second RU4)
in the upper right unicuspid tooth row. Supernu-
merary unicuspids previously have been reported
for Blarina (Choate, 1968), but this appears to be
a fairly rare dental abnormality in soricids. It is
much more common for individuals to lack uni-
cuspids (Choate, 1968, 1970).
Cryptotis merriami Choate, 1970
(fig. 10)
Cryptotis nigrescens merriami J. R. Choate, 30 De-
cember 1970, Univ. Kansas Publ. Mus. Nat. Hist.
19:277; E. R. Hall, 3 April 1981, The Mammals of
North America 1:63.
Cryptotis nigrescens: G. G. Goodwin, 29 May 1942,
Bull. Amer. Mus. Nat. Hist. 79:1 17; W. H. Burt and
R. A. Stirton, 22 September 1 96 1 , Misc. Publ. Mus.
Zool., Univ. Michigan 117:21.
Cryptotis parva orophila: J. R. Choate, 30 December
1970, Univ. Kansas Publ. Mus. Nat. Hist. 19:262
(part); E. R. Hall, 3 April 1981, The Mammals of
North America 1 :6 1 (part).
Cryptotis nigrescens nigrescens: J. R. Choate, 30 De-
cember 1970, Univ. Kansas Publ. Mus. Nat. Hist.
19:279 (part); E. R. Hall, 3 April 1981, The Mam-
mals of North America 1:63 (part).
Holotype— Skin and skull of adult female, U.S.
National Museum of Natural History no. 77050,
obtained by E. W. Nelson and E. A. Goldman
(collector number 8846) 21 December 1895 from
14
FIELDIANA: ZOOLOGY
Guatemala, Huehuetenango, Jacaltenango, 5400
ft.
Distribution— Cryptotis merriami occurs in the
highlands of Chiapas, Guatemala, Honduras, El
Salvador, and northern Nicaragua (fig. 1 1), with a
disjunct population in the Tilaran Highlands of
Costa Rica (fig. 14). Known elevational distribu-
tion is from 975 to 1650 m.
Description— A medium-sized Cryptotis, HB
averaging 69 (table 4); tail short, averaging 43%
(± 1, n = 23) of HB; dorsal hairs usually about 4
mm long, ranging from 3 to 5 mm long, two-band-
ed; dorsal and lateral pelage may be Olive Brown,
Mummy Brown, or Clove Brown; venter is slightly
paler than dorsum, Buffy Brown or Olive Brown.
Rostrum broad and moderately long (PL/CBL
= 43.4% ± 1.0, n = 16); interorbital area very
broad; usually only one dorsal foramen (76%, n =
21), normally located on the right frontal, medium
to very large in size; a well-developed foramen
typically present posterior to the dorsal articular
facet on one (10%, n = 10) or both (70%) sides of
the skull; normally no foramen dorsal to dorsal
articular facet (70%, n = 10); anterior process of
the petromastoid low and very narrow (fig. 1 8 A);
paroccipital process prominent; zygomatic plate
broad in proportion to CBL (12.0% ± 0.8, n =
1 6) and PL (27.5% ± 2. 1 , n = 23); anterior border
of zygomatic plate from posterior one-half of para-
style/mesostyle valley to mesostyle of M 1 , pos-
terior border from posterior one-half of mesostyle/
metastyle valley of M2 to middle of M3, and from
middle of base of maxillary process to posterior
to maxillary process; palate wide; upper tooth row
crowded; unicuspids massive; U4 normally dis-
placed medially, so that U3 and P4 in contact or
nearly so; U4 not visible in lateral view of skull;
lateral view of U3 also sometimes partially ob-
structed by P4; posterior borders of P4, M 1 , and
M2 unrecessed or only very slightly recessed; M3
with well-developed paracrista, and paracone, re-
duced precentrocrista, poorly developed and nor-
mally uncolored mesostyle and protocone, hypo-
cone absent or vestigial and incorporated into the
posterior cingulum; upper dentition quite bulbous.
Mandible large; horizontal ramus deep; coro-
noid process high (HCP/ML = 76.8% ± 3.5, n =
16) and broad, joining mandible at a steep angle;
horizontal and vertical branches of articular con-
dyle short and broad; lower sigmoid notch very
shallow; posterior border of lower incisor extends
almost to posterior border of cingulum of p4; low-
er dentition wide and long; only hypoconid in tal-
onid of m3.
•
•
2 -
•
•
v v
1 -
•
•
V
■
•
V
• •
•
• •
V
o •
V
V
o «o
V
V
V
. -
O
° o
V V
V
3 -
O
•
G
C
c
uerrero
moyensis
merriami
i i
1
1 1
-4-3-2-10 1 2 3 4
canonical axis 1
Fig. 9. Plot of specimens of C. mayensis from Guer-
rero and the Yucatan Peninsula and C. merriami on
canonical axes 1 and 2 from discriminant function anal-
ysis of mandibular variables.
Comparisons— Cryptotis mayensis— C merria-
mi is approximately the same in overall size but
has a darker pelage, which is dark brown in ap-
pearance rather than grayish brown; skull is much
wider overall, especially the rostrum, interorbital
area, braincase, and palate; upper dentition much
broader; longer mandible; shorter coronoid pro-
cess.
Cryptotis magna— C. merriami is much smaller
(HB averaging 69 vs. 86; table 4) and a much
shorter tail (averaging 43% of HB vs. 53%); skull
much smaller in all dimensions but has a relatively
broader zygomatic plate (averaging 1 2.0% of CBL
vs. 10. 1% ± 0.6); usually only one dorsal foramen;
well-developed foramen posterior to dorsal artic-
ular facet; rarely possesses foramina dorsal to dor-
sal articular facet; coronoid process lower (aver-
aging 76.8% of mandible length vs. 83.0% ± 2.4);
no entoconid in talonid of m3.
Cryptotis hondurensis—C. merriami is much
larger (HB averaging 69 vs. 61 ± 5; table 4) and
has a shorter, broader tail that is thick and rounder
at the tip (rather than tapering to a point as in C.
hondurensis); relatively shorter, broader rostrum;
wider palate; M3 less complex, lacking metacone;
dentition bulbous and upper and lower molars and
unicuspids much more massive.
Remarks— The dorsal pelage of C. merriami is
indistinctly two-banded. The proximal five-sixths
of the dorsal hairs is a medium steel gray; the distal
tip is medium brown. In two specimens from Gua-
temala (ummz 117844, 117845), the pelage is
darker overall than those from El Salvador. The
tips of the hairs of the Guatemalan specimens are
a darker brown.
WOODMAN & TIMM: SMALL-EARED SHREWS
15
5 mm
Fig. 10. Dorsal, ventral, and lateral views of the skull and lateral view of the skull and mandible of C. merriami
(mvz 130335). Reprinted with permission of the Biological Society of Washington.
Two specimens of C. merriami merit discus-
sion. A male (ummz 117111) from Santa Maria
de Ostuma, Nicaragua, is the first specimen of the
C. nigrescens complex to be identified from that
country. This specimen was identified previously
as C. parva orophila (Choate, 1970). A specimen
of unknown sex from 4.5 km NE Tilaran (ku
84365) is the first record of this species from Costa
Rica. It previously was referred to C. n. nigrescens
(Choate, 1970). Both specimens display the large
size, wide skull, broad zygomatic plate, robust
mandible, and bulbous dentition characteristic of
C. merriami. For this reason, we refer both to this
species. They clearly are not referable to C. ni-
grescens or C. parva orophila. However, both spec-
imens come from regions that are isolated geo-
graphically from the main range of C. merriami
in Chiapas and northern Central America, and
further study of larger series of specimens of this
group may prove the shrews from Nicaragua and
Costa Rica to be distinct. In addition, C. merriami
and C. nigrescens may be sympatric or parapatric
in parts of Costa Rica's Tilaran Cordillera.
To test the overall similarity of the two speci-
mens of C. merriami from Santa Maria de Ostuma
and Tilaran to C. merriami from the main part of
its range and to Costa Rican C. nigrescens, we
carried out a PCA using seven cranial variables
(ZP, IO, U1B, M2B, PL, TR, UTR) measured
from the two specimens from Santa Maria de Os-
tuma and Tilaran, 21 C. merriami, and 20 ran-
domly selected C. nigrescens from Monteverde.
All variables weighed out heavily on factor 1 , in-
dicating that it is a good estimator of overall size.
Factor 2 represented a contrast between UTR and
negatively weighted IO (table 7). A plot of factor
1 vs. factor 2 scores from this analysis (fig. 12)
shows the larger C. merriami clustering on the
right half of the graph, and the smaller C. nigres-
cens are concentrated in the left half of the graph.
The two species overlap completely on the shape
axis (factor 2). The specimen of C. merriami from
Santa Maria de Ostuma is intermediate in size,
defining the lower size limit of C. merriami. The
specimen from Tilaran plots well within the main
scatter of C. merriami.
Specimens of C. merriami from Cerro Caca-
guatique, El Salvador, are geographically isolated
from the main range of the species. However, the
lack of any morphological differentiation between
specimens from Cerro Cacaguatique and other
parts of the range argues against their being con-
sidered different taxa.
The habitat on Cerro Cacaguatique, as noted by
16
FIELDIANA: ZOOLOGY
A C. merriami
800 m contour shown
Fig. 1 1. Map of southern Mexico and northern Central America showing the distribution of C. merriami. See
also Figure 14.
Burt and Stirton (1961), was oak forest with scat-
tered pines from about 3500 ft [1065 m] up. Below
that elevation, much of the area had been cleared
for cultivation of coffee, bananas, and corn. Near
the summit at 4000 ft [1220 m], epiphytes, ferns,
and mosses occurred. Burt and Stirton (1961) re-
ported taking C. merriami with Heteromys des-
marestianus, Ototylomys phyllotis, and Peromys-
cus mexicanus along the rocky canyon slopes near
the bottom, but specimen tags indicate that the
shrews were taken from 3500 ft to the summit.
Goldman (1951) described the type locality of
C. merriami as an indigenous village at an ele-
vation of about 5400 ft [ 1 645 m] and a little below
the transition between the Humid Upper Tropical
and Lower Austral or Upper Austral life zones.
The locality is near the lower limit of cypress (Cu-
pressus), but much of the region already had been
deforested for cultivation of crops.
Few reproductive data are available for C. mer-
riami, and no pregnant or lactating females have
been recorded. Individual males collected on 23
May and 26 November had well-developed lateral
glands. Two males captured on 2 January and 1 7
December, respectively, had no evidence of glan-
dular development.
Cryptotis nigrescens (J. A. Allen, 1895)
(fig. 13)
Marina (Soriciscus) nigrescens J. A. Allen, 8 Novem-
ber 1895, Bull. Amer. Mus. Nat. Hist. 7:339.
Blarina micrura: J. A. Allen, 22 September 1 893, Bull.
Amer. Mus. Nat. Hist. 5:238.
Table 7. Factor loadings from PCA of cranial vari-
ables of C. merriami and C. nigrescens.
Variable
Factor 1
Factor 2
PL
0.932
-0.112
TR
0.920
0.053
M2B
0.920
-0.231
U1B
0.909
-0.076
IO
0.869
-0.288
UTR
0.793
0.549
ZP
0.776
0.196
WOODMAN & TIMM: SMALL-EARED SHREWS
17
z -
1 -
V
V
V
9
V
°°o
o
o
0 -
V
v v
o° ° o
-1 -
V
%
• o o
9 o
o
-2 -
?
V
o
O
C. merriam
•
Nicara
gua
-3 -
V
C. nig
rescens
T
Tilaran
i
i
1 I
factor 1
Fig. 12. Plots of principal components factor scores
of C. merriami and C. nigrescens on factor axes 1 and 2.
Blarina nigrescens: C. H. Merriam, 31 December 1895,
N. Amer. Fauna 10:31.
C[ryptotis] nigrescens: G. S. Miller, Jr., 31 October
1911, Proc. Biol. Soc. Washington 24:222.
Cryptotis nigrescens: G. G. Goodwin, 10 December
1944, Amer. Mus. Novitates 1267:1; G. G. Good-
win, 31 December 1946, Bull. Amer. Mus. Nat.
Hist. 87:288; H. W. Setzer, 29 September 1950, J.
Washington Acad. Sci. 40:300; E. R. Hall and K.
R. Kelson, 31 March 1959, The Mammals of North
America 1:63 (part); C. O. Handley, Jr., 22 Novem-
ber 1966, Checklist of the mammals of Panama, in
R. L. Wenzel and V. J. Tipton (eds.), Ectoparasites
of Panama, p. 756.
Cryptotis zeteki: H. W. Setzer, 29 September 1950, J.
Washington Acad. Sci. 40:299; G. G. Goodwin, 28
June 1954, Amer. Mus. Novitates 1677:2; E. R.
Hall and K. R. Kelson, 31 March 1959, The Mam-
mals of North America 1:62.
Cryptotis tersus: G. G. Goodwin, 28 June 1954, Amer.
Mus. Novitates 1677:1; E. R. Hall and K. R. Kel-
son, 3 1 March 1959, The Mammals of North Amer-
ica 1:64.
Cryptotis nigrescens zeteki: C. O. Handley, Jr., 22 No-
vember 1966, Checklist of the mammals of Pana-
ma, in R. L. Wenzel and V. J. Tipton (eds.), Ec-
toparasites of Panama, p. 756.
Cryptotis parva orophila: J. R. Choate, 30 December
1970, Univ. Kansas Publ. Mus. Nat. Hist. 19:262
(part).
Cryptotis nigrescens nigrescens: J. R. Choate, 30 De-
cember 1970, Univ. Kansas Publ. Mus. Nat. Hist.
19:279 (part); E. R. Hall, 3 April 1981, The Mam-
mals of North America 1:63 (part).
Holotype— Skin and skull of subadult, sex un-
known, American Museum of Natural History no.
9591/7952, collected by G. K. Cherrie (collector
number 2004) 5 September 1 89 1 from Costa Rica,
San Jose Province, San Isidro.
Distribution— Occurs in highlands above 800
m in Costa Rica and western Panama (fig. 14). In
Costa Rica, C. nigrescens is known from the Ti-
laran, Central, and Talamanca cordilleras; in Pan-
ama, it is known from the Chiriqui Cordillera. The
known elevational distribution in Costa Rica is
from 870 to 2865 m; in Panama, it is 820 to
2150 m.
Description— A small to medium Cryptotis, HB
averaging 68 (table 4); tail short, averaging 43%
(± 5, n = 70) of HB; snout in fluid-preserved
specimens short and broad; dorsal hairs approx-
imately 4.5 mm long, ranging from 4 to 5 mm,
two-banded; pelage coloration varies somewhat
geographically; dorsal and lateral pelage Mummy
Brown or Clove Brown; ventral pelage slightly pal-
er, Buffy Brown, Saccardo's Umber, Olive Brown,
or Mouse Gray.
Rostrum narrow and of normal length (PL/CBL
= 42.6% ± 0.8, n = 70); interorbital area of mod-
erate breadth; often two dorsal foramina (68%, n
= 59), small to large size, positioned close to suture
between frontals; normally no foramen posterior
to the dorsal articular facet (88%, n = 57); usually
a well-developed foramen dorsal to external capit-
ular facet on one (2%, n = 57) or both (93%) sides
of the skull, but foramen may be minute (3%);
anterior process of the petromastoid very low and
very narrow (fig. 18B); paroccipital process prom-
inent; zygomatic plate narrow in proportion to
CBL ( 1 0.5% ± 0.9, n = 75) and PL (24.8% ±2.1);
anterior border of zygomatic plate from mesostyle
to mesostyle/metastyle valley of Ml, posterior
border at posterior one-half of mesostyle/meta-
style of M2 to middle of M3, and usually at middle
of base of maxillary process, but can occur from
anterior one-half of this process to posterior to the
process; palate moderately wide; upper tooth row
not especially crowded; U4 normally in line of
unicuspid tooth row, preventing contact between
U3 and P4; U4 usually obscured by P4 in lateral
view, but often partly visible; posterior borders of
P4, Ml, and M2 unrecessed or only slightly re-
cessed; M3 with well-developed paracrista, and
paracone, reduced precentrocrista and mesostyle,
poorly developed, but often colored postcentro-
crista and metacone (but postcentrocrista short with
mesostyle and metacone closely associated), pro-
tocone present and often colored, poorly devel-
oped hypocone often present; dentition not bul-
bous.
Mandible of moderate length and breadth; cor-
onoid process low (HCP/ML = 71.7% ± 2.6, n =
70) and narrow, joins mandible at a steep angle;
18
FIELDIANA: ZOOLOGY
Fig. 13. Dorsal, ventral, and lateral views of the skull and lateral view of the skull and mandible of C. nigrescens
(ku 143389). Scale bar = 5 mm.
articular condyle relatively short and narrow; low-
er sigmoid notch very shallow; posterior border
of lower incisor almost to posterior border of cin-
gulum of p4; vestigial entoconid occasionally pres-
ent (38%, n = 50) in talonid of m3.
Comparisons— Cryptotis nigrescens is the only
member of the C. nigrescens complex sometimes
possessing an entoconid in the talonid of m3.
Cryptotis mayensis—C. nigrescens has a much
darker pelage, dark brown in appearance rather
than grayish brown; skull is less robust; rostrum
not as broad; anterior process of the petromastoid
lower; foramen associated with ventral branch of
sinus canal normally absent; foramen dorsal to
dorsal articular facet of skull normally present;
much more likely to have two dorsal foramina,
one on each frontal; relatively and absolutely nar-
rower zygomatic plate, anterior border of which
is positioned more posteriorly relative to upper
tooth row; palate relatively wider; upper dentition
not as massive, unicuspids not as broad; upper
molars relatively longer; unicuspid tooth row less
crowded; mandible relatively longer and more
slender; coronoid process shorter and narrower;
branches of articular process narrower; lower mo-
lars relatively longer; hypoconid sometimes pres-
ent on m3.
Cryptotis magna— C. nigrescens is much smaller
(HB averaging 68 vs. 86; table 4) and has a much
shorter tail (averaging 43% of HB vs. 53%); skull
much smaller in all dimensions; foramen associ-
ated with ventral branch of sinus canal normally
absent; coronoid process much lower (averaging
71.7% of ML vs. 83.0% ± 2.4); dentition not bul-
bous.
Cryptotis merriami—C. nigrescens has a gen-
erally darker pelage; skull is not nearly as robust,
much narrower overall, especially the rostrum, in-
terorbital area, braincase, and palate; foramen as-
sociated with ventral branch of sinus canal nor-
mally absent; anterior process of the petromastoid
lower; relatively and absolutely narrower zygo-
matic plate; upper dentition absolutely smaller,
narrower, less massive; unicuspid row generally
less crowded; M3 slightly more developed, pos-
sessing minute postcentrocrista and metacone;
mandible shorter and more slender; coronoid pro-
cess shorter and narrower; branches of articular
WOODMAN & TIMM: SMALL-EARED SHREWS
19
10
100 km
• C. nigrescens
A C. merriami
800 m contour shown
&
84
82
Fig. 14. Map of Costa Rica and western Panama showing the distribution of C. nigrescens.
process narrower; lower molars narrower; hypo-
conid sometimes present on m3.
Cryptotis hondurensis—C. nigrescens is larger
(HB averaging 68 vs. 61 ± 5; table 4); shorter tail
(averaging 41% of HB vs. 46%); foramen associ-
ated with ventral branch of sinus canal normally
absent; M3 less complex, lacking metacone; cor-
onoid process of mandible lower; vestigial ento-
conid sometimes present on m3.
Remarks— Viewed anteriorly, the pelage of C.
nigrescens is very reflective. The resulting sheen
distorts the hues, making colors appear paler, es-
pecially on the venter. Dorsal pelage of C. nigres-
cens appears two-banded. The proximal three-
quarters of the hairs is medium steel gray; the
remaining distal portion is dark brown. In general,
specimens from Costa Rica have slightly darker
pelage, from Mummy Brown to Bone Brown and
Clove Brown dorsally with a Buffy Brown or Olive
Brown venter. Specimens from western Panama
are Mummy Brown dorsally with a venter of Buffy
20
Brown, Saccardo's Umber, or Sepia in the Chiriqui
highlands. The darkest specimens are two shrews
(usnm 520692, 520693) from Fish Camp, Bocas
del Toro Province, Panama, which are Blackish
Brown 2 dorsally and Deep Grayish Olive ven-
trally. They are in contrast to the slightly paler
specimens from other localities in western Pana-
ma. These two specimens from Fish Camp also
have very long tails (52-53% of HB). However,
both are young animals with nearly unworn den-
tition, and this may account for the differences
between them and other C. nigrescens from Pan-
ama.
There is a fair amount of variation between the
large series of C. nigrescens from Monteverde,
Costa Rica, and those from near San Felix (in-
cluding Cerro Bollo), Panama. Specimens from
Monteverde and from San Felix differ greatly in
size (table 1). In addition, the presence of a hy-
poconid on m3 is much more prevalent in the
Panamanian series (7 1 %, n = 1 4) than in the Mon-
FIELDIANA: ZOOLOGY
teverde specimens (25%, n = 32), and two dorsal
foramina are present more often in specimens from
near San Felix (94%, n = 1 6) than in those from
Costa Rica (58%, n = 43). Further collecting and
study of Panamanian and Costa Rican Cryptotis
may show that C. nigrescens is a complex of species.
Specimens of C. nigrescens from Cinchona and
Monteverde, Costa Rica, and from Santa Clara
and Boquete Trail on Cerro Punta, Panama, pre-
viously were identified as C. parva orophila
(Choate, 1970). The small size of Costa Rican C.
nigrescens and the simple M3 can make the two
species difficult to separate. Cryptotis nigrescens
can be distinguished by its much darker pelage
coloration, on both venter and dorsum, and by its
relatively longer tail (43% ± 5 of HB in C. ni-
grescens vs. 33% ± 4, n = 11). In addition, C.
nigrescens has a longer zygomatic plate, the pos-
terior margins of P4, M 1 , and M2 are not as deeply
recessed, M3 possesses a reduced metacone, and
there is sometimes a hypoconid on m3. Our cor-
rection of these identifications means that C. parva
presently is not known south of Costa Rica.
Setzer (1950) gave the location of Cerro Punta,
the type locality of C. zeteki (a junior synonym of
C. nigrescens) as 8°42'N, 82°48'W. Based on re-
cent maps and gazetteers, the community is lo-
cated at 8°34'N, 81 °50'W.
Hall's (1981, p. 63) map of the distribution of
C. nigrescens implies that the species is found
throughout Costa Rica, including low elevations
along both coasts (with the exception of the Nicoya
and Osa peninsulas), and throughout much of cen-
tral and eastern Panama. Cryptotis nigrescens, as
we understand it today, inhabits only higher ele-
vations, and it is restricted to the Tilaran, Central,
and Talamanca Cordilleras of Costa Rica and the
Chiriqui Cordillera of Panama. At Monteverde
the species occurs in cloud forest, in forest frag-
ments, and in pastures from 870 to 1660 m, and
in western Panama it has been taken in cloud for-
est, disturbed cloud forest, elfin woodland, and
drier Pacific slope forest from 1275 to 1856 m.
The known upper elevational range for C. nigres-
cens is 2865 m. This probably approaches the true
upper range for the species, because collections of
shrews from higher elevations do not include this
species. Cryptotis gracilis is known from 2435 to
3536 m in the Talamancan and Chiriqui high-
lands, and C.jacksoni. a member of the C. gracilis
complex, has been taken from 2350 to 3 1 80 m on
Volcan Irazu (Woodman, 1992). That this is not
a simple case of competitive exclusion is suggested
by the fact that C nigrescens and C. gracilis over-
lap in their elevational range and that both have
been collected from the same locality in Panama
(R. Pine, pers. comm.).
In the Monteverde area, we found C. nigrescens
in a variety of habitat types including Tropical
Premontane Rain Forest, Tropical Premontane
Moist Forest, Tropical Premontane Wet Forest,
and Tropical Lower Montane Wet Forest. Com-
mon small mammals in these habitats include
Peromyscus nudipes, Heteromys desmarestianus,
Scotinomys teguina, and Reithrodontomys creper
and Oryzomys albigularis at the higher elevations.
A second, undescribed species of small-eared shrew
occurs at the higher elevations in the Monteverde
Cloud Forest. This is a much larger shrew and is
a member of the C. gracilis complex. At this point,
we cannot say whether these two species are truly
syntopic, although they are sympatric in this area.
Cryptotis nigrescens probably is abundant at Mon-
teverde but is infrequently observed and difficult
to capture. During several weeks of fieldwork in
1989, we had over 1,570 trapnights using a com-
bination of Sherman live traps, Museum Specials,
and common snap traps baited with a mixture of
suet, seeds, raisins, and vanilla, and over 500 trap-
nights using pitfalls with drift fences. Of a total of
24 shrews we obtained, none were captured in
baited traps, 8 were taken in our pitfalls, 5 more
were captured in pitfalls set by entomologists in a
study apart from ours, 3 were brought in by do-
mestic cats, and 8 were encountered dead on a
trail or road by local residents prior to our arrival.
Few reproductive data are available for C. ni-
grescens. A female (usnm 516623) from east of
Cerro Pando was recorded as "recently lactating"
on 16 March 1973, and a lactating female (usnm
5 1 6620) was collected at Santa Clara, western Pan-
ama, on 25 March 1972. We captured an adult
female (ku 143389) with three embryos (crown-
rump length = 4 mm) at Monteverde on 12 May
1 989 and another with an enlarged uterus on 2 1
May (ku 143396), suggesting that it had recently
given birth. Two females (usnm 54 1 044, 54 1 030),
pregnant with a single embryo each (crown-rump
length = 15 mm, 5 mm), were taken on 2 and 4
July 1980, respectively, 24-24.5 km NNE of San
Felix, western Panama. Females without embryos
have been captured between 9 and 23 May at Mon-
teverde (n = 6) and on 1 3 and 26 June in Panama
(n = 2). Our inspection of male C. nigrescens shows
well-developed lateral glands on 20 individuals
collected from 9 May through 1 July. Individual
males without well-developed lateral glands were
collected on 26 February, 7 March, 1 1 March, 3
WOODMAN & TIMM: SMALL-EARED SHREWS
21
Fig. 15. Dorsal, ventral, and lateral views of the skull and lateral view of the skull and mandible of C. mera
(usnm 337969). Scale bar = 5 mm.
April, 28 July, and 4 July. On 9 and 18 May we
recorded individual males from Monteverde with
melanistic sheaths surrounding the testes; both an-
imals had well-developed lateral glands and en-
larged testes (3.5 x 2.5 mm and 4 x 2.5 mm,
respectively).
A young adult female (ku 143393) captured at
Monteverde on 15 May 1989 lacks digits on its
right forefoot. The vestigial first digit is the only
one bearing a claw, and this is poorly formed. The
shape of the foot suggests that the digits were never
fully developed during ontogeny rather than lost
later due to an accident.
Cryptotis mera Goldman, 1912
(fig. 15)
Cryptotis merus E. A. Goldman, 20 September 1912,
Smiths. Misc. Coll. 60(2): 17; E. A. Goldman, 24
April 1920, Smiths. Misc. Coll. 69(5): 171; H. W.
Setzer, 29 September 1950, J. Washington Acad.
Sci. 40:300; G. G. Goodwin, 28 June 1954, Amer.
Mus. Novitates 1677:2.
Cryptotis mera: E. R. Hall and K. R. Kelson, 3 1 March
1959, The Mammals of North America 1:61.
Cryptotis nigrescens mera: C. O. Handley, Jr., 22 No-
vember 1966, Checklist of the mammals of Pana-
ma, in R. L. Wenzel and V. J. Tipton (eds.), Ec-
toparasites of Panama, p. 756.
Cryptotis nigrescens nigrescens: J. R. Choate, 30 De-
cember 1970, Univ. Kansas Publ. Mus. Nat. Hist.
19:279 (part); E. R. Hall, 3 April 1981, The Mam-
mals of North America 1:63 (part).
Holotype— Skin and skull of an adult male,
U.S. National Museum of Natural History no.
178976, collected by E. A. Goldman (original
number 2 1 669) 2 May 1912 from Panama, Darien
Province, Mount Pirri [Cerro Pirre], near head of
Rio Limon, 4500 ft.
Distribution— Cerro Tacarcuna and Cerro Mali
in the eastern Serrania de Darien, and Cerro Pirre
in the Serrania de Pirre (fig. 1 7). The species un-
doubtedly occurs in Colombia, but all specimens
currently known are from the Panamanian side of
the border. The recorded elevational distribution
of the species is 1370-1525 m.
22
FIELDIANA: ZOOLOGY
Description— A medium-sized Cryptotis, HB
averaging 69 (table 4); tail short, averaging 39%
(± 2, n = 6) of HB; dorsal hairs approximately 4
mm long, ranging from 3.5 to 4.5 mm, indistinctly
two-banded; dorsal and lateral pelage Mummy
Brown; ventral pelage slightly paler, Saccardo's
Umber or Mouse Gray.
Rostrum of normal length (PL/CBL = 42.4% ±
1 . 1 , n = 5); interorbital area of moderate breadth;
often two dorsal foramina (66%, n = 6), not nor-
mally equal in size, positioned close to suture be-
tween frontals; normally no foramen posterior to
the dorsal articular facet ( 1 00%, n = 4); usually a
well-developed foramen dorsal to dorsal articular
facet on both sides of the skull (75%, n = 4), but
foramen may be minute (25%); anterior process
of petromastoid very low and very narrow (fig.
18B); paroccipital process prominent; zygomatic
plate narrow in proportion to CBL (1 1.0% ± 0.7,
n = 5) and PL (26.0% ± 1.3, n = 6); anterior border
of zygomatic plate at mesostyle/metastyle valley
of Ml, posterior border at metastyle of M2 to
middle of M3, and from anterior edge to posterior
edge of maxillary process; palate moderately wide;
upper tooth row crowded; unicuspids massive; U4
normally displaced medially, so U3 and P4 in con-
tact or nearly so; U4 usually not visible in lateral
view; lateral view of U3 also often partly ob-
structed by P4; posterior borders of P4, M 1 , and
M2 only slightly recessed; M3 with well-developed
paracrista and paracone, reduced precentrocrista,
mesostyle and postcentrocrista poorly developed
and normally uncolored, protocone present and
often colored, hypocone poorly developed, ap-
pearing as a posterior cingulum; dentition quite
bulbous.
Mandible of moderate length and breadth; cor-
onoid process high (HCP/ML = 76.3% ± 4.8, n
= 6), but narrow, joins mandible at fairly steep
angle; articular condyle relatively short and broad;
lower sigmoid notch very shallow; posterior bor-
der of lower incisor almost to posterior border of
cingulum of p4; only hypoconid in talonid of m3.
Comparisons— Cryptotis mayensis — C. mera
has a much darker pelage, dark brown in appear-
ance rather than grayish brown; skull is much
shorter, but interorbital area much broader; fo-
ramen associated with ventral branch of sinus ca-
nal absent; palate relatively wider; anterior process
of the petromastoid lower; foramen associated with
ventral branch of sinus canal normally absent; fo-
ramen dorsal to dorsal articular facet of skull nor-
mally present; much more likely to have two dor-
sal foramina, one on each frontal; mandible more
slender; coronoid process shorter and much nar-
rower; lower molars relatively longer.
Cryptotis magna— C. mera is much smaller (HB
averaging 69 vs. 86; table 4) and has a much short-
er tail (averaging 39% of HB vs. 53%); skull much
smaller in all dimensions; coronoid process lower
(averaging 76.3% of mandible length vs. 83.0 ±
2.4); no entoconid in talonid of m3.
Cryptotis merriami—C. mera is much smaller
in nearly all cranial dimensions; foramen associ-
ated with ventral branch of sinus canal absent;
relatively broader palate; anterior process of the
petromastoid lower; mandible not as deep; coro-
noid process much narrower.
Cryptotis hondurensis—C. mera is larger (HB
averaging 69 vs. 6 1 ± 5; table 4) and has a shorter
tail (averaging 39% of HB vs. 46%); foramen as-
sociated with ventral branch of sinus canal absent;
M3 less complex, lacking metacone; dentition bul-
bous.
Cryptotis nigrescens—C. mera has a relatively
wider skull, especially in the interorbital area, and
palate; relatively longer zygomatic plate; more
crowded upper tooth row; relatively shorter, deep-
er mandible; no hypoconid on m3.
Remarks— Choate (1970) considered C. mera
a junior synonym of C. nigreseens nigrescens. At
the time of his study, however, he had a total of
only 29 specimens at hand. Recent collection of
large series of C. nigrescens from Costa Rica and
western Panama permitted us to make more ex-
tensive comparisons, and we recognize C. mera as
a distinct species.
Dorsal pelage of C. mera is two-banded and is
generally similar to that of C. nigrescens from
western Panama in color.
Cryptotis mera is known from cloud forest on
two mountain areas isolated from each other by
lowlands below 200 m. Specimens from the Ser-
rania de Pirre have a shorter coronoid process
(HCP/ML = 71%, n = 2) than those from the
Serrania de Darien (79% ± 3, n — 4), but are
otherwise indistinguishable. Populations from
these two areas may prove to be distinct once ad-
ditional specimens are available, but at this time
we consider them to be conspecific.
The holotype and two topotypes of C. mera were
collected between 4500 and 5000 ft [1370 and
1 525 m] in the Serrania de Pirre. Goldman (1912)
described the region as covered with unbroken
forest and having a seasonal but very heavy annual
rainfall. Although little actual rain falls during the
dry season, clouds normally envelop the upper
elevations, providing moisture year-round.
WOODMAN & TIMM: SMALL-EARED SHREWS
23
Fig. 16. Dorsal, ventral, and lateral views of the skull and lateral view of the skull and mandible of C. colombiana
(fmnh 69816). Scale bar = 5 mm.
Reproductive data for C. mera are scarce. Two
females (usnm 337966, 337969) from Cerro Mali
and Cerro Tacarcuna were lactating on 12 Feb-
ruary and 1 1 March 1 964, respectively.
Although Goldman (1912) originally published
this species name as "merits," the correct spelling
is "mera." Cryptotis is feminine, dictating that the
feminine ending be used on all adjectival species
names used in combination with it (Woodman,
1993).
Cryptotis colombiana, new species
(fig. 16)
Holotype— Skin and skull of adult female, Field
Museum of Natural History no. 69816, collected
15 October 1950 by Philip Hershkovitz (original
no. 4723). Skull nearly complete, but with a punc-
ture hole in the dorsal braincase; skin in very good
condition, but missing a small patch of fur on left
venter, some fur matted.
Type Locality— Colombia; Central Cordillera;
Antioquia Dept., Sonson; 15 km E of Rio Negrito;
1750 m.
Distribution— At present, known only from
type locality (fig. 1 7).
Diagnosis— A small to medium-sized Cryptotis
with very dark fur, a short tail, broad rostrum,
palate, and interorbital area, foramen dorsal to
dorsal articular facet but not posterior to this facet,
petromastoid with broad anterior process and with
large foramen in tympanic process, narrow zygo-
matic plate, crowded tooth row, bulbous and un-
recessed dentition, simple M3, long mandible with
a broad coronoid process that joins the horizontal
ramus at nearly a right angle, short and broad
articular condyle with no notch between the ar-
ticular facets, shallow lower sigmoid notch, and
lacking entoconid in talonid of m3.
Description of Holotype— A small to medi-
um-sized Cryptotis, HB = 76 (table 4); tail short,
36% of HB; dorsal hairs approximately 4-5 mm
long, two-banded; dorsal and lateral pelage Olive
Brown to Fuscous; ventral pelage only slightly pal-
er, Bufiy Brown to Hair Brown.
Rostrum broad and of normal length (PL/CBL
= 44%); interorbital area broad; two large dorsal
foramina, close to sagittal suture; no foramen pos-
24
FIELDIANA: ZOOLOGY
# C. mera
Jf C. colombiana
1000 m contour shown
Fig. 17. Map showing the distribution of C. mera along the Panama/Colombia border and the type locality of
C. colombiana in the Central Cordillera of Colombia.
terior to the dorsal articular facet; a moderately
well-developed foramen dorsal to dorsal articular
facet on both sides of the skull; anterior process
of the petromastoid high and wide (fig. 18C); large
foramen on posterior edge of tympanic process of
petromastoid; paroccipital process low; zygomatic
plate narrow in proportion to CBL (10.1%) and
PL (22.7%); anterior border of zygomatic plate at
posterior of mesostyle/metastyle valley of Ml,
posterior border at parastyle of M3 and anterior
to posterior edge of base of maxillary process; pal-
ate wide; upper tooth row crowded; U4 reduced
and peglike; U4 in line with other unicuspids, but
too small to prevent contact between U3 and P4;
U4 and corner of U3 obscured by P4 in lateral
view; posterior borders of P4, Ml, and M2 only
slightly recessed; M3 with well-developed pa-
racrista and paracone, reduced precentrocrista that
joins the posterior cingulum; protocone of M3
poorly developed and uncolored; hypocone of M3
absent or part of posterior cingulum; dentition bul-
bous.
Mandible relatively long and of moderate
breadth; coronoid process broad and low (HCP/
ML = 69.6%) joining mandible at nearly a right
angle; articular condyle distinctive: short and
broad, lacking a lingual notch between the dorsal
and ventral articular surfaces; lower sigmoid notch
very shallow; posterior border of lower incisor al-
most to posterior border of posterior cingulum of
p4; only hypoconid in talonid of m3.
Comparisons— Cryptotis colombiana is larger
than any of the four species previously referred to
C. nigrescens, and it differs from those species, C.
hondurensis, and C. magna in having a shorter
tail, a much broader and higher anterior process
of the petromastoid (fig. 1 8); a large foramen on
posterior edge of tympanic process of petromas-
toid; a lower, less prominent paroccipital process;
and no distinct notch between the articular facets
of the articular condyle of the mandible.
Cryptotis mayensis—C. colombiana has a much
darker pelage, dark brown in appearance rather
than grayish brown; a moderately well-developed
WOODMAN & TIMM: SMALL-EARED SHREWS
25
I 1
1 mm
Fig. 18. Ventral view of the right tympanic region of the skulls of C. merriami (A, ummz 1 17845), C. nigrescens
(B, ku 143385), and C. colombiana (C, fmnh 69816), illustrating the location and relative development of the anterior
process of the petromastoid (ap) and the paroccipital process (pp), and the foramen on the posterior edge of the
tympanic process of the petromastoid (pf).
foramen dorsal to the dorsal articular facet, but
no foramen posterior to this facet; relatively nar-
rower zygomatic plate; shorter unicuspid tooth row
and longer molariform tooth row; broader inter-
orbital area and much broader palate; smaller uni-
cuspids; much shorter and much narrower coro-
noid process; longer mandible.
Cryptotis magna — C. colombiana is much
smaller (HB = 76 vs. 86; table 4) and has a much
1 -
° ~1 -
o
o
-3 -
o
• •
oo "o *# #
• • •
O C. mera •
• C. nigrescens
v C. colombiana
-3
factor 1
Fig. 19. Plot of principal components factor scores
of C. colombiana, C. mera, and C. nigrescens on factor
axes 1 and 2.
shorter tail (36% of HB vs. 53%); skull much
smaller in all dimensions; coronoid process much
lower (averaging 69.6% of ML vs. 83.0% ± 2.4);
no entoconid in talonid of m3.
Cryptotis merriami— C. colombiana has a mod-
erately well-developed foramen dorsal to the dor-
sal articular facet, but no foramen posterior to this
facet; wider palate; relatively narrower zygomatic
plate; longer molariform tooth row; much shorter
and narrower coronoid process; longer mandible.
Cryptotis hondurensis—C. colombiana is much
larger (HB = 76 vs. 61 ± 5; table 4) and has a
relatively shorter tail (36% of HB vs. 46%); fora-
men associated with ventral branch of sinus canal
normally absent; narrower zygomatic plate (10.1
vs. 1 1 .5); M3 less complex, lacking metacone; cor-
onoid process of mandible lower; dentition bul-
bous.
Cryptotis nigrescens— C. colombiana differs in
its much broader interorbital area and palate; longer
molariform tooth row; somewhat broader coro-
noid process; dentition bulbous.
Cryptotis mera—C. colombiana has a broader
interorbital area and somewhat broader palate;
relatively narrower zygomatic plate; much longer
molariform tooth row; shorter, broader coronoid
process; longer mandible.
Remarks— Cryptotis colombiana is the only
species of the C. nigrescens group that is known
to occur in South America, although C. mera un-
26
FIELDIANA: ZOOLOGY
doubtedly occurs in Colombia along the Pana-
manian border. Based on the structure of the man-
dible and characteristics of the skull and teeth,
both species clearly are more closely related to the
C. nigrescens group than to any of the South Amer-
ican species. Cryptotis colombiana and C. mera
are considerably smaller than any other described
South American Cryptotis, and they are also the
only two South American shrews with a short,
broad articular condyle, and a nearly right angle
junction between the coronoid process and the
body of the mandible. All other described species
of Cryptotis from South America have a high, nar-
row articular process and a mandible with a wide
internal angle between the coronoid process and
the horizontal ramus. In addition, Cryptotis co-
lombiana is distinctive in being the only species
known to lack a notch between the two articular
facets of the articular condyle and a high, very
broad anterior process of the petromastoid. The
occurrence of C colombiana on the Central Cor-
dillera of Colombia suggests that it, or another
closely related species, is likely to occur at high
elevations in the Western Cordillera, which is lo-
cated between the known geographic ranges of C.
colombiana and C. mera.
To test the overall similarity of C colombiana
to that of the other, geographically closest mem-
bers of the C nigrescens complex, we carried out
PCA using eight cranial variables (CBL, IO, U1B,
M2B, PL, TR, UTR, MTR) measured from the
holotype of C. colombiana, 5 C. mera, and 22 C.
nigrescens from the vicinity of San Felix and Cerro
Bollo in the easternmost part of that species' known
range. Factor axis 1 from the PCA estimated over-
all size, and factor 2 represented a contrast be-
tween a heavily weighted MTR and negatively
weighted U 1 B (table 8). In a plot of factor scores
on factor axis 1 vs. factor axis 2 (fig. 19), C. colom-
biana is the largest of the three species, whereas
its closest geographic neighbor, C. mera, is the
smallest. Individuals of C. nigrescens, which occur
furthest to the northwest geographically, overlap
both of the other two species, but in general C.
nigrescens is intermediate in size. Along the shape
axis (factor 2), the three species overlap complete-
ly, reinforcing an impression of conservation of
form within the C. nigrescens complex.
No reproductive data are available for C colom-
biana.
Other species of mammals collected by Hersh-
kovitz along an elevational transect from 1 500 to
2100 m at the type locality of C colombiana in-
clude Dasyprocta punctata, Akodon sp., Neacomys
Table 8. Factor loadings from PCA of cranial vari-
ables of C. colombiana, C. mera. and C. nigrescens.
Variable
Factor 1
Factor 2
UTR
0.948
-0.012
TR
0.918
0.257
PL
0.895
0.070
CBL
0.866
0.177
M2B
0.765
-0.265
IO
0.708
-0.360
U1B
0.695
-0.528
MTR
0.509
0.754
tenuipes, Oryzomys albigularis, O. alfaroi, Rei-
throdontomys mexicanus, Rhipidomys latimanus,
Thomasomys laniger, and Sciurus pucheranii.
Etymology— The specific epithet colombiana
is derived from Colombia, the country of origin
of the holotype and the only region in which this
shrew is known to occur.
Acknowledgments
We thank Guillermo Canessa M., Miguel Ro-
driguez R., and Carlos Salas A., Servicio de Vida
Silvestre, San Jose, for issuing permits in Costa
Rica to make our work possible. William Aspinal
and the Tropical Science Center provided per-
mission to work in the Monteverde Cloud Forest
Reserve. Mr. and Mrs. John Campbell of Mon-
teverde kindly permitted us and numerous other
biologists to work on their farm. Steve Ashe, Rob-
ert Brooks, Barb Clauson, Cathy Langtimm, Rich-
ard Leschen, and Richard and Meg LaVal assisted
us in the field in Costa Rica and obtained numer-
ous specimens for us that greatly facilitated our
work. Special thanks go to Fernando and Lilian
Cervantes for opening up their home in Mexico
City to us during our work there and for facilitating
specimen loans. William Lopez-Forment C. gra-
ciously guided fieldwork in Mexico. Thor Holmes
skillfully prepared many of the delicate specimens
used in our work. We thank Amy Lathrop for the
cranial drawings used as Figures 4, 1 3, 1 5, and 1 6
and Anne Musser for Figure 10. Elizabeth Montes
G. and Adrian Nieto M. improved the Spanish
translation of our abstract. We thank the following
curators and collections managers for loans or for
permission to examine specimens under their care:
GuyG. Musser (amnh), Robert C. Dowler (asnhc),
Ticul Alvarez and Sergio Ticul Alvarez (encb),
Bruce D. Patterson (fmnh), Fernando Cervantes
R. (ibunam), Reinaldo Aguilar (iNBio), Linda J.
WOODMAN & TIMM: SMALL-EARED SHREWS
27
Barkley, Sarah B. George, and John E. Heyning
(lacm), Maria E. Rutzmoser (mcz), Jerry R. Choate
(mhp), Elmer C. Birney (mmnh), Pablo Sanchez
(mncr), James L. Patton and Barbara R. Stein
(mvz), Mark D. Engstrom (rom), George D.
Schrimper (sui), Jose Ramirez-Pulido (uami), Phil
Myers (ummz), Gustavo A. Cruz (unah), and Mi-
chael D. Carleton, Alfred L. Gardner, Linda K.
Gordon, and Robert Fisher (usnm). Jaime E. Pe-
faur (Universidad de los Andes, Merida) and Al-
berto Cadena (Universidad Nacional de Colom-
bia, Bogota) graciously searched their collections
for additional specimens of C. colombiana. Bruce
D. Patterson confirmed identifications of mam-
mals collected with C. colombiana at the type lo-
cality. Robert C. Dowler, Mark D. Engstrom, Phil-
ip Hershkovitz, and Ronald H. Pine shared their
unpublished information and pertinent specimens
with us from their ongoing investigations. Mark
D. Engstrom, Robert S. Hoffmann, Norman A.
Slade, and an anonymous reviewer provided help-
ful comments that substantially improved our
manuscript. Portions of this project were funded
by the E. Raymond and Mary Hall Fund, the Glen
C. Rinker Fund, and the Panorama Society Fund
of the University of Kansas Museum of Natural
History, National Geographic Society, Organiza-
tion for Tropical Studies, Rice Foundation of Chi-
cago, and University of Kansas General Research
Fund. Woodman was supported by the KU Fel-
lowship Program for Latin American Studies.
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FIELDIANA: ZOOLOGY
Woodman, N. 1992. Biogeographical and evolution-
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Specimens Examined
Specimens in the following list marked with an
asterisk (*) are cranial remains recovered from owl
pellets. Those marked with double asterisks (**)
are from archeological or paleontological sites.
Cryptotis colombiana (1).— COLOMBIA: AN-
TIOQUIA: Sonson, 15 km E of Rio Negrito, 1750
m (1 fmnh— holotype).
Cryptotis hondurensis (3). -HONDURAS:
FRANCISCO MORAZAN: 12 km WNW El Za-
morano, Cerro Uyuca, 1680 m (1 ku— holotype);
near San Juancito mines, 1700 m (1 fmnh); La
Rosario, San Juancito, La Tigra National Park (1
unah).
Cryptotis magna (49). -MEXICO: O AX AC A:
La Esperanza, 1430 m (1 uami); Santiago Comal-
tepec, 1 km N La Esperanza, 1525 m (1 ibunam);
Santiago Comaltepec, 1 1 km SW La Esperanza,
2000 m (25 ibunam); Vista Hermosa, 1600 m (2
ku); 3.5 km N Vista Hermosa, 1360 m (1 ku); 2.3
km S Vista Hermosa, 1560 m (1 ku); 12 km S
Vista Hermosa, 1920 m (4 ku); 21 km S Vista
Hermosa, 2080 m (1 ku); 28.6 km S Vista Her-
mosa, 2350 m (1 ku); 31.6 km S Vista Hermosa,
Cerro Pelon, 2650 m (1 ku); 3.4 mi SSW Vista
Hermosa, 6200 ft (1 ku); 6.5 mi SSW Vista Her-
mosa, 7100 ft (3 ku); 12 mi SSW Vista Hermosa,
9300 ft (1 ku); Llano de las Flores, 2800-2900 m
(4 ku); Atepec, 13 km W Atepec, 2820 m (1
ibunam); Mt. Zempoaltepec (1 usnm).
Cryptotis mayensis (122).— BELIZE: CAYO
DISTRICT: Baking Pot (1 rom); MEXICO:
CAMPECHE: 60 km SE of Dzibalchen (19°10'N,
89°20'W) ( 1 rom, 1 asnhc); 7.5 km W of Escarcega
(1 asnhc); La Tuxpana Champoton [La Tuxpefia]
(1 usnm); QUINTANA ROO: 2 km SE of Lago
Chichencanab [spelled variously "Laguna Chick-
ankanaab" (Alvarez and Martinez, 1967:205);
"Laguna Chichancanab" (Choate, 1970:277; U.S.
Board on Geographic Names, 1956)] (1 encb); 6
km S, 1.5 km W of Tres Garantias (1 asnhc);
YUCATAN: Actun Chacaljas ( 1 cranium** amnh);
Actun Coyok, 5-90 cm (2 crania, 8 right mandi-
bles** amnh); Actun Has, 0-85 cm (7 crania**
amnh); Actun Lara, 0-1 30 cm (5 crania** amnh);
Actun Oxkintok, 30-75 cm (3 left mandibles
amnh); Actun Spukil, owl perch (63 crania* amnh);
Actun Spukil, excavations (3 crania, 9 right man-
dibles** amnh); Chichen Itza (1 fmnh, 1 usnm—
holotype); Chichen Itza ["along the rim of the Xto-
16k cenote" (Hatt, 1938:334)] (1 amnh); SW of
Dzilam de Bravo (6* ibunam); 2.5 km NW Dzitya
(1 sui); Loltun (2* asnhc); Lolton [Loltun], surface
of cavern floor (3 crania, 4 left mandibles* amnh);
6 km S Merida (1 ku); 13 km W Peto (1 ku*);
Xbac ( 1 usnm).
Additional Records— GUATEMALA: PE-
TEN: Uaxactun (Choate, 1970:277); MEXICO:
YUCATAN: Buctzotz, Calotmul, Izamal, Naba-
lam, Senotillo, Temax, Tzalam, Valladolid, Xbac
(Gaumer, 1917); Uxmal, Mayan ruins (Alvarez
and Martinez, 1967:205; Choate, 1970:277).
Cryptotis cf. mayensis (39). -MEXICO:
GUERRERO: Zumpango del Rio District, Canon
del Zopilote, 1 1.5 km S of the Rio Mescala bridge
near Mescala; Cueva de Macuiltzingo (39*
ibunam).
Cryptotis mera (7). -PANAMA: DARIEN:
Cerro Tacarcuna, 4800 ft (3 usnm); Cerro Mali,
4700 ft (1 usnm); Mount Pirri [Cerro Pirre], E
slope near head of Rio Limon, 4500-5000 ft (3
usnm— including holotype).
Cryptotis merriami (29). -COSTA RICA:
GUANACASTE: 4.5 km NE of Tilaran (1 ku).
EL SALVADOR: MORAZAN: Mt. Cacagua-
tique, north slope, 3800-4000 ft (3 mvz); SAN
MIGUEL: Mt. Cacaguatique, 3500-4000 ft (8 mvz,
2 ummz). GUATEMALA: ALTA VERAPAZ: La
Primavera (1 amnh); Tucuru, Hacienda Concep-
tion, 1 1 00 m ( 1 ummz); San Pedro Carcha ( 1 encb);
HUEHUETENANGO: Barillas, Hacienda Santa
Gregoria ( 1 ummz); Jacaltenango, 5400 ft (4 usnm—
including holotype). HONDURAS: EL PARAI-
SO: Yuscaran, Monserrat [Cerro de Moncerrato]
WOODMAN & TIMM: SMALL-EARED SHREWS
29
Cloud Forest (1 mcz); FRANCISCO MORAZAN:
La Tigre National Park, San Juancito, La Rosario
(1 unah); LEMPIRA: Las Flores Gracias (1 amnh);
SANTA BARBARA: San Jose (1 amnh). MEX-
ICO: CHIAPAS: Volcan Kagchina, 3.5 km N Las
Margaritas, 1500 m (2 mhp*). NICARAGUA:
MAT AG ALP A: 9 mi N of Matagalpa, Santa Mar-
ia de Ostuma, 1400 m (1 ummz).
Additional Records — EL SALVADOR:
AHUACHAPAN: 2 mi NW Apaneca (Burt and
Stirton, 1961). GUATEMALA: ALTA VERA-
PAZ: La Primavera, 3200 ft (Choate, 1970:279).
MEXICO: CHIAPAS: Cueva Los Llanos, 9 km S
Las Margaritas, 1500 m (Choate, 1970:279).
Cryptotis nigrescens (11 9). -COST A RICA:
ALAJUELA: Cinchona, 1600 m (1 ku); Monte-
verde Cloud Forest Reserve, 1 580-1 600 m (3 ku);
Monteverde Cloud Forest Reserve, Penas Blancas
Valley, 870 m (2 ku); ALAJUELA/GU ANA-
CASTE border: Monteverde Cloud Forest Re-
serve, 1580 m (1 ku); CART AGO: [Volcan] Irazu,
8000-9400 ft (1 amnh); GUANACASTE/PUN-
TARENAS border: near Monteverde, Cerro Ami-
gos, 1750-1790 m (3 ku); PUNTARENAS: Coto
Brus [Canton], Sabalito District, Las Tablas, Rio
Coton, 1700 m(l mncr); Monteverde, 1345-1600
m (3 FMNH, 1 INBio, 24 KU, 3 LACM, 1 MMNH, 7
ummz); Monteverde Cloud Forest Reserve, 1530—
1660 m (5 ku); San Luis, 1200 m [ca. 2.5 km S
Monteverde] ( 1 ku); 1 miSWofFincaLasCruces,
San Vito, 4000 ft (3 lacm); SAN JOSE: San Isidro
(1 amnh— holotype); 9 mi N of San Isidro del
General (Pan American Highway), 4800 ft (1
ummz). PANAMA: BOCAS DEL TORO: Fish
Camp, 4900 ft (8°58'N, 82°40'W) (2 usnm); El
Volcan [Hato del Volcan] (4 usnm); 19 km NNW
of El Volcan, E of Cerro Pando, 8°56'30"N,
82°42' 1 5"W, 6400 ft (1 usnm); Volcan de Chiriqui,
Rio Candela, 6000 ft (2 amnh); 25 km NNE of
San Felix, 1425-1500 m (5 usnm); BOCAS DEL
TORO/CHIRIQUI border: Cerro Bollo, 3.5 km E
Escopeta, which latter is at ca. 23 km NNE San
Felix, 1800-1856 m (14 usnm); CHIRIQUI: Cerro
Punta (1 usnm); Cerro Punta, 6500 ft (1 usnm—
holotype of C. zeteki); Cerro Punta, 2700 ft (2
usnm); Cerro Punta, Boquete Trail, 6800 ft (1
usnm); 24 km NNE of San Felix, 1275-1350 m
(6 usnm); 24.5 km NNE of San Felix, 1325-1350
m (2 usnm); 14.5 km NW of El Volcan, Finca
Santa Clara, 8°51'30"N, 82°44'45"W, 1200-2150
m (12 usnm); Santa Clara, Volcan de Chiriqui,
1 700 m (3 usnm); Santa Clara, on the Pan Amer-
ican road 1 5 mi from Costa Rica, 4200 ft (1 amnh—
holotype of C. tersus).
30
FIELDIANA: ZOOLOGY
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