HOLMY LIBRARY Iff WRWU. HALL KBq 71995 UNlvtRSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY HB01W95 590.5 FI n.s. BIX H I H 1 i l AJL % a OOlOgy 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. Literature Cited Alvarez, T., and A. Martinez G. 1 967. New records of Cryptotis mayensis from the Yucatan Peninsula, Mexico. The Southwestern Naturalist, 12: 204-205. Bee, J. W., D. Murariu, and R. S. Hoffmann. 1980. Histology and histochemistry of specialised integu- mentary glands in eight species of North American shrews (Mammalia: Insectivora). Travaux du Mu- seum d'Histoire Naturelle Grigure Antipa, 22: 547- 569. Burt, W. H., andR. A. Stirton. 1961. The mammals of El Salvador. Miscellaneous Publications of the Mu- seum of Zoology, University of Michigan, 117: 1-69. Choate, J. R. 1 968. Dental abnormalities in the short- tailed shrew, Blarina brevicauda. Journal of Mam- malogy, 49: 251-258. . 1970. Systematics and zoogeography of Middle American shrews of the genus Cryptotis. University of Kansas Publications, Museum of Natural History, 19: 195-317. Corbet, G. B., and J. E. Hill. 1991. A World List of Mammalian Species. Oxford University Press, Ox- ford, 243 pp. Dowler, R. C, and M. D. Engstrom. 1988. Distri- butional records of mammals from the southwestern Yucatan Peninsula of Mexico. Annals of Carnegie Mu- seum, 57: 159-166. Eadie, W. R. 1 938. The dermal glands of shrews. Jour- nal of Mammalogy, 19: 171-174. Gaumer, G. F. 1917. Monografia de los mamiferos de Yucatan. Departamento de Talleres Graficos de la Se- cretaria de Fomento, Mexico, xli + 332 pp. Goldman, E. A. 1912. New mammals from eastern Panama. Smithsonian Miscellaneous Collections, 60(2): 1-18. . 1951. Biological investigations in Mexico. Smithsonian Miscellaneous Collections, 115: i-xiv + 1-176. Hall, E. R. 1981. The Mammals of North America, 2nd ed. John Wiley & Sons, New York, 1: 1-600 + 90. Hall, E. R., and K. R. Kelson. 1959. The Mammals of North America. The Ronald Press Co., New York, 1: 1-546 + 79. Hatt, R. T. 1938. Notes concerning mammals col- lected in Yucatan. Journal of Mammalogy, 19: 333- 337. Honacki, J. H., K. E. KlNMAN, AND J. W. Koeppl. 1982. Mammal Species of the World. Allen Press, Inc., and The Association of Systematics Collections, Lawrence, Kans., x + 694 pp. Iskjaer, C, N. A. Slade, J. E. Childs, G. E. Glass, and G. W. Korch. 1989. Body mass as a measure of body size in small mammals. Journal of Mammalogy, 70: 662-667. Lopez- FormentC, W.,andG. UrbanoV. 1977. Res- tos de pequefios mamiferos recuperados en regurgi- taciones de lechuza, Tyto alba, en Mexico. Anales del Instituto de Biologia, Universidad Nacional Auton- oma de Mexico, Serie Zoologia, 48(1): 231-242. Merriam, C. H. 1895. Revision of the shrews of the American genera Blarina and Notiosorex. North American Fauna, 10: 5-34. Murariu, D. 1976. Les glandes tegumentaires de cer- tains insectivores (Mammalia— Insectivora) de Rou- mainie. Anatomie, histologic et histochimie. Travaux du Museum d'Histoire Naturelle Grigure Antipa, 17: 387^13. Ridgway, R. 1912. Color Standards and Color No- menclature. Privately published by the author, Wash- ington, D.C., 43 pp. + 53 pi. Robertson, P. B., and E. A. Rickart. 1975. Cryptotis magna. Mammalian Species, 61: 1-2. Rudd, R. L. 1955. Age, sex, and weight comparisons in three species of shrews. Journal of Mammalogy, 36: 323-339. Setzer, H. W. 1950. Two new shrews of the genus Cryptotis from Panama. Journal of the Washington Academy of Sciences, 40: 299-300. U.S. Board on Geographic Names. 1956. Gazetteer no. 15. Mexico. U.S. Government Printing Office, Washington, D.C., 4 + 750 pp. Wilson, D.E., and D.M.Reeder, eds. 1993. Mammal Species of the World, 2nd ed. Smithsonian Institution Press, Washington, D.C., 1,206 pp. 28 FIELDIANA: ZOOLOGY Woodman, N. 1992. Biogeographical and evolution- ary relationships among Central American small-eared shrews of the genus Cryptotis (Mammalia: Insectivora: Soricidae). Ph.D. diss.. University of Kansas, Law- rence, xi + 386 pp. . 1993. The correct gender of mammalian ge- neric names ending in -otis. Journal of Mammalogy, 74: 544-546. Woodman, N., and R. M. Timm. 1992. A new species of small-eared shrew, genus Cryptotis (Insectivora: So- ricidae), from Honduras. Proceedings of the Biological Society of Washington, 105: 1-12. . Submitted. Geographic variation and biogeo- graphical relationships among large-footed shrews of the Cryptotis goldmani complex (Insectivora: Sorici- dae), with the description of a new species. Journal of Zoology. 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 HECKMAN l—l BINDERY INC. |§| ^JAN9^^