A new species of Cryptotis (Mammalia, Eulipotyphla, Soricidae) from the Sierra de Perijá, Venezuelan-Colombian Andes

Journal of Mammalogy, 96(4):800–809, 2015 DOI:10.1093/jmammal/gyv085 Published online July 31, 2015

A new species of Cryptotis (Mammalia, Eulipotyphla, Soricidae) from the Sierra de Perijá, Venezuelan-Colombian Andes Marcial Quiroga-Carmona* and Neal Woodman Departamento de Biología, Facultad Experimental de Ciencias y Tecnología, Universidad de Carabobo, Valencia 2005, Venezuela (MQC) Centro de Ecología, Instituto Venezolano de Investigaciones Científicas, Km 11 Carretera Panamericana, apartado postal 21827, Caracas 1020-A, Venezuela (MQC) United States Geological Survey, Patuxent Wildlife Research Center, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA (NW) * Correspondent: [email protected] The Sierra de Perijá is the northern extension of the Cordillera Oriental of the Andes and includes part of the border between Colombia and Venezuela. The population of small-eared shrews (Mammalia, Eulipotyphla, Soricidae, Cryptotis) inhabiting the Sierra de Perijá previously was known from only a single skull from an individual collected in Colombia in 1989. This specimen had been referred to alternatively as C. thomasi and C. meridensis, but a more precise definition of the known Colombian and Venezuelan species of Cryptotis has since excluded the Sierra de Perijá population from any named species. The recent collection of a specimen from the Venezuelan slope of Sierra de Perijá prompted us to re-evaluate the taxonomic status of this population and determine its relationship with other Andean shrews. Our examination of the available specimens revealed that they possess a unique suite of morphological and morphometrical characters, and we describe the Sierra de Perijá population as a new species in the predominantly South American C. thomasi species group. Recognition of this new species adds to our knowledge of this genus in South America and to the biodiversity of the Sierra de Perijá. La Sierra de Perijá es la extensión más septentrional de la Cordillera Oriental de los Andes, e incluye parte de la frontera entre Colombia y Venezuela. La población de musarañas de orejas pequeñas (Mammalia, Eulipotyphla, Soricidae, Cryptotis) que habita esta serranía fue previamente conocida solo por el cráneo de un ejemplar colectado en Colombia en 1989. Este fue identificado alternativamente como C. thomasi y C. meridensis; sin embargo, la definición más precisa de las especies de Cryptotis de Colombia y Venezuela ha excluido a la población de la Sierra de Perijá de cualquier especie conocida para la región. El hallazgo reciente de un ejemplar en la vertiente venezolana de la Sierra de Perijá, impulsó la reevaluación del estado taxonómico de esta población y la determinación de sus relaciones con otras musarañas andinas. Nuestra revisión de los ejemplares disponibles reveló que poseen un conjunto de características morfológicas y morfométricas únicas, por lo tanto nosotros describimos la población de Cryptotis presente en la Sierra de Perijá como una especie nueva perteneciente al grupo de especies C. thomasi. El reconocimiento de esta nueva especie aumenta nuestro conocimiento sobre este género en América del Sur y la biodiversidad de la Sierra de Perijá. Key words:  Blarinini, Cryptotis thomasi species group, neotropical shrews, Soricinae, Soricomorpha, South America Published by Oxford University Press on behalf of the American Society of Mammalogists. This work is written by (a) US Government employee(s) and is in the public domain in the US.

Small-eared shrews (Mammalia, Eulipotyphla, Soricidae) of the genus Cryptotis Pomel, 1848 are endemic to the New World (Choate 1970), and the genus includes the only species of the family that occur in South America. Although some North American species occur in lowland habitats (Guevara et al. 2014), all South American Cryptotis inhabit montane environments above approximately 1,200 m (Woodman and

Péfaur 2008; Quiroga-Carmona and Molinari 2012; Moreno and Albuja 2014), mostly in the northern Andes from northern Peru (Cadenillas et al. 2010) to Colombia and Venezuela. In addition, 2 isolated species occur outside of the Andes in the Sierra de Aroa (Quiroga-Carmona and Molinari 2012) and Serranía del Litoral of northern Venezuela (Quiroga-Carmona 2013). 800



QUIROGA-CARMONA AND WOODMAN—A NEW CRYPTOTIS FROM THE SIERRA DE PERIJÁ 801

Species of Cryptotis are often partitioned among 4 character-based groups that may correspond to monophyletic clades (Choate 1970; Woodman and Timm 1993, 1999; Woodman 2002). The 14 known South American species are all members of either the mostly Central American C. nigrescens species group (3 species) or the mostly Andean C. thomasi species group (11 species—Woodman and Péfaur 2008; QuirogaCarmona and Molinari 2012; Quiroga-Carmona 2013; Moreno and Albuja 2014). The Sierra de Perijá is the northernmost extension of the Andes in northeastern Colombia and northwestern Venezuela. Shrews were unknown from this poorly explored portion of the Cordillera Oriental of the Northern Andes until 1989, when an individual was fortuitously found dead on a trail in cloud forest near Finca el Suspiro, Departamento del Cesar, Colombia (Duarte and Viloria 1992). At the time, all of the larger shrews from eastern Colombia and Venezuela were considered to be C. thomasi (e.g., Eisenberg 1989), and this specimen was thought to mark an extension of the northern limit of that species’ geographic range. A second specimen, also referred to C. thomasi, was subsequently reported to have been collected in March 2006, between 2,850 and 3,100 m near Corregimiento San José de Oriente, also in Depto. César (Corredor-Carrillo and Muñoz-Saba 2007). Based on size and skull characters (e.g., shape of the mandible) visible in photographs of the Finca el Suspiro specimen (Duarte and Viloria 1992: page 240, figures 2–5), Woodman (2002) suggested that the population belonged to the C. thomasi species group, but noted that other characters (lack of an obvious foramen on the tympanic process of the petromastoid, deeper inferior sigmoid notch, relatively shorter posterior mandible and short zygomatic plate, broad palate and interorbital region) did not conform entirely with either C. thomasi or C. tamensis. He restricted the distribution of C. thomasi to elevations above 2,700 m in the southern Cordillera Oriental, with C. tamensis occupying the central portion of that mountain range (Woodman 1996, 2002). These actions left the identity of the Sierra de Perijá population in doubt. More recently, a third specimen of Cryptotis was found in páramo surrounding Pico Tétari on the Venezuelan slope of the Sierra de Perijá. Our comparisons of this individual and the Finca el Suspiro specimen with extant species of the C. thomasi species group occurring in Colombia and Venezuela (C. aroensis, C. medellinius, C. meridensis, C. tamensis, C. squamipes, and C. venezuelensis) confirm that the Sierra de Perijá population is morphologically distinct and is not referable to any known species. Herein, we describe this population as a new species, contrast it with other species from the region, and provide details of its natural history.

Materials and Methods Specimens examined in our study (Appendix I) are housed in the following institutions: American Museum of Natural History, New York (AMNH); Natural History Museum, London (BMNH); Colección de Vertebrados de la Universidad de los

Andes, Mérida (CVULA); Museo de la Estación Biológica de Rancho Grande, Maracay (EBRG); Field Museum, Chicago (FMNH); Instituto de CienciasNaturales, Universidad Nacional de Colombia, Bogotá (ICN); University of Kansas Natural History Museum, Lawrence (KU); Museo de Biología de la Universidad del Zulia, Maracaibo (MBLUZ); Museo de Biología de la Universidad Central de Venezuela, Caracas (MBUCV); Museum of Comparative Zoology, Cambridge (MCZ); Museo de Historia Natural La Salle, Caracas (MHNLS); Museum National d’Histoire Naturelle, Paris (MNHN); Museo de la Universidad de Antioquia, Medellín (MUA); Royal Ontario Museum, Toronto (ROM); National Museum of Natural History, Washington (USNM). Our investigations of Cryptotis from the Sierra de Perijá were limited to 2 specimens (MBLUZ–105, MHNLS–12354) in systematic collections. A 3rd specimen, reported by CorredorCarrillo and Muñoz-Saba (2007), does not appear to have been accessioned or cataloged in the collection of the ICN, as was intended (C. Cárdenas-González, in litt. Universidad Nacional de Colombia, Bogotá, Cundinamarca). We initially made comparisons of the available specimens with all known species of the C. thomasi species group in Colombia and Venezuela. We focused our analyses, however, on the 3 species (C. meridensis, C. tamensis, and C. thomasi) that occupy adjacent portions of the eastern Andes, because these species were most likely to be conspecific with the population in Sierra de Perijá. Unless otherwise stated, general comparative statements regarding specific characters are relative to all other species in the genus Cryptotis. Dental nomenclature follows Choate (1970) and Reumer (1984), cranial terminology follows Gaughran (1954), and capitalized color names are those of Smithe (1975). External measurements (Table 1) were recorded from museum labels or field notes and are reported to the nearest mm or g. These include: head-and-body length (HB); hindfoot length (HL); ear length (EL); tail length (TL); and body mass (BM). Craniomandibular variables (Table 1) follow Woodman and Timm (1993) and Quiroga-Carmona and Molinari (2012) and include: condylobasal length (CBL), cranial breadth (CB), breadth of zygomatic plate (ZP), interorbital breadth (IO), breadth across 1st unicuspids (U1B), breadth across 2nd molars (M2B), palatal length (PL), length of upper toothrow (TR), length of unicuspid toothrow (UTR), length of 3rd unicuspid (U3), length of 4th unicuspid (U4), space between 3rd unicuspid and 4th premolar (U3–P4), length of molariform toothrow (MTR), length of mandible (ML), height of coronoid process (HCP), height of coronoid valley (HCV), height of articular condyle (HAC), distance from the articular condyle to posterior edge of 3rd lower molar (AC3), length of lower toothrow (TRM), and length of 1st lower molar (Lm1). These measurements were taken to the nearest 0.01 mm using a digital caliper (CBL, CB) or an ocular micrometer in a binocular stereomicroscope and rounded to the nearest 0.1 mm. To examine morphometrical relationships among shrews from Sierra de Perijá and adjacent regions, we carried out principal component analyses (PCA) on 7 log10-transformed

802

JOURNAL OF MAMMALOGY

Table 1.—Measurements of selected Cryptotis from the eastern Andes of Colombia and Venezuela. Statistics are mean ± SD and range. Measurements follow Woodman and Timm (1993), Quiroga-Carmona and Molinari (2012), and Quiroga-Carmona (2013). Dashes: mean data not available. AC3, articular condyle to m3; CB, cranial breadth; CBL, condylobasal length; HAC, height of articular condyle; HB, head-andbody length; HCP, height of coronoid process; HCV, height of coronoid valley; IO, interorbital breadth; Lm1, length of first lower molar; M2B, breadth across second molars; ML, length of mandible; MTR, length of molariform toothrow; PL, palatal length; TL, tail length; TR, length of upper toothrow; TRM, length of lower toothrow; U1B, breadth across first unicuspids; UTR, length of unicuspid toothrow; ZP, breadth of zygomatic plate. Measurements

Sierra de Perijá

C. meridensis

C. tamensis

C. thomasi

n = 54   89 ± 5 76–102   34 ± 3 25–41

n = 15

n = 20

Paratype

86 ± 4 81–91

87 ± 4 79–96

68

36 ± 2 32–39

24 ± 2 20–27

36

n = 58   21.7 ± 0.8 20.5–23.5 (n = 43)

n = 16

n = 38

Holotype, paratype

21.7 ± 0.5 20.7–22.6 (n = 23)

21.5 ± 0.5 20.8–22.9 (n = 13)

—, —

10.5 ± 0.3 10.0–11.4 (n = 45)

10.5 ± 0.2 10.1–10.8 (n = 11)

10.6 ± 0.2 10.2–11.0 (n = 17)

10.2, —

2.0 ± 0.2 1.4–2.4

2.1 ± 0.1 1.9–2.3

2.0 ± 0.2 1.7–2.4

1.5, —

5.1 ± 0.2 4.6–5.4

5.0 ± 0.2 4.8–5.3

5.0 ± 0.2 4.5–5.4

5.7, —

2.9 ± 0.1 2.6–3.1 (n = 37)

3.0 ± 0.1 2.7–3.1

2.8 ± 0.1 2.6–2.9

2.9, —

6.5 ± 0.3 6.0–7.1

6.4 ± 0.1 6.1–6.7

6.2 ± 0.2 5.8–6.5

7.0, —

9.7 ± 0.4 8.8–10.4

9.5 ± 0.2 9.1–9.8

9.3 ± 0.3 8.7–9.8

10.2, —

8.3 ± 0.4 7.3–9.0

8.3 ± 0.3 7.9–8.8

8.2 ± 0.2 7.7–8.7

8.8, —

2.9 ± 0.2 2.6–3.2

2.9 ± 0.1 2.6–3.1

2.8 ± 0.1 2.4–3.0

3.0, —

5.9 ± 0.2 5.5–6.3

5.9 ± 0.1 5.6–6.1

5.8 ± 0.1 5.5–6.1

6.2, —

7.1 ± 0.4 6.3–7.9

7.0 ± 0.2 6.9–7.4

7.1 ± 0.3 6.5–7.7

7.5, 7.4

4.9 ± 0.2 4.5–5.4

4.6 ± 0.2 4.3–4.8

4.6 ± 0.2 4.3–4.9

4.9, 4.8

3.3 ± 0.1 3.1–3.6

3.1 ± 0.1 3.0–3.2

3.1 ± 0.1 2.8–3.4

3.0, 3.0

4.5 ± 0.2 4.0–5.1

4.4 ± 0.1 4.2–4.6

4.3 ± 0.2 3.8–4.7

4.4, 4.3

External measurements  HB

 TL

Skull measurements  CBL

 CB

 ZP

 IO

 U1B

 M2B

 PL

 TR

 UTR

 MTR

 ML

 HCP

 HCV

 HAC



QUIROGA-CARMONA AND WOODMAN—A NEW CRYPTOTIS FROM THE SIERRA DE PERIJÁ 803 Table 1.—Continued

Measurements

Sierra de Perijá

C. meridensis

C. tamensis

C. thomasi

5.7 ± 0.3 5.1–6.5 (n = 33)

5.4 ± 0.1 5.2–5.7

5.7 ± 0.2 5.0–6.0

5.8, 5.3

6.6 ± 0.3 6.1–7.2 (n = 56)

6.6 ± 0.2 6.3–6.9 (n = 33)

6.4 ± 0.2 6.0–6.7

6.8, 6.5

2.0 ± 0.1 1.7–2.1 (n = 33)

2.0 ± 0.1 1.8–2.1

1.9 ± 0.1 1.7–2.0

1.9, 2.0

 AC3

 TRM

 Lm1

cranial variables (ZP, PL, IO, TR, UTR, MTR, and M2B) from 58 C. meridensis, 16 C. tamensis, 38 C. thomasi, and the specimen from Finca el Suspiro. To control for size differences and remove possible allometric effects, we performed a 2nd PCA in which each variable was standardized for size by subtracting the geometric mean of all variables following Mosimann and James (1979).

Table 2.—Factor loadings for the first 2 factor axes from principal components analysis (PCA) of 7 log10-transformed cranial measurements from Cryptotis meridensis, C. tamensis, C. thomasi, and a specimen from the Sierra de Perijá. Variables are listed in descending order by their loadings on the first axis. IO, interorbital breadth; M2B, breadth across second molars; MTR, length of molariform toothrow; PL, palatal length; TR, length of upper toothrow; UTR, length of unicuspid toothrow; ZP, breadth of zygomatic plate.

Results

Component loadings

Morphology.—Several qualitative skull characters confirm that the population of Cryptotis in the Sierra de Perijá belongs to the C. thomasi species group. These include the larger posterior element (contrasted with the anterior element) of the ectoloph M1; the relatively slender anterior unicuspids with their concavely curved postero-occlusal borders (when viewed laterally); the long, low-crowned lower premolars; and the relatively low coronoid process of the mandible and the wide angle at which its anterior border joints the relatively long horizontal ramus of the mandible. Within the C. thomasi species group, the Sierra de Perijá population is distinguished from its congeners by a unique combination of external and craniomandibular characters that includes a relatively long tail; a cranium with a narrow zygomatic plate, broad interorbital region, broad posterior palate, large U4 not visible in lateral view, small foramen on the posterior edge of the tympanic process of the petromastoid, and simple M3; and a mandible with a low coronoid process, relatively short posterior portion, and shallow lower sigmoid notch. Morphometrics.—Our PCA of 7 cranial variables yielded a model in which the 1st factor axis, which accounts for > 53% of total variation, is a size axis dominated by lengths of the toothrow and length and width of the palate, but with little contribution from interorbital length or from breadth of zygomatic plate (Table 2). The 2nd factor axis, which represents > 14% of the variation, is dominated by breadth of the zygomatic plate. In a plot of factor scores on these 2 axes (Fig. 1A), C. meridensis, C. tamensis, and C. thomasi mostly overlap, reflecting the generally conservative cranial shape within the C. thomasi species group (Woodman 2002). C. meridensis, however, exhibits a greater size range along the 1st axis than either C. tamensis or C. thomasi, and it represents all of the largest individuals among these 3 species. The specimen from Finca el Suspiro

Variable MTR TR PL M2B UTR IO ZP Eigenvalues Total variance explained (%)

Axis 1

2

0.905 0.891 0.885 0.847 0.761 0.617 −0.291 4.158 59.394

0.111 0.022 −0.061 −0.051 −0.039 0.434 0.898 1.015 14.497

in the Sierra de Perijá (MBLUZ–105) plots among the larger C. meridensis along the 1st factor axis, and it is larger than any C. tamensis or C. thomasi. Along the 2nd axis, it plots lower than any other specimen, a result of its short zygomatic plate. Together, the 2 axes separate this individual from all 3 of the other species in the plot. In our 2nd PCA of 7 cranial variables controlled for size, the 1st factor axis accounts for nearly 47% of the variation in the model and represents the lengths of toothrow and palate contrasted with a negatively weighted breadth of zygomatic plate (Table 3). The 2nd factor axis, with > 19% of the variation, is a contrast between length of the unicuspid toothrow and negatively weighted interorbital breadth. The plot of factor scores on these 2 axes (Fig. 1B) again shows a large region of overlap among most C. meridensis, C. tamensis, and C. thomasi. C. meridensis again exhibits a greater range of variation along the 1st axis, and it includes most of the individuals with the highest values along this axis (i.e., individuals with longer palate and toothrow combined with narrower zygomatic plate). The specimen from the Sierra de Perijá has the highest value, plotting most closely with C. meridensis and well away from either C. tamensis or C. thomasi. Along the 2nd axis, the Sierra

804

JOURNAL OF MAMMALOGY Table 3.—Factor loadings for the first 2 factor axes from principal components analysis (PCA) of 7 log10-transformed cranial measurements with the geometric mean subtracted to remove size. Variables are listed in descending order by their loadings on the first axis. IO, interorbital breadth; M2B, breadth across second molars; MTR, length of molariform toothrow; PL, palatal length; TR, length of upper toothrow; UTR, length of unicuspid toothrow; ZP, breadth of zygomatic plate. Component loadings Variable ZP PL TR MTR M2B IO UTR Eigenvalues Total variance explained (%)

Axis 1

2

−0.976 0.766 0.745 0.721 0.672 0.084 0.459 3.284 46.913

0.081 0.109 0.296 −0.304 −0.319 −0.848 0.610 1.391 19.876

Fig. 1.——Principal components analyses (PCA) of Cryptotis meridensis, C. tamensis, C. thomasi, and Cryptotis from the Sierra de Perijá: A) plots of scores on factor axes 1 and 2 from PCA of 7 log10transformed cranial measurements (Table 2); B) plots of scores on factor axes 1 and 2 from PCA of 7 log10-transformed cranial measurements with the geometric mean subtracted to account for possible allometry (Table 3).

de Perijá specimen plots relatively low (i.e., shorter unicuspid toothrow, broader interorbital region), but overlaps with the ranges of all 3 species. Our morphological and morphometrical study of the specimens of Cryptotis from Sierra de Perijá indicates that they represent a population of shrews in the C. thomasi species group, but one that is distinct from all described species.

Systematic Biology Family Soricidae G. Fischer, 1814 Subfamily Soricinae G. Fischer, 1814 Genus Cryptotis Pomel, 1848 Cryptotis perijensis, new species Perijá Shrew; Musaraña de la Sierra de Perijá Figs. 2 and 4. Cryptotis thomasi: Duarte and Viloria, 1992:240 (part). Cryptotis meridensis meridensis: Linares, 1998:106 (part).

Fig. 2.—A) Dorsal and B) ventral views of the skull and C) lateral view of the skull and mandible of the holotype of Cryptotis perijensis (MBLUZ–105). White bar = 5 mm.

Holotype.—MBLUZ–105, an adult of unknown sex, collected on 22 March 1989, by A. Viloria (22-III-1989). The basisphenoid and temporal of the cranium are damaged and angular processes of both mandibles are missing (Fig. 2). Type locality.—Near Finca el Suspiro, Departamento del Cesar, Colombia, 2,000 m (10°21ʹN, 72°57ʹW; Fig. 3).



QUIROGA-CARMONA AND WOODMAN—A NEW CRYPTOTIS FROM THE SIERRA DE PERIJÁ 805

Fig. 3.—Distribution of selected species of Cryptotis in Colombia and Venezuela: C. aroensis (black circles), C. medellinius (black squares), C. meridensis (white triangles), C. perijensis (stars), C. squamipes (black pentagons), C. tamensis (black triangles), C. thomasi (white squares), and C. venezuelensis (white circles). Localities for C. perijensis correspond (from north to south) to the holotype, a specimen reported by CorredorCarrillo and Muñoz-Saba (2007), and the paratype. Distributions of C. medellinius, C. squamipes, and C. thomasi are based on Woodman (2002) and Woodman and Péfaur (2008).

Paratype.—MHNLS–12354, an adult male preserved in ethanol (Fig. 4) with the crushed skull partially removed; collected January 2009 by F. Rojas-Runjaicin páramo at the base of Pico Tétari, Estado Zulia, Venezuela, 3,200 m (10°02ʹ N, 72°57ʹ W; Fig. 3). Diagnosis.—C. perijensis is a member of the C. thomasi species group of small-eared shrews that is distinguished from other Colombian and Venezuelan members of the group by its relatively long tail (shorter in C. medellinius, C. meridensis, C. squamipes, C. tamensis, C. thomasi); narrow zygomatic plate (broader in C. medellinius, C. meridensis, C. squamipes, C. tamensis, C. thomasi); broad interorbital region (narrower in C. medellinius, C. meridensis, C. squamipes, C. tamensis, C. thomasi); broad posterior palate (narrower in C. squamipes, C. tamensis, C. thomasi); large, lingually displaced U4, which is not easily visible in lateral view (U4 visible in C. medellinius, C. meridensis, C. tamensis); minute foramen on the posterior edge of the tympanic process of petromastoid (larger in C. tamensis; huge in C. medellinius, C. thomasi); simple M3 (complex in C. thomasi); relatively low coronoid process (higher in C. meridensis, C. squamipes); relatively short posterior portion of the mandible (longer in C. meridensis, C. thomasi); deeper inferior sigmoid notch (shallower in C. meridensis, C. tamensis, C. thomasi). Description.—A medium-sized (HB = 68) small-eared shrew with a long tail (LT = 36; LT/HB = 53%); Dark Gray dorsal pelage with Lead-Gray luster and comprised of bicolored hairs (Gray bases, Dark Brown tips); ventral pelage slightly paler than dorsum (Fig. 4). Although the cranium is incomplete, a number of cranial dimensions (Table 1) indicate that the skull of C. perijensis is large for the genus, with a long rostrum, relatively broad interorbital

Fig. 4.—A) Ventral, B) lateral, and C) dorsal external views of the paratype of Cryptotis perijensis (MHNLS–12354) at the time it was found at the base of Pico Tétari. Black bar = 40 mm.

region (IO/PL = 55.9%), and moderately broad posterior palate (M2B/PL = 68.6%). There are 2 large dorsal foramina on the frontals. The zygomatic plate is short (ZP = 1.5 mm; ZP/PL = 14.7%); the anterior border is aligned with the metastyle of M1 or parastyle

806

JOURNAL OF MAMMALOGY

of M2 and the posterior border with the posterior mesostyle/metastyle valley of M3. In lateral view, the posterior edge of the maxillary process is aligned with the mesostyle of M3 and separated from the posterior edge of the zygomatic plate by an extension of the palate. The anterior process of the petromastoid is low and thin-walled, and the posterior edge of the tympanic process of the petromastoid has a minute foramen. The upper dentition is bulbous and robust. The unicuspid toothrow is moderately long (UTR = 3 mm; UTR/ PL = 29.4%), and the unicuspids (U1–U3) are relatively slender in lateral view, with concavely curved postero-occlusal borders. The unicuspids possess well developed, broadened cingulae and well-developed posterolingual cuspules. The U4 is large (36% of the area of U3), displaced lingually and not readily visible in lateral view of the cranium (because U3–P4 is reduced: 0.15 mm). The anterior element of the ectoloph of M1 is smaller than posterior element. The M3 is simple, with reduced postcentrocrista and lacking metacone and hypocone. The mandible is long, with an elongated posterior region (AC3/ML = 71.6–77.3%). The coronoid process is relatively short (HCP/ML = 64.9–65.3), its anterior border forming a wide angle with the horizontal ramus of the mandible. The inferior sigmoid notch is moderately deep. Lower premolars are long and low-crowned. The talonid of m3 lacks an entoconid. Etymology.—The specific epithet perijensis [Perij(á)+ensis] is a toponym referring to the type region in the Sierra de Perijá. Nomenclatural statement.—A life science identifier (LSID) number was obtained for the new species Cryptotis perijensis: urn:lsid:zoobank.org:act:4810D7EE-7664-40DB-81DEC98B455EE3E7. Distribution.—Known solely from the northern half of the Sierra de Perijá in eastern Colombia and western Venezuela (Fig. 3). Comparisons.—Measurements and indices for C. perijensis are provided in Table 1 and the preceding species description. Unless otherwise indicated, measurements and indices in the following accounts are those for the species to which C. perijensis is being compared and derive from Woodman (2002), QuirogaCarmona and Molinari (2012), and Quiroga-Carmona (2013). Cryptotis aroensis Quiroga-Carmona and Molinari, 2012.— This species has a larger body size (HB = 79 ± 2) than C. perijensis; relatively shorter tail (TL/HB = 46 ± 3%); paler, grayish brown pelage; longer zygomatic plate (ZP = 1.9 ± 0.1; ZP/ PL = 21.3 ± 1.2%); narrower interorbital region (IO = 5.0 ± 0.1); shorter palate (PL = 9.1  ±  0.1); narrower posterior palate (M2B = 5.5 ± 0.1; M2B/PL = 61.0 ± 0.1%); more complex M3. C. aroensis is endemic to the Sierra de Aroa, Venezuela (Fig. 3). Cryptotis medellinius Thomas, 1921.—This species has larger body size (C. medellinius: HB = 85 ± 7); relatively shorter tail (TL/HB = 43 ± 6%); longer zygomatic plate (ZP = 2.0 ± 0.2; ZP/PL = 21.2 ± 2.1%); smaller U4 (U4 = 40 ± 9% area of U3); huge foramen on the posterior edge of the tympanic process of each petromastoid; higher coronoid process (HCP/ ML  =  68.5 ± 2.5%). C. medellinius inhabits the northern Cordillera Central and northern Cordillera Occidental of Colombia (Fig. 3).

Cryptotis meridensis (Thomas, 1898).—This species has paler, chocolate brown pelage; larger body size (HB = 89 ± 5); relatively shorter tail (TL/HB = 38 ± 4%); broader zygomatic plate (ZP = 2.0 ± 0.2; ZP/PL = 21.2 ± 2.0%); posterolingual cuspules on the cingulae of U1–3 typically minute or absent; smaller U4 (U4 = 19 ± 7% area of U3); U4 lacking on one or both sides in 25% of individuals; higher coronoid process (HCP/ML = 69.8 ± 2.5%); longer posterior mandible (AC3/ ML  =  81.0 ± 2.6%). C. meridensis is endemic to the Cordillera de Mérida, Venezuela (Fig. 3). Cryptotis squamipes (Allen, 1912).—This species has a broader zygomatic plate (ZP = 2.1 ± 0.1; ZP/PL = 22.2 ± 0.6%) and interorbital region (IO = 5.4 ± 0.2); posterolingual cuspules on U1–U3 typically minute or absent; typically smaller U4 (U4 = 39 ± 10% area of U3); more complex M3; higher coronoid process (HCP/ML = 71.4 ± 5.2%); minute entoconid occasionally present in talonid of m3. C. squamipes occurs in the southern Andes of Colombia (Fig. 3). Cryptotis tamensis Woodman, 2002.—This species has a larger body size (HB = 86 ± 4); relatively shorter tail (TL/ HB = 42 ± 3%); paler Mummy Brown to Clove-Brown pelage; broader zygomatic plate (ZP = 2.1 ± 0.1; ZP/PL = 21.9 ± 1.4%); narrower interorbital area (IO = 5.0 ± 0.2; IO/PL = 53.0 ± 1.7%); smaller U4 (U4 = 29 ± 8% of area of U3). C. tamensis inhabits the Macizo del Tamá between Colombia and Venezuela and the Nudo de Pamplona in Colombia (Fig. 3). Cryptotis thomasi (Merriam, 1897).—This species has a larger body size (HB = 87 ± 4); shorter tail (TL = 24 ± 2; TL/ HB = 29 ± 4%); broader zygomatic plate (ZP = 2.0 ± 0.2; ZP/ PL = 21.3 ± 1.7%); narrower interorbital area (IO = 5.0 ± 0.2; IO/PL = 52.5 ± 2.1%); narrower palate (M2B = 6.2 ± 0.2); U4 aligned with unicuspid row and visible in the lateral view of the cranium; huge foramen on the posterior edge of the tympanic process of both petromastoids; more complex M3; entoconid often (approximately 30% of specimens) present and obvious in talonid of m3. Cryptotis thomasi is known from above 2,500 m in the central Cordillera Oriental of Colombia (Fig. 3). Cryptotis venezuelensis Quiroga-Carmona, 2013.—This species has a larger body size (HB = 72–84); broader zygomatic plate (ZP = 2.2 ± 0.2; ZP/PL =22.9 ± 1.4%); narrower interorbital region (IO = 5.0 ± 0.1; IO/PL = 51.7 ± 0.8%) and posterior palate (M2B = 6.4 ± 0.1; M2B/PL = 65.4 ± 1.6%); relatively lower coronoid process of the mandible (HCP/ ML = 57.7–61.5%); relatively shorter posterior mandible (AC3/ LM = 65.4%); more complex M3; conspicuous entoconid in talonid of m3. C. venezuelensis is endemic to the Serranía del Litoral, Venezuela (Fig. 3). Remarks.—Cryptotis is the second most diverse soricid genus in the New World (after Sorex). Recognition of C. perijensis increases the number of recognized species to 42, of which 15 occur in South America. The presence of members of 2 distinct species groups strongly suggests that the genus colonized South America at least twice, with subsequent diversification within each group. However, the evolutionary and biogeographical history of the genus remains to be adequately deciphered.



QUIROGA-CARMONA AND WOODMAN—A NEW CRYPTOTIS FROM THE SIERRA DE PERIJÁ 807

Corredor-Carrillo and Muñoz-Saba (2007) reported “Cryptotis thomasi” (= C. perijensis) among the 12 species of mammals they documented at 2,850–3,100 m near Corregimiento San José de Oriente, Cesar Province, Colombia, in early March 2006. Other species at the site included Sturnira erythromos, Cerdocyon thous, Odocoileus virginianus, Sciurus granatensis, Cavia porcellus, Cuniculus paca, C. taczanowskii, Akodon bogotensis, Nephelomys albigularis, and Sylvilagus brasiliensis. The dominant higher-elevation vegetation associations in the Sierra de Perijá include high-Andean cloud forest (up to approximately 2,900 m), which has a single level of stratification above the understory, and sub-páramo (approximately 2,900– 3,500 ml) dominated by a mixture of bamboo, scrub, terrestrial herbs, and low stem-rosettes (Arellano-P. and Rangel-Ch. 2007; Rangel-Ch. and Arellano-p. 2007; Rangel-Ch. 2009). The few localities of collection for C. perijensis indicate its occurrence in Montane and Lower Montane Wet Forest life zones of the traditional Holdridge system (Holdridge 1947; Ewel et al. 1965; IGAC 1988). Duarte and Viloria (1992) described the habitat in which the Finca el Suspiro specimen was found as moderately disturbed cloud forest. This indicates some level of tolerance of C. perijensis for physical modification of its habitat as has been noted for C. tamensis (Woodman 2002). Sparse information accompanying the Pico Tétari specimen and the published record of an individual from near Corregimiento San José de Oriente (Corredor-Carrillo and Muñoz-Saba 2007) suggest that C. perijensis also inhabits open páramo or sub-páramo vegetation. These generalized habitats and elevational distribution correspond with those described for other species of Cryptotis present in the Andes (Woodman and Péfaur 2008; Moreno and Albuja 2014). Most of the botanical communities in the Sierra de Perijá, particularly those of the sub-Andean and Andean zones, have strong biogeographical affinities with other mountain ranges of the Northern Andes (e.g., Cordillera de Mérida, Macizo del Tamá and Sierra Nevada de Santa Marta.). In contrast, a large endemic floristic component is present in the páramo zone (see Rivera-Diaz and Fernandez-Alonzo 2003). Recorded endemism is rare among the faunal communities and is restricted to a species of hummingbird, the Perijá Metaltail (Metallura iracunda), and several avian subspecies (Hernández-Camacho et al. 1992). C. perijensis is the 1st endemic mammal species described from this mountain range. In terms of its biota, the Sierra de Perijá is one of the least known regions of the northern Andes (Rangel-Ch. 2007). The region currently faces serious threats including loss of vegetation coverage a result of agriculture and livestock activities. Official protection is limited to the Parque Nacional Sierra de Perijá on the Venezuelan slope (Morales et al. 2007; Rodríguez et al. 2010). Poor roads and security issues make this region logistically difficult to access for studies of wildlife. In consequence, the breadth of its natural biological diversity is incompletely known (Viloria 1990, 2005; Morales et al. 2007; Rangel-Ch. 2007). These circumstances further encumber proper conservation management and policy (Primack et al.

2001). It is hoped that recognition of C. perijensis as a species endemic to the Sierra de Perijá will prompt additional scientific and conservation interest in this neglected region of the Andes.

Acknowledgments We thank the Fundación Provita for partial sponsorship of this work through Iniciativa de Especies Amenazadas (IEA). We thank the following curators and collections managers for loans or for permission to examine specimens under their care: R. Voss (AMNH); P. Jenkins, L. Tomsett, and R. Portela-Miguez (BMNH); P. Soriano and J. Murillo (CVULA); F. Bisbal and J. Sánchez (EBRG); L. R. Heaney, B. D. Patterson, J. Phelps, and W. T. Stanley (FMNH); A. Cadena and H. Lopez-Arévalo (ICN); R. M. Timm (KU); R. Calchi and D. Prieto (MBLUZ); C. Ferreira and M. Salazar (MBUCV); J. M. Chupasko and M. Omura (MCZ); A. Ferrer (MHNLS); G. Pothet, M. Tranier, and C. Denys, Museum National d’HistoireNaturelle, Paris (MNHN); C. Cuartas-Calles and C. Delgado-V. (MUA); J. Eger and B. Lim (ROM). Special thanks to F. Rojas-Runjaic (MHNLS) for providing the paratype specimen (and its photographs), and J. Molinari for allowing work in his laboratory during this research. G. D’Elía and R. D. Fischer provided valuable comments on previous versions of this manuscript. Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the United States government.

Literature Cited Allen, J. A. 1912. Mammals from western Colombia. Bulletin of the American Museum of Natural History 31:71–95. Arellano- P., H., and J. O. Rangel-Ch. 2007. Cima y topoclima. Los ecosistemas de la Alta Montaña de Perijá. Pp. 19–41 in Colombia, diversidad biótica V, la alta montaña de la Serranía de Perijá (J. O. Rangel-Ch., ed.). Universidad Nacional de Colombia, Bogotá, Colombia. Cadenillas, R., et al. 2010. Mamíferos menores del distrito de La Granja, Cajamarca, Perú; con ampliación de la distribución de Histiotus velatus y del género Cryptotis (abstract). Libro de Resúmenes del II Congreso de la Sociedad Peruana de Mastozoología, Arequipa, Perú. Choate, J. R. 1970. Systematics and zoogeography of Middle American shrews of the genus Cryptotis. University of Kansas Publications, Museum of Natural History 19:195–317. Corredor-Carrillo, D. A., and Y. Muñoz-Saba. 2007. Mamíferos de la alta montaña de Perijá. Pp. 221–233 in Colombia, diversidad biótica V, la alta montaña de la Serranía de Perijá (J. O. Rangel-Ch, ed.). Universidad Nacional de Colombia, Bogotá, Colombia. Duarte, M. A., and A. L. Viloria. 1992. Nuevo hallazgo de Cryptotis thomasi (Merriam, 1897) (Mammalia: Insectivora) en la Sierra de Perijá, Noreste de Colombia. Acta Científica Venezolana 43:240–242. Eisenberg, J. E. 1989. Mammals of the neotropics. The northern Neotropics. University of Chicago Press, Chicago, Illinois. Ewel, J. J., A. Madriz, and J. A. Tosi. 1965. República de Venezuela. Mapa Ecológico. Ministerio de Agricultura y Cría, Dirección de Investigación, Caracas, Venezuela.

808

JOURNAL OF MAMMALOGY

Gaughran, G. R. L. 1954. A comparative study of the osteology and myology of the cranial and cervical regions of the shrew, Blarina brevicauda, and the mole, Scalopus aquaticus. University of Michigan Museum of Zoology, Miscellaneous Publications 80:1–82. Guevara, L., V. Sánchez-Cordero, L. León-Paniagua, and N.  Woodman. 2014. A new species of small-eared shrew (Mammalia, Eulipotyphla, Cryptotis) from the Lacandona rain forest, México. Journal of Mammalogy 95:759–753. Hernández-Camacho, J., A. Hurtado-Guerra, R. OrtizQuijano, and T. Walschburger. 1992. Centros de Endemismos de Colombia. Pp. 175–190 in La Diversidad Biológica en Iberoamérica I (G. Halffter, ed.). Acta Zoológica Mexicana, nueva serie. Instituto de Ecología, Xalapa, México. Holdridge, L. R. 1947. Determination of world plant formations from simple climatic data. Science 105:367–368. IGAC. 1988. Suelos y bosques de Colombia. Instituto Geográfico Agustín Codazzi, Santafé de Bogotá, Colombia. Linares, O. J. 1998. Mamíferos de Venezuela. Sociedad Conservacionista Audubon de Venezuela, Caracas, Venezuela. Merriam, C. H. 1897. Descriptions of five new shrews from Mexico, Guatemala, and Colombia. Proceedings of the Biological Society of Washington 11:227–230. Morales, M., et al. 2007. Atlas de páramos de Colombia. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia. Moreno, C. P. A., and V. L. Albuja. 2014. Una nueva especie de Musaraña del género Cryptotis Pomel 1848 (Mammalia: Soricomorpha: Soricidae) de Ecuador y estatus taxonómico de Cryptotis equatoris Thomas (1912). Papéis Avulsos de Zoologia 54:403–418. Mosimann, J. E., and F. C. James. 1979. New statistical methods for allometry with applications to Florida red-winged black birds. Evolution 32:444–459. Quiroga-Carmona, M. 2013. Una nueva especie de musaraña del género Cryptotis (Soricomorpha: Soricidae) de la Serranía del Litoral en el norte de Venezuela. Mastozoología Neotropical 20:123–137. Quiroga-Carmona, M., and J. Molinari. 2012. Description of a new shrew of the genus Cryptotis (Mammalia: Soricomorpha: Soricidae) from the Sierra de Aroa, an isolated mountain range in northwestern Venezuela, with remarks on biogeography and conservation. Zootaxa 3441:1–20. Primack, R., R. Rozzi, and P. Feinsinger. 2001. Diseño de áreas protegidas. Pp. 469–477 in Fundamentos de Conservación Biológica. Perspectivas Latinoamericanas (R. Primack, R. Rozzi, P. Feinsinger, R. Dirzo, and F. Massardo, eds.). Fondo de Cultura Económica, México, D.F. Rangel-Ch., J. O. 2007. La Alta Montaña de Perijá: consideraciones finales. Pp. 417–436 in Colombia, Diversidad Biótica V, La Alta Montaña de la Serranía de Perijá (J. O. Rangel-Ch., ed.). Universidad Nacional de Colombia, Bogotá, Colombia. Rangel-Ch., J. O. 2009. Ecosistemas zonales de la Serranía de Perijá. Pp. 633–660 in Colombia, Diversidad Biótica VIII, Media y Baja montaña de la Serranía de Perijá (J. O. Rangel-Ch., ed.). Universidad Nacional de Colombia, Bogotá, Colombia. Rangel-Ch., J. O., and H. Arellano-P. 2007. Los ecosistemas de la Alta Montaña de Perijá. Pp. 329–346 in Colombia, Diversidad Biótica V, La Alta Montaña de la Serranía de Perijá (O. RangelCh., ed.). Universidad Nacional de Colombia, Bogotá, Colombia. Reumer, J. W. F. 1984. Ruscinian and early Pleistocene Soricidae (Insectivora, Mammalia) from Tegelen (The Netherlands) and Hungary. Scripta Geologica 73:1–173.

Rivera-Diaz, O., and J. L. Fernandez-Alonzo. 2003. Análisis corológico de La flora endémica de La Sierra de Perijá, Colombia. Anales del Jardín Botánico de Madrid 60:347–369. Rodríguez, J. P., F. Rojas-Suarez, and D. Giraldo-Hernández. 2010. Libro Rojo de los ecosistemas terrestres de Venezuela. Primera edición. Provita, Lenovo, Shell de Venezuela, Caracas, Venezuela. Smithe, F. B. 1975. Naturalist’s color guide. Parts I and II. American Museum of Natural History, New York. Thomas, O. 1898. On seven new small mammals from Ecuador and Venezuela. Annals and Magazine of Natural History, Series 7:451–457. Thomas, O. 1921. New Cryptotis, Thomasomys and Oryzomys from Colombia. Annals and Magazine of Natural History 8:354–357. Viloria, A. L. 1990. La sierra de Perijá y su problemática políticoecológica. Revista Nacional de Ciencias Sociales 2:49–51. Viloria, A. L. 2005. Señales y amenazas de deterioro ambiental en el territorio Barí: una cronología. El Guacharo, Boletín divulgativo de la Sociedad Venezolana de Espeleología 60:14–20. Woodman, N. 1996. Taxonomic status of the enigmatic Cryptotis avia (Mammalia: Insectivora: Soricidae), with comments on the distribution of the Colombian small-eared shrew, Cryptotis colombiana. Proceedings of the Biological Society of Washington 109:409–418. Woodman, N. 2002. A new species of small-eared shrew from Colombia and Venezuela (Mammalia: Soricomorpha: Soricidae: Genus Cryptotis). Proceedings of the Biological Society of Washington 115:249–272. Woodman, N., and J. Péfaur. 2008. Order Soricomorpha Gregory, 1910. Pp. 177–187 in Mammals of South America. 1. Marsupials, xenarthrans, shrews, and bats (A. L. Gardner, ed.). University of Chicago Press, Chicago, Illinois. Woodman, N., and R. M. Timm. 1993. 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. Fieldiana: Zoology (New Series) 1452:1–30. Woodman, N., and R. M. Timm. 1999. Geographic variation and evolutionary relationships among broad-clawed shrews of the Cryptotis goldmani-group (Mammalia: Insectivora: Soricidae). Fieldiana: Zoology (New Series) 1497:1–35.

Appendix I: Specimens Examined Cryptotis aroensis Quiroga-Carmona and Molinari, 2012 (n = 3). VENEZUELA. Yaracuy: Sierra de Aroa, Sector Las Cumaraguas, 1,730 m (CVULA I–8548—holotype; I–8546, I–8547—paratypes). Cryptotis meridensis (Thomas, 1898) (n = 117). VENEZUELA. Mérida: [no locality] (BMNH no number). near Laguna Mucubají, 3,600 m, 3.25 km ESE of Apartaderos (USNM 579287–579291, 579293, 579295–579298). Páramo de Mucubají, near Laguna de Mucubají, 3,600 m (CVULA 333–335, 337, 359, 421, 766, 874, 1168, 1486, 1487–1496, 1650, 1762, 8549; MHNLS 11220). near Laguna Negra, 3,500 m, 5.75 km ESE of Apartaderos (USNM 579273–579286). Laguna Negra, Páramo de Mucubají, 3,460 m (MBUCV 1882–1885, 2773; MHNLS 11221). Páramo de [La] Culata, 9,000 ft, Río Mucujún (FMNH 21837, 21838). Montes de La Culata, 2,000 m, Mérida (BMNH 98.7.1.13). Río Mucujún, 9,000– 12,500 ft (FMNH 21839, 21840, 21842, 21844; BMNH 29.11.7.16; USNM 260756). Montes del Valle, 2,125–2,165 m, Mérida (BMNH 98.7.1.11, 98.7.1.12, 98.5.15.5—holotype). Páramo Tambor (FMNH



QUIROGA-CARMONA AND WOODMAN—A NEW CRYPTOTIS FROM THE SIERRA DE PERIJÁ 809

21845). near Santa Rosa, 1,980–1,990 m, 1 km N, 2 km W of Mérida (USNM 385110–385112). near La Mucuy, 2,450 m, 2.9 km E of Tabay (USNM 579305). 5.5 km E and 2 km S Tabay, 2,630 m (EBRG 3639, 3640, 3641). near Middle Refugio, 2 km S, 5.5 km E of Tabay, 2,630 m (USNM 385106). Mérida (BMNH 98.7.1.10; MNHN 1900542, 1900-543; USNM 94165). La Aguada, near Laguna La Fría, 3,600 m, 7 km SE of Mérida (USNM 579299–579304). Near Loma Redonda cable car station, 4,100 m, 8.8 km SE of Mérida (USNM 579306). El Tambor, 8,800 ft (AMNH 96156–96158); Páramo de los Conejos, 9,600 ft (AMNH 96159). La Coromoto, 3,160–3,175 m, 4 km S, 6.5 km E of Tabay (USNM 385101, 385104). Near Laguna Verde, 3,533–3,545 m, 7.5 km E, 6 km S of Tabay (USNM 385102, 385103, 385105). km 36 on Apartaderos–Barinas road, 10 km E Santo Domingo, 1,670 m (CVULA 821).El Salvajal, 8 km W Mérida, 2,000 m (CVULA 2458). Monte Zerpa, 6 km N Mérida, 2,160 m (CVULA 1211, 5888, 6890, 6891). Escagüey, 1.5 km SSW Mucurubá, 2,200 m (CVULA 6899, 6900). La Carbonera, 14 km SE La Azulita, 2,250 m (CVULA 7912). Páramo de Mariño, 7 km W Tovar, NE Laguna Brava, 2,300 m (CVULA 2352). Asentamiento Monterrey, 8 km NNE Mérida, 2,300 m (CVULA 3502, 7078). Páramo de San José, 4.7 km SSE San José, 3,100 m (CVULA 8537–8545). 7.5 km E and 6 km S Tabay, 3,545 m (EBRG 3642). Táchira: Cerro Alto Duque, 2.5 km SE El Cobre, 2,500 m (MBLUZ 167). Páramo El Zumbador, 10 km SSW El Cobre, 2,750 m (CVULA 5744). Cryptotis tamensis Woodman, 2002 (n = 21). COLOMBIA. Norte de Santander, Páramo de Tamá, head of Río Táchira (FMNH 18571, 18614, 18615); Páramo de Tamá (FMNH 18572, 18608–18611, 18613,

18621; MCZ 21004; USNM 260747). VENEZUELA. Táchira: Buena Vista, 2,380–2,415 m (USNM 418566, 418567—holotype, 418569, 418570); Páramo de Tamá (FMNH 18617, 18620). Buena Vista, 41 km SW San Cristóbal, near Páramo del Tamá, 2,460 m (EBRG 3643, 3644, 11743—topotypes). Cryptotis thomasi (Merriam, 1897) (n = 41). COLOMBIA: [no specific locality] (BM 54.1.11.4; MCZ 27596). Cundinamarca: Represa del Neusa (ICN 9659). Páramo de Bogotá, 2,900 m (AMNH 37381; MNHN 1962-1068). La Selva, near Bogotá, (BM 97.5.21.2— holotype, 99.10.3.4; USNM 80903). Boquerón, near Bogotá (BM 99.10.3.1). Páramo de Monserrate, 3,200–3,300 m (ICN 9649, 9650, 9652, 9658; ROM 51870). San Francisco, 3,000–3,500 m (AMNH 71354, 71355; FMNH 71023–71029, 71035). San Cristóbal, 2,800– 2,900 m (FMNH 71030–71034, 71036, 71037). Reserva Biológica Carpanta, 3,000 m (ICN 10995, KU 157765). Páramo de Choachí, 3,000 m (AMNH 38405, MCZ 20092, 27597, 27598). Páramo el Verjón (MCZ 19995, 20091—[holotype of C. avia]). Chipaque (USNM 251960). Páramo de Chisacá, 3,100 m (ICN 5223). Cryptotis venezuelensis Quiroga-Carmona, 2013 (n = 3). VENEZUELA. Aragua: Sector Cerro Geremba, Monumento Natural Pico Codazzi, 2,283 m (EBRG 27336—holotype; 27337—paratype). Vargas: El Junquito, 2,100 m (MBUCV 402—paratype).

Submitted 6 January 2015. Accepted 11 May 2015. Associate Editor was Ryan W. Norris.